Blue Radiation Interacts with Green Radiation to Influence Growth and Predominantly Controls Quality Attributes of Lettuce


Meng, Q., J. Boldt, and E.S. Runkle. 2020. Blue radiation interacts with green radiation to influence growth and predominately controls quality attributes of lettuce. Journal of the American Society for Horticultural Sciences 145(2):75-87.

With recent shifts in modern agriculture to more urban environments, indoor farming has become increasingly popular. Such an environment allows growers to control everything from air flow to water. Arguably the most important aspect that can be controlled is lighting, specifically with the use of light-emitting diodes (LEDs) that allow for customizable wavelengths for varying stages in a plant’s life cycle. Research has shown that different wavelengths can produce different results. For instance, exposing certain varieties of lettuce to blue radiation (400-500 nm) has been tied to a significant reduction in biomass weight, as well as an increase in the production of secondary metabolites. Growers will often combine blue with red and far-red radiation (600-800 nm) to achieve desired results. Green radiation (500-600 nm), however, does not have much of a history of being used by growers, despite its ability to penetrate deep into the leaves. This has huge implications on its ability to drive photosynthesis and has recently been studied as a substitute for blue radiation. Previous research on this subject has shown an increase in biomass of several lettuce varieties, but the authors believe this could have been attributed to the shifting levels of blue radiation. To combat this, they designed a new experiment to keep levels of blue radiation constant and substitute red radiation for green.


In this experiment, researchers focused on ‘Rouxai’ red leaf lettuce and tested the effects of varying wavelengths using LED lighting. During the light quality treatment phase of the experiment, each treatment had a 20 hour photoperiod and a total photosynthetic photon flux density (PPFD) of 180 μmol m-2 s-1 . They exposed the lettuce to nine different treatments of lighting, including combinations of blue and red radiation, as well as introducing green radiation at a photon flux density (PFD) of 60 μmol m-2 s-1 in place of a reduction in red radiation. Researchers measured biomass accumulation including fresh and dry mass, different morphological features such as plant diameter and leaf number, and coloration of the foliage. They found that at 20 μmol m-2 s-1 of blue radiation, the presence of 60 μmol m-2 s-1 green radiation increased the fresh mass of lettuce, but had negative effects on the weight at any higher levels of blue radiation. Additionally, an increase in blue radiation with 60 μmol m-2 s-1 of green radiation decreased leaf diameter, width, and length. With an increase in blue radiation, lettuce is known to have an increase in red color of the foliage. Without green radiation, the foliage became saturated at 20 μmol m-2 s-1 of blue radiation. However, with 60 μmol m-2 s-1 of green radiation, the saturation point increased to 60 μmol m-2 s-1 of blue radiation. This has interesting implications for growers wishing to increase the coloration of their foliage.


This study provides valuable information regarding the role green radiation may have in a plant’s life cycle. With an increase in fresh mass at low levels of blue radiation, incorporating green radiation at the right stages could potentially increase yield for growers. Additionally, the presence of green light has been shown to vary depending on the species and age of the crop, which implies further research is needed on the subject. Having this research and knowledge that green radiation influences photosynthesis and other varying characteristics of lettuce growth is critical for growers looking to optimize their lighting in controlled environments.

Effects of continuous or end-of-day far-red light on tomato plant growth, morphology, light absorption, and fruit production


Kalaitzoglou, P., W. van Ieperen, J. Harbinson, M. van der Meer, S. Martinakos, K. Weerheim, C.C.S. Nicole, and L.F.M. Marcelis. 2019. Effects of continuous or end-of-day far-red light on tomato plant growth, morphology, light absorption, and fruit production. Frontiers in Plant Science 10: 322


LEDs are becoming increasingly common in modern controlled environment horticultural systems. The potential energy-savings and flexibility of LEDs makes them attractive lighting options compared to traditional high-pressure sodium or metal halide bulbs. Questions remain regarding the effects of light wavelengths emitted by LEDs on plant growth. Plants respond to differing wavelengths of light by changing physical characteristics and growth. Shading causes a low red light (R) to far-red light (FR) ratio (R:FR), resulting in low levels of the active form of a key plant photoreceptor, phytochrome. This low phytochrome stationary state (PSS) leads to a variety of shade avoidance responses that affect plant morphology and development. In contrast, LEDs used for greenhouse lighting often emit low levels or zero far-red light, causing plants to have higher PSS values than sunlight. Little is known about the effects of changing R:FR ratios on photosynthesis and plant growth in greenhouses using LEDs. Using tomato as a model crop, researchers at Wageningen University investigated how tomato morphology changes in response to higher than sunlight R:FR ratio supplied by LEDs and what effect these changes have on plant light absorption and growth (Kalaitzoglou et al., 2019). In addition, the researchers were interested if a short end-of-day FR treatment (EOD-FR) could compensate for any negative effects of growing plants without FR light during daytime.


The researchers conducted two experiments (EXP1 & EXP2). In both experiments, greenhouse chambers were divided into 15 equal compartments, each containing 20 tomato plants. Each compartment was illuminated by a combination of red (95%) and blue (5%) LEDs that supplied approximately 150 mmol
m-2 s-1 of photosynthetically active radiation (PAR) over the course of a 16 hour day. Additional FR LEDs were installed to provide five different treatments based on FR intensities. In four treatments, both FR LEDs and red/blue LEDs were on at the same time during the day, resulting in plant PSS values of 0.70, 0.73, 0.80, 0.88. The fifth treatment was a 15 minute end of day FR cycle (EOD-FR) following the end of the photoperiod. To investigate the interaction between the effects of FR and solar radiation on plant morphology the first experiment (EXP1) used a blackout screen to block incoming sunlight, while the second experiment (EXP2) exposed the plants to broadband solar radiation from morning to afternoon. In addition, EXP1 lasted only four weeks after transplanting while EXP2 was extended to 16 weeks to allow for tomato fruit development.

In both experiments, researchers measured plant growth and morphology traits such as plant height, petiole angle, and leaf area. In EXP2, fruit traits including total fruit weight (g/plant), number of fruits, and number of open flowers at 4 weeks were recorded. Additional measured traits included leaf ligh absorbance and chlorophyll and carotenoid content. To simulate light absorption for each treatment, researchers constructed a 3D plant model using GroIMP software (Hemmerling et al., 2008). Researchers used the model to estimate the effects of changes in plant morphology in response to FR light treatments on plant light absorption.

Supplemental Figure S5. (Kalaitzoglou et al., 2019)


Increasing R:FR ratio to levels above sunlight had a negative impact on tomato plant growth. In both experiments, morphological parameters such as plant height and leaf area decreased as PSS values became higher (Supplemental Figure S5). The researchers concluded that lower levels of plant light absorption in high PSS treatments were primarily caused by the decrease in leaf area, ultimately reducing plant growth. Similar results were observed for fruit characteristics in which fruit size and fruit number were greater in treatments with increasing FR compared to the 0.88 PSS treatment (no FR) and EOD-FR treatment. FR treatments also stimulated early flower and fruit maturity. Interestingly, while leaf PAR absorbance and chlorophyll content were lower in low PSS treatments, net photosynthesis was higher. Researchers attributed this result to the Emerson effect in which a higher rate of photosynthesis occurs when plants are exposed to a simultaneous mixture of red and far-red light. In both studies, EOD-FR treatments were not enough to offset the negative effects of growing plants with low levels or zero FR light throughout the day. In conclusion, the results of the study indicated that the presence of FR light increased fruit yield and accelerated flowering and FR LEDs could be a beneficial addition to greenhouses to improve tomato fruit production.

LEDs for photons, physiology and food

Original paper: P. M. Pattison, J. Y. Tsao , G. C. Brainard, and B. Bugbee 2018. LEDs for photons, physiology and food. Nature. 563:493-500.

Compared to traditional lighting, LED lighting offers greater light control, improved performance, and decreased energy consumption. Due to these facts, LED lighting is beginning to be used for an array of new applications to improve human health and localize food production in controlled environments. For the first time in history, the use of LED lighting enables humans to engineer lighting of environments to elicit specific responses.

Four main features separate LED lighting from traditional lighting – light spectral control, light intensity control, control of light distribution in space, and ready integration with other technologies. LEDs for photons, physiology, and food outlines some of the applications and research avenues that LED lighting will enable in both humans and plants.

Lighting impacts both humans and plants greatly. In humans, light affects daily rhythms of sleep and wakefulness, body temperature, alertness, psychomotor performance, neurocognitive responses, and the secretion of hormones. Among the open questions posed regarding lighting for human health and productivity are the nature of the detailed pathways within the melanopsin-based photoreceptor system, interactions between the retinohypothalamic and primary optical tracts, the relationship between the dose of light and physiological regulation in everyday environments, and how to frame our understanding of the positive and negative effects of light. Light-emitting diodes will enable more precise and effective lighting research to be conducted relating to the aforementioned questions, which will enable LED lighting to be increasingly tailored to enhance human health and productivity.

Plants not only require light as fuel for photosynthesis but also use light as a signal to direct plant morphology and metabolite profile. Light sources and color filters have long been used to investigate plant responses to light. However, prior to LED lighting, many of these studies have been limited, mainly because they were conducted at low light levels on single leaves. LED lighting now enables research to be conducted at higher light intensities at the plant canopy level. Additionally, LED lighting allows light intensity, spectrum, and timing of light application to be precisely controlled, taking plant-light response research to new levels.

LED lighting has not only enhanced our understanding of plant-light responses but has also made it cost-effective to grow certain plants indoors for food. To demonstrate the efficacy of indoor agriculture, the authors calculate the grams of dry mass produced per mole of photons for various crops. In doing so, the authors conclude that the photon cost (% of dry market price) is 1% for microgreens, 5% for lettuce, 18% for tomatoes, 103% for general vegetables (i.e. broccoli), and 10,000% for staple crops (i.e. rice).

The main parameters driving the increased photon cost for the above mention crops are:

  1. Fraction of photons absorbed by the plant: Microgreens can be grown at a very high density, thus the fraction of photons absorbed by the plant is very high. However, as plant size increases, plant spacing must also increase. Increased spacing between plants leads to reduced radiation captured and thus reduces the fraction of photons absorbed by plants, as some of these photons will inevitability be lost in space between plants.
  1. Quantum yield (moles of carbon fixed per mole of photons absorbed): The more a particular crop benefits from increased light levels will dictate its quantum yield. Lettuce benefits from higher light levels than microgreens and tomato benefits from higher light than lettuce, thus the quantum efficiency of lettuce is lower than that of microgreens and the that of tomato is lower than lettuce.
  1. Harvest index (moles of carbon in edible product per mole of carbon in plant biomass): Microgreens and lettuce have a very high harvest index as the entire aboveground portion of the plants are edible. Alternatively, tomato stems and leaves are not edible, reducing harvest index. Other general vegetables and staple crops harvest index is further reduced, as these crops generally posses even less edible plant biomass. Thus, for crops with low harvest index, photons are being captured by non-edible plant biomass, leading to increased photon cost per dry mass.

Based on these parameters the authors concluded that “electric light input is a small cost for microgreens, a high cost for general vegetables, and an unacceptable cost for staple crops”. Currently, most indoor farms are focused on growing leafy greens. However, as LED lighting efficiency and technology continues to increase, more general vegetables will be attempted to be grown indoors. Nevertheless, according to this report, even if LEDs were 100% efficient, growing staple crops indoors would not be cost-effective.

It is clear that LED lighting will continually replace traditional lighting and become the standard light source for humans and plants. By 2035, it is estimated that 86% of electrical lighting installs in the U.S. will be LED, which will save roughly US$52 billion per year in direct energy costs. Research into physiological responses to light will allow lighting systems to be optimized and the full potential of LED lighting to be reached, which include improving human health and productivity, increasing the feasibility of local food production in controlled environments, and decreased energy consumption.

Hydroponics vs. Soil Cultivation: Functional and Taste Compound Comparison

Original Paper
Tamura Y, Mori T, Nakabayashi R, Kobayashi M, Saito K, Okazaki S, Wang N and Kusano M (2018) Metabolomic Evaluation of the Quality of Leaf Lettuce Grown in Practical Plant Factory to Capture Metabolite Signature. Front. Plant Sci.9:665.
doi: 10.3389/fpls.2018.00665


Cultivation of certain crops is moving out of the field. Indoor production has taken the form of greenhouses, tunnels, and plant factories. These growing methods been collectively deemed controlled environment agriculture (CEA). The attraction is in the name – control. Moving crops out of the field helps remove risk of unpredictable weather and can allow for optimized conditions for crop production. It even enables year-round growing that provides a steady source of fresh food to the public and income to the growers instead of the seasonal flux of traditional agriculture.

With food moving indoors under controlled conditions, crops are receiving different types of input in terms of nutrients, lighting, day/night cycling, temperatures, disease and pest stresses, and other variables. Some crops are growing differently and looking different as well. It all depends on the control conditions.

As plant growth and development changes due to these controlled environments, the metabolic processes dictating that growth and development are probably varying as well. As a result, there may be changes in the plant’s profile of chemical compounds, or metabolites, which take part in and are produced by plant metabolism. These compounds are integral to the structure and general function of the plant as well as its defense against pests and disease. Again, with cultivation conditions changing, the metabolite compositions of the plants are likely changing simultaneously.

The Experiment

If we want to know if or how CEA is changing the metabolite profiles of our food compared to field cultivation, we need to isolate each element of the “control” to determine what changes are being caused by which conditions. To this end, a group from RIKEN in Japan that studies metabolite profiles (an analytical chemistry practice called metabolomics) chose to compare compounds of lettuce grown in a hydroponic system (plant roots growing directly into water with an added nutrient solution) within a Keystone Technology Inc. (Japan) plant factory to lettuce grown in a similarly-controlled growth chamber except planted traditionally in soil (Table 1).

Table 1. Plant factory conditions for hydroponic cultivation treatment compared to growth chamber conditions for the soil treatment.

This group chose two lettuce cultivars, ‘Black Rose’ and ‘Red Fire’, with one head of each cultivar grown per treatment – hydroponics and soil – for a total of 4 heads of lettuce in the experiment. Tamura et al. observed smaller and more pigmented leaves from the soil-grown lettuce compared to the hydroponic production. To detect the metabolites present in the lettuce they used precise instruments (gas and liquid chromatography mass spectrometry). They included samples from leaves on the outside of the head and the middle to account for variation in metabolite production in different parts of the plant.


Analysis resulted in 133 identified compounds and 185 unidentified. Based on the relative abundances of all 318 metabolites, they were able to clearly separate samples of hydroponically grown lettuce from those grown in soil.

Upon further study, they determined that hydroponic lettuce had higher amounts of amino acids (protein building blocks) than the soil-cultivated lettuce. On the other hand, lettuce grown in soil contained more sugars and compounds that contribute to taste and possible health-benefits, such as sesquiterpenes and organic acids. Particularly, glutamate, a metabolite contributing to the umami (or savory) taste profile of a food, was significantly higher in ‘Red Fire’ lettuce grown hydroponically. However, a sugar, sucrose, and a compound associated with bitterness, lactucopicrin-15-oxalate, were both significantly lower in the hydroponic lettuce.

Conclusions and Considerations

This study is valuable due to it being the first of its kind—applying metabolomics to understand how our crops are changing in CEA systems. These results need to be validated by another experiment in which the conditions other than soil/hydroponics are identical. Previous work by Li and Kubota (2009) demonstrated that differences in light intensity and quality can affect metabolite production in a CEA setting.

Additionally, different fertilization regimes largely influence the amount of nitrogen plants can access to produce amino acids. With the hydroponic lettuce receiving almost 3x the fertilizer compared to the lettuce in soil, a higher amino acid content in hydroponic lettuce cannot be completely attributed to hydroponic production itself. Therefore, the differences in control conditions presented in Table 1 above are confounded with the soil/hydroponic treatments, making interpretation of results complicated. This also points to the importance of collaboration across scientific disciplines to ensure the most effective and efficient experiments are conducted.


Li, Q., and Kubota, C. (2009). Effects of supplemental light quality on growth and phytochemicals of baby leaf lettuce. Environ. Exp. Bot. 67, 59–64. doi: 10.1016/j.envexpbot.2009.06.011

A small amount of UV radiation is needed for tomato plant health

Original paper: Kubota, C., T. Eguchi, and M. Kroggel. 2017. UV-B radiation dose requirement for suppressing intumescence injury on tomato plants. Scientia Horticulturae. 226:366-371.


UV-B radiation (300-320 nm) has the shortest wavelength of the sunlight spectrum in the natural environment. Too strong UV radiation can cause issues such as leaf burn, but too little UV radiation also becomes problematic for certain species of plants including tomato. The sensitivity varies among varieties of tomato and a particular rootstock tomato used for grafting, for example, is a very sensitive one that must be grown under light including UV-B.  A typical disorder caused by lack of UV-B radiation are leaf tumors called intumescence (or oedema). In severe cases, plants cannot grow and usually die. This becomes problematic when plants are grown under protected environments (such as tunnels) covered with UV-blocking plastic material or under sole source electric lighting that does not emit UV radiation (such as LEDs). A research group led by Chieri Kubota (currently at the Ohio State University) identified the needed amount of UV radiation to maintain plants without causing such disorder.  This group also worked on other innovative approaches to mitigate intumescence injury, including a discovery of an effective LED lighting protocol to mitigate the intumescence injury.


In their experiment, they grew rootstock tomato ‘Beaufort’ plants as their experimental plant material for its high sensitivity to induce intumescence under UV-B deficient light environment.  The lamps that they tested were red and blue LEDs or cool-white T5 fluorescent lamps. While red and blue LEDs do not emit UV radiation at all, T5 fluorescent lamps do emit small amount of UV-B radiation. When ‘Beaufort’ plants were grown under LEDs, plants developed massive amounts of intumescence, but the injury was less severe under T5 fluorescent lamps due to the UV radiation.  UV-B supplementation was examined for the plants under LEDs at varied hours of UV-B exposure (1.5, 3.0 and 6.0 hours) at a very low intensity of UV-B of 0.12 W m-2 (or 0.31 μmol m-2 s-1). The UV-B exposure was conducted during the night time (every night), rather than day time. Kubota’s group showed that an increasing amount (dose) of UV-B exposure drastically improved ‘Beaufort’ tomato plant health.  The highest daily dose of UV-B they examined was 2.6 kJ m-2 (or 6.7 mmol m-2). This UV-level is less than one-tenth of what you can expect outdoors during the summer.  Under this dose of 2.6 kJ m-2, plants still exhibited a minor injury of intumescence. So the group used a mathematical approach (linear regression) to estimate the dose of UV-B that would have eliminated the intumescence injury, which was 5 kJ m-2 (or 14 mmol m-2), about twice of the highest examined dose in this study, but still far below the natural UV level of sunlight.


The impact of this study includes possible contributions to the design of plant growing facilities that use LED lighting.  The linear dose response they described in this study is particularly useful as one can choose a combination of UV-B intensity and exposure time suitable for their production setting.  A lower intensity for long exposure and a higher intensity for shorter exposure could yield the same effective dose.  Kubota group also showed that night-time application of UV-B is effective and this is helpful for growers who do not want to expose workers to UV-B light while they are in the facility.  The known/Knowing the dose response also makes the application methods more flexible, including using a movable light source over the plant canopy. The finding will be helpful for Indoor farming as well as greenhouse crop production.