Photosynthetic adaptation strategies in peppers under continuous lighting: insights into photosystem protection.

Energy efficient lighting strategies have received increased interest from controlled environment producers. Long photoperiods (up to 24 h - continuous lighting (CL)) of lower light intensities could be used to achieve the desired daily light integral (DLI) with lower installed light capacity/capital costs and low electricity costs in regions with low night electricity prices. However, plants grown under CL tend to have higher carbohydrate and reactive oxygen species (ROS) levels which may lead to leaf chlorosis and down-regulation of photosynthesis. We hypothesize that the use of dynamic CL using a spectral change and/or light intensity change between day and night can negate CL-injury. In this experiment we set out to assess the impact of CL on pepper plants by subjecting them to white light during the day and up to 150 µmol m-2 s-1 of monochromatic blue light at night while controlling the DLI at the same level. Plants grown under all CL treatments had similar cumulative fruit number and weight compared to the 16h control indicating no reduction in production. Plants grown under CL had higher carbohydrate levels and ROS-scavenging capacity than plants grown under the 16h control. Conversely, the amount of photosynthetic pigment decreased with increasing nighttime blue light intensity. The maximum quantum yield of photosystem II (Fv/Fm), a metric often used to measure stress, was unaffected by light treatments. However, when light-adapted, the operating efficiency of photosystem II (ΦPSII) decreased and non-photochemical quenching (NPQ) increased with increasing nighttime blue light intensity. This suggests that both acclimated and instantaneous photochemistry during CL can be altered and is dependent on the nighttime light intensity. Furthermore, light-adapted chlorophyll fluorescence measurements may be more adept at detecting altered photochemical states than the conventional stress metric using dark-adapted measurements.

Continuous lighting can improve yield and reduce energy costs while increasing or maintaining nutritional contents of microgreens.

Microgreens represent a fast growing segment of the edible greens industry. They are prized for their colour, texture, and flavour. Compared to their mature counterparts, microgreens have much higher antioxidant and nutrient content categorizing them as a functional food. However, current production practices in plant factories with artificial light are energy intensive. Specifically, the lack of sunlight within the indoor structure means all of the light must be provided via energy consuming light fixtures, which is energy intensive and costly. Plant growth is usually increased with the total amount of light provided to the plants - daily light integral (DLI). Long photoperiods of low intensity lighting (greater than 18h) providing the desired/target DLI can reduce the capital costs for light fixtures and electricity costs. This is achieved by moving the electricity use from peak daytime hours (high price) to off-peak hours (low price) during the night in regions with time-based pricing scheme and lowering the electricity use for air conditioning, if plant growth is not compromised. However, lighting with photoperiods longer than tolerance thresholds (species/cultivar specific) usually leads to plant stress/damage. Therefore, we investigated the effects of continuous 24h white light (CL) at two DLIs (~14 and 21 mol m-2 d-1) on plant growth, yield, and antioxidant content on 4 types of microgreens - amaranth, collard greens, green basil, and purple basil to see if it compromises microgreen production. It was found that amaranth and green basil had larger fresh biomass when grown under CL compared to 16h when the DLIs were the same. In addition, purple basil had higher biomass at higher DLI, but was unaffected by photoperiods. Plants grown under the CL treatments had higher energy-use-efficiencies for lighting (10-42%) than plants grown under the 16h photoperiods at the same DLI. Notably, the electricity cost per unit of fresh biomass ($ g-1) was reduced (8-38%) in all microgreens studied when plants were grown under CL lighting at the same DLIs. Amaranth and collard greens also had higher antioxidant content. Taken together, growing microgreens under CL can reduce electricity costs and increase yield while maintaining or improving nutritional content.

The Power of Far-Red Light at Night: Photomorphogenic, Physiological, and Yield Response in Pepper During Dynamic 24 Hour Lighting.

Supplemental light is needed during the winter months in high latitude regions to achieve the desired daily light integral (DLI) (photoperiod × intensity) for greenhouse pepper (Capsicum annuum) production. Peppers tend to have short internodes causing fruit stacking and higher labor time for plant maintenance when grown under supplemental light. Far-red light can increase internode length, and our previous study on tomatoes (Solanum lycopersicum) also discovered monochromatic blue light at night during continuous lighting (CL, 24 h) increased stem elongation. Furthermore, the use of low-intensity, long photoperiod lighting can reduce light fixture costs and overall electricity costs due to lower power prices during the night. Therefore, we investigated the use of blue and/or far-red light during the night period of CL to increase stem elongation. Three pepper cultivars with different internode lengths/growing characteristics ('Maureno,' 'Gina,' and 'Eurix') were used to investigate the effects on plant morphology in a short experiment, and one cultivar 'Maureno' was used in a long experiment to assess the impact on fruit yield. The five lighting treatments that were used are as follows: 16 h of white light during the day followed by either 8 h of darkness (16W - control), white light (24W), blue light only (16W + 8B), blue + far-red light (16W + 8BFR), or far-red light only (16W + 8FR). Calculated nighttime phytochrome photostationary state (PSS) was 0.833, 0.566, 0.315, and 0.186 for 24W, 16W + 8B, 16W + 8BFR, and 16W + 8FR respectively. All five treatments had the same DLI in photosynthetically active radiation (PAR) and far-red light. The 16W + 8BFR and 16W + 8FR treatments significantly increased internode length compared to 16W and 24W but neither was more impactful than the other. The 16W + 8B treatment also increased internode length but to a lesser extent than 16W + 8BFR and 16W + 8FR. This indicates that a nighttime PSS of 0.315 is sufficient to maximize stem elongation. Both 16W + 8B and 16W + 8BFR drove photosynthesis during the nighttime supporting a similar yield compared to 16W. Therefore, 16W + 8BFR is the most potential lighting strategy as it can lead to a greater reduction in the light fixture and electrical costs while maintaining yield and enhancing internode length.

Light Spectra and Root Stocks Affect Response of Greenhouse Tomatoes to Long Photoperiod of Supplemental Lighting.

Plant biomass and yield are largely dictated by the total amount of light intercepted by the plant (daily light integral (DLI)-intensity × photoperiod). It is more economical to supply the desired DLI with a long photoperiod of low-intensity light because it uses fewer light fixtures, reducing capital costs. Furthermore, heat released by the light fixtures under a long photoperiod extended well into the night helps to meet the heating requirement during the night. However, extending the photoperiod beyond a critical length (>17 h) may be detrimental to production and lead to leaf chlorosis and a reduction in leaf growth and plant vigor in greenhouse tomato production. It is known that red light can increase leaf growth and plant vigor, as can certain rootstocks, which could compensate for the loss in plant vigor and leaf growth from long photoperiods. Therefore, this study investigated the response of tomatoes grafted onto different rootstocks to a long photoperiod of lighting under red and other light spectra. Tomato plants 'Trovanzo' grafted onto 'Emperator' or 'Kaiser' were subjected to two spectral compositions-100% red or a mix of red (75%), blue (20%), and green (5%) light for 17 h or 23 h. The four treatments supplied similar DLI. Leaf chlorosis appeared in all plants under 23 h lighting regardless of spectral compositions between 20 and 54 days into the treatment. The yield for 23 h mixed lighting treatment was lower than both 17 h lighting treatments. However, the 23 h red lighting treatment resulted in less leaf chlorosis and the plants grafted onto 'Emperator' produced a similar yield as both 17 h lighting treatments. Therefore, both spectral compositions and rootstocks affected the response of greenhouse tomatoes to long photoperiods of lighting. With red light and proper rootstock, the negative yield impact from long photoperiod lighting can be eliminated.

Continuous Light Does Not Compromise Growth and Yield in Mini-Cucumber Greenhouse Production with Supplemental LED Light.

Continuous lighting (CL, 24 h) can reduce the light intensity/light capital costs used to achieve the desired amount of light for year-round greenhouse vegetable production in comparison to short photoperiods of lighting. However, growth under CL has led to leaf injury characterized by chlorosis unless a thermoperiod or alternating light spectrum during CL is used. To date, there is no literature relating to how cucumbers (Cucumissativus) respond to CL with LEDs in a full production cycle. Here, we evaluated a mini-cucumber cv. "Bonwell" grown under 4 supplemental lighting strategies: Treatment 1 (T1, the control) was 16 h of combined red light and blue light followed by 8 h of darkness. Treatment 2 (T2) had continuous (24 h) red light and blue light. Treatment 3 (T3) was 16 h of red light followed by 8 h of blue light. Treatment 4 (T4) was 12 h of red light followed by 12 h of blue light. All treatments had a supplemental daily light integral (DLI) of ~10 mol m-2 d-1. Plants from all treatments showed similar growth characteristics throughout the production cycle. However, plants grown under all three CL treatments had higher chlorophyll concentrations from leaves at the top of the canopy when compared to T1. The overall photosynthetic capacity, light use efficiency, and photosynthetic parameters related to light response curves (i.e., dark respiration, light compensation point, quantum yield, and photosynthetic maximum), as well as the quantum yield of photosystem II (PSII; Fv/Fm) were similar among the treatments. Plants grown under all CL treatments produced a similar yield compared to the control treatment (T1). These results indicate that mini-cucumber cv. "Bonwell" is tolerant to CL, and CL is a viable and economical lighting strategy for mini-cucumber production.

Alternating Red and Blue Light-Emitting Diodes Allows for Injury-Free Tomato Production With Continuous Lighting.

Plant biomass is largely dictated by the total amount of light intercepted by the plant [daily light integral (DLI) - intensity × photoperiod]. Continuous light (CL, 24 h lighting) has been hypothesized to increase plant biomass and yield if CL does not cause any injury. However, lighting longer than 18 h causes leaf injury in tomato characterized by interveinal chlorosis and yield is no longer increased with further photoperiod extension in tomatoes. Our previous research indicated the response of cucumbers to long photoperiod of lighting varies with light spectrum. Therefore, we set out to examine greenhouse tomato production under supplemental CL using an alternating red (200 µmol m-2 s-1, 06:00-18:00) and blue (50 µmol m-2 s-1, 18:00-06:00) spectrum in comparison to a 12 h supplemental lighting treatment with a red/blue mixture (200 µmol m-2 s-1 red + 50 µmol m-2 s-1 blue, 06:00-18:00) at the same DLI. Our results indicate that tomato plants grown under supplemental CL using the red and blue alternating spectrum were injury-free. Furthermore, parameters related to photosynthetic performance (i.e., Pnmax, quantum yield, and Fv/Fm) were similar between CL and 12 h lighting treatments indicating no detrimental effect of growth under CL. Leaves under CL produced higher net carbon exchange rates (NCER) during the subjective night period (18:00-06:00) compared to plants grown under 12 h lighting. Notably, 53 days into the treatment, leaves grown under CL produced positive NCER values (photosynthesis) during the subjective night period, a period typically associated with respiration. At 53 days into the growth cycle, it is estimated that leaves under CL will accumulate approximately 800 mg C m-2 more than leaves under 12 h lighting over a 24 h period. Leaves grown under CL also displayed similar diurnal patterns in carbohydrates (glucose, fructose, sucrose, and starch) as leaves under 12 h lighting indicating no adverse effects on carbohydrate metabolism under CL. Taken together, this study provides evidence that red and blue spectral alternations during CL allow for injury-free tomato production. We suggest that an alternating spectrum during CL may alleviate the injury typically associated with CL production in tomato.

Effect of elevated CO2 and spectral quality on whole plant gas exchange patterns in tomatoes.

In controlled environment plant production facilities, elevating either light or CO2 levels generally has led to increased biomass and yield due to enhanced canopy photosynthesis. Today, advancements in light-emitting diodes (LEDs) have made this technology a viable option for both supplementary lighting in greenhouses and a sole lighting source in controlled environment chambers. Our study used tomato plants grown under both ambient CO2 (AC) and elevated CO2 (EC) conditions then exposed them to various CO2 and lighting treatments during both whole plant and leaf level measurements. Plants grown under EC reached the first flower developmental stage 8 days sooner and were approximately 15cm taller than those grown under AC. However, under AC plants had more leaf area while their dry weights were similar. Of note, under EC chlorophyll a and b were lower, as were carotenoids per unit leaf area. Whole plant analyses, under all CO2 challenges, showed that plants exposed to high-pressure sodium (HPS), red-blue LED, and red-white LED had similar photosynthesis, respiration, and daily carbon gain. Under different light qualities, day-time transpiration rates were similar among CO2 conditions. Day-time water-use efficiency (WUE) was higher in plants grown and exposed to EC. Similarly, WUE of plants grown under AC but exposed to short-term elevated CO2 conditions was higher than those grown and tested under AC during all light treatments. Under all CO2 conditions, plants exposed to red-white and red-blue LEDs had lower WUE than those exposed to HPS lighting. Assessing alterations due to CO2 and light quality on a whole plant basis, not merely on an individual leaf basis, furthers our understanding of the interactions between these two parameters during controlled environment production. Principle component analyses of both whole plant and leaf data indicates that increasing CO2 supply has a more dramatic effect on photosynthesis and WUE than on transpiration.

Effects of Light Quality and Intensity on Diurnal Patterns and Rates of Photo-Assimilate Translocation and Transpiration in Tomato Leaves.

Translocation of assimilates is a fundamental process involving carbon and water balance affecting source/sink relationships. Diurnal patterns of CO2 exchange, translocation (carbon export), and transpiration of an intact tomato source leaf were determined during 14CO2 steady-state labeling under different wavelengths at three pre-set photosynthetic rates. Daily patterns showed that photosynthesis and export were supported by all wavelengths of light tested including orange and green. Export in the light, under all wavelengths was always higher than that at night. Export in the light varied from 65-83% of the total daily carbon fixed, depending on light intensity. Photosynthesis and export were highly correlated under all wavelengths (r = 0.90-0.96). Export as a percentage of photosynthesis (relative export) decreased as photosynthesis increased by increasing light intensity under all wavelengths. These data indicate an upper limit for export under all spectral conditions. Interestingly, only at the medium photosynthetic rate, relative export under the blue and the orange light-emitting diodes (LEDs) were higher than under white and red-white LEDs. Stomatal conductance, transpiration rates, and water-use-efficiency showed similar daily patterns under all wavelengths. Illuminating tomato leaves with different spectral quality resulted in similar carbon export rates, but stomatal conductance and transpiration rates varied due to wavelength specific control of stomatal function. Thus, we caution that the link between transpiration and C-export may be more complex than previously thought. In summary, these data indicate that orange and green LEDs, not simply the traditionally used red and blue LEDs, should be considered and tested when designing lighting systems for optimizing source leaf strength during plant production in controlled environment systems. In addition, knowledge related to the interplay between water and C-movement within a plant and how they are affected by environmental stimuli, is needed to develop a better understanding of source/sink relationships.

The Effect of Spectral Quality on Daily Patterns of Gas Exchange, Biomass Gain, and Water-Use-Efficiency in Tomatoes and Lisianthus: An Assessment of Whole Plant Measurements.

Advancements in light-emitting diode (LED) technology have made them a viable alternative to current lighting systems for both sole and supplemental lighting requirements. Understanding how wavelength specific LED lighting can affect plants is thus an area of great interest. Much research is available on the wavelength specific responses of leaves from multiple crops when exposed to long-term wavelength specific lighting. However, leaf measurements do not always extrapolate linearly to the complexities which are found within a whole plant canopy, namely mutual shading and leaves of different ages. Taken together, both tomato (Solanum lycopersicum) leaves under short-term illumination and lisianthus (Eustoma grandiflorum) and tomato whole plant diurnal patterns of plants acclimated to specific lighting indicate wavelength specific responses of both H2O and CO2 gas exchanges involved in the major growth parameters of a plant. Tomato leaves grown under a white light source indicated an increase in transpiration rate and internal CO2 concentration and a subsequent decrease in water-use-efficiency (WUE) when exposed to a blue LED light source compared to a green LED light source. Interestingly, the maximum photosynthetic rate was observed to be similar. Using plants grown under wavelength specific supplemental lighting in a greenhouse, a decrease in whole plant WUE was seen in both crops under both red-blue (RB) and red-white (RW) LEDs when compared to a high pressure sodium (HPS) light. Whole plant WUE was decreased by 31% under the RB LED treatment for both crops compared to the HPS treatment. Tomato whole plant WUE was decreased by 25% and lisianthus whole plant WUE was decreased by 15% when compared to the HPS treatment when grown under RW LED. The understanding of the effects of wavelength specific lighting on both leaf and whole plant gas exchange has significant implications on basic academic research as well as commercial greenhouse production.

Scabin, a Novel DNA-acting ADP-ribosyltransferase from Streptomyces scabies.

A bioinformatics strategy was used to identify Scabin, a novel DNA-targeting enzyme from the plant pathogen 87.22 strain of Streptomyces scabies Scabin shares nearly 40% sequence identity with the Pierisin family of mono-ADP-ribosyltransferase toxins. Scabin was purified to homogeneity as a 22-kDa single-domain enzyme and was shown to possess high NAD(+)-glycohydrolase (Km (NAD) = 68 ± 3 µm; kcat = 94 ± 2 min(-1)) activity with an RSQXE motif; it was also shown to target deoxyguanosine and showed sigmoidal enzyme kinetics (K0.5(deoxyguanosine) = 302 ± 12 µm; kcat = 14 min(-1)). Mass spectrometry analysis revealed that Scabin labels the exocyclic amino group on guanine bases in either single-stranded or double-stranded DNA. Several small molecule inhibitors were identified, and the most potent compounds were found to inhibit the enzyme activity with Ki values ranging from 3 to 24 µm PJ34, a well known inhibitor of poly-ADP-ribosyltransferases, was shown to be the most potent inhibitor of Scabin. Scabin was crystallized, representing the first structure of a DNA-targeting mono-ADP-ribosyltransferase enzyme; the structures of the apo-form (1.45 Å) and with two inhibitors (P6-E, 1.4 Å; PJ34, 1.6 Å) were solved. These x-ray structures are also the first high resolution structures of the Pierisin subgroup of the mono-ADP-ribosyltransferase toxin family. A model of Scabin with its DNA substrate is also proposed.