Multi-Omics Analyses of Lettuce (Lactuca sativa) Reveals Primary Metabolism Reorganization Supporting Distinct Features of Secondary Metabolism Induced by Supplementing UV-A Radiation.

UV can serve as an effective light spectrum for regulating plant secondary metabolites, while relevant studies on UV-A are much less extensive than those on UV-B. A comprehensive understanding of the selective effects of UV-A on different secondary metabolites and the specific features of primary metabolism that drive these effects is still lacking. To address this knowledge gap, we conducted a study to analyze the dynamic changes in the metabolome and transcriptome of lettuce leaves irradiated with red plus UV-A light (monochromatic red light as control). Generally, UV-A promoted the synthesis of most phenylpropanoids and terpenoids originating from the shikimate and methylerythritol phosphate (MEP) pathway in plastids but sacrificed the synthesis of terpenoids derived from the mevalonate (MVA) pathway, particularly sesquiterpenes. Increased precursors supply for the shikimate and MEP pathway under UV-A was directly supported by the activation of the Calvin-Benson cycle and phosphoenolpyruvate transport. Whereas, along with phosphoenolpyruvate transport, the TCA cycle was restrained, causing deprivation of the MVA pathway precursor. In addition, UV-A also activated the plastidic oxidative branch of the pentose phosphate pathway, photorespiration, and malate shuttle, to ensure a sufficient supply of nitrogen, circulation homeostasis of the Calvin-Benson cycle, and energy balance, thus indirectly supporting UV-A-induced specific secondary metabolic output. This study provides a comprehensive framework for understanding the flexible primary-secondary metabolism interactions that are able to produce specific metabolites favorable for adaptation to environmental stimuli.

Quality and Metabolomics Analysis of Houttuynia cordata Based on HS-SPME/GC-MS.

Houttuynia cordata is a medicinal and edible plant with a wide biological interest. Many parts were discarded due to various modes of consumption, resulting in resource waste. In this study, a comprehensive study was conducted on various edible indicators and medicinal components of Houttuynia cordata to understand its edible and medicinal value. The edible indexes of each root, stem, and leaf were determined, and the metabolites of different parts were investigated using the headspace solid-phase micro-extraction technique (HS-SPME-GC-MS). The differential metabolites were screened by orthogonal partial least squares discriminant analysis (OPLS-DA) and clustering analysis. The results of the study showed that the parts of Houttuynia cordata with high edibility values as a vegetable were mainly the roots and leaves, with the highest vitamin C content in the roots and the highest total flavonoids, soluble sugars, and total protein in the leaves. The nutrient content of all the stems of Houttuynia cordata was lower and significantly different from the roots and leaves (p < 0.05). In addition, 209 metabolites were isolated from Houttuynia cordata, 135 in the roots, 146 in the stems, 158 in the leaves, and 91 shared metabolites. The clustering analysis and OPLS-DA found that the parts of Houttuynia cordata can be mainly divided into above-ground parts (leaves and stems) and underground parts (roots). When comparing the differential metabolites between the above-ground parts and underground parts, it was found that the most important medicinal component of Houttuynia cordata, 2-undecanone, was mainly concentrated in the underground parts. The cluster analysis resulted in 28 metabolites with up-regulation and 17 metabolites with down-regulation in the underground parts. Most of the main components of the underground part have pharmacological effects such as anti-inflammatory, anti-bacterial and antiviral, which are more suitable for drug development. Furthermore, the above-ground part has more spice components and good antioxidant capacity, which is suitable for the extraction of edible flavors. Therefore, by comparing and analyzing the differences between the edible and medicinal uses of different parts of Houttuynia cordata as a medicinal and food plant, good insights can be obtained into food development, pharmaceutical applications, agricultural development, and the hygiene and cosmetic industries. This paper provides a scientific basis for quality control and clinical use.

Dynamic Responses of Ascorbate Pool and Metabolism in Lettuce to Light Intensity at Night Time under Continuous Light Provided by Red and Blue LEDs.

To understand the dynamic changes of hydroponic lettuce growth, ascorbate (AsA) pool and metabolism under two different dark period light intensities (LL, 20 µmol·m-2·s-1; CL, 200 µmol·m-2·s-1) of continuous light and normal light (NL, 0 µmol·m-2·s-1) provided by red (R) and blue (B) LEDs, the chlorophyll fluorescence parameters, ascorbate pool size, AsA metabolism-related enzyme activities, and H2O2 contents of lettuce were measured at 0, 8, 16, 24, 32, 40, 48, 56, 64, and 72 h after light treatment and the lettuce growth parameters were measured on the 9th day after light treatment. The results showed that compared with the NL, CL treatment for 9 days significantly increased the biomass, dry matter content, and specific leaf weight of lettuce, but had no significant effect on the leaf area and root-to-shoot ratio; LL had no significant effect on lettuce biomass, but it would reduce the root-shoot ratio. Compared with the NL, the AsA content of CL increased significantly within 8 h after light treatment (at the end of first dark period), and then maintained at a relatively stable level with a slight increase; there was no significant difference in AsA contents between NL and LL showing the same circadian rhythm characteristics. Overall, the activities of L-galactono-1,4-lactone dehydrogenase (GalLDH), ascorbate peroxidase(APX), monodehydroascorbate reductase (MDHAR), and glutathione reductase (GR) under CL were the highest among the three treatments, and the differences with the other two treatments reached significant levels at several time points; there was almost no significant difference in the activities of GalLDH, APX, MDHAR, and GR between NL and LL; there was no significant difference in the activities of dehydroascorbate reductase (DHAR) under different treatments. Compared with the NL, CL caused a sharp decrease of PSⅡ maximal photochemical efficiency (Fv/Fm) in lettuce within 0-8 h after treatment, which then stabilized at a relatively stable level; the Fv/Fm value under the LL was almost the same as the NL. Except for 32 h, the H2O2 content of lettuce under CL was the highest among the three treatments during the entire experimental period, and was significantly higher than that of NL at several time points; the H2O2 content of LL was almost the same as NL. In summary, lettuce biomass, AsA contents, AsA metabolism-related enzyme activities, chlorophyll fluorescence parameters, and H2O2 contents were regulated by the dark period light intensities of continuous light rather than continuous light signals.

LED Light Quality of Continuous Light before Harvest Affects Growth and AsA Metabolism of Hydroponic Lettuce Grown under Increasing Doses of Nitrogen.

To study the effects of light quality of continuous light before harvest on the growth and ascorbic acid (AsA) metabolism of lettuce (Lactuca sativa L.) grown under relative high nitrogen level, lettuce plants grown under different nitrogen levels (8, 10 and 12 mmol·L-1) were subjected to continuous light with different red: blue light ratios (2R:1B and 4R:1B) before harvest. The results showed that the shoot fresh weight of lettuce under 12 mmol·L-1 nitrogen level was significantly higher than that under other treatments. There were no significant differences in shoot dry weight, root fresh weight, root dry weight, soluble sugar content, nitrate content and AsA content in leaves among the treatments at different nitrogen levels. The content of AsA in leaves was significantly higher than that in petioles before and after continuous light. Under the same nitrogen level, the fresh weight of lettuce under continuous light quality 4R:1B was significantly higher than that under other treatments. The content of AsA in lettuce leaves increased in different degrees after continuous light before harvest. High yield and AsA content could be obtained by 72 h continuous light with red and blue light 4R:1B at 12 mmol·L-1 nitrogen level. After continuous light, the content of AsA increased significantly due to the increase of the ratio of red light and nitrogen level, which increased the activities of L-galactono-1,4-lactone dehydrogenase (GalLDH) and dehydroascorbic acid reductase (DHAR) involved in AsA synthesis and in the recycling of DHAR to AsA respectively.

Light Quality Affected the Growth and Root Organic Carbon and Autotoxin Secretions of Hydroponic Lettuce.

Light is a crucial environmental signal and photosynthetic energy for plant growth, development, and primary and secondary metabolism. To explore the effects of light quality on the growth and root exudates of hydroponic lettuce (Lactuca sativa L.), white LED (W, control) and four the mixtures of red (R) and blue (B) LED with different R/B light intensity ratios (R/B = 2, 2R1B; R/B = 3, 3R1B; R/B = 4, 4R1B; and R/B = 8, 8R1B) were designed. The results showed that the biomass of lettuce under 8R1B and W treatments was higher than that under other light quality treatments. The photosynthetic rate (Pn) under red and blue light was significantly higher than that of white light. Total root length, root surface area, and root volume were the highest under 8R1B. 4R1B treatment significant increased root activity by 68.6% compared with W. In addition, total organic carbon (TOC) content, TOC content/shoot dry weight, TOC content/root dry weight, and TOC content/root surface area were the highest under 4R1B. Moreover, 8R1B treatment reduced the concentration of benzoic acid and salicylic acid, and the secretion ability of benzoic acid and salicylic acid by per unit root surface area and accumulation by per unit shoot dry weight. In addition, 2R1B and 3R1B reduced the secretion ability of gallic acid and tannic acid by per unit root surface area and accumulation by per unit shoot dry weight. In conclusion, this study showed that the secretion of autotoxins could be reduced through the mediation of red and blue light composition of LEDs in a plant factory. In terms of autotoxin secretion reduction efficiency and yield performance of lettuce, 8R1B light regime is recommended for practical use.

Regulation of Ascorbate Accumulation and Metabolism in Lettuce by the Red:Blue Ratio of Continuous Light Using LEDs.

Ascorbate (AsA), an antioxidant that cannot be synthesized and stored by the human body, plays an essential role in the proper functioning of both plants and humans. With the goal of increasing the AsA level in lettuce, the effects of different ratios of red (R) to blue (B) light (75R:25B, 50R:50B, and 25R:75B) on AsA pool sizes as well as the transcript levels and activities of key enzymes involved in AsA metabolism were constantly monitored for 12 days under continuous light (200 µmol⋅m-2⋅s-1) from LEDs. The results showed that lettuce biomass was positively correlated with the ratio of red light, while the AsA pool size had a positive correlation with the ratio of blue light during the whole experiment. The 25R:75B treatment increased the expression of genes involved in AsA biosynthesis (GMP, GME, GGP, GPP, GLDH) and regeneration (APX, MDHAR, DHAR, and GR) on day 3 but only significantly elevated the activities of enzymes involved in AsA regeneration (APX, MDHAR, DHAR, and GR) subsequently. AsA regeneration enzymes (MDHAR, DHAR and GR) had greater correlations with the AsA level than the AsA synthesis enzyme (GLDH). Thus, it is concluded that a high ratio of blue light elevated the AsA level mainly by promoting AsA regeneration rather than biosynthesis. Taken together, altering the red:blue ratio of continuous light from high to low before harvest is recommended for lettuce cultivation to achieve both high yield and high quality.

Morphological and Physiological Stress Responses of Lettuce to Different Intensities of Continuous Light.

In this study, specific dynamic changes in growth, oxidative stress, ascorbate metabolism, and chlorophyll fluorescence were monitored during 12 days in lettuce plants exposed to continuous light (CL) of different intensities: low light (LL, 100 µmol·m-2·s-1), medium light (ML, 200 µmol·m-2·s-1), and high light (HL, 300 µmol·m-2·s-1). Lettuce plants grown under CL of higher light intensity gained greater biomass, dry weight ratio, root/shoot ratio, and specific leaf FW, but not leaf area. Both the reactive oxygen species (ROS) production and the lipid peroxidation degree, measured in terms of the malondialdehyde (MDA) levels, were progressively enhanced by increasing the light intensity of CL. Overall, the pool sizes of ascorbate (AsA) and glutathione, as well as the activities of enzymes involved in AsA metabolism, had positive correlations with light intensity under CL. Ascorbate peroxidase and dehydroascorbate reductase presented the maximal and minimal responses to light intensity, respectively, among all the studied enzymes. After 6 days under CL, ML and HL intensity caused reversible photoinhibition, represented by lower values of maximum quantum efficiency (F v /F m), effective quantum yield (ΦPSII), and photochemical quenching (qP) and a higher value of non-photochemical quenching (qN). However, this photoinhibition recovered on day 12 with increasing of F v /F m, ΦPSII, and qP. Taken together, under ML and HL conditions, greater AsA level could help maintain photosynthetic efficiency by elevating excess excitation energy dissipation, though ROS accumulation and lipid peroxidation could not be prevented in the long-term. Likewise, there was no dark period under LL condition, but no photooxidative stress was observed in lettuce. Thus, it is concluded that photooxidative stress induced by CL can be attributed to excessive daily light integral instead of circadian asynchrony.