Photosystems and antioxidative system of rye, wheat and triticale under Pb stress.

Lead (Pb2+) pollution in the soil sub-ecosystem has been a continuously growing problem due to economic development and ever-increasing anthropogenic activities across the world. In this study, the photosynthetic performance and antioxidant capacity of Triticeae cereals (rye, wheat and triticale) were compared to assess the activities of antioxidants, the degree of oxidative damage, photochemical efficiency and the levels of photosynthetic proteins under Pb stress (0.5 mM, 1 mM and 2 mM Pb (NO3)2). Compared with triticale, Pb treatments imposed severe oxidative damage in rye and wheat. In addition, the highest activity of major antioxidant enzymes (SOD, POD, CAT, and GPX) was also found to be elevated. Triticale accumulated the highest Pb contents in roots. The concentration of mineral ions (Mg, Ca, and K) was also high in its leaves, compared with rye and wheat. Consistently, triticale showed higher photosynthetic activity under Pb stress. Immunoblotting of proteins revealed that rye and wheat have significantly lower levels of D1 (photosystem II subunit A, PsbA) and D2 (photosystem II subunit D, PsbD) proteins, while no obvious decrease was noticed in triticale. The amount of light-harvesting complex II b6 (Lhcb6; CP24) and light-harvesting complex II b5 (Lhcb5; CP26) was significantly increased in rye and wheat. However, the increase in PsbS (photosystem II subunit S) protein only occurred in wheat and triticale exposed to Pb treatment. Taken together, these findings demonstrate that triticale shows higher antioxidant capacity and photosynthetic efficiency than wheat and rye under Pb stress, suggesting that triticale has high tolerance to Pb and could be used as a heavy metal-tolerant plant.

A Comprehensive Evaluation of Salt Tolerance in Tomato (Var. Ailsa Craig): Responses of Physiological and Transcriptional Changes in RBOH's and ABA Biosynthesis and Signalling Genes.

Salinity is a ubiquitous stressor, depleting osmotic potential and affecting the tomato seedlings' development and productivity. Considering this critical concern, we explored the salinity response in tomato seedlings by evaluating them under progressive salt stress duration (0, 3, 6, and 12 days). Intriguingly, besides the adverse effect of salt stress on tomato growth the findings exhibited a significant role of tomato antioxidative system, RBOH genes, ABA biosynthesis, and signaling transcription factor for establishing tolerance to salinity stress. For instance, the activities of enzymatic and non-enzymatic antioxidants continued to incline positively with the increased levels of reactive oxygen species (O2•-, H2O2), MDA, and cellular damage, suggesting the scavenging capacity of tomato seedlings against salt stress. Notably, the RBOH transcription factors activated the hydrogen peroxide-mediated signalling pathway that induced the detoxification mechanisms in tomato seedlings. Consequently, the increased gene expression of antioxidant enzymes and the corresponding ratio of non-enzymatic antioxidants AsA-GSH suggested the modulation of antioxidants to survive the salt-induced oxidative stress. In addition, the endogenous ABA level was enhanced under salinity stress, indicating higher ABA biosynthesis and signalling gene expression. Subsequently, the upregulated transcript abundance of ABA biosynthesis and signalling-related genes suggested the ABA-mediated capacity of tomato seedlings to regulate homeostasis under salt stress. The current findings have revealed fascinating responses of the tomato to survive the salt stress periods, in order to improve the abiotic stress tolerance in tomato.

Salicylic Acid Protects Photosystem II by Alleviating Photoinhibition in Arabidopsis thaliana under High Light.

Salicylic acid (SA) is considered to play an important role in plant responses to environmental stresses. However, the detailed protective mechanisms in photosynthesis are still unclear. We therefore explored the protective roles of SA in photosystem II (PSII) in Arabidopsis thaliana under high light. The results demonstrated that 3 h of high light exposure resulted in a decline in photochemical efficiency and the dissipation of excess excitation energy. However, SA application significantly improved the photosynthetic capacity and the dissipation of excitation energy under high light. Western blot analysis revealed that SA application alleviated the decrease in the levels of D1 and D2 protein and increased the amount of Lhcb5 and PsbS protein under high light. Results from photoinhibition highlighted that SA application could accelerate the repair of D1 protein. Furthermore, the phosphorylated levels of D1 and D2 proteins were significantly increased under high light in the presence of SA. In addition, we found that SA application significantly alleviated the disassembly of PSII-LHCII super complexes and LHCII under high light for 3 h. Overall, our findings demonstrated that SA may efficiently alleviate photoinhibition and improve photoprotection by dissipating excess excitation energy, enhancing the phosphorylation of PSII reaction center proteins, and preventing the disassembly of PSII super complexes.

Phytochemical screening and allelopathic potential of phytoextracts of three invasive grass species.

Undoubtedly, it is important to remain vigilant and manage invasive grasses to prevent their spread and mitigate their negative impact on the environment. However, these aggressive plants can also play a beneficial role in certain contexts. For example, several invasive grasses provide valuable forage for livestock and have disease control potential. Therefore, a research experiment was conducted to explore the pros and cons of this approach, not only for surrounding vegetation but also for human and animal disease control. The study is primarily focused on developing livestock feed, plant-derived herbicides, and an understanding of the phytotoxic effects of invasive species. All plant parts of Cenchrus ciliaris L., Polypogon monspeliansis L., and Dicanthium annulatum (Forssk.) Stapf, were tested for their phyto-chemical screening, proximate, and toxicity analysis which was caused by the methanolic extract of these grass species. Qualitative phytochemical screening tests were performed for proximate composition analysis and toxicity assessment essays. The phytochemical analysis revealed the positive results for alkaloids, flavonoids, coumarins, phenols, saponins, and glycosides, while negative for tannins. Comparison of proximate analysis intimated maximum moisture (10.8%) and crude fat (4.1%) in P. monspeliensis, whereas maximum dry matter (84.1%), crude protein (13.95%), crude fiber (11%), and ash (7.2%) in D. annulatum. Five (10, 100, 500, 100, 10,000 ppm) and three (10, 1000, 10,000 ppm) different concentrations of methanolic extract prepared from C. ciliaris, P. monspeliansis, and D. annulatum were used respectively for root inhibition and seed germination essay. Furthermore, three different concentrations (10, 30, 50 mg) of plant fine powder were used for sandwich method test. There was a significant decline in the growth rate of experimental model radish seeds (P > 0.005), and results from sandwich method tests showed suppressed growth of root hairs, inhibiting the anchoring of the radish seed. In comparison, results manifest that; P. monspeliansis indicated an upsurge of inhibition (66.58% at 10,000 ppm), D. annulatum revealed soar germination (75.86% in controlled conditions), and C. ciliaris exhibited dramatic shoot up of inhibition because of sandwich method test (14.02% at 50 mg). In conclusion, although grasses are toxic, it is important to consider the beneficiary account.

Corrigendum: Foliar application of putrescine alleviates terminal drought stress by modulating water status, membrane stability, and yield- related traits in wheat (Triticum aestivum L.).

[This corrects the article DOI: 10.3389/fpls.2022.1000877.].

Expression profiling of TaARGOS homoeologous drought responsive genes in bread wheat.

Drought tolerant germplasm is needed to increase crop production, since water scarcity is a critical bottleneck in crop productivity worldwide. Auxin Regulated Gene involved in Organ Size (ARGOS) is a large protein family of transcription factors that plays a vital role in organ size, plant growth, development, and abiotic stress responses in plants. Although, the ARGOS gene family has been discovered and functionalized in a variety of crop plants, but a comprehensive and systematic investigation of ARGOS genes in locally used commercial wheat cultivars is still yet to be reported. The relative expression of three highly conserved TaARGOS homoeologous genes (TaARGOS-A, TaARGOS-B, TaARGOS-D) was studied in three drought-tolerant (Pakistan-2013, NARC-2009 and NR-499) and three sensitive (Borlaug-2016, NR-514 and NR-516) wheat genotypes under osmotic stress, induced by PEG-6000 at 0 (exogenous control), 2, 4, 6, and 12 h. The normalization of target genes was done using ß-actin as endogenous control, whereas DREB3, as a marker gene was also transcribed, reinforcing the prevalence of dehydration in all stress treatments. Real-time quantitative PCR revealed that osmotic stress induced expression of the three TaARGOS transcripts in different wheat seedlings at distinct timepoints. Overall, all genes exhibited significantly higher expression in the drought-tolerant genotypes as compared to the sensitive ones. For instance, the expression profile of TaARGOS-A and TaARGOS-D showed more than threefold increase at 2 h and six to sevenfold increase after 4 h of osmotic stress. However, after 6 h of osmotic stress these genes started to downregulate, and the lowest gene expression was noticed after 12 h of osmotic stress. Among all the homoeologous genes, TaARGOS-D, in particular, had a more significant influence on controlling plant growth and drought tolerance as it showed the highest expression. Altogether, TaARGOSs are involved in seedling establishment and overall plant growth. In addition, the tolerant group of genotypes had a much greater relative fold expression than the sensitive genotypes. Ultimately, Pakistan-2013 showed the highest relative expression of the studied genes than other genotypes which shows its proficiency to mitigate osmotic stress. Therefore, it could be cultivated in arid and semi-arid regions under moisture-deficient regimes. These findings advocated the molecular mechanism and regulatory roles of TaARGOS genes in plant growth and osmotic stress tolerance in contrasting groups of wheat genotypes, accompanied by the genetic nature of identified genotypes in terms of their potential for drought tolerance.

N-Acetylcysteine Priming Alleviates the Transplanting Injury of Machine-Transplanted Rice by Comprehensively Promoting Antioxidant and Photosynthetic Systems.

The stress of transplanting injury adversely affects rice growth and productivity worldwide. N-acetylcysteine (NAC), the precursor of glutathione, is a potent ROS scavenger with powerful antioxidant activity. Previous studies on the application of NAC in plants mainly focused on alleviating the stress of heavy metals, UV-B, herbicides, etc. However, the role of NAC in alleviating transplanting injury is still not clear. A barrel experiment was carried out to explain the mechanism of NAC regulating the transplanting injury to machine-transplanted rice during the recovery stage. The results showed that NAC priming shortened the time of initiation of tillering and increased the tiller numbers within 3 weeks after transplanting. In addition, NAC priming increased the chlorophyll content, net photosynthetic rate, and sucrose content, thereby improving the dry weight at the recovery stage, especially root dry weight. At the same time, NAC priming significantly increased the activity of ascorbate peroxidase (APX), glutathione reductase (GR), catalase (CAT), and superoxide dismutase (SOD). In addition, it also regulated flavonoids and total phenols contents to reduce hydrogen peroxide (H2O2) and malondialdehyde (MDA) contents, especially at the initial days after transplanting. These results suggest that NAC priming improves the tolerance of rice seedlings against transplanting injury by enhancing photosynthesis and antioxidant systems at initial days after transplanting, thereby promoting the accumulation of dry matter and tillering for higher yield returns.

Foliar application of putrescine alleviates terminal drought stress by modulating water status, membrane stability, and yield- related traits in wheat (Triticum aestivum L.).

Drought stress is one of the major limitations to the growth and yield productivity of cereal crops. It severely impairs the early growing and grain -filling stages of wheat. Therefore, cost- effective and eco-friendly approaches for alleviating drought stress in cereal crops are in high demand. Polyamines, such as putrescine, have a significant effect on improving crop yield under drought- stress conditions. Therefore, the current study was executed with the aim of exploring the significance of putrescine in alleviating drought stress and improving yield- related traits in wheat. Two distinct wheat cultivars (Fakhar-e-Bhakkar and Anaj-2017) were treated with the foliar application of different concentrations (control, 0.5, 1.0, and 1.5 PPM) of putrescine (put) under two moisture conditions (well- watered and terminal drought stress). The results demonstrate that the imposition of terminal drought stress significantly reduces different physiological and yield- related traits of both wheat cultivars. The reduction of relative water content (RWC%), membrane stability index (MSI), leaf area, tillers per plant, biomass yield, number of spikelets per spike, 100-grain weight, grain yield per plant, and straw yield was greater in Anaj-2017 than in Fakhar-e-Bhakkar cultivar. The results further explain that the foliar application of increased concentrations of putrescine from 0.0 to 1.0 PPM gradually improved physiological and yield traits, whereas these traits declined with the application of putrescine at the highest dose (1.5 PPM). The exogenous application of 1.0 PPM putrescine improved the relative water content (19.76%), specific leaf area (41.47%), and leaf area ratio (35.84%) compared with the controlled treatment. A higher grain yield (28.0 g plant-1) and 100-grain weight (3.8 g) were obtained with the foliar application of 1.0 PPM putrescine compared with controlled treatments. The findings of this study confirm the protective role of putrescine against terminal drought stress. It is therefore recommended to use putrescine at a concentration of 1.0 PPM, which could help alleviate terminal drought stress and attain better wheat yield.

Exogenous spermidine modulates polyamine metabolism and improves stress responsive mechanisms to protect tomato seedlings against salt stress.

Salt stress negatively affects plant growth, development, and crop productivity causing serious economic loss to agricultural production. Here, we investigated the exogenous application of spermidine (Spd) on tomato seedlings grown under salt stress. Salt stress reduced plant growth, biomass accumulation and chlorophyll contents, thus negatively affecting photosynthesis. Alternatively, Spd application effectively reduced the salinity-induced adverse effects in tomato seedlings by activating the H2O2 mediated signaling involving the enhanced expression of RBOH1 and salt stress-responsive genes SlMYB102, SlHKT1, SlWRKY1 and SlDREB2, and improving detoxification through higher antioxidative activity and osmolyte (proline) accumulation under salt stress. It was further confirmed by significantly lower amount of H2O2, malondialdehyde and electrolyte leakage, and better ion homeostasis (Na+/K+ ratio) and photosynthetic performance of Spd-treated seedlings under salt stress. Furthermore, Spd application modulated endogenous polyamines and enhanced the biosynthesis of endogenous Spd and spermine from putrescine. Altogether, these results confirm the important role of Spd against salt stress and suggest that the increased endogenous Spd content in plants could regulate a number of stress-responsive mechanisms to protect tomato seedlings against salt stress. These results provide a good direction for further elucidation of the detailed interplay between polyamine metabolism and H2O2-mediated signaling, which would help to improve abiotic stress tolerance in plants.

Comparative analysis of farmer practices and high yield experiments: Farmers could get more maize yield from maize-soybean relay intercropping through high density cultivation of maize.

Intercropping is a high-yield, resource-efficient planting method. There is a large gap between actual yield and potential yield at farmer's field. Their actual yield of intercropped maize remains unclear under low solar radiation-area, whether this yield can be improved, and if so, what are the underlying mechanism for increasing yield? In the present study, we collected the field management and yield data of intercropping maize by conducting a survey comprising 300 farmer households in 2016-2017. Subsequently, based on surveyed data, we designed an experiment including a high density planting (Dense cultivation and high N fertilization with plough tillage; DC) and normal farmer practice (Common cultivation; CC) to analyze the yield, canopy structure, light interception, photosynthetic parameters, and photosynthetic productivity. Most farmers preferred rotary tillage with a low planting density and N fertilization. Survey data showed that farmer yield ranged between 4-6 Mg ha-1, with highest yield recorded at 10-12 Mg ha-1, suggesting a possibility for yield improvement by improved cropping practices. Results from high density experiment showed that the two-years average yield for DC was 28.8% higher than the CC. Compared to CC, the lower angle between stem and leaf (LA) and higher leaf area index (LAI) in DC resulted in higher light interception in middle canopy and increased the photosynthetic productivity under DC. Moreover, in upper and lower canopies, the average activity of phosphoenolpyruvate (PEP) carboxylase was 70% higher in DC than CC. Briefly, increase in LAI and high Pn improved both light interception and photosynthetic productivity, thereby mediating an increase in the maize yield. Overall, these results indicated that farmer's yields on average can be increased by 2.1 Mg ha-1 by increasing planting density and N fertilization, under plough tillage.

Maize/soybean strip intercropping produces higher crop yields and saves water under semi-arid conditions.

Sustainable increases in crop production require efficient use of resources, and intercropping can improve water use efficiency and land productivity at reduced inputs. Thus, in a three-year field experiment, the performance of maize/soybean strip intercropping system differing with maize plant density (6 maize plants m-2, low, D1; 8 maize plants m-2, medium, D2; and 10 maize plants m-2, high, D3) was evaluated in comparison with sole maize or soybean cropping system. Results revealed that among all intercropping treatments, D2 had a significantly higher total leaf area index (maize LAI + soybean LAI; 8.2), total dry matter production (maize dry matter + soybean dry matter; 361.5 g plant-1), and total grain yield (maize grain yield + soybean grain yield; 10122.5 kg ha-1) than D1 and D3, and also higher than sole maize (4.8, 338.7 g plant-1, and 9553.7 kg ha-1) and sole soybean (4.6, 64.8 g plant-1, and 1559.5 kg ha-1). The intercropped maize was more efficient in utilizing the radiation and water, with a radiation use efficiency of 3.5, 5.2, and 4.3 g MJ-1 and water use efficiency of 14.3, 16.2, and 13.3 kg ha-1 mm-1, while that of intercropped soybean was 2.5, 2.1, and 1.8 g MJ-1 and 2.1, 1.9, and 1.5 kg ha-1 mm-1 in D1, D2, and D3, respectively. In intercropping, the land and water equivalent ratios ranged from 1.22 to 1.55, demonstrating that it is a sustainable strategy to improve land and water use efficiencies; this maximization is likely associated with the species complementarities for radiation, water, and land in time and space, which resulted in part from competition avoidance responses that maximize the economic profit (e. g., 1300 US $ ha-1 in D2) over sole maize (798 US $ ha-1) or sole soybean (703 US $ ha-1). Overall, these results indicate that optimizing strip intercropping systems can save 20-50% of water and land, especially under the present scenario of limited resources and climate change. However, further research is required to fully understand the resource capture mechanisms of intercrops in intercropping.

Metabolic Disturbance Induced by the Embryo Contributes to the Formation of Chalky Endosperm of a Notched-Belly Rice Mutant.

Grain chalkiness is a key quality trait of the rice grain, whereas its underlying mechanism is still not thoroughly understood because of the complex genetic and environmental interactions. We identified a notched-belly (NB) mutant that has a notched-line on the belly of grains. The line dissects the endosperm into two distinct parts, the upper translucent part, and the bottom chalky part in the vicinity of the embryo. Using this mutant, our previous studies clued the negative influence of embryo on the biochemical makeup of the endosperm, suggesting the need for the in-depth study of the embryo effect on the metabolome of developing endosperm. This study continued to use the NB mutant to evolve a novel comparison method to clarify the role of embryo in the formation of a chalky endosperm. Grain samples of the wild-type (WT) and NB were harvested at 10, 20, and 30 days after fertilization (DAF), and then divided into subsamples of the embryo, the upper endosperm, and the bottom endosperm. Using non-targeted metabolomics and whole-genome RNA sequencing (RNA-seq), a nearly complete catalog of expressed metabolites and genes was generated. Results showed that the embryo impaired the storage of sucrose, amino acid, starch, and storage proteins in the bottom endosperm of NB by enhancing the expression of sugar, amino acids, and peptide transporters, and declining the expression of starch, prolamin, and glutelin synthesis-related genes. Importantly, the competitive advantage of the developing embryo in extracting the nutrients from the endosperm, transformed the bottom endosperm into an "exhaustive source" by diverting the carbon (C) and nitrogen (N) metabolism from synthetic storage to secondary pathways, resulting in impaired filling of the bottom endosperm and subsequently the formation of chalky tissue. In summary, this study reveals that embryo-induced metabolic shift in the endosperm is associated with the occurrence of grain chalkiness, which is of relevance to the development of high-quality rice by balancing the embryo-endosperm interaction.

Growth and development of soybean under changing light environments in relay intercropping system.

BACKGROUND: Maize-soybean relay-intercropping (MSR) is a famous system of crop production in developing countries. However, maize shading under this system directly affects the light quality and intensity of soybean canopy. This is a challenging scenario in which to implement the MSR system, in terms of varieties selection, planting pattern, and crop management since the duration of crop resource utilization clearly differs. METHODS: Therefore, this experiment aimed to elucidate the effect of leaf excising treatments from maize top to fully clarify the needs and balance of light quality and intensity of intercrop-soybean under MSR in field conditions. The effects of different leaf excising treatments (T0, no removal of leaves; T2, removal of two topmost leaves; T4, removal of four topmost leaves; T6, removal of six topmost leaves from maize plants were applied at first-trifoliate stage (V1) of soybean) on photosynthetically active radiation transmittance (PART), red to far-red ratio (R:FR), morphological and photosynthetic characteristics and total biomass production at second-trifoliate stage (V2), fifth-trifoliate stage (V5), and flowering-stage (R1) of soybean were investigated through field experiments for 2-years under MSR. RESULTS: As compared to T0, treatment T6 increased the PART and R:FR ratio at soybean canopy by 77% and 37% (V2), 70% and 34% (V5), and 41% and 36% (R1), respectively. This improved light environment in T6 considerably enhanced the leaf area index, SPAD values and photosynthetic rate of soybean plants by 66%, 25% and 49% at R1, respectively than T0. Similarly, relative to control, T6 also increased the stem diameter (by 29%) but decreased the plant height (by 23%) which in turn significantly increased stem breaking strength (by 87%) by reducing the lodging rate (by 59%) of soybean plants. Overall, under T6, relay-cropped soybean produced 78% of sole soybean seed-yield, and relay-cropped maize produced 81% of sole maize seed-yield. Our findings implied that by maintaining the optimum level of PART (from 60% to 80%) and R:FR ratio (0.9 to 1.1), we can improve morphological and photosynthetic characteristics of soybean plants in MSR. Therefore, more attention should be paid to the light environment when considering the sustainability of MSR via appropriate planting pattern selection.