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BACKGROUND: Plant biostimulants are diverse substances and microorganisms used to enhance plant growth. The global market for biostimulants is projected to increase 12 % per year and reach over $2,200 million by 2018. Despite the growing use of biostimulants in agriculture, many in the scientific community consider biostimulants to be lacking peer-reviewed scientific evaluation. SCOPE: This article describes the emerging definitions of biostimulants and reviews the literature on five categories of biostimulants: i. microbial inoculants, ii. humic acids, iii. fulvic acids, iv. protein hydrolysates and amino acids, and v. seaweed extracts. CONCLUSIONS: The large number of publications cited for each category of biostimulants demonstrates that there is growing scientific evidence supporting the use of biostimulants as agricultural inputs on diverse plant species. The cited literature also reveals some commonalities in plant responses to different biostimulants, such as increased root growth, enhanced nutrient uptake, and stress tolerance.

The use of bioeffectors, formally known as plant biostimulants, has become common practice in agriculture and provides a number of benefits in stimulating growth and protecting against stress. A biostimulant is loosely defined as an organic material and/or microorganism that is applied to enhance nutrient uptake, stimulate growth, enhance stress tolerance or crop quality. This review is intended to provide a broad overview of known effects of biostimulants and their ability to improve tolerance to abiotic stresses. Inoculation or application of extracts from algae or other plants have beneficial effects on growth and stress adaptation. Algal extracts, protein hydrolysates, humic and fulvic acids, and other compounded mixtures have properties beyond basic nutrition, often enhancing growth and stress tolerance. Non-pathogenic bacteria capable of colonizing roots and the rhizosphere also have a number of positive effects. These effects include higher yield, enhanced nutrient uptake and utilization, increased photosynthetic activity, and resistance to both biotic and abiotic stresses. While most biostimulants have numerous and diverse effects on plant growth, this review focuses on the bioprotective effects against abiotic stress. Agricultural biostimulants may contribute to make agriculture more sustainable and resilient and offer an alternative to synthetic protectants which have increasingly falling out of favour with consumers. An extensive review of the literature shows a clear role for a diverse number of biostimulants that have protective effects against abiotic stress but also reveals the urgent need to address the underlying mechanisms responsible for these effects. Graphical abstract Biostimulants have protective effects against abiotic stress.

Background Fulvic acid (FA) is a kind of plant growth regulator, which can promote plant growth, play an important role in fighting against drought, improve plant stress resistance, increase production and improve quality. However, the function of FA in tea plants during drought stress remain largely unknown. Results Here, we examined the effects of 0.1 g/L FA on genes and metabolites in tea plants at different periods of drought stress using transcriptomics and metabolomics profiles. Totally, 30,702 genes and 892 metabolites were identified. Compared with controlled groups, 604 and 3331 differentially expressed metabolite genes (DEGs) were found in FA-treated tea plants at 4 days and 8 days under drought stress, respectively; 54 and 125 differentially expressed metabolites (DEMs) were also found at two time points, respectively. Bioinformatics analysis showed that DEGs and DEMs participated in diverse biological processes such as ascorbate metabolism ( GME , AO, ALDH and L-ascorbate), glutathione metabolism ( GST , G6PDH , glutathione reduced form and CYS-GYL), and flavonoids biosynthesis ( C4H , CHS , F3’5’H , F3H , kaempferol, quercetin and myricetin). Moreover, the results of co-expression analysis showed that the interactions of identified DEGs and DEMs diversely involved in ascorbate metabolism, glutathione metabolism, and flavonoids biosynthesis, indicating that FA may be involved in the regulation of these processes during drought stress. Conclusion The results indicated that FA enhanced the drought tolerance of tea plants by (i) enhancement of the ascorbate metabolism, (ii) improvement of the glutathione metabolism, as well as (iii) promotion of the flavonoids biosynthesis that significantly improved the antioxidant defense of tea plants during drought stress. This study not only confirmed the main strategies of FA to protect tea plants from drought stress, but also deepened the understanding of the complex molecular mechanism of FA to deal with tea plants to better avoid drought damage.

Background Enriching the soil with organic matter such as humic and fulvic acid to increase its content available nutrients, improves the chemical properties of the soil and increases plant growth as well as grain yield. In this study, we conducted a field experiment using humic acid (HA), fulvic acid (FA) and recommended dose (RDP) of phosphorus fertilizer to treat Hordeum vulgare seedling, in which four concentrations from HA, FA and RDP (0.0 %, 50 %, 75 % and 100%) under saline soil conditions . Moreover, some agronomic traits (e.g. grain yield, straw yield, spikes weight, plant height, spike length and spike weight) in barley seedling after treated with different concentrations from HA, FA and RDP were determined. As such the beneficial effects of these combinations to improve plant growth, N, P, and K uptake, grain yield, and its components under salinity stress were assessed. Results The findings showed that the treatments HA + 100% RDP (T1), HA + 75% RDP (T2), FA + 100% RDP (T5), HA + 50% RDP (T3), and FA + 75% RDP (T6), improved number of spikes/plant, 1000-grain weight, grain yield/ha, harvest index, the amount of uptake of nitrogen (N), phosphorous (P) and potassium (K) in straw and grain. The increase for grain yield over the control was 64.69, 56.77, 49.83, 49.17, and 44.22% in the first season, and 64.08, 56.63, 49.19, 48.87, and 43.69% in the second season,. Meanwhile, the increase for grain yield when compared to the recommended dose was 22.30, 16.42, 11.27, 10.78, and 7.11% in the first season, and 22.17, 16.63, 11.08, 10.84, and 6.99% in the second season. Therefore, under salinity conditions the best results were obtained when, in addition to phosphate fertilizer, the soil was treated with humic acid or foliar application the plants with fulvic acid under one of the following treatments: HA + 100% RDP (T1), HA + 75% RDP (T2), FA + 100% RDP (T5), HA + 50% RDP (T3), and FA + 75% RDP (T6). Conclusions The result of the use of organic amendments was an increase in the tolerance of barley plant to salinity stress, which was evident from the improvement in the different traits that occurred after the treatment using treatments that included organic amendments (humic acid or fulvic acid).

Aims Long-term application of organic materials has been shown to significantly enhance the content of soil organic matter (SOM), underscoring the critical need to examine the components of soil organic carbon for a deeper understanding of SOM functionalities. Thus, the structural changes and microbial driving mechanisms of water extractable organic matter (WEOM), fulvic acid (FA) and humic acid (HA) were investigated in black soil by a long-term fertilization experiment. Methods This 33-year experiment comprises five treatments: no fertilizer (CK), chemical fertilizer (NPK), chemical fertilizer with low-rate straw (NPKJ1), chemical fertilizer with high-rate straw (NPKJ2), and chemical fertilizer with organic manure (NPKM). We also conducted a detailed study of WEOM, FA, HA, and the microbial community structure in both the 0–20 cm and 20–40 cm soil layers. Results Our findings indicate that organic material application primarily sourced WEOM, FA, and HA from microbial metabolism and plant-derived origins, exhibiting humus and aromatization characteristics with high molecular weight. WEOM was rich in fulvic acid-like and humic acid-like compounds, while FA and HA contained more protein-like components. Organic material use altered WEOM, FA, and HA structures by impacting soil microbial biomass carbon (MBC) and fungal/bacterial biomass. In 0–20 cm soil layer, SOM content was mainly influenced by humus, especially the HA fraction, whereas in 20–40 cm soil layer, it was predominantly affected by WEOM. Conclusions The present study emphasizes that the application of organic materials can influence the structure of microbial communities, thereby affecting the composition of WEOM, FA, and HA, consequently influencing the organic matter content in different soil layers.

The heterotrophic denitrification requires the participation of electrons which are derived from direct electron donor (usually nicotinamide adenine dinucleotide (NADH)), and the electrons are transferred via electron transport system in denitrifiers and then consumed by denitrifying enzymes. Despite the reported electron transfer ability of humic substances (HS), the influences of fulvic acid (FA), an ubiquitous major component of HS, on promoting NADH generation, electron transfer, and consumption in denitrification process have never been reported. The presence of FA, compared with the control, was found not only significantly improved the total nitrogen (TN) removal efficiency (99.9 % versus 74.8 %) but remarkably reduced the nitrite accumulation (0.2 against 43.8 mg/L) and N 2 O emission (0.003 against 0.240 mg nitrogen/mg TN removed). The mechanisms study showed that FA increased the metabolism of carbon source via glycolysis and tricarboxylic acid (TCA) cycle pathways to produce more available NADH. FA also facilitated the electron transfer activities from NADH to denitrifying enzymes via complex I and complex III in electron transport system, which improved the reduction of nitrate and accelerated the transformations of nitrite and N 2 O, and lower nitrite and N 2 O accumulations were therefore observed. In addition, the consumption of electrons in denitrification was enhanced due to FA stimulating the synthesis and the catalytic activity of key denitrifying enzymes, especially nitrite reductase and N 2 O reductase. It will provide an important new insight into the potential effect of FA on microbial denitrification metabolism process and even nitrogen cycle in nature niches.

Background HS-2 ® Pro, a humic product, is produced from composted thinnings of coniferous trees and is commercially available as a biostimulant. In this study, it was fractionated into humic acid (HA), hydrophobic fulvic acid (FA), hydrophilic FA, and highly hydrophilic FA to identify humic fractions with biostimulant effects. Each fraction was characterized using diffuse reflectance infrared Fourier transform (DRIFT) spectroscopy, high-performance size-exclusion chromatography (HPSEC), and reversed-phase high-performance liquid chromatography (RP-HPLC) with π–π interactions, and their root elongation-promoting and antioxidant activities were evaluated. Results The humic product comprised approximately 70% of its weight on an ash-free basis in the HA fraction, 15% in the hydrophobic FA fraction, 2.5% in the hydrophilic FA fraction, and 4.7% in the highly hydrophilic FA fraction. DRIFT spectroscopy showed that the composition of functional groups varied considerably among the fractions. The estimation of the size distribution of the molecular associations in each fraction by HPSEC indicated that the constituents of the HA fraction are forming larger molecular associations, and the constituents of the FA fractions occur as relatively small aggregates and unbound molecules. The RP-HPLC analysis revealed that each humic fraction was composed of many constituents with different degrees of hydrophilicity/hydrophobicity. The root elongation-promoting activity evaluated from the elongation of the primary roots of komatsuna seedlings was highest in the hydrophilic and highly hydrophilic FA fractions, followed by the HA fraction, whereas the hydrophobic FA fraction showed no significant activity. The antioxidant activity evaluated based on the 2,2-diphenyl-1-picrylhydrazyl radical scavenging activity was higher in the HA and hydrophobic FA fractions and lower in the hydrophilic and highly hydrophilic FA fractions. Conclusion The present study revealed that the constituents of the hydrophilic FA, highly hydrophilic FA, and HA fractions were responsible for the root elongation-promoting activity of the humic product, whereas the constituents of the HA and hydrophobic FA fractions were responsible for the antioxidant activity. Graphical Abstract

Salt stress damage to plants has been becoming a global concern for agriculture. The application of potassium fulvic acid (PFA) is a promising strategy to alleviate the damage to plants and improve soil quality. However, the study of PFA on plant growth and rhizosphere microbial community remains limited. In this study, microcosmic experiments were conducted to verify the effect of PFA on citrus. Trifoliate orange ( Poncirus trifoliata ), the most important citrus rootstock, was used to evaluate the effect of PFA on salt damage. The results showed that PFA significantly increased the contents of chlorophyll a, chlorophyll b and carotenoid by 30.09%, 17.55% and 27.43%, and effectively avoided the yellowing and scorching of leaves under salt stress. Based on the results of two-way ANOVA, the mitigation of salt stress on trifoliate seedlings primarily attributed to the enhancement of protective enzyme activities, K + /Na + ratio and the contents of soluble sugar, soluble protein and proline. Moreover, PFA enhanced neutral protease (S-NPT), sucrase (S-SC) and urease (S-UE) of rhizosphere soil and improved soil nutrition status. The abundance of Bacillus , a kind of rhizosphere beneficial bacteria, was improved by PFA under salt stress, which was mainly associated with the increased activities of S-NPT, S-SC and S-UE. Overall, the application of PFA showed great potential for the alleviation of salt damage on citrus.

Fulvic acid (FA) can significantly alleviate cadmium (Cd) stress, but the specific metabolic response of FA to Cd toxicity is still not clarified. In the present study, we used untargeted metabolomic [gas chromatography–mass spectrometry (GC–MS) and liquid chromatography–mass spectrometry (LC–MS)] analysis to profile cucumber metabolism in response to Cd stress after spray application of FA. Our results showed that 331 differentially enriched metabolites (DEMs) were identified in leaf materials. These DEMs were enriched in 21 shared pathways in comparative groups of “Cd treatment vs. the control treatment” and “FA + Cd treatment vs. the Cd treatment.” Specifically, treatment with FA significantly enhanced the organic acid content (citric acid, isocitric acid, 2-oxoglutaric acid, fumaric acid, and malic acid), which would contribute to provide sufficient substrates for the tricarboxylic acid (TCA) cycle and amino acid biosynthesis, thereby ensuring the normal production of energy and amino acid. At the same time, FA significantly increased the amino acid content (aspartate, citrulline, histidine, leucine, and phenylalanine). The accumulation of organic acid and amino acid can act as chelating agents for heavy metal ions and as scavengers of reactive oxygen species (ROS), thereby reducing intracellular oxidative damage. Furthermore, the application of FA improves antioxidant enzymes and accelerates ROS clearance. The improved contents of organic acid and amino acid, and the increased activity of antioxidant enzymes both played a central role in the reduction of malondialdehyde (MDA, 14.08%), hydrogen peroxide (H 2 O 2 , 61.70%) contents, and superoxide anion radical (O 2 − , 30.41%) production rate in plants under Cd stress. Taken together, the present study demonstrates the effects of FA on the antioxidant capacity and carbohydrate and amino acid metabolism of cucumber seedlings exposed to Cd stress, which provides comprehensive insights into the regulation of plants’ response to Cd toxicity with FA was applied in cucumber.

Background Fulvic acid enhances plant growth and interacts synergistically with phosphate fertilizer to alleviate the agricultural production problem of low phosphorus fertilizer utilization efficiency. However, the underlying mechanism of its action remains poorly understood. In this study, we investigated the impact of fulvic acid application with varying concentrations (0, 40, 60, 80 and 120 mg/L) on rice performance in plants grown in a hydroponic system subjected to low phosphorus stress. The rice growth phenotypes, biomass, root morphology, phosphorus uptake, and the impact of fulvic acid on the rhizosphere environment of rice, were assessed. Results The findings showed that adding appropriate concentrations of exogenous fulvic acid could promote the growth performance of rice under low phosphorus stress. Particularly at T1 (40 mg/L) and T2 (60 mg/L) over the control effectively increased rice biomass by 25.42% and 24.56%, respectively. Fulvic acid treatments stimulated root morphogenesis, up-regulated phosphate transporter genes, and facilitated phosphorus absorption and accumulation. Especially T1 (20.52%), T2 (18.10%) and T3 (20.48%) treatments significantly increased phosphorus uptake in rice, thereby alleviating low phosphorus stress. Additionally, fulvic acid elevated organic acids concentration in roots and up-regulated plasma membrane H + -ATPase genes, promoting organic acids secretion. This metabolic alteration can also alleviate low phosphorus stress in rice. Conclusions The effect of exogenous fulvic acid on physiological indicators is concentration-dependent under low phosphorus stress, enhances rice performance and reduces reliance on phosphorus fertilizer. This provides new insights to shed light on the mechanism of alleviating low phosphorus stress in rice through fulvic acid application, an eco-friendly tool.