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Protein hydrolysates (PH) and humic substances (HS) are biostimulants that promote crop growth. Despite the widening of knowledge and insights provided by research activities over the past thirty years, there are still uncertainties concerning the possible direct absorption by plants of organic nitrogen as amino acids (AA) delivered as PH and the role of HS. This study aimed to assess the impact of PH treatment on the amino acid profile of maize seedlings and the influence of HS on amino acid uptake. Maize seedlings were treated with either PH from the yeast of the sugarcane fermentation process or a synthetic mixture of amino acids mimicking the PH composition using dual-labeled amino acids (13C, 15N glutamic acid) combined with commercially available humic substances. The amino acid profile was analyzed using HPLC, and plant tissues were examined for 15N using nuclear magnetic resonance (NMR) and mass spectrometry. The application of PH stimulated maize growth, with a more significant effect observed in the presence of humic substances. The treatments significantly altered the plants' total amino acid content and composition profile. Maize seedlings actively uptake amino acids, representing 21% of the total nitrogen composition. The ascertained improvement of amino acid uptake stimulated by humic substances is associated with their effect on the differential expression of amino permease transporters and plasma membrane H+-ATPase. PH and amino acids enhance maize growth by significantly increasing amino acid uptake. The addition of humic substances further improved the biostimulant effects. [ABSTRACT FROM AUTHOR]

Background: Traditional agriculture is on the front line of climate change, being most impacted by the increase in the intensity and frequency of extreme events, such as floods, drought and rising temperatures. Local ecological knowledge is a recognized keystone of successfully managed socioecological systems, but loss of soil fertility, water scarcity, incidence of diseases and decreased production due to climate change are linked to the greater vulnerability experienced by traditional farmers. Plant biostimulants are natural products used to stimulate nutrient uptake and efficiency by crops, increase tolerance to abiotic/biotic stress and improve quality without negative impacts on the environment if obtained from renewed sources. Humic substances are some of the most used plant biostimulants in agriculture and play a central role in plant adaptation. Materials and methods: We reviewed and discussed a sample set of papers (n = 52) about humic substances to mitigate abiotic stress in crops using data basis from Web of Science (Clarivate Analytics), Scopus—IBM (International Business Machines Corporation), and Scielo (Scientific Electronic Library Online). Results: The predominance of authors in the global south is notable, but it is not a coincidence, since this is where the effects of climate change will have the greatest impact. The mechanisms involved in the stress mitigation involve the activation of signaling factors, gene response induction, the accumulation of osmoprotective and anti-oxidant compounds, the induction of antioxidative metabolism, ion homeostasis, membrane transport and adjustment of hormonal balance. The intriguing question is: how can a complex mixture of molecules affect so many distinct effects on plants responsible for plant adaptation? Conclusions: The complexity of humic substances challenges our knowledge method, but supramolecular chemistry may provide answers that enable us to broaden our understanding of the plant defense mechanisms modulated by these substances. [ABSTRACT FROM AUTHOR]

Background: Huanglongbing (HLB) is a devastating citrus disease. Before callose deposition, the bacterial infection causes oxidative stress, starting cell damage. Humic substances are among the most efficient antioxidants found in nature. Furthermore, it is also previously reported that humic substances can induce a phenylpropanoid metabolism contributing to the production of antimicrobial compounds. It has been noted by technicians and growers in the Brazil's main citrus belt that orchards continuously treated with humic substances can live well with the symptoms of HLB. Methods: We treated two young citrus orchards (Citrus sinensis cultivars Baía and Pera) with soluble humic substances (HS) isolated from peat either combined or not with a plant growth-promoting bacteria consortium. The activity of key enzymes related to the response against biotic stresses, such as peroxidase (POX), β-1,3-glucanase (PR) and phenylalanine ammonia-lyase (PAL), as well as the differential transcription level of target genes linked to plant stress response by qRT-PCR was monitored for six months. Results: No differences were obtained between sole humic substances and their application combined with plant growth-promoting bacteria, indicating that the microbial consortium had no modulatory effect on HS-treated plants. The treatments promoted the activities of POX, PAL and β-1,3-glucanase, which remained significantly higher concerning the control throughout the evaluation period. In addition, treatments positively regulated the transcription levels of CsPR-7, CsPR-3 and CsPR-11 genes. The experimental data were qualitatively similar to those found in commercial orange orchards treated continuously for 12 and 6 years with humic substances, where the activity and transcription levels were also more significant than for untreated plants. Conclusion: Our results indicate that humic substances can trigger immune-mediated responses in plants and they can be used as a natural chemical priming agent to mitigate disease symptoms and contribute to more resilient citrus cultivation. [ABSTRACT FROM AUTHOR]

The effect of humic substances (HSs) in combination with or without plant-growth-promoting bacteria on soybean vegetative growth and root nodulation was examined in this study. Seeds were inoculated with Bradyrhizobium japonicum SEMIA 5079 in the presence of HSs from leonardite and Herbaspirillum seropedicae HRC54. Additional HSs and H. seropedicae application at the substrate surface was conducted at the V3 stage. The experiment was carried out in a greenhouse using pots filled with a top layer of an Oxisol soil, and plants were harvested at the R1 stage. The HS and H. seropedicae treatments significantly promoted plant shoot and root growth. The number and weight of soybean nodules were higher in the treated plants when compared to a control. The plant nodulation process was affected by the treatments that included activities of malate dehydrogenase (MDH), nitrate reductase (NR) and plasma membrane H+-ATPase (MHA). At low concentrations, the HSs and H. seropedicae increased the nodule quantity, size and weight, favoring plant growth. Combining humic substances and plant-growth-promoting bacteria (PGPB) could be a promising approach to promoting soybean nodulation and increasing crop production. [ABSTRACT FROM AUTHOR]

Soil organic matter (SOM) has a critical role in regulating soil phosphorus (P) dynamics and producing phytoavailable P. However, soil P dynamics are often explained mainly by the effects of soil pH, clay contents, and elemental compositions, such as calcium, iron, and aluminum. Therefore, a better understanding of the mechanisms of how SOM influences phytoavailable P in soils is required for establishing effective agricultural management for soil health and enhancement of soil fertility, especially P-use efficiency. In this review, the following abiotic and biotic mechanisms are discussed; (1) competitive sorption between SOM with P for positively charged adsorption sites of clays and metal oxides (abiotic reaction), (2) competitive complexations between SOM with P for cations (abiotic reaction), (3) competitive complexations between incorporation of P by binary complexations of SOM and bridging cations with the formation of stable P minerals (abiotic reaction), (4) enhanced activities of enzymes, which affects soil P dynamics (biotic reaction), (5) mineralization/immobilization of P during the decay of SOM (biotic reaction), and (6) solubilization of inorganic P mediated by organic acids released by microbes (biotic reaction). [ABSTRACT FROM AUTHOR]

In weathered tropical soils, the inherent pedoclimatic characteristics strengthen the role of soil organic matter (SOM) pools within the carbon cycle for both forest and agricultural ecosystems. The fast SOM turnover and the humid climate hasten the impact of land use and emphasize the importance of water extractable organic matter (WEOM) on soil organic carbon (SOC) dynamics. The goal of this study was to determine the short‐term dynamics of bulk SOM and WEOM pool in weathered Brazilian soil, as conditioned by both land use and type of plant inputs. To this aim we used the appraisal of the natural δ13C‐OC signature to follow the variation of SOC pools and to overcome the analytical issues related to low OC concentration of WEOM fractions. In a one‐year laboratory incubation, soil samples from a natural forest and an adjacent site under sugarcane monoculture were amended with either forest litter or sugarcane straw to evaluate SOM and WEOM dynamics. The bulk SOM was mainly affected by inherited conditions rather than on the organic amendments. The incorporation of sugarcane residues in the forest site produced a partial replacement of the original SOC, while the amendments to the cultivated soil promoted a stable increase in SOC. The analysis of WEOM highlighted its influence as a reactive pool in the short‐term SOM dynamics. The δ13C variation of WEOM from cultivated soil corroborated the significant ponderal rate of OC input from forest litter that accounted for 22% of total dissolved carbon. Unexpected evidence on the shift of isotopic dilution between SOC pools indicates that the evaluation of δ13C to determine the origin of the SOC fraction should be conservatively adopted. The results outline that the steady state of SOM in forest systems may undergo a rapid decline with limited counteracting effects of exogenous OM inputs. Conversely, OC depleted agricultural soils may benefit from organic amendments, thus acting as an effective sink for SOC accumulation. [ABSTRACT FROM AUTHOR]

Herbaspirillum seropedicae is an endophytic bacterium that can fix nitrogen and synthesize phytohormones, which can lead to a plant growth-promoting effect when used as a microbial inoculant. Studies focused on mechanisms of action are crucial for a better understanding of the bacteria-plant interaction and optimization of plant growth-promoting response. This work aims to understand the underlined mechanisms responsible for the early stimulatory growth effects of H. seropedicae inoculation in maize. To perform these studies, we combined transcriptomic and proteomic approaches with physiological analysis. The results obtained eight days after inoculation (d.a.i) showed increased root biomass (233 and 253%) and shoot biomass (249 and 264%), respectively, for the fresh and dry mass of maize-inoculated seedlings and increased green content and development. Omics data analysis, before a positive biostimulation phenotype (5 d.a.i.) revealed that inoculation increases N-uptake and N-assimilation machinery through differentially expressed nitrate transporters and amino acid pathways, as well carbon/nitrogen metabolism integration by the tricarboxylic acid cycle and the polyamine pathway. Additionally, phytohormone levels of root and shoot tissues increased in bacterium-inoculated-maize plants, leading to feedback regulation by the ubiquitin-proteasome system. The early biostimulatory effect of H. seropedicae partially results from hormonal modulation coupled with efficient nutrient uptake-assimilation and a boost in primary anabolic metabolism of carbon–nitrogen integrative pathways. [ABSTRACT FROM AUTHOR]

The use of humic substances in agriculture as a biostimulant emerged as one of the promising methods to promote sustainable production. Different molecular, biochemical, and physiological processes are triggered, resulting in nutrient efficiency use and protection against abiotic stress. Understanding plant changes promoted by humic substances is essential for innovative and tailored biostimulation technologies. Cell metabolites are the final target of the response chain, and the metabolomic approach can be helpful in unveiling pathways related to plant response. This study aimed to evaluate a global metabolic alteration of rice leaves induced by humic acids (HA) applied in a hydroponics system. Using 1H NMR and GC-TOF/MS analysis, we observed a significant decrease in all main metabolites classes in leaves treated with HA, including lipids, organic acids, amino acids, and carbohydrates. Metabolites in higher concentrations in HA-treated plants are candidates as markers of HA bioactivity, including amino acids, intermediates of tricarboxylic acid cycle, and lipids, and aromatic compounds related to plant-stress response. [ABSTRACT FROM AUTHOR]

Background: Innovative technologies are required in agricultural production to eliminate the environmental risk generated by the intensive use of fertilizers and pesticides. Soil organic matter is a keystone for the transition towards sustainable production. However, it is not an easy task to increase soil organic matter in highly weathered soils without considerable resources and energy. Here, we highlighted the role of biological inputs in plant adaptation to low fertility and water scarcity. The direct use of humic acids and plant growth-promoting bacteria on plants can modify the root architecture systems, including surface area and roots length, thus allowing greater soil exploration. Material and methods: Within a socio-historical perspective of concepts and research methods, a case study is presented on the effects of humic acids applied together with plant-growth promoting bacteria, as an efficient tool for supporting the transition to more suitable production system. We implemented this natural ecological approach onto a polyculture system with different tropical fruits (banana, passion fruit, papaya and pineapple) and evaluated crop yields. Results: We observed increases of around 50 and 90% in banana and papaya yield, respectively, and 25% in passion fruit productions, with significantly greater yields maintained over four production cycles. No effect was observed in ananas production probably due to the large shading level in the area. Conclusion: The biostimulant formulated with endophytic diazotrophic bacteria and humic acids represents a low-cost technology that enhances crop yields and can play an important role in promoting a transition process towards sustainable agriculture. [ABSTRACT FROM AUTHOR]

Background: The reliance on chemical inputs to support high yields is the Achilles' heel of modern crop production. The soil organic matter management is as old as agriculture itself. Recently, the use of soluble humic substances as plant growth promoters has been brought to attention due to their effects on nutrient uptake and water use efficiency. Humic substances applied directly at low concentrations can trigger different molecular, biochemical, and physiological processes in plants. However, how humic substances exert this plethoric regulatory action remains unclear. The objective of this study was to evaluate changes in the transcription level of genes coding cell receptors, phosphatases, synthesis, and function of different plant hormones and transcription factors. Materials and methods: After seven days of humic acid treatment, we used RNAseq in maize root seedlings. The level of gene transcription was compared with control plants. Results: Plant kinase receptors and different phosphatases were regulated by humic acids. Likewise, genes related to plant hormones (auxin, gibberellin, ethylene, cytokinin, abscisic acid, brassinosteroids, jasmonic and salicylic acids) were transcript in differential levels in maize root seedlings as well as the expression of a hundred of transcription factors modifying the signal transduction pathway via alterations of the subsequent gene response. Conclusion: We showed a general mechanism for simultaneously regulating the activity of several hormones where humic acids act as a key regulatory hub in plant responses integrating hormonal signalling and response pathways. [ABSTRACT FROM AUTHOR]

A plethora of bacteria–fungal interactions occur on the extended fungal hyphae network in soil. The mycosphere of saprophytic fungi can serve as a bacterial niche boosting their survival, dispersion, and activity. Such ecological concepts can be converted to bioproducts for sustainable agriculture. Accordingly, we tested the hypothesis that the well-characterised beneficial bacterium Serratia marcescens UENF-22GI can enhance plant growth-promoting properties when combined with Trichoderma longibrachiatum UENF-F476. The cultural and cell interactions demonstrated S. marcescens and T. longibrachiatum mutual compatibility. Bacteria cells were able to attach, forming aggregates to biofilms and migrating through the fungal hyphae network. Long-distance bacterial migration through growing hyphae was confirmed using a two-compartment Petri dishes assay. Fungal inoculation increased the bacteria survival rates into the vermicompost substrate over the experimental time. Also, in vitro indolic compound, phosphorus, and zinc solubilisation bacteria activities increased in the presence of the fungus. In line with the ecophysiological bacteria fitness, the bacterium–fungal combination boosted tomato and papaya plantlet growth when applied into the plant substrate under nursery conditions. Mutualistic interaction between mycosphere-colonizing bacterium S. marcescens UENF-22GI and the saprotrophic fungi T. longibrachiatum UENF-F467 increased the ecological fitness of the bacteria alongside with beneficial potential for plant growth. A proper combination and delivery of mutual compatible beneficial bacteria–fungal represent an open avenue for microbial-based products for the biological enrichment of plant substrates in agricultural systems. [ABSTRACT FROM AUTHOR]

Background: The bacterial-spot disease caused by different Xanthomonas species is one of the major tomato diseases that reduce crop production and quality. Pesticides indiscriminate usage has resulted in an increase in resistant bacterial strains as well as contamination of farmers, consumers and the environment. Plant growth-promoting bacteria and humic acids can act as elicitors of plant defence mechanism causing extensive transcriptional and metabolic reprogramming which, in turn, produce a range of plant chemical defences. The purpose of this study was to study how humic acids and plant growth-promoting bacteria, when applied to the substrate, affected the severity of bacterial spot symptoms in tomato leaves. Materials and methods: One-month-old Micro-Tom tomato (Solanum lycopersicum L.) were transferred to 3 L pots filled with a sterile mixture of sand and vermiculite (2:1, v:v) and treated or not (control) with 250 mL of 4.5 mmol C. L−1 of humic acids, Herbaspirillum seropedicae (108 CFU. mL−1) and the combination of humic acids plus H. seropedicae. One day after substrate treatment, the leaves were inoculated (or not) with X. euvesicatoria (Xe). The area below the disease progression curve based on severity scores and the number of symptomatic leaflets was used to assess phytopathogen virulence. The concentration of oxalic, citric and succinic acids in leaf extracts were determined using HPLC analysis. Results: Sole or combined H. seropedicae (BAC) and humic acids (HA) application promoted shoot and root growth related to control when plants were challenged with Xe pathogen. For plants inoculated with Xe, more significant plant-growth promotion results were obtained for HA + BAC treatment. The first visible symptoms were observed 16 days after inoculation with 2 × 104 CFU. g−1 of Xe cells in leaves of control plants. HA and BAC applied alone or combined reduced disease severity. Only plants treated with HA were able to reduce disease incidence (number of the leaflets with symptoms). Organic acids, such as oxalic, citric and succinic acids, rose in Xe-inoculated leaves. The reduced amount of organic acids in diseased leaves treated with HA + BAC may be linked to a decrease in disease progression. Conclusion: Humic acids and H. seropedicae increased growth by modulating the content of organic acids in leaf tissue, attenuating the symptoms of the bacterial spot disease. [ABSTRACT FROM AUTHOR]

Background: Salinity is one of the major environmental threats to agriculture, limiting plant growth and reducing crop yield. The use of humic substances to alleviate salt stress in plants is well reported, but the mechanisms remain unclear. This work aimed to apply humic acids on seedlings to acclimate plants to tolerate further salt stress exposition as a pre-treatment. Materials and methods: Two independent experiments with mono (maize) and dicot (tomato) seedlings were carried out. Maize was primed by humic acids (4 mM C) and further submitted to moderate salinity exposition (60 mM NaCl). The acclimation period of maize seedlings was characterized by ion balance and transcriptomic analysis of salt response genes. The tomato seedlings were also primed by humic acids (4 mM C) and exposed further to salinity (200 mM NaCl), and we measured only physiological aspect, including the activity of plasma membrane proton pumps and net photosynthesis rate. Results: Seedlings primed by humic acids minimized the salinity stress by changing ion balance, promoting plasma membrane proton pumps activity and enhancing photosynthesis rate and plant growth. We showed for the first time that maize seedlings treated with humic acids had a high transcription level of salt responsive genes and transcription factors even before the salt exposition. Conclusion: Humic acids previously activate cellular and molecular salt defence machinery, anticipating the response and reducing salinity stress. This is a key knowledge to manipulate manufactured biostimulants based on humic substances towards a maximized crop protection. [ABSTRACT FROM AUTHOR]

Background: Salinity is one of the major environmental threats to agriculture, limiting plant growth and reducing crop yield. The use of humic substances to alleviate salt stress in plants is well reported, but the mechanisms remain unclear. This work aimed to apply humic acids on seedlings to acclimate plants to tolerate further salt stress exposition as a pre-treatment. Materials and methods: Two independent experiments with mono (maize) and dicot (tomato) seedlings were carried out. Maize was primed by humic acids (4 mM C) and further submitted to moderate salinity exposition (60 mM NaCl). The acclimation period of maize seedlings was characterized by ion balance and transcriptomic analysis of salt response genes. The tomato seedlings were also primed by humic acids (4 mM C) and exposed further to salinity (200 mM NaCl), and we measured only physiological aspect, including the activity of plasma membrane proton pumps and net photosynthesis rate. Results: Seedlings primed by humic acids minimized the salinity stress by changing ion balance, promoting plasma membrane proton pumps activity and enhancing photosynthesis rate and plant growth. We showed for the first time that maize seedlings treated with humic acids had a high transcription level of salt responsive genes and transcription factors even before the salt exposition. Conclusion: Humic acids previously activate cellular and molecular salt defence machinery, anticipating the response and reducing salinity stress. This is a key knowledge to manipulate manufactured biostimulants based on humic substances towards a maximized crop protection. [ABSTRACT FROM AUTHOR]

The demand for biostimulants has been growing at an annual rate of 10 and 12.4% in Europe and Northern America, respectively. The beneficial effects of humic substances (HS) as biostimulants of plant growth have been well-known since the 1980s, and they can be supportive to a circular economy if they are extracted from different renewable resources of organic matter including harvest residues, wastewater, sewage sludge, and manure. This paper presents an overview of the scientific outputs on application methods of HS in different conditions. Firstly, the functionality of HS in the primary and secondary metabolism under stressed and non-stressed cropping conditions is discussed along with crop protection against pathogens. Secondly, the advantages and limitations of five different types of HS application under open-fields and greenhouse conditions are described. Key factors, such as the chemical structure of HS, application method, optimal rate, and field circumstances, play a crucial role in enhancing plant growth by HS treatment as a biostimulant. If we can get a better grip on these factors, HS has the potential to become a part of circular agriculture. [ABSTRACT FROM AUTHOR]

Purpose: The use of humic substances (HS) in agriculture is beneficial and has positive environmental impacts. However, to optimize the use of HS possible links between their structural characteristics and bioactivity must be shown. The goal of this study is to evaluate the bioactivity of different humic fractions extracted from vermicompost (VC) in rice plants and to shed light to possible structure-function relationships.Materials and methods: Humic-like fractions were obtained from cattle manure vermicompost processed by African nightcrawlers (Eudrilus eugeniae spp.). Humic-like acid fraction using only water as extractor (HLAw), HLA fraction extracted following the International Humic Substances Society (IHSS) recommended method, and the solid residue (humified residual (HR)) after extraction of HLA were characterized using complementary chemical, physic, and spectroscopic technics (elemental composition, UV-Vis and Fourier transform infrared spectroscopy (FTIR) spectroscopies, 13C-CP MAS NMR, and MEV). Biological activity of the three HS was conducted in growth chambers and measured in roots using WinRhizo Arabidopsis software. Principal component analysis (PCA) was used to find a grouping pattern between the structural variables evaluated and the obtained root parameters.Results and discussion: Differences were found in elemental composition among HS with larger C/N ratio in HR than in HLA and HLAw. HLA and HLAw FTIR spectra showed carboxyl band at 1714.66 cm−1 better resolved than in HR. Bands at 1642 cm−1 (amide I) and 1510 cm−1 (lignin), were better resolved in HLA. 13C-NMR showed the following order of aromaticity: HLA > HLAw > HR. For HLAw bioactivity, the structures CAlkyl-H,R, CC=O, and CCOO-H,R correlated with the number and growth of smaller root. The aromatic CAr-H,R, CAr-O,N, and aliphatic CAlkyl-O,N, CAlkyl-O, and CAlkyl-di-O structures in HLA, correlated with larger roots growth. HR also stimulated root growth and development in rice plants.Conclusions: Aliphatic and oxygenated structures in HLAw showed a relation with induction of initial root emissions, whereas the presence of aromatic compounds in HLA was related with root growth stimulation activity. Higher concentration of HLAw was necessary to produce an equivalent stimulus compared with HLA; it could indicate that, although both fractions showed similar types of structures in their composition, differences in the predominant structures may be determining different effects on the root. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is a limiting factor for plant growth, especially in highly weathered tropical soils. Plants have several mechanisms to overcome low P availability in soil, such as humic substances, that reduce phosphate (Pi) adsorption on oxide surfaces and enhance soil P availability. However, the direct influence of humic substances on Pi transporters in root cells or the distribution of P species in leaves remains unclear. Tomato seedlings were grown in a sand-vermiculite mixture with low or high P concentrations (10 or 100 mg kg-1 KH2PO4, respectively) and humic acids (0 or 48 mg C L-1) isolated from vermicompost. Plant responses were evaluated in the fifth week by measuring root and shoot weights and P concentration, and differential expression in the roots of the high-affinity Pi transporter genes LePT1 and LePT2. In addition, the distribution of P species in the leaves was assessed using 31P-NMR. Humic acids increased the root biomass and changed the distribution of P species in the leaves. Inorganic phosphate was the major compound in plants supplied with a high P concentration, whereas in plants supplied with a low P concentration, Pi was only identified in plants not treated with humic acids. Glycerophosphodiester and phosphorylcholine accumulated in plants treated with humic acid, indicating a modified metabolic pathway for economical P consumption at low P concentrations. High transcript accumulation of LePT2 was observed in roots treated with humic acids at both P concentrations. Our results show that humic substances are strategically involved in plant adaptation to P availability. [ABSTRACT FROM AUTHOR]

BACKGROUND Recycling of phosphorus (P) from organic residues ( ORs) is important to develop environmentally friendly agriculture. The use of this P source depends on phosphatase enzymes, which can be affected by a chain of parameters during maturation of ORs. In this study the phosphatase activity levels throughout vermicomposting of filter cake ( FC) and cattle manure ( CM) were correlated with different physical and chemical parameters in an effort to increase the knowledge about recycling of P from ORs. RESULTS FC presented higher total nitrogen content ( TNC), total organic carbon ( TOC), humic acid ( HA) content, water-soluble P ( WSP), phosphatase activities and nanopore volume than CM during vermicomposting. Decreases in TOC of CM resulted from carbohydrate mineralization, which was not observed for FC. CM showed increased hydrophobic index during vermicomposting while FC showed a slight decrease. CONCLUSION Phosphatase activities correlated positively with TOC, pH and WSP and negatively with HA content for both vermicomposts. Nanopore volume was negatively correlated with phosphatase activities for FC but not for CM. No correlations between hydrophobicity and phosphatase activities were found for FC. Increased hydrophobicity throughout vermicomposting of CM could be partially associated with decreases in phosphatase levels. © 2015 Society of Chemical Industry [ABSTRACT FROM AUTHOR]

HS were applied to garlic and stimulated plant growth and nutrition. The authors investigated the chemical features of HA and found that HA alleviated the nutritional stress of plants by improving their nutrient use efficiency more than plants fertilized with high NPK. Keywords: humic substances (HS); biostimulants; microbiome; soil quality (SQ); plant nutrition and metabolism EN humic substances (HS) biostimulants microbiome soil quality (SQ) plant nutrition and metabolism 1 2 2 02/10/22 20220209 NES 220209 The intense crop fertilization along with to climate change have a negative synergistic impact on soil fertility by decreasing the soil organic matter content and biological activity. [Extracted from the article]

Increasing evidences have indicated that humic substances can induce plant growth and productivity by functioning as an environmental source of auxinic activity. Here we comparatively evaluate the effects of indole-3-acetic acid (IAA) and humic acids (HA) isolated from two different soils (Inseptsol and Ultisol) and two different organic residues (vermicompost and sewage sludge) on root development and on activities of plasmalemma and tonoplast H+ pumps from maize roots. The data show that HA isolated from these different sources as well as low IAA concentrations (10−10 and 10−15 M) improve root growth through a markedly proliferation of lateral roots along with a differential activation not only of the plasmalemma but also of vacuolar H+-ATPases and H+-pyrophosphatase. Further, the vacuolar H+-ATPase had a peak of stimulation in a range from 10−8 to 10−10 M IAA, whereas the H+-pyrophosphatase was sensitive to a much broader range of IAA concentrations from 10−3 to 10−15 M. It is proposed a complementary view of the acid growth mechanism in which a concerted activation of the plasmalemma and tonoplast H+ pumps plays a key role in the root cell expansion process driven by environment-derived molecules endowed with auxinic activity, such as that of humic substances. [ABSTRACT FROM AUTHOR]

Phosphorus (P) deficiency is a major constraint in highly weathered tropical soils. Although phosphorous rock reserves may last for several hundred years, there exists an urgent need to research efficient P management for sustainable agriculture. Plant hormones play an important role in regulating plant growth, development, and reproduction. Humic substances (HS) are not only considered an essential component of soil organic carbon (SOC), but also well known as a biostimulant which can perform phytohormone-like activities to induce nutrient uptake. This review paper presents an overview of the scientific outputs in the relationship between HS and plant hormones. Special attention will be paid to the interaction between HS and plant hormones for nutrient uptake under P-deficient conditions. [ABSTRACT FROM AUTHOR]

Background: The use of humic substances as plant biostimulants has been increasingly attracting farmers and stunning researchers. The ability of these substances to enhance root growth by changing root architecture is often linked to their hormonal activities, such as auxin effects and nitric oxide production. Humeomics accesses the molecular constituents of humic substances, revealing the importance of alkyl components because of their conformations and chemical activities. Here, we describe the alkamides present in humic acids and compare their bioactivities using plasma membrane H+-ATPase activity level as a biochemical marker. Methods: Humic acids isolated from vermicompost were analyzed using 13C and 15N nuclear magnetic resonance spectroscopy. The unbound fraction was extracted with ethyl acetate and submitted to gas chromatography coupled to mass spectrometry to detect the presence of N-isopropyldecanamide. We synthesized N-isopropyldecanamide and treated maize seedlings for 7 and 15 days with different concentrations. The root growth and plasma membrane H+-ATPase activity were monitored. Nitric oxide accumulation in the lateral roots was imaged using 4,5-diaminofluorescein diacetate. The results were compared with those obtained for seedlings treated with humic acids isolated from vermicompost. Results: The amide functional group produced the only nitrogen signal in the 15N humic acid resonance spectrum and similar alkamide moieties were found in the unbound humic extract through comparisons using gas chromatography coupled to mass spectrometry. The synthesis of N-isopropyldecanamide had few steps and produced a high yield (86%). The effects of N-isopropyldecanamide on root growth were concentration dependent. High concentrations (10−4 M) enhanced root growth after 15 day of diminishing shoot biomass. However, low concentrations (10−8 M and 10−6 M) promoted root growth at 7 and 15 days, similar to the humic acid-induced plasma membrane H+-ATPase activity. Both N-isopropyldecanamide and humic acids enhanced nitric oxide accumulation during lateral root emergence. Conclusion: We described for the first time the effects of N-isopropyldecanamide on the plasma membrane H+-ATPase activity in maize seedling roots and compared its effects with those caused by humic acids. N-Isopropyldecanamide was detected in the unbound fraction of the humic supramolecular assembly, indicating that the putative hormone-like effects of these substances result also from the presence of this new class of plant regulators, in addition to other molecules. [ABSTRACT FROM AUTHOR]

Background: The use of plant-based biostimulants to increase crop yield and enhance protection has grown in popularity. We previously manufactured an inoculant combining humic acids with plant growth-promoting bacteria and observed a significant effect on crop yields. However, electrophysiology studies elucidating the mechanisms responsible for enhanced nutrient and water uptake are scarce and are generally restricted to in vitro models. Materials and methods: In this study, a suspension of humic acids isolated from vermicompost along with Herbaspirillum seropedicae was applied to maize seedlings. We measured the root H+ flux with an ion-selective vibrating probe system. Furthermore, the transcription of plasma membrane H+ ATPase, aquaporin and high- or low-affinity nitrate transporters was measured. Results: Inoculation activated the extracellular H+ flux, thus changing the pH and membrane potential of maize root cells and altering the electrochemical potential generated by P–H+-ATPase at the biochemical and molecular level. The overexpression of aquaporins was also observed; however, nitrate transporters were repressed by the inoculants. Conclusion: We demonstrate an increase in the H+ efflux in maize root seedlings inoculated with H. seropedicae and concomitant changes in membrane voltage. The inoculation of H. seropedicae in the presence of humic acids decreased the H+ flux and surface acidification without changes in the aquaporin transcription level. However, the H+ flux in root seedlings in inoculated plants (with or without humic acids) was larger in respect to control plants. These results support the increased water and nitrogen efficiency in plants inoculated with humic acids and H. seropedicae. [ABSTRACT FROM AUTHOR]