Phosphorus Acquisition Efficiency and Transcriptomic Changes in Maize Plants Treated with Two Lignohumates.

Lignohumates are increasing in popularity in agriculture, but their chemistry and effects on plants vary based on the source and processing. The present study evaluated the ability of two humates (H1 and H2) to boost maize plant performance under different phosphorus (P) availability (25 and 250 µM) conditions in hydroponics, while understanding the underlying mechanisms. Humates differed in chemical composition, as revealed via elemental analysis, phenol and phytohormone content, and thermal and spectroscopic analyses. H1 outperformed H2 in triggering plant responses to low phosphorus by enhancing phosphatase and phytase enzymes, P acquisition efficiency, and biomass production. It contained higher levels of endogenous auxins, cytokinins, and abscisic acid, likely acting together to stimulate plant growth. H1 also improved the plant antioxidant capacity, thus potentially increasing plant resilience to external stresses. Both humates increased the nitrogen (N) content and acted as biostimulants for P and N acquisition. Consistent with the physiological and biochemical data, H1 upregulated genes involved in growth, hormone signaling and defense in all plants, and in P recycling particularly under low-P conditions. In conclusion, H1 showed promising potential for effective plant growth and nutrient utilization, especially in low-P plants, involving hormonal modulation, antioxidant enhancement, the stimulation of P uptake and P-recycling mechanisms.

Chemical Structure and Biological Activity of Humic Substances Define Their Role as Plant Growth Promoters.

Humic substances (HS) are dominant components of soil organic matter and are recognized as natural, effective growth promoters to be used in sustainable agriculture. In recent years, many efforts have been made to get insights on the relationship between HS chemical structure and their biological activity in plants using combinatory approaches. Relevant results highlight the existence of key functional groups in HS that might trigger positive local and systemic physiological responses via a complex network of hormone-like signaling pathways. The biological activity of HS finely relies on their dosage, origin, molecular size, degree of hydrophobicity and aromaticity, and spatial distribution of hydrophilic and hydrophobic domains. The molecular size of HS also impacts their mode of action in plants, as low molecular size HS can enter the root cells and directly elicit intracellular signals, while high molecular size HS bind to external cell receptors to induce molecular responses. Main targets of HS in plants are nutrient transporters, plasma membrane H+-ATPases, hormone routes, genes/enzymes involved in nitrogen assimilation, cell division, and development. This review aims to give a detailed survey of the mechanisms associated to the growth regulatory functions of HS in view of their use in sustainable technologies.

Preliminary Study on the Activity of Phycobiliproteins against Botrytis cinerea.

Phycobiliproteins (PBPs) are proteins of cyanobacteria and some algae such as rhodophytes. They have antimicrobial, antiviral, antitumor, antioxidative, and anti-inflammatory activity at the human level, but there is a lack of knowledge on their antifungal activity against plant pathogens. We studied the activity of PBPs extracted from Arthrospiraplatensis and Hydropuntiacornea against Botrytiscinerea, one of the most important worldwide plant-pathogenic fungi. PBPs were characterized by using FT-IR and FT-Raman in order to investigate their structures. Their spectra differed in the relative composition in the amide bands, which were particularly strong in A. platensis. PBP activity was tested on tomato fruits against gray mold disease, fungal growth, and spore germination at different concentrations (0.3, 0.6, 1.2, 2.4, and 4.8 mg/mL). Both PBPs reduced fruit gray mold disease. A linear dose-response relationship was observed for both PBPs against disease incidence and H. cornea against disease severity. Pathogen mycelial growth and spore germination were reduced significantly by both PBPs. In conclusion, PBPs have the potential for being also considered as natural compounds for the control of fungal plant pathogens in sustainable agriculture.

Bioactivity of Size-Fractionated and Unfractionated Humic Substances From Two Forest Soils and Comparative Effects on N and S Metabolism, Nutrition, and Root Anatomy of Allium sativum L.

Humic substances (HS) are powerful natural plant biostimulants. However, there is still a lack of knowledge about the relationship between their structure and bioactivity in plants. We extracted HS (THE1-2) from two forest soils covered with Pinus mugo (1) or Pinus sylvestris (2). The extracts were subjected to weak acid treatment to produce size-fractionated HS (high molecular size, HMS1-2; low molecular size, LMS1-2). HS were characterized for total acidity, functional groups, element and auxin (IAA) contents, and hormone-like activity. HS concentrations ranging from 0 to 5 mg C L-1 were applied to garlic (Allium sativum L.) plantlets in hydroponics to ascertain differences between unfractionated and size-fractionated HS in the capacity to promote mineral nutrition, root growth and cell differentiation, activity of enzymes related to plant development (invertase, peroxidase, and esterase), and N (nitrate reductase, glutamine synthetase) and S (O-acetylserine sulphydrylase) assimilation into amino acids. A positive linear dose-response relationship was determined for all HS in the range 0-1 mg C L-1, while higher HS doses were less effective or ineffective in promoting physiological-biochemical attributes of garlic. Bioactivity was higher for size-fractionated HS according to the trend LMS1-2>HMS1-2>THE1-2, with LMS2 and HMS2 being overall more bioactive than LMS1 and HMS1, respectively. LMS1-2 contained more N, oxygenated functional groups and IAA compared to THE1-2 and HMS1-2. Also, they exhibited higher hormone-like activities. Such chemical properties likely accounted for the greater biostimulant action of LMS1-2. Beside plant growth, nutrition and N metabolism, HS stimulated S assimilation by promoting the enrichment of garlic plantlets with the S amino acid alliin, which has recognized beneficial properties in human health. Concluding, this study endorses that i) treating THE with a weak acid produced sized-fractionated HS with higher bioactivity and differing in properties, perhaps because of novel molecular arrangements of HS components that better interacted with garlic roots; ii) LMS from forest soils covered with P. mugo or P. sylvestris were the most bioactive; iii) the cover vegetation affected HS bioactivity iv); HS stimulated N and S metabolism with relevant benefits to crop nutritional quality.

Degradative Ability of Mushrooms Cultivated on Corn Silage Digestate.

The current management practice of digestate from biogas plants involves its use for land application as a fertilizer. Nevertheless, the inadequate handling of digestate may cause environmental risks due to losses of ammonia, methane and nitrous oxide. Therefore, the key goals of digestate management are to maximize its value by developing new digestate products, reducing its dependency on soil application and the consequent air pollution. The high nitrogen and lignin content in solid digestate make it a suitable substrate for edible and medicinal mushroom cultivation. To this aim, the mycelial growth rate and degradation capacity of the lignocellulosic component from corn silage digestate, undigested wheat straw and their mixture were investigated on Cyclocybe aegerita, Coprinus comatus, Morchella importuna, Pleurotus cornucopiae and Pleurotus ostreatus. The structural modification of the substrates was performed by using attenuated total reflectance-Fourier transform infrared (ATR-FTIR) spectroscopy. Preliminary in vitro results demonstrated the ability of P. ostreatus, P. cornucopiae and M. importuna to grow and decay hemicellulose and lignin of digestate. Cultivation trials were carried out on C. aegerita, P. cornucopiae and P. ostreatus. Pleurotus ostreatus showed the highest biological efficiency and fruiting body production in the presence of the digestate; moreover, P. ostreatus and P. cornucopiae were able to degrade the lignin. These results provide attractive perspectives both for more sustainable digestate management and for the improvement of mushroom cultivation efficiency.

Automated image analysis and hyperspectral imagery with enhanced dark field microscopy applied to biochars produced at different temperatures.

Biochar from agricultural biomasses and solid wastes represents a win-win solution for a rational waste management. Its sustainable usage requires the identification and standardization of biochar characteristics. The aim of this work was to identify the physical-chemical and spatial characteristics of biochars from pruning residues (PR), poultry litter (PL), and anaerobic cattle digestate (CD) at two pyrolysis temperatures (350 °C and 550 °C). The biochar characterization was carried out by applying emerging imaging techniques, 2D automated optical image analysis and hyperspectral enhanced dark-field microscopy (EDFM), and by SEM analysis. As predictable, the feedstock composition and the pyrolysis temperature strongly influence the physical structures of the biochar samples. Irrespective of charring temperature, PR biochar was mainly characterized by a broken and fragmented structure with an irregular and rough particle surface, completely different from the original PR wood cell. The EDFM imaging analysis evidenced the thermal degradation of PR vegetal products, composed primarily of hemicellulose, cellulose and lignin. On the contrary, small and regular particles with a smooth surface were produced by the PL pyrolysis, especially at 550 °C, due to the lower PL morphological homogeneity in comparison with the other biomasses. Finally, CD charring at both temperatures was characterized by changes in chemical composition, suggested by a lower pixel intensity. In conclusion, the emerging imaging techniques used in this study proved to be very effective in analyzing some properties of biochars, and can, therefore be considered as promising experimental strategies for detecting the feedstock and pyrolysis temperature of biochar.

Effects of Two Protein Hydrolysates Obtained From Chickpea (Cicer arietinum L.) and Spirulina platensis on Zea mays (L.) Plants.

Two liquid protein hydrolysates obtained from chickpea (Cicer arietinum L.) (CA) and Spirulina platensis (SP) were analyzed via FT-IR and SERS spectroscopy. Their hormone-like activities and contents in indole-3-acetic acid (IAA), isopentenyladenosine (IPA), nitrogen (N), carbon (C), sulfur (S), phenols, amino acids, and reducing sugars were determined. CA and SP showed different chemical compositions in N, C, sugars, amino acid, and TP contents, which were generally higher in CA. The two products exhibited (IAA)-like and gibberellin (GA)-like activities and contained the hormones IAA and IPA. Specifically, CA held higher (∼3.6 fold) IAA-like activity than SP, while its GA-like activity was comparable to SP. The content in IAA was similar between hydrolysates, but CA contained ∼6 fold more IPA. CA and SP were further supplied at two different dosages (0.1 and 1 mL L-1) for 2 days to maize (Zea mays L.) plants grown in hydroponics. They positively influenced plant growth and accumulation of N-compounds (proteins, chlorophylls and phenols), with a more pronounced effect observed in plants treated with CA. Furthermore, they increased the activity of two enzymes, i.e., peroxidase and esterase, which are established markers for plant growth, differentiation and organogenesis-related processes. Peroxidase activity in particular, was enhanced by ∼1.6 and ∼2.3 fold in leaves and roots of CA-treated plants, respectively. Greater accumulation of macro (Ca, Mg, and K) and micro (Cu, Zn) elements was also evident in plants supplied with these products. In conclusion, our data indicate that both CA and SP exert positive effects in maize plants. However, CA appeared to be more efficient than SP to improve plant nutrition and growth parameters in some respects, likely by virtue of its higher content in phytochemicals (hormones, phenols, amino acids, reducing sugars) that may act as signaling molecules, and more pronounced IAA-like activity.

Stimulated Adsorption of Humic Acids on Capped Plasmonic Ag Nanoparticles Investigated by Surface-Enhanced Optical Techniques.

The adsorption of humic substances on Ag nanoparticles (AgNPs) is of crucial environmental importance and determines the toxicity of these NPs and the structure of adsorbed organic matter. In this work, the adsorption of two standard soil and leonardite International Humic Substances Society humic acids was studied on AgNPs of different sizes, shapes (spherical and star-like), and interfacial chemical compositions. Surface-enhanced optical (Raman and fluorescence) spectroscopies were used to follow the specific chemical groups involved in this adsorption. By means of the latter optical techniques, information regarding the binding mechanism and the macromolecular aggregation can be deduced. The influence of the surface chemical composition induced by the different functionalizations of the interfaces of these NPs is highly important regarding the chemical interactions of these complex organic macromolecules. The surface functionalization with positively charged alkyl diamines led to a large increase in the adsorption as well as a strong structural rearrangement of the macromolecule once adsorbed onto the surface.

Spectroscopic-Chemical Fingerprint and Biostimulant Activity of a Protein-Based Product in Solid Form.

A solid biostimulant (AA309) obtained through thermobaric hydrolysis applied on trimmings and shavings of bovine hides tanned with wet-blue technology was chemically characterized, and its effects in maize (Zea mays L.) were evaluated. AA309 contained 13.60% total nitrogen (N), mainly in organic forms (13.40%), and several amino acids, especially lysine, phenylalanine, glycine, aspartate, and isoleucine. AA309 was further analyzed using Fourier Transform Infrared (FT-IR) spectroscopy, which revealed the presence of amide I and amide II bands, indicative of peptide structures. When supplied to maize plants for 15 days at two N dosages (2.1 or 4.2 mg/kg), AA309 induced positive physiological responses, likely because of its content in amino acids functioning as signaling molecules. The low dosage was the most effective in improving leaf (+24%) and root (+98%) dry weight, photosynthetic activity (+70%), and accumulation of N (+80%), proteins (+65⁻75%) and antioxidants (+60%). Spectroscopic analyses (Solid-state Cross-Polarization Magic Angle Spinning Carbon-13 Nuclear Magnetic Resonance, CP/MAS 13C⁻NMR, and High resolution-magic angle spinning nuclear magnetic resonance, HR-MAS NMR) on plant tissues revealed the increase in proteins, lignin structures and cutin in AA309-treated plants compared to untreated plants. Our results indicate that AA309 could be used as a valuable biostimulant in agriculture.

Evaluation of Seaweed Extracts From Laminaria and Ascophyllum nodosum spp. as Biostimulants in Zea mays L. Using a Combination of Chemical, Biochemical and Morphological Approaches.

Seaweed extracts can be employed as biostimulants during crop cultivation owing to their positive effects on plant performance. Therefore, in this study one extract from Laminaria (A) and five extracts from Ascophyllum nodosum (B-F) were assayed on maize (Zea mays L.) plants supplied for 2 days with 0.5 mL L-1 of single products to evaluate their capacity to stimulate root growth and morphology, nutrition, and sugars accumulation. Firstly, extracts were chemically characterized via Fourier transform infrared (FT-IR) and FT-Raman spectroscopies, and their content in carbon, nitrogen, phenolic acids and hormones (indole-3-acetic acid, IAA, and Isopentenyladenosine, IPA) was quantified. The auxin like- and gibberellic acid -like activities of all extracts were also determined. FT-IR and FT-Raman spectra provided complementary information depicting distinct spectral pattern for each extract. Bands assigned to alginic and uronic acids were dominant in FT-IR spectra, while those corresponding to polyaromatic rings were evident in FT-Raman spectra. In general, extracts stimulated root growth, nutrition, esterase activity, and sugar content. However, they showed high variation in chemical features, which may explain their different capacity in triggering physiological responses in maize. Among A. nodosum extracts for instance, E was the most efficient in promoting root morphology traits, likely because of its elevate content in IAA (32.43 nM), while F extract was the highest in phenol content (1,933 mg L-1) and the most successful in improving plant nutrition. On the other hand, C extract was very effective in stimulating root elongation, but did not influence plant nutrition. B and D extracts induced similar positive effects on plants, although they greatly varied in chemical composition. Laminaria extract (A) differed from A. nodosum extracts, because of its low content in total phenols and the presence of both IAA- and GA-like activity. We conclude that all seaweed extracts acted as biostimulants in maize, but their chemical properties appeared crucial in predicting the physiological response preferentially elicited by individual seaweed extracts.

Size fractionation as a tool for separating charcoal of different fuel source and recalcitrance in the wildfire ash layer.

Charcoal is a heterogeneous material exhibiting a diverse range of properties. This variability represents a serious challenge in studies that use the properties of natural charcoal for reconstructing wildfires history in terrestrial ecosystems. In this study, we tested the hypothesis that particle size is a sufficiently robust indicator for separating forest wildfire combustion products into fractions with distinct properties. For this purpose, we examined two different forest environments affected by contrasting wildfires in terms of severity: an eucalypt forest in Australia, which experienced an extremely severe wildfire, and a Mediterranean pine forest in Italy, which burned to moderate severity. We fractionated the ash/charcoal layers collected on the ground into four size fractions (>2, 2-1, 1-0.5, <0.5mm) and analysed them for mineral ash content, elemental composition, chemical structure (by IR spectroscopy), fuel source and charcoal reflectance (by reflected-light microscopy), and chemical/thermal recalcitrance (by chemical and thermal oxidation). At both sites, the finest fraction (<0.5mm) had, by far, the greatest mass. The C concentration and C/N ratio decreased with decreasing size fraction, while pH and the mineral ash content followed the opposite trend. The coarser fractions showed higher contribution of amorphous carbon and stronger recalcitrance. We also observed that certain fuel types were preferentially represented by particular size fractions. We conclude that the differences between ash/charcoal size fractions were most likely primarily imposed by fuel source and secondarily by burning conditions. Size fractionation can therefore serve as a valuable tool to characterise the forest wildfire combustion products, as each fraction displays a narrower range of properties than the whole sample. We propose the mineral ash content of the fractions as criterion for selecting the appropriate number of fractions to analyse.

Biological Activity of Vegetal Extracts Containing Phenols on Plant Metabolism.

The influence of vegetal extracts derived from red grape, blueberry fruits and hawthorn leaves on Zea mays L. plant growth and the activity of phenylalanine ammonia-lyase (PAL), a key enzyme of the phenylpropanoid pathway, was investigated in laboratory experiments. The extracts were characterized using FT-IR and Raman spectroscopies in order to obtain a pattern of the main functional groups. In addition, phenols content was determined by HPLC, whereas the content of indoleacetic acid and isopentenyladenosine hormones was determined by ELISA test and the auxin and gibberellin-like activities by plant-bioassays. The treated maize revealed increased root and leaf biomass, chlorophyll and sugars content with respect to untreated plants. Hawthorn, red grape skin and blueberry at 1.0 mL/L induced high p-coumaric content values, whilst hawthorn also showed high amounts of gallic and p-hydroxybenzoic acids. PAL activity induced by hawthorn at 1.0 mL/L had the highest values (11.1-fold UNT) and was strongly and linearly related with the sum of leaf phenols. Our results suggest that these vegetal extracts contain more than one group of plant-promoting substances.

Snow vole (Chionomys nivalis Martins) affects the redistribution of soil organic matter and hormone-like activity in the alpine ecosystem: ecological implications.

In alpine environments, colonies of snow vole (Chionomys nivalis Martins) cause strong pedoturbation, which may affect humification process and soil organic matter (SOM) cycling, with repercussions on the hormone-like activity of organics. We investigated the effect of snow vole pedoturbation on the chemical and spectroscopic features of soil organic fractions, and the potential hormone-like activity of humic and fulvic acids (HA, FA). The study site was located on the high-mountain environment of the Majella massif (central Italy). Pedoturbated and regular soils were morphologically described and characterized for pH and content of total organic carbon, total extractable carbon, HA, and FA. Both HA and FA were extracted and investigated using attenuated total reflectance/Fourier transform infrared (ATR/FTIR), nuclear magnetic resonance with high-resolution magic angle spinning (HRMAS-NMR), and (1)H-(13)C heteronuclear single quantum coherence (HSQC). HA and FA were also tested for their auxin-like and gibberellin-like activities. Results provide evidences that bioturbated and regular soils contain a poorly decomposed SOM, but HA and FA with a well-defined molecular structure. The HA and FA from both bioturbated and regular soils show a hormone-like activity with a different allocation along the soil profile. In the regular soil, the highest auxin-like activity was shown by HA and FA from Oe1 horizon, while gibberellin-like activity was expressed by FA from Oe2 horizon. Burrowing activity determines a redistribution of organics throughout the profile with a relatively high auxin-like activity in the FA from straw tunnel wall (STW) and gibberellin-like activity in the HA from vole feces (VF). The relative high presence of carboxylic acids, amides, proteins, and amino acids in the FA from STW and the aromatic moieties in the HA from VF put evidences for their different behavior. The fact that snow vole activity has modified the chemical and biological properties of SOM in these soils otherwise considered governed only by low temperature has important ecological implications such as the preservation of soil fertility and vegetal biodiversity.

Spontaneous aggregation of humic acid observed with AFM at different pH.

Atomic force microscopy in contact (AFM-C) mode was used to investigate the molecular dynamics of leonardite humic acid (HA) aggregate formed at different pH values. HA nanoparticles dispersed at pH values ranging from 2 to 12 were observed on a mica surface under dry conditions. The most clearly resolved and well-resulted AFM images of single particle were obtained at pH 5, where HA appeared as supramolecular particles with a conic shape and a hole in the centre. Those observations suggested that HA formed under these conditions exhibited a pseudo-amphiphilic nature, with secluded hydrophobic domains and polar subunits in direct contact with hydrophilic mica surface. Based on molecular simulation methods, a lignin-carbohydrate complex (LCC) model was proposed to explain the HA ring-like morphology. The LCC model optimized the parameters of ß-O-4 linkages between 14 units of 1-4 phenyl propanoid, and resulted in an optimized structure comprising 45-50 linear helical molecules looped spirally around a central cavity. Those results added new insights on the adsorption mechanism of HA on polar surfaces as a function of pH, which was relevant from the point of view of natural aggregation in soil environment.

Capsicum chinensis L. growth and nutraceutical properties are enhanced by biostimulants in a long-term period: chemical and metabolomic approaches.

Two biostimulants, one derived from alfalfa plants (AH) and the other obtained from red grape (RG), were chemically characterized using enzyme linked immuno-sorbent assays, Fourier transform infrared (FT-IR) and Raman spectroscopies. Two doses (50 and 100 mL L(-1) for RG, and 25 and 50 mL L(-1) for AH) of biostimulants were applied to Capsicum chinensis L. plants cultivated in pots inside a tunnel. The experimental design consisted of the factorial combination of treatment (no biostimulant, plus AH, plus RG) at three doses (zero, low, and high) and two time-course applications (at the second and fourth week after transplantation) and the effects were recorded at flowering and maturity. Both biostimulants contained different amounts of indoleacetic acid and isopentenyladenosine; the AH spectra exhibited amino acid functional groups in the peptidic structure, while the RG spectra showed the presence of polyphenols, such as resveratrol. These results revealed that at flowering, RG and AH increased the weights of fresh leaves and fruits and the number of green fruits, whereas at maturity, the biostimulants most affected the fresh weight and number of red fruits. At flowering, the leaves of the treated plants contained high amounts of epicatechin, ascorbic acid, quercetin, and dihydrocapsaicin. At maturity, the leaves of the treated plants exhibited elevated amounts of fructose, glucose, chlorogenic, and ferulic acids. Moreover, green fruits exhibited a high content of chlorogenic acid, p-hydroxybenzoic acid, p-coumaric acid and antioxidant activity, while both AH- and RG-treated red fruits were highly endowed in capsaicin. The (1)H high-resolution magic-angle spinning (HRMAS)-nuclear magnetic resonance (NMR) spectra of red fruits revealed that both products induced a high amount of NADP(+), whereas RG also increased glucose, fumarate, ascorbate, thymidine and high molecular weight species. Our results suggested that AH and RG promoted plant growth and the production of secondary metabolites, such as phenols.

Effect of a peat humic acid on morphogenesis in leaf explants of Pyrus communis and Cydonia oblonga . Metabolomic analysis at an early stage of regeneration.

Plant regeneration is a critical step in most in vitro breeding techniques. This paper studies the effects of a low-molecular-weight humic acid (HA) on morphogenesis from pear and quince leaf explants. Variable HA amounts [0 (control), 1, 5, 10, and 20 mg C L(-1)] were added to the regeneration media. A dose-response effect was observed in pear for root and shoot production; it was improved at HA 1 mg C L(-1) and considerably reduced at the highest amounts. HA was, instead, ineffective in quince. The (1)H HR-MAS NMR analyses of calli in the induction phase showed more evident metabolite (asparagine, alanine, and γ-aminobutyric acid) signals in quince than in pear. The assignment of overlapped signals in both genotypes was supported by the 2D NMR analyses. Spectroscopic characterization suggested also an enhancement of asparagine contents in morphogenic calli of pear with respect to the control and higher HA amount treatments.

Effect of commercial lignosulfonate-humate on Zea mays L. metabolism.

Lignosulfonate-humate a and lignosulfonate-humate b, derived by an industrial process from lignin, were studied chemically and biologically, and their effects on maize metabolism compared with the responses induced by humic substances obtained from leonardite. Lignosulfonate-humate a and lignosulfonate-humate b elicited hormonelike activity and leonardite displayed giberellin properties. To improve our understanding of their biological action, lignosulfonate-humate a, lignosulfonate-humate b and leonardite were supplied to maize plants and their effect was studied on growth, nitrogen metabolism and photosynthesis. All products increased root and leaf growth. Glutamine-synthetase, glutamate-synthase enzyme activities and protein content were all increased. The treatments also increased chlorophyll content, glucose, fructose and rubisco enzyme activity, suggesting a positive role of lignosulfonate-humate a, lignosulfonate-humate b and leonardite in the photosynthetic process. In addition, an increase in phenol content was observed. In light of these results, being environmentally friendly products, lignosulfonate-humate a and lignosulfonate-humate b could be used to increase crop yield.

High molecular size humic substances enhance phenylpropanoid metabolism in maize (Zea mays L.).

A high molecular weight humic fraction (>3,500 Da) was characterized chemically by DRIFT and 1H NMR spectroscopy, and was applied to Zea mays L. plants to evaluate its effect on phenylpropanoid metabolism. The activity and gene expression of phenylalanine (tyrosine) ammonia-lyase (PAL/TAL), and the concentrations of phenolics and their amino acid precursors phenylalanine and tyrosine were assayed. Maximum induction of PAL/TAL activity and expression was obtained when the concentration of added humic substance was 1 mg C/l hydroponic solution. Phenylalanine and tyrosine significantly decreased (-16% and -22%, respectively), and phenolic compounds increased in treated plants. The effects of the humic substance could be ascribed partly to indoleacetic acid (27 nmol/mg C) in the humic fraction. Our results suggest that this humic fraction induces changes in phenylpropanoid metabolism. This is the first study that shows a relationship between humic substances and the phenylpropanoid pathway.

Humic substances biological activity at the plant-soil interface: from environmental aspects to molecular factors.

Humic substances (HS) represent the organic material mainly widespread in nature. HS have positive effects on plant physiology by improving soil structure and fertility and by influencing nutrient uptake and root architecture. The biochemical and molecular mechanisms underlying these events are only partially known. HS have been shown to contain auxin and an "auxin-like" activity of humic substances has been proposed, but support to this hypothesis is fragmentary. In this review article, we are giving an overview of available data concerning molecular structures and biological activities of humic substances, with special emphasis on their hormone-like activities.