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.

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It is widely reported that some humic substances behave as exogenous auxins influencing root growth by mechanisms that are not yet completely understood. This study explores the hypothesis that the humic acids' effects on root development involve a nitric oxide signaling. Maize seedlings were treated with HA 20 mg C L(-1), IAA 0.1 nM, and NO donors (SNP or GSNO), in combination with either the auxin-signaling inhibitor PCIB, the auxin efflux inhibitor TIBA, or the NO scavenger PTIO. H(+)-transport-competent plasma membrane vesicles were isolated from roots to investigate a possible link between NO-induced H(+)-pump and HA bioactivity. Plants treated with either HA or SNP stimulated similarly the lateral roots emergence even in the presence of the auxin inhibitors, whereas NO scavenger diminished this effect. These treatments induced H(+)-ATPase stimulation by threefold, which was abolished by PTIO and decreased by auxin inhibitors. HA-induced NO synthesis was also detected in the sites of lateral roots emergence. These data depict a new scenario where the root development stimulation and the H(+)-ATPase activation elicited by either HA or exogenous IAA depend essentially on mechanisms that use NO as a messenger induced site-specifically in the early stages of lateral root development.

The apparent high molecular mass of humic acids (HAs) hardly seems compatible with their direct effects in plant physiology. However, previous evidence has indicated that HAs are non-covalent associations of relatively small molecules, which can be broken down by the action of organic acids. The aim of this work was to evaluate the effects of organic acids on the structure of HAs by spectroscopy and on their bioactivity by following the responses of maize root growth. Changes in the exudation of organic acids from maize seedlings treated with HAs at 50 mg C L 21 were evaluated by high-performance liquid chromatography. The results are in agreement with the concept that HAs are chemical aggregates that acquire characteristics typical of low-molecular-mass humic substances when exposed to organic acids exuded by the roots. Maize seedlings grown in solutions supplemented with HAs plus citric acid at 0.0005, 0.005 and 0.05 mM exhibited significant changes in their root area, primary root length, number of lateral roots and lateral root density and increases in plasma membrane H + -ATPase activity. Furthermore, the root exudation profile of plants treated with HAs exhibited an increase in the efflux of oxalic and citric acids, with a concurrent decrease in malic and succinic acids. These data reveal a crosstalk between HAs and plants where the exudation of organic acids from the roots influences and is influenced by bioactive molecules released from HAs during root development.

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ABSTRACT:Since the beginning of Human civilization, the soil organic matter has been used as plant growth promoter and/or regulator.Indeed, early in plant science history, even before the auxin concept has been established, the term "auximones" was coined to describe plant growth promoting humic acids derived from peat. Despite of this, until the end of the 20 th century, humic substances remained as some of the most neglected environment signals in plant physiology research. However, this scenario has changed in last decade with the discovery that the major systems of energy transduction of the plant cell membranes, the proton pumps, can be tightly orchestrated by humic substances just as elicited by a hormonal signaling. Differential activations of both plasma membrane (PM H + -ATPase) and vacuolar pumps (V-ATPase and H + -PPase) are modulated by humic substances triggering ion signatures related to specific patterns of plant growth and development. Phytohormones have been found to be associated with this humus bioactivity, and nitric oxide acting as a second messenger in a signaling pathway in which plants can sense the soil environment to cope with specific conditions. In this review, we discuss some of the most influential data available in literature, which have shaped this underexplored interface between the chemistry of the organic matter and the plant physiology. The key role of organic matter in the sustainable agriculture will also be highlighted from a biochemical perspective of the plant cell responses to biofertilization, specially in tropical environments.

P or que será que em pleno século 21 ainda utilizamos esterco animal na agricultura, e não apenas os fertilizantes minerais? Será somente pelo custo relativamente menor? Ou será resultado dos efeitos benéficos sobre as características físicas, químicas e biológicas do solo? Pode ser por ambos os motivos, e acrescidos a esses, ainda existem os efeitos benéficos encontrados na fisiologia das plantas, que também ajudam a explicar a permanência dos adubos orgânicos de origem animal ou vegetal. Vários efeitos fisiológicos são relatados há séculos, muito embora não sejam completamente elucidados.A matéria orgânica do solo é constituída por compostos de carbono em diferentes graus de associação com as fases minerais do solo originados a partir da decomposição de resíduos vegetais e animais. Além de ser fonte de nutrientes, a matéria orgânica apresenta cargas de superfície que contribuem para o aumento da capacidade de troca de cátions (CTC) do solo e, devido a sua alta reatividade, regula a disponibilidade de vários nutrientes, em especial os micronutrientes, bem como a atividade de elementos potencialmente fitotóxicos como Al 3+ e Mn 2+ , em solos ácidos, e metais pesados. Nos ambientes tropicais, a matéria orgânica do solo tem importância elevada. É amplamente reconhecida por seus efeitos benéficos à física e química dos solos devido a melhor agregação e retenção de água, maior CTC e disponibilidade de nutrientes. Além destes existem os aspectos biológicos que estão relacionados com microrganismos benéficos encontrados na matéria orgânica. No cultivo de hortaliças em geral e, mais especificamente para produção de tomate e demais hortaliças mais exigentes em nutrientes, a matéria orgânica e suas frações possuem papel fundamental.Existem visões diferentes acerca das características moleculares da matéria orgânica. O debate existe devido às características dos métodos utilizados para extrair a matéria orgânica do solo. A utilização de extratores alcalinos fortes é utilizada pelos grupos tradicionais que ZANDONADI DB; SANTOS MP; MEDICI LO; SILVA J. 2014. Ação da matéria orgânica e suas frações sobre a fisiologia de hortaliças. Horticultura Brasileira 32: 14-20. Ação da matéria orgânica e suas frações sobre a fisiologia de hortaliças RESUMOOs adubos orgânicos fornecem nutrientes, melhoram as condições do solo e também apresentam bioatividade, ou seja, efeitos estimulantes nas plantas. Efeitos como indução de crescimento e melhora na qualidade nutricional vêm justificando a crescente comercialização de bioestimulantes e fertilizantes de base orgânica. Neste trabalho, a bioatividade é discutida com base na fisiologia vegetal. A literatura disponível comprova ações fisiológicas destas substâncias, principalmente no crescimento de raízes e aumento na absorção de nutrientes. Contudo, as abordagens das pesquisas têm focado nos aspectos básicos relacionados às frações extraídas em laboratório, que não representam necessariamente a realidade da matéria orgânica em seu estado natural no solo e sua bioatividade. Por outro lado, co...

A standard protocol to evaluate the effects of biostimulants on plant physiology is still lacking. The proton pumps present in the vacuolar and plasma membranes are the primary agents responsible for the regulation of the electrochemical gradient that energizes the nutrient uptake system and acid growth mechanism of plant cells. In this study, two of these enzymes were characterized as biochemical markers of biostimulant activity. A simple and fast protocol based on the degree of root acidification using a pH sensitive dye and the Micro-Tom tomato as a plant model is proposed as an efficient methodology to prove the efficacy of biostimulants that are claimed to improve nutrient acquisition and root growth. The results agree with the data from more conventional, expensive and time-consuming proton pump assays. A direct correlation was found between plasmalemma proton-adenosine triphosphatase (H + -ATPase) activation and the amount of rhizosphere acidification observed in the bromocresol gel. Moreover, roots of the diageotropica (dgt) Micro-Tom plants, defective in auxin responses, barely acidify bromocresol purple gel even in the presence of indole-3-acetic acid (IAA, 1 μM). The biostimulant TEA (vermicompost water extract, 25 %) enhances proton extrusion by 40 % in wild type (WT) plants, but no effect was induced in dgt plants. These results reinforce the notion that the class of biostimulant known as humic substances stimulates plant proton pumps and promotes root growth by exerting an auxin-like bioactivity and establish the usefulness of an economically and technically feasible assay to certify this kind of biostimulant.