Nicola Tomasi

Release of plant-borne flavonoids into the rhizosphere and their role in plant nutrition

Plants release a multitude of organic compounds into the rhizosphere, some of which are flavonoids. These products of secondary metabolism are mainly studied for their antioxidant properties and for their role in the establishment of rhizobium-legume symbiosis; however, it has been recently demonstrated that flavonoids can also affect nutrient availability through soil chemical changes. This review will give an overview of the types and amounts of flavonoids released by roots of different plant species, as well as summarize the available knowledge on root exudation mechanisms. Subsequently, factors influencing their release will be reported, and the methodological approaches used in the literature will be critically described. Finally, the direct contribution of plant-borne flavonoids on the nitrogen, phosphorous and iron availability into the rhizosphere will be discussed.

Microbial interactions in the rhizosphere: beneficial influences of plant growth-promoting rhizobacteria on nutrient acquisition process. A review

Plant growth-promoting rhizobacteria (PGPR) are soil bacteria that are able to colonize rhizosphere and to enhance plant growth by means of a wide variety of mechanisms like organic matter mineralization, biological control against soil-borne pathogens, biological nitrogen fixation, and root growth promotion. A very interesting feature of PGPR is their ability of enhancing nutrient bioavailability. Several bacterial species have been characterized as P-solubilizing microorganisms while other species have been shown to increase the solubility of micronutrients, like those that produce siderophores for Fe chelation. The enhanced amount of soluble macro- and micronutrients in the close proximity of the soil-root interface has indeed a positive effect on plant nutrition. Furthermore, several pieces of evidence highlight that the inoculation of plants with PGPR can have considerable effects on plant at both physiological and molecular levels (e.g., induction of rhizosphere acidification, up- and downregulation of genes involved in ion uptake, and translocation), suggesting the possibility that soil biota could stimulate plants being more efficient in retrieving nutrients from soil and coping with abiotic stresses. However, the molecular mechanisms underlying these phenomena, the signals involved as well as the potential applications in a sustainable agriculture approach, and the biotechnological aspects for possible rhizosphere engineering are still matters of discussion.

Plant-borne flavonoids released into the rhizosphere: impact on soil bio-activities related to plant nutrition. A review

Plants produce and release in the surrounding soil, the so-called rhizosphere, a vast variety of secondary metabolites. Among them, flavonoids are the most studied, mainly for their role in the establishment of rhizobium–legume symbiosis; on the other hand, some studies highlight that they are also important in the plant strategies to acquire nutrients from the soil, for example, by acting on its chemistry. The scope of this review is to give a quick overview on the types and amounts of plant-released flavonoids in order to focus on their effects on soil activities that in turn can influence nutrient availability and so plant mineral nutrition; emphasis is given to the different nutrient cycles, soil enzyme, and soil bacteria activities, and their influence on soil macrofauna and roots of other plants. Finally, the possible outcome of the climate change on these processes is discussed.

Plasma membrane H+-ATPase-dependent citrate exudation from cluster roots of phosphate-deficient white lupin.

White lupin (Lupinus albus L.) is able to grow on soils with sparingly available phosphate (P) by producing specialized structures called cluster roots. To mobilize sparingly soluble P forms in soils, cluster roots release substantial amounts of carboxylates and concomitantly acidify the rhizosphere. The relationship between acidification and carboxylate exudation is still largely unknown. In the present work, we studied the linkage between organic acids (malate and citrate) and proton exudations in cluster roots of P-deficient white lupin. After the illumination started, citrate exudation increased transiently and reached a maximum after 5 h. This effect was accompanied by a strong acidification of the external medium and alkalinization of the cytosol, as evidenced by in vivo nuclear magnetic resonance (NMR) analysis. Fusicoccin, an activator of the plasma membrane (PM) H+-ATPase, stimulated citrate exudation, whereas vanadate, an inhibitor of the H+-ATPase, reduced citrate exudation. The burst of citrate exudation was associated with an increase in expression of the LHA1 PM H+-ATPase gene, an increased amount of H+-ATPase protein, a shift in pH optimum of the enzyme and post-translational modification of an H+-ATPase protein involving binding of activating 14-3-3 protein. Taken together, our results indicate a close link in cluster roots of P-deficient white lupin between the burst of citrate exudation and PM H+-ATPase-catalysed proton efflux.

Genome-wide microarray analysis of tomato roots showed defined responses to iron deficiency

BackgroundPlants react to iron deficiency stress adopting different kind of adaptive responses. Tomato, a Strategy I plant, improves iron uptake through acidification of rhizosphere, reduction of Fe3+ to Fe2+ and transport of Fe2+ into the cells. Large-scale transcriptional analyses of roots under iron deficiency are only available for a very limited number of plant species with particular emphasis for Arabidopsis thaliana. Regarding tomato, an interesting model species for Strategy I plants and an economically important crop, physiological responses to Fe-deficiency have been thoroughly described and molecular analyses have provided evidence for genes involved in iron uptake mechanisms and their regulation. However, no detailed transcriptome analysis has been described so far.ResultsA genome-wide transcriptional analysis, performed with a chip that allows to monitor the expression of more than 25,000 tomato transcripts, identified 97 differentially expressed transcripts by comparing roots of Fe-deficient and Fe-sufficient tomato plants. These transcripts are related to the physiological responses of tomato roots to the nutrient stress resulting in an improved iron uptake, including regulatory aspects, translocation, root morphological modification and adaptation in primary metabolic pathways, such as glycolysis and TCA cycle. Other genes play a role in flavonoid biosynthesis and hormonal metabolism.ConclusionsThe transcriptional characterization confirmed the presence of the previously described mechanisms to adapt to iron starvation in tomato, but also allowed to identify other genes potentially playing a role in this process, thus opening new research perspectives to improve the knowledge on the tomato root response to the nutrient deficiency.

Secretion activity of white lupin’s cluster roots influences bacterial abundance, function and community structure

White lupin (Lupinus albus L. cv. Amiga) reacts to phosphate deficiency by producing cluster roots which exude large amounts of organic acids. The detailed knowledge of the excretion physiology of the different root parts makes it a good model plant to study plant-bacteria interaction. Since the effect of the organic acid exudation by cluster roots on the rhizosphere microflora is still poorly understood, we investigated the abundance, diversity and functions of bacteria associated with the cluster roots of white lupin, with special emphasis on the influence of root proximity (comparing root, rhizosphere soil and bulk soil fractions) and cluster root growth stages, which are characterized by different excretion activities. Plants were grown for five weeks in microcosms, in the presence of low phosphate concentrations, on acidic sand inoculated with a soil suspension from a lupin field. Plate counts showed that bacterial abundance decreased at the stage where the cluster root excretes high amounts of citrate and protons. In vitro tests on isolates showed that the frequencies of auxin producers were highest in juvenile and mature cluster roots and significantly decreased in senescent cluster roots. However, no significant difference in the frequency of auxin producers was found between cluster and non cluster roots. The diversity and structure of bacterial communities were investigated by DGGE of 16S rDNA and 16S rRNA. The diversity and community structure were mostly influenced by root proximity and, to a lesser extent, by cluster root stage. The richness of bacterial communities decreased with root proximity, whereas the proportion of active populations increased. The high citrate and proton excretion occurring at the mature stage of cluster roots had a strong impact on the structure and richness of the bacterial communities, both in the root and in the rhizosphere soil.

Beneficial effects of silicon on hydroponically grown corn salad (Valerianella locusta (L.) Laterr) plants.

Soil-less cultivation of horticultural crops represents a fairly recent innovation to traditional agriculture which has several advantages including higher water-use efficiency. When plants are grown with this system, their roots come in contact with nutrients solely via the hydroponic solution. Although its beneficial effects have been widely demonstrated, silicon (Si) is mostly omitted from the composition of nutrient solutions. Therefore, the objective of this study was to assess the beneficial effect of Si addition to hydroponic solution on quali-quantitative aspects of edible production of two cultivars of corn salad (Valerianella locusta (L.) Laterr.) grown in soil-less floating system. Impacts on shelf life of this food were also studied. Results show that the supply of Si increased the edible yield and the quality level reducing the nitrate concentration in edible tissues. This result might be attributed to changes either in the metabolism (such as the nitrate assimilation process) or to the functionality of root mechanisms involved in the nutrient acquisition from the outer medium. In fact, our results show for the first time the ability of Si to modulate the root activity of nitrate and Fe uptake through, at least in part, a regulation of gene expression levels of the proteins involved in this phenomenon. In addition, the presence of Si decreased the levels of polyphenoloxidase gene expression at harvest and, in post-harvest, slowed down the chlorophyll degradation delaying leaf senescence and thus prolonging the shelf life of these edible tissues. In conclusion, data showed that the addition of Si to the nutrient solution can be a useful tool for improving quali-quantitatively the yield of baby leaf vegetable corn salad as well as its shelf life. Since the amelioration due to the Si has been achieved only with one cultivar, the recommendation of its inclusion in the nutrient solution does not exclude the identification of cultivars suitable for this cultivation system and the comprehension of agronomical and environmental factors which could limit the Si benefits.

Influence of hydroponic and soil cultivation on quality and shelf life of ready‐to‐eat lamb's lettuce (Valerianella locusta L. Laterr)

BACKGROUND Nowadays, there is an increasing interest in the hydroponic floating system to cultivate leafy vegetables for ready-to-eat salads. It is reasonable that different growing systems could affect the quality and shelf life of these salads. RESULTS The quality and shelf life of ready-to-eat lambs lettuce grown in protected environment in soil plot or in soil-less system over hydroponic solution with or without the addition of 30 µmol L⁻¹ silicon were evaluated. Minimum effects were observed on colour, firmness and microbial counts. Hydroponic cultivation largely affected plant tissue hydration, leading to weight loss and structural modifications during refrigerated storage. The shelf life of lambs lettuce was limited by the development of visually detectable unpleasant sensory properties. Shelf life, calculated by survival analysis of consumer acceptability data, resulted about 7 days for soil-cultivated salad and 2 days for the hydroponically grown ones. The addition of silicon to the hydroponic solution resulted in an interesting strategy to increase plant tissue yield and reduce nitrate accumulation. CONCLUSIONS Although hydroponic cultivation may have critical consequences on product quality and shelf life, these disadvantages could be largely counterbalance by increased yield and a reduction of nitrate accumulation when cultivation is performed on nutritive solutions with supplemental addition of silicon.

Copper accumulation in vineyard soils: Rhizosphere processes and agronomic practices to limit its toxicity.

Viticulture represents an important agricultural practice in many countries worldwide. Yet, the continuous use of fungicides has caused copper (Cu) accumulation in soils, which represent a major environmental and toxicological concern. Despite being an important micronutrient, Cu can be a potential toxicant at high concentrations since it may cause morphological, anatomical and physiological changes in plants, decreasing both food productivity and quality. Rhizosphere processes can, however, actively control the uptake and translocation of Cu in plants. In particular, root exudates affecting the chemical, physical and biological characteristics of the rhizosphere, might reduce the availability of Cu in the soil and hence its absorption. In addition, this review will aim at discussing the advantages and disadvantages of agronomic practices, such as liming, the use of pesticides, the application of organic matter, biochar and coal fly ashes, the inoculation with bacteria and/or mycorrhizal fungi and the intercropping, in alleviating Cu toxicity symptoms.

Nitrate transport in cucumber leaves is an inducible process involving an increase in plasma membrane H+-ATPase activity and abundance

BackgroundThe mechanisms by which nitrate is transported into the roots have been characterized both at physiological and molecular levels. It has been demonstrated that nitrate is taken up in an energy-dependent way by a four-component uptake machinery involving high- and low- affinity transport systems. In contrast very little is known about the physiology of nitrate transport towards different plant tissues and in particular at the leaf level.ResultsThe mechanism of nitrate uptake in leaves of cucumber (Cucumis sativus L. cv. Chinese long) plants was studied and compared with that of the root. Net nitrate uptake by roots of nitrate-depleted cucumber plants proved to be substrate-inducible and biphasic showing a saturable kinetics with a clear linear non saturable component at an anion concentration higher than 2 mM. Nitrate uptake by leaf discs of cucumber plants showed some similarities with that operating in the roots (e.g. electrogenic H+ dependence via involvement of proton pump, a certain degree of induction). However, it did not exhibit typical biphasic kinetics and was characterized by a higher Km with values out of the range usually recorded in roots of several different plant species. The quantity and activity of plasma membrane (PM) H+-ATPase of the vesicles isolated from leaf tissues of nitrate-treated plants for 12 h (peak of nitrate foliar uptake rate) increased with respect to that observed in the vesicles isolated from N-deprived control plants, thus suggesting an involvement of this enzyme in the leaf nitrate uptake process similar to that described in roots. Molecular analyses suggest the involvement of a specific isoform of PM H+-ATPase (CsHA1) and NRT2 transporter (CsNRT2) in root nitrate uptake. At the leaf level, nitrate treatment modulated the expression of CsHA2, highlighting a main putative role of this isogene in the process.ConclusionsObtained results provide for the first time evidence that a saturable and substrate-inducible nitrate uptake mechanism operates in cucumber leaves. Its activity appears to be related to that of PM H+-ATPase activity and in particular to the induction of CsHA2 isoform. However the question about the molecular entity responsible for the transport of nitrate into leaf cells therefore still remains unresolved.