Studies on the influence of foliar nutrient sprays on the root exudation pattern in four crop plants

Summary Studies conducted to examine the exudation pattern of amino acids and sugars in four crop plants,viz sorghum, sunnhemp, ragi, and tomato indicated that in all, 17 known amino acids and 4 sugars were exuded and that the number and nature of the exuded amino acids and sugars differed with the plant species and with the age of plant. Glutamic and aspartic acids were found to be present in the exudates of all the plant species at all stages of plant growth examined. The quantities of amino acids and sugars differed with plant species and the maximum quantity of the chemicals was exuded during the early stages of plant growth. Glutamic acid among amino acids, and glucose among sugars, were always present in higher concentrations than the others, in the exudates in all the four crop plants.Foliar application of nitrogen in the form of NaNO₃ and phosphorus as Na₂HPO₄, was found to alter the exudation pattern of amino acids and sugars and such influence differed in different plant species. There was a general increase in the total concentration of amino acids and a decrease in sugar content in the exudates after treatment of the foliage with N, while a decrease in the amino acid content and increase in total sugars with P-treatment was observed.     

Composition of root exometabolites of the symbiotically effective pea cultivar triumph and its parental forms

The qualitative and quantitative composition of low-molecular exometabolites in roots of pea (Pisum sativum L.) was studied with a cultivar Triumph and its parental forms (a symbiotically effective variety k-8274 and a modern highly productive cv. Classic). A relationship between root exudation and the ability of cultivars to establish symbiosis was analyzed. In the early stages of plant growth, the roots of cv. Triumph exhibited low exudation of organic acids, sugars, and amino acids. The quantitative composition of organic acids in the root exudates of cv. Triumph was close to that of cv. k-8274, whereas the composition of sugars and amino acids was similar to that of cv. Classic. In the field experiment, the effect of inoculation with a mixture of rhizobium strains and mycorrhizal fungus on plant growth was more evident in cv. Triumph than in cvs. Classic and k-8274. The results suggest that the high symbiotic potential of cv. Triumph is related to exudation of pyruvic and succinic acids that were the major components of root exometabolites both in Triumph and k-8274 cultivars.     

Tomato root exudates and their effect on the growth and antifungal activity of Pseudomonas strains

The study of the effect of the root exometabolites of tomato plants on the growth and antifungal activity of the plant growth-promoting Pseudomonas strains showed that the antifungal activity of plant growth-promoting rhizobacteria in the plant rhizosphere may depend on the sugar and organic acid composition of root exudates.     

Root Exudates of Tomato Plants and Their Effect on the Growth and Antifungal Activity of Pseudomonas Strains

The study of the effect of the root exometabolites of tomato plants on the growth and antifungal activity of plant growth–promoting Pseudomonas strains showed that the antifungal activity of plant growth–promoting rhizobacteria in the plant rhizosphere may depend on the sugar and organic acid composition of root exudates.     

温室CO2施肥对黄瓜幼苗根系生长及分泌物和伤流液组成的影响

为探讨温室蔬菜CO2施肥的根际效应,以黄瓜幼苗为试材,研究了CO2施肥(上午施肥/上、下午施肥;施肥浓度/对照浓度(950±50)/(350±50)μmol/molCO2)对根系生长及分泌物和伤流液组成的影响。结果表明,CO2施肥明显促进黄瓜幼苗根系发育,根系生物量显著增加;单株根系分泌物中氨基酸、糖、有机酸和酚酸总量增加,但单位鲜重根系分泌量却呈现增幅减少、无变化甚至降低趋势,说明单株分泌量增加主要由根系生长量的增加所引起。CO2施肥促进幼苗对养分的吸收,伤流液中矿质元素、ZT浓度增加,但GA、ABA和IAA浓度降低;与上午CO2施肥相比,上、下午均CO2施肥的效果更明显。CO2施肥促进了黄瓜幼苗根系发育及其代谢活性,为地上部的旺盛生长创造了条件。     

Characterization of Root Exudates at Different Growth Stages of Ten Rice (Oryza sativa L.) Cultivars

Abstract: Plant root exudates play important roles in the rhizosphere. We tested three media (nutrient solution, deionized water and CaSO₄ solution) for three periods of time (2, 4 and 6 h) for collecting root exudates of soil‐grown rice plants. Nutrient culture solution created complications in the analyses of exudates for total organic C (TOC) by the wet digestion method and of organic acids by HPLC due to the interference by its components. Deionized water excluded such interference in analytical analyses but affected the turgor of root cells; roots of four widely different rice cultivars excreted 20 to 60 % more TOC in deionized water than in 0.01 M CaSO₄. Furthermore, the proportion of carbohydrates in TOC was also enhanced. Calcium sulfate solution maintained the osmotic environment for root cells and did not interfere in analytical procedures. Collection for 2 h avoided under‐estimation of TOC and its components exuded by rice roots, which occurred during prolonged exposure. By placing plants in 0.01 M CaSO₄ for 2 h, root exudates of soil‐grown traditional, tall rice cultivars (Dular, B40 and Intan), high‐yielding dwarf cultivars (IR72, IR52, IR64 and PSBRc 20), new plant type cultivars (IR65598 and IR65600) and a hybrid (Magat) were collected at seedling, panicle initiation, flowering and maturity and characterized for TOC and organic acids. The exudation rates were, in general, lowest at seedling stage, increased until flowering but decreased at maturity. Among organic acids, malic acid showed the highest concentration followed by tartaric, succinic, citric and lactic acids. With advancing plant growth, exudation of organic acids substituted exudation of sugars. Root and shoot biomass were positively correlated with carbon exudation suggesting that it is driven by plant biomass. As root exudates provide substrates for methanogenesis in rice fields, large variations in root exudation by cultivars and at different growth stages could greatly influence CH₄ emissions. Therefore, the use of high‐yielding cultivars with lowest root excretions, for example IR65598 and IR65600, would mediate low exudate‐induced CH₄ production. The screening of exciting rice cultivars and breeding of new cultivars with low exudation rates could offer an important option for mitigation of CH₄ emission from rice agriculture to the atmosphere.     

济南地区日光温室土壤养分的分布状况和累积规律

对济南地区不同种植年限日光温室内外及不同蔬菜作物的土壤养分差异和频数分布进行比较,分析了日光温室土壤养分随种植年限的累积特征,并对不同蔬菜温室土壤养分与种植年限进行曲线拟合,研究了引起土壤盐渍化和酸化的原因.结果表明:济南地区日光温室土壤碱解氮、速效磷、速效钾、有机质、电导率显著高于棚外土壤,增幅分别为135.3％、475.2％、290.1％、97.7％、188.7％,pH值较棚外土壤降低0.31;各土壤养分的频数分布曲线均呈正态分布.不同蔬菜温室土壤养分含量各有差别,其中,碱解氮和电导率为番茄＞甜椒＞黄瓜,有机质和pH为黄瓜＞甜椒＞番茄,速效磷为黄瓜＞番茄＞甜椒,速效钾为番茄＞黄瓜＞甜椒.日光温室土壤有酸化的趋势,但不明显.温室土壤盐渍化程度加重主要由碱解氮和速效钾的累积所致.pH的降低与碱解氮的累积关系密切.日光温室各土壤养分随种植年限的累积规律基本一致,1～2年为养分的快速累积期,3～4年的累积速率变缓,以后处于一个稳定水平,整体上表现为土壤系统的动态平衡.除pH的累积为负向外,其他均为正向累积.不同蔬菜温室土壤养分与种植年限呈极显著相关,且可用2次或3次曲线进行拟合.     

Differential inactivation of seed exudate stimulation of Pythium ultimum sporangium germination by Enterobacter cloacae influences biological control efficacy on different plant species

This study was initiated to understand whether differential biological control efficacy of Enterobacter cloacae on various plant species is due to differences in the ability of E. cloacae to inactivate the stimulatory activity of seed exudates to Pythium ultimum sporangium germination. In biological control assays, E. cloacae was effective in controlling Pythium damping-off when placed on the seeds of carrot, cotton, cucumber, lettuce, radish, tomato, and wheat but failed to protect corn and pea from damping-off. Seeds from plants such as corn and pea had high rates of exudation, whereas cotton and cucumber seeds had much lower rates of exudation. Patterns of seed exudation and the release of P. ultimum sporangium germination stimulants varied among the plants tested. Seed exudates of plants such as carrot, corn, lettuce, pea, radish, and wheat were generally more stimulatory to P. ultimum than were the exudates of cotton, cucumber, sunflower, and tomato. However, this was not directly related to the ability of E. cloacae to inactivate the stimulatory activity of the exudate and reduce P. ultimum sporangium germination. In the spermosphere, E. cloacae readily reduced the stimulatory activity of seed exudates from all plant species except corn and pea. Our data have shown that the inability of E. cloacae to protect corn and pea seeds from Pythium damping-off is directly related to its ability to inactivate the stimulatory activity of seed exudates. On all other plants tested, E. cloacae was effective in suppressing damping-off and inactivating the stimulatory activity of seed exudates.     

Biocontrol activity and root colonization by Pseudomonas corrugata strains CCR04 and CCR80 against Phytophthora blight of pepper

Previously, we selected Pseudomonas corrugata strains CCR04 and CCR80 as rhizobacteria suppressive to Phytophthora blight of pepper caused by Phytophthora capsici. In this study, we investigated soil microbial activity in pepper plants root-drenched with strains CCR04 and CCR80 in relation to their biocontrol activity, root colonization by using bacterial population counts and scanning electron microscopy, biofilm formation and cell motility as well as cell sensitivity to hydrogen peroxide (H₂O₂). As a result, strains CCR04 and CCR80 more effectively suppressed disease expression in pepper plants through root colonization than did Paenibacillus polymyxa AC-1 (positive control), Escherichia coli DH5α (negative control) or MgSO₄ solution (untreated control). Strains CCR04 and CCR80 had efficient biofilm formation and cell motility (swimming and swarming activities) abilities and responded to certain tested compounds (amino acids, organic acids and sugars), which can be found in root exudates. Strains CCR04 and CCR80 and the positive control strain AC-1 were relatively insensitive to H₂O₂, a reactive oxidative species at concentration up to 20 mM, unlike the negative control strain DH5α. Taken together, these results suggest that P. corrugata CCR04 and CCR80 can effectively inhibit P. capsici infection of pepper plants through successful colonization of plant roots. This bacterial colonization may be facilitated by the biofilm formation ability and cell motility in addition to reduced sensitivity to H₂O₂ and probably the production of antimicrobial compounds. These findings highlight the potential of strains CCR04 and CCR80 as biocontrol agents for the management of Phytophthora blight of pepper.     

Effect of Pseudomonas fluorescens on the root exudates of two tomato mutants differently sensitive to Fe chlorosis

Plants with different Fe-mobilization properties are known to differ in the amount and kind of Fe-reducing and Fe-chelating compounds exuded by their roots. Although rhizosphere bacteria are known to affect the exudation of organic compounds by the plant roots, their effect on the root exudates of plants differing in Fe-mobilization properties is not known. We studied the effect of Pseudomonas fluorescens, on the exudation of sugars and organic and amino acids by roots of an iron chlorosis-resistant (T3238FER) and a chlorosis-susceptible (T3238fer) tomato mutant. Under sterile conditions two tomato mutants grew equally well and did not differ in the total amount of sugars and organic acid exuded by their roots. More amino acids, however, were exuded by the roots of T3238FER than T323fer. Mutants differed in the amount of oxalic acid and the amino acids Ala, Asp, Gaba, Gln, Gly, His, Hyl, Ile, Leu, Lys, Phe, Pro, and Val exuded by their roots into sterile rooting media. Addition of P. fluorescens to the rooting medium did not affect the growth of T3238FER but stimulated the root growth of chlorosis-susceptible T3238fer, reduced the amounts of glucose, arabinose and fructose but increased the amount of sucrose, reduced the amounts of fumaric, malic and oxalic acid but increased the amounts of citric and succinic acid in the rooting media of both mutants. P. fluorescens resulted in the following changes in the amino acids in the rooting media: reduced the amounts of Gly, Leu, and Lys in T3238FER, and of Asp, Gln, Hyp, and Ile in T3238fer, and increased the amounts of Cys, Glu, His, Hyp, Ile, Phe and Tyr in T3238FER and of Ala, Glu, His, Phe, and Ser in T323fer—in cases more than 40-fold. These differential effects of P. fluorescens in altering the pattern of organic and amino acids compounds with some Fe-chelating properties detected in the rooting medium of these two mutants may indicate that the differences in Fe-chlorosis susceptibility of these tomato mutants may be the result of, or modified by, the interactions between plant roots and rhizosphere microorganisms. We postulate that the Fe-chlorosis susceptibility in plants may be the product of the interactions between soil microorganisms and plant roots, and may not be solely related to the plant per se.     

Simultaneous Growth on Citrate Reduces the Effects of Iron Limitation during Toluene Degradation in Pseudomonas

Rhizoremediation has been suggested as an attractive bioremediation strategy for the effective breakdown of pollutants in soil. The presence of plant root exudates such as organic acids, sugars, and amino acids that may serve as carbon sources or biosynthetic building blocks and the limited bioavailability of iron may influence the degradation of pollutants in the rhizosphere. To test the effect of such compounds on hydrocarbon degradation, trace concentrations of yeast extract or mixtures of organic acids and amino acids were added to continuous cultures of Pseudomonas putida mt2 and P. putida WCS358 (TOL) growing on toluene. By addition of these compounds increased growth yields and higher specific growth rates on toluene were obtained. The effects of iron limitation on the substrate utilization pattern of both strains were tested by growing the strains on a mixture of toluene and the readily degradable carbon source citrate while the iron concentration was varied. Simultaneous use of both substrates under carbon-limited as well as iron-limited conditions was observed. Growth yields were less reduced and iron requirement was lower during iron-limited growth in the toluene + citrate grown cultures compared to cultures in which toluene was used as the sole carbon source. The kinetic properties of the cells for toluene degradation were less hampered by the lack of iron when citrate was used as an additional carbon source. The results indicate that the availability of low concentrations of natural organic compounds, such as produced in the rhizosphere, may positively influence the degradative performance of hydrocarbon-degrading bacteria.     

Root exudates of plants

Summary Comparisons have been made of the amount and composition of seed and seedling exudates in barley, wheat, cucumber, and bean. Except in the case of wheat a greater proportion of the total nitrogen content in the seed exudates was formed by protein and peptide nitrogen than by nitrogen of free amino acids. In contrast, the greater part of the total nitrogen in the exudates of seedlings was formed by free amino acid nitrogen, except in the case of barley. Peptides represented 8 to 26 per cent of the protein and peptide fraction in seed exudates, the highest amount being found in bean. On an equal weight basis, the spectrum of amino acids released from seeds and seedlings differed little between barley, wheat and cucumber; greater differences were observed in bean exudates. Seedlings exuded reducing substances to a considerably greater extent than seeds. The spectrum of reducing sugars in seed and root exudates differed greatly, especially in the case of keto sugars. Differences in the organic acid spectra were small, except for bean plants, the seedling exudate of which contained more organic acids with a richer spectrum than seed exudate. Both seeds and seedlings of cucumber exuded a small quantity and a poor spectrum of organic acids.     

Biochemical Changes in Root Exudate and Xylem Sap of Tomato Plants Infected with Meloidogyne incognita

Under two monoxenic culture techniques of growing plants (filter paper and silica sand cultures), sugar in root exudate from Meloidogyne incognita-infected tomato increased 133 to 836% over controls. In contrast, amino acids were moderately reduced 52 to 56%. Chromatographic analysis showed that galled root exudate contained three sugars, twelve amino acids, and three organic acids, whereas healthy root exudate contained four sugars, fifteen amino acids, and four organic acids. Polysaccharide was responsible for the large increase of sugars in galled root exudates. The concn and the absolute amount of total sugars in the infected plant xylem sap were greater than in healthy plant xylem sap up to 6 wk after inoculation, whereas amino acids were moderately lower than in controls throughout the test period. Chromatographic analysis showed that xylem sap from both healthy and infected plants at 4 wk after inoculation contained four sugars and five organic acids. We identified 18 and 17 amino acids in the healthy and infected plant xylem sap, respectively. The concn of sugar increased as the nematode inoculum increased at 2, 4 and 6 wk after inoculation. The amino acids in all samples from the infected plant moderately decreased with an increase of nematode inoculum. We suggest that changes in total sugars and amino acids, of infected plant xylem sap and root exudate are a probable mechanism by which tomato plants are predisposed to Fusarium wilt.     

Tomato seed and root exudate sugars: composition, utilization by Pseudomonas biocontrol strains and role in rhizosphere colonization

The role of tomato seed and root exudate sugars as nutrients for Pseudomonas biocontrol bacteria was studied. To this end, the major exudate sugars of tomato seeds, seedlings and roots were identified and quantified using high-performance liquid chromatographic (HPLC) analysis. Glucose, fructose and maltose were present in all studied growth stages of the plant, but the ratios of these sugars were strongly dependent on the developmental stage. In order to study the putative role of exudate sugar utilization in rhizosphere colonization, two approaches were adopted. First, after co-inoculation on germinated tomato seeds, the root-colonizing ability of the efficient root-colonizing P. fluorescens strain WCS365 in a gnotobiotic quartz sand-plant nutrient solution system was compared with that of other Pseudomonas biocontrol strains. No correlation was observed between the colonizing ability of a strain and its ability to use the major exudate sugars as the only carbon and energy source. Secondly, a Tn5lacZ mutant of P. fluorescens strain WCS365, strain PCL1083, was isolated, which is impaired in its ability to grow on simple sugars, including those found in exudate. The mutation appeared to reside in zwf, which encodes glucose-6-phosphate dehydrogenase. The mutant grows as well as the parental strain on other media, including tomato root exudate. After inoculation of germinated sterile tomato seeds, the mutant cells reached the same population levels at the root tip as the wild-type strain, both alone and in competition, indicating that the ability to use exudate sugars does not play a major role in tomato root colonization, despite the fact that sugars have often been reported to represent the major exudate carbon source. This conclusion is supported by the observation that the growth of mutant PCL1083 in vitro is inhibited by glucose, a major exudate sugar, at a concentration of 0.001%, which indicates that the glucose concentration in the tomato rhizosphere is very low.     

Interactions of Bacillus spp. and plants--with special reference to induced systemic resistance (ISR)

Biological control of soil-borne pathogens comprises the decrease of inoculum or of the disease producing activity of a pathogen through one or more mechanisms. Interest in biological control of soil-borne plant pathogens has increased considerably in the last few decades, because it may provide control of diseases that cannot or only partly be managed by other control strategies. Recent advances in microbial and molecular techniques have significantly contributed to new insights in underlying mechanisms by which introduced bacteria function. Colonization of plant roots is an essential step for both soil-borne pathogenic and beneficial rhizobacteria. Colonization patterns showed that rhizobacteria act as biocontrol agents or as growth-promoting bacteria form microcolonies or biofilms at preferred sites of root exudation. Such microcolonies are sites for bacteria to communicate with each other (quorum sensing) and to act in a coordinated manner. Elicitation of induced systemic resistance (ISR) by plant-associated bacteria was initially demonstrated using Pseudomonas spp. and other Gram-negative bacteria. Several strains of the species Bacillus amyloliquefaciens, B. subtilis, B. pasteurii, B. cereus, B. pumilus, B. mycoides, and B. sphaericus elicit significant reductions in the incidence or severity of various diseases on a diversity of hosts. Elicitation of ISR by these strains has been demonstrated in greenhouse or field trials on tomato, bell pepper, muskmelon, watermelon, sugar beet, tobacco, Arabidopsis sp., cucumber, loblolly pine, and two tropical crops (long cayenne pepper and green kuang futsoi). Protection resulting from ISR elicited by Bacillus spp. has been reported against leaf-spotting fungal and bacterial pathogens, systemic viruses, a crown-rotting fungal pathogen, root-knot nematodes, and a stem-blight fungal pathogen as well as damping-off, blue mold, and late blight diseases. This progress will lead to a more efficient use of these strains which is worthwhile approach to explore in context of biocontrol strategies.     

Root exudates of plants

Summary The effect of temperature on the exudation of sugars and amino acids from germinating seeds of maize and cucumber and the effect of the so called cold shock on root exudation of plant seedling was studied. After the first forty-eight hours the exudation of these compounds from germinating seeds generally rises in proportion to the temperature. However, certain relative changes were observed in the composition of the exudates. Exudation of maltose and fructose with arabinose, for example is lower at 28°C than at 19°C.When plants previously growing under favourable temperature conditions were exposed for three days to a lower temperature, there was a several times higher exudation from the roots of maize and a marked rise in exudation from roots of cucumber in comparison with control plants. In the exudates of maize affected by cold shock 3 new oligosaccharides and fructose with saccharose in addition to previously detected substances were found.     

Fitness-related traits in a parasitoid fly are mediated by effects of plants on its host

Plants can affect parasitoids directly, by reducing or enhancing their ability to locate hosts, or indirectly by affecting the fitness of herbivores and thus of parasitoids. Tritrophic interactions between three host plants (cucumber, tomato, sweet pepper), a polyphagous herbivore Trichoplusia ni and a generalist parasitoid Compsilura concinnata were assessed. Plants had a strong effect on T. ni larval survival, as well as on C. concinnata fitness-related traits: cucumber-fed hosts yielded parasitoids with shorter larval development time and females had heavier pupal weights than parasitoids from host larvae that were fed tomato. Furthermore, C. concinnata was more efficient at finding cucumber-fed than tomato-fed T. ni . These results suggest that C. concinnata has different efficiency and potential as a biocontrol agent on the different crops. This highlights the importance of assessing tritrophic interactions in systems where an inundative biological control agent may be released against generalist targets on more than one crop plant.     

Spatial and temporal variation in citrate and malate exudation and tissue concentration as affected by P stress in roots of white lupin

Exudation of carboxylates represents one the most efficient strategies used by P-starved white lupin (Lupinus albus L.) to acquire phosphorus from sparingly soluble sources. This exudation occurs through proteoid root clusters, with citrate being the predominant organic acid released. The occasional detection of malate in whole root exudates suggests that this acid would also be released, but from tissues other than root clusters. To investigate the spatial and temporal pattern of exudation, citrate and malate exudation and concentration were measured in whole roots and root sections of white lupin, from seedling emergence to plant senescence due to P starvation. Both organic acids were detected in whole root exudates of P-stressed plants, and they were released at similar rates throughout the experiment. Malate was predominantly exuded from apices of both seedling taproots and proteoid roots, whereas citrate exudation was restricted to proteoid root clusters. Studies directed to address the association between carboxylate exudation and concentration in proteoid root clusters showed a non-linear response for citrate, within the range of 7 to 23 mol g^–1 fresh weight. This association was further assessed by altering citrate concentration in the whole root. Adding P to 24-day-old P-starved plants reduced citrate concentration and exudation to the level of the control P-fed plants, demonstrating that citrate exudation and concentration are associated. Malate exudation and concentration did not correlate significantly. Results indicate that citrate release by P-starved white lupin would occur whenever a certain threshold of citrate concentration is attained, and that the sites, the rates and the span of transient exudation depend on the physiological age of the tissue.     

Quantitative and qualitative effects of phosphorus on extracts and exudates of sudangrass roots in relation to vesicular-arbuscular mycorrhiza formation

A comparison was made of water-soluble root exudates and extracts of Sorghum vulgare Pers. grown under two levels of P nutrition. An increase in P nutrition significantly decreased the concentration of carbohydrates, carboxylic acids, and amino acids in exudates, and decreased the concentration of carboxylic acids in extracts. Higher P did not affect the relative proportions of specific carboxylic acids and had little effect on proportions of specific amino acids in both extracts and exudates. Phosphorus amendment resulted in an increase in the relative proportion of arabinose and a decrease in the proportion of fructose in exudates, but did not have a large effect on the proportion of individual sugars in extracts. The proportions of specific carbohydrates, carboxylic acids, and amino acids varied between exudates and extracts. Therefore, the quantity and composition of root extracts may not be a reliable predictor of the availability of substrate for symbiotic vesicular-arbuscular mycorrhizal fungi. Comparisons of the rate of leakage of compounds from roots with the growth rate of vesicular-arbuscular mycorrhizal fungi suggest that the fungus must either be capable of using a variety of organic substrates for growth, or be capable of inducing a much higher rate of movement of specific organic compounds across root cell membranes than occurs through passive exudation as measured in this study.