Re-sorption of organic components by roots of Zea mays L. and its consequences in the rhizosphere

The re-sorption of carbon compounds from the rhizosphere was investigated using ¹⁴C-labelled glucose, mannose and citric acid. Uptake in roots of 5-day-old, intact Zea mays plants in sterile solution culture was determined over a period of 48 hours. Under optimal growth conditions significant re-absorption of glucose and mannose occurred with the uptake rates being 70.5 and 40.2 g compound g^-1 root DW h^-1, respectively. For glucose and mannose approximately 25% of the ¹⁴C label taken up by the root was recovered inside the plant as low-MW compounds and 33% polymerized into high MW compounds. 42% was respired as ¹⁴C-CO₂. Citric acid by comparison showed little accumulation within plant tissues (11.4%) with most being respired and recovered as ¹⁴C-CO₂ in KOH traps (88%). The uptake rate for citric acid was 34.8 g g^-1 root DW h^-1. Over the 48-hour period a net efflux (i.e. exudation) of labelled plus unlabelled C was observed at a rate of 608 g C g^-1 root DW h^-1 (equivalent to 1520 g glucose/mannose). Of the C released as root exudates, a minimum estimate of the amount of C taken back into the plant was therefore 9.5%. The two main C fluxes within the rhizosphere, namely release of C by the root and uptake by the microorganisms, have been well documented in recent years. It is now apparent however that a third flux term, re-sorption of C by roots, can also be identified. This may play an important but previously overlooked role within the rhizosphere, and further work is needed to determine its significance.A comparison between exudate release in static (permitting accumulation of C) and flowing culture (C removed as it is released) was also made with the respective rates being 15.36 and 45.18 mg C g^-1 root DW in 2 days. The relative important of re-sorption in natural environments and laboratory experiments is discussed.     

Re-sorption of organic compounds by roots of Zea mays L. and its consequences in the rhizosphere

The influx and efflux of sugar-C and the cycling of C within intact maize roots (Zea mays L.) was studied in sterile solution culture. Using metabolic inhibitors it was shown that roots could take up sugars against the concentration gradient probably via H⁺-ATPase dependent plasmalemma proton cotransporters. In contrast to this, no evidence was found for an ATPase mediated efflux of sugars from the root. All parts of the root were capable of taking up exogenous sugars. Examination of sugar exudation sites along the root slowed efflux at all locations, with the amount of efflux linearly correlated with internal cellular concentration. The results clearly indicated that the influxefflux mechanisms are linked both spatially, temporally and with respect to the sugars capable of transportation. The turnover of C within the root was found to be extremely rapid with turnover of the soluble sugar pool being 0.8 to 15 times daily depending on root spatial location. The results strongly suggest that the recapture of sugars from outside the root plays an important role in regulating the amount of C lost to the soil which in turn will reduce both pathogen attraction and the size of the rhizosphere microbial population and will also increase the plant's C efficiency.     

Influx and efflux of amino acids from Zea mays L. roots and their implications for N nutrition and the rhizosphere

The aim of the study was to investigate the ability of Zea mays L. roots to regulate the amount of free amino acids present in the rhizosphere. The active uptake of amino acids was shown to conform to Michaelis-Menten kinetics. Comparison of amino acid-N and NO₃-N kinetic parameters and soil solution concentrations showed that root uptake of free amino acids from soil may contribute significantly to a plant's N budget. The influx of amino acids also helps to minimize net C/N losses to the soil, and is therefore important in regulating the size of the rhizosphere microbial population. Experimental data and a computer simulation model of amino acid influx/efflux in a sterile solution culture, showed that roots were capable of re-sorping over 90% of the amino acids previously lost into solution as a result of passive diffusion.     

Influx and efflux of organic acids across the soil-root interface of Zea mays L. and its implications in rhizosphere C flow

The influx and efflux of organic acids across the root-soil interface were investigated in intact, sterile maize (Zea mays L.) roots under a variety of experimental conditions. Under nutrient-sufficient conditions the efflux of organic acids was shown to constitute < 1% of the total C lost across the root-soil interface. Under severe nutrient stress, however, the rates of malate and citrate efflux from the root increased 33 and 12 fold respectively. Influx experiments indicated that roots could not directly reabsorb citrate-Fe³⁺ or other metal complexes from solution. Influx of citrate was observed only at high external citrate concentration ( 1 mM) or from solutions with low ionic strengths. It was postulated that citrate influx is of little importance in a soil environment.     

Bacterial communities associated with the rhizosphere of transgenic Bt 176 maize (Zea mays) and its non transgenic counterpart

The effect of transgenic Bt 176 maize on the rhizosphere bacterial community has been studied with a polyphasic approach by comparing the rhizosphere of Bt maize cultivated in greenhouse with that of its non transgenic counterpart grown in the same conditions. In the two plants the bacterial counts of the copiotrophic, oligotrophic and sporeforming bacteria, and the community level catabolic profiling, showed no significant differences; differences between the rhizosphere and bulk soil bacterial communities were evidenced. Automated ribosomal intergenic spacer analysis (ARISA) showed differences also in the rhizosphere communities at different plant ages, as well as between the two plant types. ARISA fingerprinting patterns of soil bacterial communities exposed to root growth solutions, collected from transgenic and non transgenic plants grown in hydroponic conditions, were grouped separately by principal component analysis suggesting that root exudates could determine the selection of different bacterial communities.     

玉米幼苗根系分泌物对芘污染的响应

根际袋土培试验研究了玉米幼苗根系分泌物中的可溶性糖、低分子量有机酸和氨基酸对不同芘污染水平(50、200、800mgkg-1,记为T1、T2、T3)的响应差异,探讨芘胁迫下植物根系的生理生态效应。结果表明,较低浓度芘可适当地刺激玉米的生长,高浓度芘处理抑制了玉米的生长,并且抑制作用随芘处理浓度的提高而增强;芘对玉米根系的影响要大于对茎叶的影响。芘胁迫下促进了根系分泌可溶性糖、低分子量有机酸和氨基酸增多。T1、T2和T3处理根系分泌物中可溶性糖、低分子量有机酸、氨基酸含量分别是T0处理的1.14、1.81、1.35倍,1.24、4.31、2.94倍,1.58、5.56、5.40倍。不同芘污染水平下,乙酸分泌量表现为T2T3T1,酒石酸和柠檬酸分泌量表现为T3T2T1,草酸分泌量表现为T3≈T2T1。芘处理对根系分泌氨基酸种类的影响不大,而对各氨基酸分泌量的变化幅度影响较大;芘胁迫处理对于18种常见氨基酸组分的分泌量的影响各不相同。不同芘污染水平下,天门冬氨酸、丝氨酸和丙氨酸分泌量表现为T3T2T1,苏氨酸、谷氨酸、脯氨酸、甘氨酸、胱氨酸、缬氨酸、甲硫氨酸、异亮氨酸、亮氨酸、酪氨酸、苯丙氨酸、赖氨酸、组氨酸、γ-氨基丁酸、鸟氨酸分泌量表现为T2T3T1。     

根分泌物对根际矿物营养及根际微生物的效应

综述了根系分泌物对植物生长的生理生态学效应,并就根系分泌物的定义、产生机制、组成成分和影响因素等方面进行了讨论。指出根系分泌物在缓解低矿物营养胁迫对植株造成的伤害及决定根际微生物的种群密度和数量方面起着重要的作用;根系分泌物的产生机制多样,组成成分复杂,影响因素繁多。对根分泌物的深入研究有助于进一步了解植物体与土壤间进行的生理生化过程及其调控机制。     

玉米幼苗根际土壤微生物活性对芘污染的响应

用根际袋法土培试验研究了玉米幼苗根际与非根际土壤微生物量碳、微生物熵、代谢熵和土壤酶活性对不同芘污染水平(50、200、800mg·kg-1,记为T1、T2、T3)的响应差异。结果表明,较低浓度芘可适当的刺激玉米幼苗的生长,而较高浓度芘则抑制幼苗生长,其抑制作用随芘处理浓度的提高而增强;芘对玉米根系的影响要大于对茎叶的影响。玉米幼苗能够明显促进土壤中芘的去除。根际和非根际土壤中芘的去除率分别为56.67%-76.18%和32.64%-70.44%,根际土壤中芘的平均去除率比非根际土壤高16.06%。同处理中根际土壤芘含量显著低于非根际土壤,随着芘处理浓度的提高其差异更加显著。根际土壤微生物量碳、微生物熵、多酚氧化酶活性、脱氢酶活性和磷酸酶活性均高于非根际土壤,代谢熵低于非根际土壤,且其差异随芘处理浓度的提高而增大。在不同芘污染水平下,微生物量碳、微生物熵和脱氢酶活性根际和非根际土壤为T1T2T3,代谢熵为T3T2T1;多酚氧化酶活性根际土壤为T2T1T3,非根际土壤为T1T2T3;磷酸酶活性根际土壤为T3T1T2,非根际土壤为T1T2T3。土壤中残余芘含量与土壤微生物量碳、微生物熵、多酚氧化酶、脱氢酶和磷酸酶活性呈显著负相关,与代谢熵呈显著正相关。     

Organic acids in the rhizosphere of Banksia integrifolia L.f.

Water leachates of the proteoid root layer of a mature stand of Banksia integrifolia were analysed for low molecular weight (LMW) organic acids by GC, HPLC and colorimetric techniques. Large amounts of organic acids (2500 g in 100 mL of leachate) were found in the proteoid root layer compared to the surrounding soil and leaf litter (230 g in 100 mL of leachate). Citric acid represented approximately 50% of the total organic acids leached, malic acid approximately 18%, and aconitic acid constituted approximately 17%. Concentration of citric acid in the proteoid root layer may enhance the availability of phosphorus for plant uptake.     

Role of root derived organic acids in the mobilization of nutrients from the rhizosphere

The role of organic acids in the mobilization of plant nutrients from the rhizosphere was assessed in seven contrasting soil types. The results indicated that malate was poor at mobilizing micronutrients from all the test soils, whilst citrate was capable of mobilizing significant quantities. Citrate was also capable of mobilizing P from one soil which possessed a large Ca-P fraction. This mobilization of P was due to both the complexing action of the citrate anion and due to the dissolution properties of the protons released from citric acid upon equilibrium with the soil solution. The reaction of citrate with cations was found to be near instantaneous with significant absorption to the solid phase in some soils at low concentrations. Soil decomposition studies indicated that citrate was rapidly broken down in organic soils but was more resistant to degradation in subsoil horizons. It was concluded that organic acids can be expected to be of little consequence in nutrient mobilization from high pH soils, whilst in acid soils they may be involved both in a more general mechanism for micronutrient uptake or as a potential Al detoxification mechanism.Abbreviations C₆H₈O₇ Citric acid and H-citrate indicates - C₆H₅O₇Na₃ whilst Na-citrate indicates     

Bacterial diversity of the rhizosphere of maize (Zea mays) grown in tropical soil studied by temperature gradient gel electrophoresis

The bacterial diversity and population dynamics in the rhizosphere of two maize cultivars (Nitroflint and Nitrodent) grown in tropical soils was studied, by traditional cultivation techniques and 16S rRNA gene-based molecular analysis of DNA directly extracted from soil and rhizosphere samples. Rhizosphere and soil samples were taken at three different plant growth stages. Total aerobic bacterial counts were determined. Fingerprints of the most dominant bacterial population were generated by TGGE separation of 16S rRNA gene fragments amplified from total community DNA using eubacterial specific primers. To reduce the complexity of TGGE fingerprints or to analyse less abundant populations, primers specific for different phylogenetic groups have been used. A comparison of the cfu obtained for rhizosphere of both cultivars indicated significant differences only for rhizosphere and soil samples taken 40 days after sowing. However, a comparison of TGGE patterns indicated that the composition of the bacterial community analysed at different plant growth stages for both cultivars was similar. A comparison of -, -proteobacterial and actinomycete TGGE patterns of both cultivars confirmed this observation. The eubacterial TGGE profiles reflected strong seasonal population shifts in the bacterial rhizosphere community of both maize cultivars which could be also observed in the TGGE patterns of - and -proteobacteria and to a lesser extent for actinomycetes. The rhizosphere effect was much more pronounced for young roots compared to samples taken from mature maize plants. The rhizosphere fingerprints showed a reduced complexity for young plants with up to five dominating bands while for mature plants patterns similar to those of soil were observed. Sequencing of dominant clones indicated that the dominant population found at all plant growth stages can be assigned to Arthrobacter populations.     

Potential errors caused by roots in analyses of rhizosphere soil

Roots contain high concentrations of many elements, and have the potential to interfere with measurements of chemical change in rhizosphere soil. To assess potential interferences, maize (Zea mays L.) roots (free of soil) and soil (free of roots) were extracted separately with several extractants commonly used to assess the status of soil nutrients. The maize roots were grown within filter envelopes which prevented direct contact with soil, but permitted passage of mineral nutrients and water from the adjacent soil. Water, ammonium acetate (pH 7), DTPA (pH 7.3), Morgan's solution (pH 4.8), and dilute HCl were used as extractants. Most elements were released readily into soluble forms from roots killed by freezing to lyse the cells. Significantly lower amounts were extracted from fresh roots, with the greatest differences between fresh and killed roots for the extractants H₂O and DTPA, which were the mildest in terms of acidity and salt concentration. Extraction of P from the fresh roots by H₂O and HCL was particularly low. Contamination of rhizosphere samples with root materials would almost certainly prevent the accurate measurement of water-soluble P, K, Mn, Zn, Cu, and Na in the slightly alkaline soil used in this experiment. Large errors would be likely also for P, Mn, and Cu extracted by ammonium acetate. The DTPA extractant is normally used only for micronutrient metals or heavy metals, and the small amounts of these elements released by roots should not contribute to significant error. With Morgan's solution, errors would likely be large only for P. Dilute HCl is a reasonably strong extractant for many elements in soil, and major errors from roots contained in rhizosphere samples are unlikely. The relatively high probability of errors in extractions of soluble elements from rhizosphere soil is unfortunate, because these elements are among the most readily available to plants and the most likely to be altered by the normal activities of roots.     

Wetting and drying cycles in the maize rhizosphere under controlled conditions. Mechanics of the root-adhering soil

Mechanical properties of the topsoil (sandy Podsol and silty Luvisol, FAO) adhering to maize (Zea mays L.) roots and its bulk soil counterpart were studied as a function of soil texture and final soil water suction at harvest, with three soil water suction values of approximately 30, 50 and 60 kPa. Two scales of observation were also selected: the whole soil:root system and the root-adhering soil aggregates. Three methods were used to characterize the stability of the soil:root system: mechanical shaking in air, and dispersion by low-power ultrasonication, with or without preliminary immersion of the soil:root system in water. Soil disruption kinetics, which were fitted with first-order kinetics equations, were analyzed and discussed. For example, silty soil ultrasonication kinetics, without preliminary water-immersion, could be divided into two parts: the first faster part, which was characterized by a mean rate K value of 6.8–7.2 mJ^-1, is attributed to soil slaking, whereas the second slower part, which was characterized by a mean rate K value of 1.5–1.6 mJ^-1, was attributed to the rupture of the `firmly root-adhering soil' from the roots. A clear plant effect was observed for both aggregate tensile strength and friability, with higher aggregate strength for the root-adhering silty soil (450–500 kPa) than for its bulk silty soil counterpart (410–420 kPa), and lower friability (coefficient of variation of the aggregate strength) for the root-adhering silty soil (e.g. 67% at a soil water suction value of 30 kPa) than for its bulk silty soil counterpart (e.g. 49% at asoil water suction value of 30 kPa). These effects were attributed to root exudation, which was significantly higher for the driest silty topsoil than for the wetter ones. In conclusion, the mechanical properties of the silty topsoil adhering to the maize roots are attributed to both physical and biological interactions occurring in the maize rhizosphere. This revised version was published online in June 2006 with corrections to the Cover Date.     

Graviresponsiveness and abscisic-acid content of roots of carotenoid-deficient mutants of Zea mays L.

The abscisic-acid (ABA) content of roots of the carotenoid-deficient w-3, vp-5, and vp-7 mutants of Z. mays was analyzed using gas chromatography-mass spectrometry with an analysis sensitivity of 6 ng ABA g^–1 fresh weight (FW). Roots of normal seedlings of the same lines were characterized by the following amounts of ABA (as ng ABA g^–1 FW,±standard deviation): w-3, 279±43; vp-5, 237±26; vp-7, 338±61. We did not detect any ABA in roots of any of the mutants. Thus, the lack of carotenoids in these mutants correlated positively with the apparent absence of ABA. Primary roots of normal and mutant seedlings were positively gravitropic, with no significant differences in the curvatures of roots of normal as compared with mutant seedlings. These results indicate that ABA 1) is synthesized in maize roots via the carotenoid pathway, and 2) is not necesary for positive gravitropism by primary roots of Z. mays.Abbreviation ABA abscisic acid     

Enhanced formation of aerenchyma and induction of a barrier to radial oxygen loss in adventitious roots of Zea nicaraguensis contribute to its waterlogging tolerance as compared with maize (Zea mays ssp. mays)

Enhancement of oxygen transport from shoot to root tip by the formation of aerenchyma and also a barrier to radial oxygen loss (ROL) in roots is common in waterlogging‐tolerant plants. Zea nicaraguensis (teosinte), a wild relative of maize (Zea mays ssp. mays), grows in waterlogged soils. We investigated the formation of aerenchyma and ROL barrier induction in roots of Z. nicaraguensis, in comparison with roots of maize (inbred line Mi29), in a pot soil system and in hydroponics. Furthermore, depositions of suberin in the exodermis/hypodermis and lignin in the epidermis of adventitious roots of Z. nicaraguensis and maize grown in aerated or stagnant deoxygenated nutrient solution were studied. Growth of maize was more adversely affected by low oxygen in the root zone (waterlogged soil or stagnant deoxygenated nutrient solution) compared with Z. nicaraguensis. In stagnant deoxygenated solution, Z. nicaraguensis was superior to maize in transporting oxygen from shoot base to root tip due to formation of larger aerenchyma and a stronger barrier to ROL in adventitious roots. The relationships between the ROL barrier formation and suberin and lignin depositions in roots are discussed. The ROL barrier, in addition to aerenchyma, would contribute to the waterlogging tolerance of Z. nicaraguensis.     

基于菌植互作的植物根际细菌钝化镉作用和机制研究进展

土壤重金属镉(Cd)污染严重危害农产品安全生产,植物根际细菌在钝化土壤Cd和帮助作物抵御Cd胁迫方面发挥重要作用。本文首先概括在修复Cd污染土壤中得到广泛应用的植物根际细菌种类,并从根际细菌直接吸附Cd、调整土壤理化特性、调控土壤微生物群落和其他作用4方面阐述了植物根际细菌对Cd的钝化作用,其次从菌植互作角度阐述植物根系分泌物与根际细菌群落相互影响对土壤Cd的钝化作用。最后展望重金属胁迫下植物根际钝化Cd核心菌群的构建,以在新兴学科与技术的快速发展中探明植物根系-微生物互作体系的分子机制,深入开展植物根际细菌钝化修复重金属污染土壤的理论研究和实践。     

Temperature dependence of trans plasma membrane electron transport in Zea mays L. roots

Intact Zea mays L. cv. Golden Bantam seedlings which were not cold adapted were exposed to various temperatures. Trans plasma membrane potential difference was measured in a temperature range from 0 to 40 °C using intracellular microelectrodes. The depolarization caused by electron transfer across the PM to artificial external electron acceptors was investigated. Active membrane potential increased with temperature in the range from 0 to 15 °C but was independent of temperature above 20 °C. Depolarization caused by the non-membrane-permeating electron acceptors hexacyanoferrate III (HCF III) and hexabromoiridate IV (HBIIV) took place over the whole temperature range investigated. The effect of HBI IV increased up to 10 °C whereas the HCF III effects increased up to 25 °C.     

Role of proteinaceous amino acids released in root exudates in nutrient acquisition from the rhizosphere

The role of proteinaceous amino acids in rhizosphere nutrient mobilization was assessed both experimentally and theoretically. The degree of adsorption onto the soil's solid phase was dependent on both the amino acid species and on soil properties. On addition of amino acids to both soil and freshly precipitated Fe(OH)₃, no detectable mobilization of nutrients (K, Na, Ca, Mg, Cu, Mn, Zn, Fe, S, P, Si and Al) was observed, indicating a very low complexation ability of the acidic, neutral and basic amino acids. This was supported by results from a solution equilibria computer model which also predicted low levels of amino acid complexation with solutes present in the soil solution. On comparison with the Fe(OH)₃ and equilibria data obtained for the organic acid, citrate, it was concluded that amino acids released into the rhizosphere have a limited role in the direct acquisition of nutrients by plants. The effectiveness of root exudates such as amino acids, phytosiderophores and organic acids in nutrient mobilization from the rhizosphere is discussed with reference to rhizosphere diffusion distances, microbial degradation, rate of complexation and the root's capacity to recapture exudate-metal complexes from the soil.     

Action of malformin A1 on gravitropic curvature in primary roots of maize (Zea mays L.)

Malformins, a small family of cyclic pentapeptides, are active plant growth regulators isolated from the fungusAspergillus niger. We purified malformin A₁ from the crude malformin A mixture, and studied its action in the gravitropic response of maize roots. Intact primary roots that had been pretreated vertically with malformin A₁ were placed in a humidified box in the horizontal position. Positive curvature (downward) was inhibited in the pretreated roots compared with the control. In addition, we measured the lateral transport of IAA in primary roots. Roots pretreated with malformin A, did not show asymmetric distribution of IAA between the upper and lower sides of the elongation zone. Malformin A, also stimulated ethylene production in maize root segments. Our results had suggested that malformin A₁ might inhibit the lateral transport of IAA across the roots from the upper to the lower side because of an increased level of ethylene. Therefore, we placed more IAA on the upper side at the initial phase of gravistimulation. These results were consistent with malformin A₁-pretreated roots showing inhibited positive gravitropic curvature.