A new concept of root exudation1

After discussing numerous models for exudation from the xylem of roots, we present a new biphasic exudation model based on osmoregulation of the root symplast by stretch-activated ion channels (SA channels). We tested some features of the model in maize roots. (1) Using a microdrop recorder we showed that bathing the roots in 50 mmol m^?3 gadolinium ions, known to inhibit some SA channels, inhibited xylem exudation by over 80% after 24h application. (2) Measuring xylem exudation from single roots into an attached micropipette revealed the capacity of the roots to perform strong autonomous exudation pulses. (3) In partially encased roots, the rhizodermis exuded water concurrently to xylem exudation. These results were regarded as supporting our model. An interesting observation with the microdrop recorder, which does not address the theory, is that addition of a variety of inorganic ions to distilled water as the roots' bathing medium instantaneously and reversibly increases xylem exudation, evidently nonosmotically.     

Evaluating the role of root citrate exudation as a mechanism of aluminium resistance in maize genotypes

Organic anion exudation by roots as a mechanism of aluminium (Al) resistance has been intensively studied lately. In the present study, we evaluated qualitative and quantitative aspects of root exudation of organic anions in maize genotypes of distinct sensitivity to Al in response to Al exposure. Maize seedlings were grown axenically in nutrient solution and root exudates were collected along the whole seminal root axis for a short period (4 h) using a divided-root-chamber technique. In root exudates collected from 10-mm long root apices, citrate accounted for 67% of the total organic anions found, followed by malate (29%), trans-aconitate (3%), fumarate (<1%), and cis-aconitate (1%). Rates of citrate exudation from root apices of two genotypes with differential resistance to Al were consistently higher in the Al resistant one, differing by a factor of 1.7 – 3.0 across a range of external Al concentrations. Furthermore, relative Al resistance of eight maize genotypes correlated significantly well with their citrate exudation rate measured at 40 M Al. Higher exudation rates were accompanied by a less inhibited root elongation. The exudation of citrate along the longitudinal axis of fully developed seminal roots showed a particular pattern: citrate was exuded mainly in the regions of root apices, either belonging to the main root or to the lateral roots in the most basal part of the main root. The involvement of citrate in a mechanism of Al resistance is evaluated in terms of protection of the root from the effects of excess Al on root elongation and on nutrient uptake along a root axis showing distinct sites of citrate exudation.     

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.     

Low-P tolerance by maize (Zea mays L.) genotypes: Significance of root growth,and organic acids and acid phosphatase root exudation

We investigated some mechanisms, which allow maize genotypes to adapt to soils which are low in available P. Dry matter production, root/shoot-ratio, root length and root exudation of organic acids and acid phosphatase were investigated in four maize genotypes grown under P-deficient and P-sufficient conditions in sterile hydroponic culture. A low-P tolerant, an acid-tolerant and a low-P susceptible genotype of maize were compared with a Swiss commercial cultivar. The study found increased root development and increased exudation of acid phosphatase under P-deficient conditions in all maize genotypes, except for the Swiss cultivar. Effects on root formation and acid phosphatase were greater for the low-P tolerant than for the low-P susceptible, and the acid soil tolerant genotypes. Organic acid contents in root tissues were increased under P deficiency and related to increased PEPC activity. However, the increase in contents was associated with an increase in exudation for the low-P tolerant genotype only. The low-P susceptible genotype was characterized by high organic acid content in roots and low organic acid exudation. The organic acids content in the phloem exudates of shoots was related to root exudation under different P supply, to the difference between lines in organic acids root content, but not to the low-P tolerance or susceptibility of maize genotypes.     

Experimental evaluation of an efflux-influx model of C exudation by individual apical root segments

The aim of this study was to evaluate if a model describing the efflux and the influx of C through the root surface could be fitted to experimental short-term kinetics of carbon (C) exudation by individual apical root segments in maize (Zea mays L.). The efflux of C was set constant or modelled by a power function of the distance from the apex to simulate the greater release of C around the root tip commonly reported in the literature. The influx was proportional to the C concentration in the external solution to simulate the active re-uptake of exudates by the root. Plants were exposed to full light or to shade to manipulate C allocation to roots. The model with a constant efflux gave satisfactory fits to the kinetics of exudation (average R(2)=0.66). The average gross efflux was then 2.1 mug C cm(-2) root surface h(-1). The model was improved if exudation was set more intense towards the root apex (average R(2)=0.74). The estimated gross efflux decreased then from 5.2 mug C cm(-2) h(-1) at the apex to 1.8 mug C cm(-2) h(-1) for the region located 5-25 cm from the root tip. The decrease in net exudation of individual roots due to the shading of plants was weak, which may indicate that the import of C by the primary roots studied was not reduced significantly. By describing the exudation of an apical root segment of variable length and diameter, the model is a first step in linking exudation to root system architecture models and to whole plant functioning.     

The Effects of Temperature on the Exudation Process of Excised Primary Roots of Zea mays

The fluid exudation rates and the ionic compositions of theexudates of excised maize roots have been determined in bathingmedia of 0.1 mM, 1.0 mM, 10.0 mM, and 50.0 mM KCl, each containing0.1 mM CaCl₂, at temperature intervals between 10 °C and30 °C.Analysis of these data in terms of an osmotic modelfor excised root exudation shows that the observed temperaturevariation in fluid exudation rate is accounted for by the observedtemperature variation in the osmotic driving force, the saltconcentration difference from xylem fluid to bathing medium.Temperature variation in the osmotic permeability of the rootand of the non-osmotic water flow are not significantly differentfrom zero.     

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

The exudation of soluble carbon compounds from Zea mays roots was investigated over a 10 day growth period under sterile and non-sterile solution culture conditions. The results showed that plants grown in sterile static solution culture, where C was allowed to accumulate, released 8 times less C than plants grown under culture conditions in which the solutions were replaced daily. The increased C loss from plant cultures in which exudates were removed daily was attributable to, (a) the reduced potential for root re-sorption of previously lost C, and (b), increasing diffusion gradients between the root and the surrounding bathing solution increasing passive leakage of exudates from the roots. In treatments where C was removed daily from the root-bathing solution, 86% of the total C lost was of a soluble low molecular weight nature, whereas, in sterile and non-sterile static cultures, allowing the accumulation of C over 10 days, this was reduced to 67.5 and 48% respectively. The main C fluxes operating in a solution culture system (efflux and influx of C by both roots and microorganisms) were examined using a computer simulation model to describe movement of soluble sugar-C in both sterile and non-sterile conditions. In sterile static cultures where C was allowed to accumulate in solution over a 10 day growth period, 98% of the C exuded was re-absorbed by the plant. Where C was removed daily from the root-bathing solution this was reduced to 86%. The predicted patterns of C accumulation were similar to those found in the experiments. Simulations showed that the pattern of accumulation and final equilibrium concentrations were dependent on the rate of exudation, the spatial characteristics of exudation, solution volume, root growth rate and the presence of a microbial population. Simulations under non-sterile conditions showed that roots can compete with microorganisms for exudates in solution indicating the possible importance of re-sorption in a soil environment. The results clearly indicate that roots are capable of regulating the net amount of C released into a solution culture with the amount of C collected being highly dependent on the experimental conditions employed. The possible implications of soluble C influx on processes operating within the rhizosphere and in experimental systems is discussed.     

Root Exudation of Phytochemicals in Arabidopsis Follows Specific Patterns That Are Developmentally Programmed and Correlate with Soil Microbial Functions

Plant roots constantly secrete compounds into the soil to interact with neighboring organisms presumably to gain certain functional advantages at different stages of development. Accordingly, it has been hypothesized that the phytochemical composition present in the root exudates changes over the course of the lifespan of a plant. Here, root exudates of in vitro grown Arabidopsis plants were collected at different developmental stages and analyzed using GC-MS. Principle component analysis revealed that the composition of root exudates varied at each developmental stage. Cumulative secretion levels of sugars and sugar alcohols were higher in early time points and decreased through development. In contrast, the cumulative secretion levels of amino acids and phenolics increased over time. The expression in roots of genes involved in biosynthesis and transportation of compounds represented in the root exudates were consistent with patterns of root exudation. Correlation analyses were performed of the in vitro root exudation patterns with the functional capacity of the rhizosphere microbiome to metabolize these compounds at different developmental stages of Arabidopsis grown in natural soils. Pyrosequencing of rhizosphere mRNA revealed strong correlations (p<0.05) between microbial functional genes involved in the metabolism of carbohydrates, amino acids and secondary metabolites with the corresponding compounds released by the roots at particular stages of plant development. In summary, our results suggest that the root exudation process of phytochemicals follows a developmental pattern that is genetically programmed.     

小麦-蚕豆间作对根系分泌低分子量有机酸的影响

通过盆栽试验收集了不同生育期单作和间作小麦、蚕豆的根系分泌物,用HPLC分析了根系分泌物中低分子量有机酸的含量和种类.结果表明: 小麦-蚕豆间作显著提高了有机酸的分泌量,在小麦分蘖期(57 d)、孕穗期(120 d)和灌浆期(142 d),间作使小麦根系有机酸分泌量分别提高155%、35.6%和92.6%;在蚕豆分枝期(57 d)和籽粒膨大期(142 d),间作使蚕豆根系有机酸分泌量提高87.4%和38.7%.小麦-蚕豆间作改变了根系分泌物中有机酸的种类,与单作小麦相比,在分蘖期,间作小麦根系分泌物中增加了乳酸;在拔节期(98 d),间作小麦根系分泌物中增加了柠檬酸,但未检测到乙酸;在蚕豆分枝期,间作蚕豆根系分泌物中增加了乙酸,但未检测到乳酸;在蚕豆籽粒膨大期,间作蚕豆根系分泌物中增加了乳酸.小麦-蚕豆间作提高了小麦根系有机酸的分泌速率,在小麦孕穗期,间作小麦分泌柠檬酸、富马酸的速率是单作小麦的179和184倍;在小麦灌浆期,间作小麦分泌乳酸的速率是单作的2.53倍.总之,小麦-蚕豆间作增加了有机酸的分泌量,改变了根系分泌物中有机酸的种类,提高了小麦根系有机酸的分泌速率.     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

Aluminum resistance in maize cannot be solely explained by root organic acid exudation. A comparative physiological study

Root apical aluminum (Al) exclusion via Al-activated root citrate exudation is widely accepted as the main Al-resistance mechanism operating in maize (Zea mays) roots. Nonetheless, the correlation between Al resistance and this Al-exclusion mechanism has not been tested beyond a very small number of Al-resistant and Al-sensitive maize lines. In this study, we conducted a comparative study of the physiology of Al resistance using six different maize genotypes that capture the range of maize Al resistance and differ significantly in their genetic background (three Brazilian and three North American genotypes). In these maize lines, we were able to establish a clear correlation between root tip Al exclusion (based on root Al content) and Al resistance. Both Al-resistant genotypes and three of the four Al-sensitive lines exhibited a significant Al-activated citrate exudation, with no evidence for Al activation of root malate or phosphate release. There was a lack of correlation between differential Al resistance and root citrate exudation for the six maize genotypes; in fact, one of the Al-sensitive lines, Mo17, had the largest Al-activated citrate exudation of all of the maize lines. Our results indicate that although root organic acid release may play a role in maize Al resistance, it is clearly not the only or the main resistance mechanism operating in these maize roots. A number of other potential Al-resistance mechanisms were investigated, including release of other Al-chelating ligands, Al-induced alkalinization of rhizosphere pH, changes in internal levels of Al-chelating compounds in the root, and Al translocation to the shoot. However, we were unsuccessful in identifying additional Al-resistance mechanisms in maize. It is likely that a purely physiological approach may not be sufficient to identify these novel Al-resistance mechanisms in maize and this will require an interdisciplinary approach integrating genetic, molecular, and physiological investigations.     

Observations on the Effects of Pressure Differences Between the Bathing Media and Exudates of Excised Maize Roots

Observation of the water fluxes from excised maize roots atvarious levels of suction showed significant differences betweencontrol roots and those which have been subjected to an evacuationpre-treatment. This involved placing excised lengths of rootin bathing medium in a flask which was evacuated (1 cmHg) for10 min; during this time air bubbles emerged from the cut ends. This pre-treatment has been shown to result in an increase inthe density of the root, which is ascribed to the replacementof air in the cortical intercellular spaces by bathing medium.The observed differences in the response to suction in the twogroups of roots is therefore explained as a decrease in theresistance to water flow of the cortex of the treated roots.     

镉胁迫下小麦根系分泌物的变化

通过水培和砂培两种方法 ,研究了镉胁迫下小麦 ( Triticum aestivm L.)根系分泌物变化。通过研究镉对小麦根系分泌氨基酸和糖 ,根系分泌其它一些次生代谢物质等的影响 ,以及根系分泌物对蔬菜种子萌发的影响 ,探讨镉胁迫下植物根系的生理生态效应。研究结果表明 :镉胁迫下 ,小麦根系分泌物无论在量上还是质上都有变化。镉胁迫下根系分泌的电解质、糖类、氨基酸以及其它一些次生代谢物质都有所变化 ,但变化情况有差异。电解质外渗率、糖类随 Cd2 + 浓度升高而增加 ,在低浓度 Cd2 + 作用下 ( 0 .5 mg/ L) ,随处理浓度的升高 ,氨基酸分泌量增加 ;当处理浓度高于相应浓度时 ,氨基酸分泌量随浓度升高而减少。随 Cd2 + 浓度升高 ,次生代谢物分泌种类减少。这说明小麦通过改变根分泌作用而缓解镉危害。由于根系分泌次生代谢物的活动比较复杂 ,因而根系的分泌作用有待于进一步研究。根系分泌物中可能含有某些物质抑制萝卜、白菜种子萌发 ,这是涉及到植物他感作用的问题 ,值得并有待于进一步研究。初步分析小麦根系在不同的环境中对镉胁迫的响应方式 ,为以后根分泌物的深入研究提供了一定的思路 ,也为镉污染区农作物的合理栽培提供新的参考资料。     

Stress-induced accumulation of wheat germ agglutinin and abscisic Acid in roots of wheat seedlings

Wheat germ agglutinin (WGA) levels in roots of 2-day-old wheat seedlings increased up to three-fold when stressed by air-drying. Similar results were obtained when seedling roots were incubated either in 0.5 molar mannitol or 180 grams per liter polyethylene glycol 6000, with a peak level of WGA after 5 hours of stress. Longer periods of osmotic treatment resulted in a gradual decline of WGA in the roots. Since excised wheat roots incorporate more [³⁵S]cysteine into WGA under stress conditions, the observed increase of lectin levels is due to de novo synthesis. Measurement of abscisic acid (ABA) levels in roots of control and stressed seedlings indicated a 10-fold increase upon air-drying. Similarly, a five- and seven-fold increase of ABA content of seedling roots was found after 2 hours of osmotic stress by polyethylene glycol 6000 and mannitol, respectively. Finally, the stress-induced increase of WGA in wheat roots could be inhibited by growing seedlings in the presence of fluridone, an inhibitor of ABA synthesis. These results indicate that roots of water-stressed wheat seedlings (a) contain more WGA as a result of an increased de novo synthesis of this lectin, and (b) exhibit higher ABA levels. The stress-induced increase of lectin accumulation seems to be under control of ABA.     

Aluminum resistance in two cultivars of Zea mays L.: Root exudation of organic acids and influence of phosphorus nutrition

Root exudation of organic acids as Al-chelating compounds and P nutrition have been suggested to play a major role in Al-resistance in higher plants. Effects of Al exposure on maize plant growth, and organic acid root content and root exudation under various levels of P nutrition were examined. Sikuani, a Colombian maize cultivar tolerant to acid soils with high Al saturation, and Corso, a Swiss cultivar, were grown in sterile hydroponic conditions for 21 days. Al-caused inhibition of root growth was lower in Sikuani than in Corso. Al effect on plant growth was decreased with increasing P content in roots. Al content in roots increased with increasing P content and was higher in Sikuani than in Corso. When exposed to Al, the contents in root apices as well as the root exudation of citric and malic acids in Corso and citric, malic and succinic acids in Sikuani increased, and were higher in Sikuani than in Corso. Increased PEP carboxylase (PEPC) activity in root apices after Al exposure partially explained the variations of organic acid content in the roots. These Al-induced changes in PEPC activity, organic acid content and exudation were reduced in plants supplied with higher P concentrations during the 21 days prior to treatment. Increased secretion of organic acids after exposure to Al appeared to be specific to Al and was not totally explained by increased root content in organic acids.     

Manganese reduction in the rhizosphere of mycorrhizal and nonmycorrhizal maize

The influence of rhizosphere microorganisms and vesicular-arbuscular (VA) mycorrhiza on manganese (Mn) uptake in maize (Zea mays L. cv. Tau) plants was studied in pot experiments under controlled environmental conditions. The plants were grown for 7 weeks in sterilized calcareous soil in pots having separate compartments for growth of roots and of VA mycorrhizal fungal hyphae. The soil was left either uninoculated (control) or prior to planting was inoculated with rhizosphere microorganisms only (MO-VA) or with rhizosphere microorganisms together with a VA mycorrhizal fungus [Glomus mosseae (Nicol and Gerd.) Gerdemann and Trappe] (MO+VA). Mycorrhiza treatment did not affect shoot dry weight, but root dry weight was slightly inhibited in the MO+VA and MO-VA treatments compared with the uninoculated control. Concentrations of Mn in shoots decreased in the order MO-VA > MO+VA > control. In the rhizosphere soil, the total microbial population was higher in mycorrhizal (MO+VA) than nonmycorrhizal (MO-VA) treatments, but the proportion of Mn-reducing microbial populations was fivefold higher in the nonmycorrhizal treatment, suggesting substantial qualitative changes in rhizosphere microbial populations upon root infection with the mycorrhizal fungi. The most important microbial group taking part in the reduction of Mn was fluorescent Pseudomonas. Mycorrhizal treatment decreased not only the number of Mn reducers but also the release of Mn-solubilizing root exudates, which were collected by percolation from maize plants cultivated in plastic tubes filled with gravel quartz sand. Compared with mycorrhizal plants, the root exudates of nonmycorrhizal plants had two fold higher capacity for reduction of Mn. Therefore, changes in both rhizosphere microbial population and root exudation are probably responsible for the lower acquisition of Mn in mycorrhizal plants.     

Sloughing of cap cells and carbon exudation from maize seedling roots in compacted sand

Sloughing of root cap cells and exudation of mucilage plays an important role in the penetration of compacted soils by roots. For the first time we have quantified the rate of sloughing of root cap cells in an abrasive growth medium that was compacted to create mechanical impedance to root growth. The number of maize ( Zea mays ) root cap cells sloughed into sand increased as a result of compaction, from 1930 to 3220 d⁻¹ per primary root. This represented a 12-fold increase in the number of cells sloughed per mm root extension (from 60 to >700). We estimated that the whole of the cap surface area was covered with detached cells in compacted sand, compared with c . 7% of the surface area in loose sand. This lubricating layer of sloughed cells and mucilage probably decreases frictional resistance to soil penetration. The total carbon deposited by the root was estimated at c . 110 μg g⁻¹ sand d⁻¹. Sloughed cells accounted for <10% of the total carbon, the vast majority of carbon being contained in mucilage exudates.