Effects of low aluminum activity in nutrient solutions on the organic acid concentrations in maize plants

The effects of low aluminum (Al) activity in nutrient solution on the concentrations of organic acids in two cultivars of maize (Zea mays L.), HS7777 Al‐sensitive and C525‐M Al‐tolerant, were studied. Aluminum stress increased total organic acid concentration in the roots and in the shoots for both cultivars. The relative increase of t‐aconitic, citric, formic, malic, and quinic acids was higher in the roots than in the shoots for both cultivars. The concentrations of c‐aconitic, isocitric, malonic, oxalic, and succinic organic acids were reduced by Al stress, principally for C525‐M. There were no consistent differences in organic acid concentrations between the cultivars to discriminate Al tolerance. The Al tolerance for C525‐M may be justified by lower Al concentrations in the root tips where cellular division takes place and/or by higher excretion of organic acids from roots to the rhizosphere for detoxification of Al by chelation.     

Aluminum tolerance of two wheat cultivars (Brevor and Atlas66) in relation to their rhizosphere pH and organic acids exuded from roots

Phytotoxicity of aluminum (Al) has become a serious problem in inhibiting plant growth on acid soils. Under Al stress, the changes of rhizosphere pH, root elongation, absorption of Al by wheat roots, organic acids exuded from roots, and some main factors related to Al-tolerant mechanisms have been studied using hydroponics, fluorescence spectrophotometry, and high performance liquid chromatography (HPLC). Two wheat cultivars, Brevor and Atlas66, differing in Al tolerance are chosen in the study. Accordingly, the rhizosphere pH has a positive effect on Al tolerance. Atlas66 (Al-tolerant) has higher capability to maintain high rhizosphere pH than Brevor (Al-sensitive) does. High pH can reduce Al3+ activity and toxicity, and increase the efficiency of exuding organic acids from the roots. More inhibition of root elongation has been found in Brevor because of the exposure of roots to Al3+ solution at low pH. Brevor accumulate more Al in roots than Atlas66 even at higher pH. Al-induced exudation of malic and citric acids has been found in Atlas66 roots, while no Al-induced organic acids have been found in Brevor. These results indicate that the Al-induced secretion of organic acids from Atlas66 roots has a positive correlation with Al tolerance. Comprehensive treatment of Al3+ and H+ indicates that wheat is adversely influenced by excess Al3+, rather than low pH.     

Mechanisms of Aluminum Tolerance in Phosphoenolpyruvate Carboxylase Transgenic Rice

Transgenic rice, in which C4-phosphoenolpyruvate carboxylase (PEPC) is overexpressed, was grown in hydroponic culture to evaluate its tolerance for high aluminum (Al) content in relation to organic acid synthesis and exudation into the rhizosphere. With regard to root growth in short-term experiments, PEPC-transgenic rice was more tolerant of Al than the wild type. At increased Al levels in the root medium, root tips of the transgenic rice accumulated less Al than those of the wild type. Aluminum-induced oxalate exudation from roots occurred at increased rates in the transgenic line. Overexpression of C4-PEPC drastically increased PEPC activity in leaves of the transgenic rice, causing higher organic acid concentrations in leaves and roots. The increased organic acid concentrations seemed to cause higher exudation rates of oxalate from roots, leading to a detoxification of Al in the root apoplast and rhizosphere.     

低分子量有机酸对高岭石中铝释放的影响

选择了几种土壤中可能存在的低分子量脂肪羧酸 ,研究了它们对高岭石中铝释放的影响。结果表明 ,有机酸可以通过络合作用促进高岭石中铝的释放。几种有机酸对体系中可溶性铝影响的大小顺序为 :草酸 >柠檬酸 >丙二酸 >苹果酸 >乳酸。草酸、柠檬酸和乳酸对可溶性铝释放的促进作用随体系pH的升高而减小 ,其中草酸体系中可溶性铝随pH的变化幅度最大。在苹果酸体系中 ,可溶性铝随pH的升高而稍有增加。体系中的可溶性铝随有机酸浓度的增加而增加 ,而交换性铝随柠檬酸浓度的增加而减少 ,先随苹果酸浓度增加而增加 ,然后又逐渐减少。与对照相比 ,柠檬酸和草酸使交换性铝的量减小 ,苹果酸和乳酸在低pH下使交换性铝明显增加 ,而苹果酸在较高pH下使交换性铝减少。有机酸影响释放出的铝在固液相间的分配比 ,苹果酸在低pH下使体系中释放出的大部分铝以交换性形态存在 ,而在较高pH下 ,大部分铝以可溶形态存在。在草酸体系中 ,释放出铝的大部分都以可溶形态存在。不同有机酸的不同表现与体系中铝的溶解平衡、铝的吸附 -解吸平衡、有机酸的吸附 -解吸平衡、有机酸的离解平衡和铝与有机酸的络合 -离解平衡有关。     

籽粒苋不同富钾基因型根系分泌物中有机酸和氨基酸的变化特点

采用溶液培养试验研究了籽粒苋(Amaranthus.spp.)不同富钾基因型在不同供钾水平条件下3个不同生长时期内根系分泌物中氨基酸和有机酸的种类及含量变化情况。结果表明,1)籽粒苋根系分泌物中氨基酸和有机酸含量随着供钾水平的升高而降低,且富钾基因型始终大于一般基因型。在低钾胁迫时,根系分泌物中有10种氨基酸和5种有机酸出现,而在供钾正常时则缺少谷氨酸、甘氨酸、苯丙氨酸、草酸和柠檬酸等;酪氨酸、-氨基丁酸和丝氨酸的含量约占氨基酸分泌总量的一半以上;苹果酸含量约占有机酸分泌总量的60%以上。2)籽粒苋生长50d时,一般基因型氨基酸和有机酸的分泌总量较生长40d时迅速降低,而富钾基因型降低速度则相对较为平稳。在3个生长期内,酪氨酸、-氨基丁酸和丝氨酸均为两类基因型根系分泌物中的主要氨基酸种类,苹果酸则是主要的有机酸类型,其在氨基酸和有机酸分泌总量中所占相对比例均随生长期的延长而升高。3)籽粒苋根系分泌物处理后的土壤速效钾含量均高于清水对照处理,富钾基因型在低钾胁迫时的根系分泌物对土壤钾的活化作用明显大于一般基因型。     

小麦和油菜Mn有效性差异的机制

By solution culture experiment,three wheat genotypes(Triticum aestivum,L.) and two oilseed rape genotypes (Brassica napus L.)differing in Mn efficiency under Mn-deficient conditions were used to study mechanisms of the difference in Mn efficiency between wheat and oilseed rape,The results showed that there were significant differences in the abilities of MnIV reduction and acidification in root rhizosphere between the two species.Compared with wheatl,oilseed rape had much higher reducing capacity and intensity of rhizosphere acidification under Mn-deficient conditions.Moreover,the higher ratio of functional leaves Mn/old leaves Mn in oilseed rape than in wheat was also an imprtant factor for the different Mn efficiencies between the two species.     

Effect of Aluminum and Silicon in Solution on the Growth of Rice (Oryza sativa L.) uspension Cells

Cells of two indica rice (Oryza sativa L.) varieties, one aluminum-tolerant (IAC3), and the other aluminum-sensitive (IR45), were grown in suspension culture in a medium with or without Al (0-800 µM), Si (0-2,000 µM), or Al and Si in combination as well as in R2 medium as control. The influence of Al or Si on cell growth was evaluated based on the growth of the cells in the treatment medium as well as in the R2 medium and compared with the effect on rice seedlings. Similar responses were observed for the cells and the seedlings; Al treatment resulted in a significant reduction of cell growth in both varieties, and the growth was reduced with the increase of the Al uptake by the cells. IAC3 was more tolerant to Al and it absorbed more Al than IR45. On the contrary, Si alone had almost no effect or exerted a negligible effect on the cell growth in both varieties and Si was hardly absorbed by the cells. There was a slight alleviative effect of Si on Al toxicity in the suspension cells of IR45, whereas, a considerable ameliorative effect in the seedlings. The content of total Al and of AI insoluble in citric acid in the cells were higher in the sole Al (800 µM) treatment than in the Al-Si combination treatment. This tendency was also observed in aluminon staining. The differences in the alleviative effects of Si and the uptake of Al and Si between the suspension cells and seedlings may have resulted from the presence of roots and morphological differences in the apoplastic tissues.     

The effects of Fe deficiency on low molecular weight organic acid exudation and cadmium uptake by Solanum nigrum L.

Abstract

The influence of Fe-deficiency on the root exudation of low molecular weight organic acids (LMWOAs), pH alteration and cadmium (Cd) accumulation and translocation were investigated in morel (Solanum nigrum L.) in hydroponic culture experiments. Tartaric, citric, malic and acetic acids were monitored because these acids were abundant and often detected as root exudates. Results showed that Fe-deficient plants excreted large amounts of LMWOAs in comparison with Fe-sufficient plants across all Cd treatments (p <0.05). In both cases the concentrations of the four organic acids were tartaric > citric > malic > acetic. The results showed that the Fe-deficient plants with higher concentrations of LMWOAs accumulated more Cd (p <0.05) and induced a decrease in solution pH compared with the Fe-sufficient plants. Cadmium accumulation in the Fe-deficient and Fe-sufficient plants had significant positive correlations with the exudation of malic and acetic acids (p <0.05 and p<0.01). Cadmium accumulation in the Fe-sufficient plants had a significant (p<0.01) positive correlation with the exudation of tartaric acid, whereas there was a negative correlation (p<0.01) between Cd accumulation and the exudation of tartaric acid in the Fe-deficient plants. No significant correlation between the exudation of citric acid and Cd accumulation was obtained. Our results indicate that Fe-deficiency could induce Cd accumulation and translocation through an increase of LMWOAs exudation and pH alteration, both of which enhance Cd bioavailability.     

低分子量有机酸对可变电荷土壤铝活化动力学的影响

从动力学角度研究了几种低分子量有机酸对2种酸性土壤中铝的活化和活化铝在土壤固/液相之间分配的影响。结果表明:对于络合能力弱的醋酸和乳酸,主要通过质子作用活化铝,且活化作用明显小于盐酸。而络合能力较强的苹果酸、草酸和柠檬酸,主要通过络合作用促进铝的释放,且这种作用随有机酸根阴离子络合能力的增强而增加。在氧化铁含量较高的砖红壤中,苹果酸、草酸和柠檬酸通过专性吸附增加土壤表面负电荷,从而增加土壤交换态铝;但在氧化铁含量较低的红壤中,草酸和柠檬酸主要通过形成可溶性铝络合物降低交换态铝。活化铝在土壤固/液相间的分配主要决定于溶液中有机阴离子与土壤固相表面对铝离子的竞争。醋酸和乳酸活化的铝主要以交换态铝存在;而草酸和柠檬酸活化的铝主要以有机酸-铝络合物存在于溶液中,特别是在氧化铁低的红壤中,这将促进铝在土壤-水体中的迁移。     

石灰性土壤中解磷优势菌的筛选研究

从石灰性土壤作物根际土样筛选出2株具有较强解P能力的优势菌株“B2”和“B67”,平板培养溶P圈D/d值分别为5.1和3·8,液体摇床培养过程中产生苹果酸、丙二酸、草酸和乳酸等多种有机酸,产酸总量分别为对照的28·25和19.78倍,pH分别较对照下降2.3和1.1,将“B2”和“B67”的培养液分别接种到4个不同地区土壤中恒温恒湿培养20d后测定结果表明,所有土壤中速效磷含量均有所提高,且接种“B2”处理分别为对照的2.64、3.04、1.71和2.20倍,接种“B67”处理分别为对照的2.12、1.90、1.35和1.78倍,且发现其溶P效果和土壤磷酸酶活性与有效活菌数变化趋势基本一致。     

The Influence of Phosphorus Sources on the Growth and Rhizosphere Soil Characteristics of Two Genotypes of Wheat (Triticum aestivum L.)

A rhizobox experiment was conducted to investigate the effects of phosphorus (P) sources on the rhizosphere soil characteristics of two wheat genotypes Xiaoyan54 (P-efficient) and Jing411 (P-inefficient), which were colonized with an arbuscular mycorrhizal fungus and grown in Cumulic Haplustoll. The four P sources included a control (no added P), OP [organic P: sodium (Na)-phytate], IP [inorganic P: monopotassium phosphate (KH₂PO₄)], and OPIP (Na-phytate plus KH₂PO₄). The results showed that when no exogenous P was added the shoot biomass of Xiaoyan54 was 28% significantly higher than Jing 411. The acid phosphatase activity (APA) in the rhizosphere soil was significantly depressed when inorganic P was added; the APA for Xiaoyan54 was higher than that of Jing411 in most of the layers regardless of the P source. Inorganic and organic P fertilizer conditions did not significantly impact the soil pH relative to the control, and the pH did not significantly differ between the two genotypes. In conclusion, when no exogenous P was added, shoot biomass of Xiaoyan54 was greater than Jing411. This increase was promoted by greater soil APA but not soil acidification. The rhizosphere soil pH was not altered across different wheat genotypes or P sources, but APA was increased in the soil of control and OP-treated plants compared with IP- and OPIP-treated plants. The APA of Xiaoyan54 was higher than Jing411 for all of the P sources. The response mechanism of rhizosphere soil acidification is different from acid phosphatase exudation in plants receiving different P sources under arbuscular mycorrhizal fungus inoculation.     

Combined Effect of Short-Term Water Deficit Stress and Aluminum Toxicity on Citrate Secretion from Soybean Roots

Abstract

In order to investigate the combined effect of drought stress and aluminum (Al) toxicity on citrate exudation in soybean, hydroponic cultivation with addition of Al and PEG-6000 was conducted to simulate Al-toxic dry soil. Results showed that 2-day exposure of soybeans to 5.5% (w/v) PEG-6000 or 100 µM AlCl₃ mainly hindered root growth, while combined exposure to PEG and Al (PEG/Al) reduced both root and shoot growth. Photosynthetic rate of first trifoliolate for the Al-tolerant genotype PI 416937 (PI) was not affected by imposition of 5.5% PEG/100 µM AlCl₃ (38–40 h), whereas photosynthetic rate for the Al-sensitive genotype YC was significantly reduced. Based on root fresh weight, Al-induced citrate exudation in the roots of soybean genotypes recovered from pre-treatment with 5.5 or 9% PEG was not altered, but was increased in the plants pretreated with 7% PEG without recovery. After 2 days of recovery from 2-day combined exposure to 5.5% PEG/100 µM AlCl₃, the Al-tolerant PI exuded more citrate than its control, but the Al-sensitive YC exuded significantly less citrate than its corresponding control. Split root experiment revealed that Al-induced citrate exudation in one half of the root system was significantly reduced by exposing the other half of the root system to 0.5 mM CaCl₂ solution containing 9% PEG or 9% PEG/50 µM AlCl₃. However, organic acid secretion was not observed in the half of the root system in the Ca solution when the other part of the root system was exposed to the Ca solution containing 50 µM AlCl₃, 9% PEG or 9% PEG/50 µM AlCl₃. This suggests that no Al- or drought-induced signals such as ABA are involved in the citrate secretion in soybean.     

Soluble root exudates of maize: Influence of potassium supply and rhizosphere microorganisms

Maize was grown for 36 days in solution culture with roots either under axenic conditions or in the presence of rhizosphere organisms. In other experiments with sterile roots the plants were grown with different concentrations of potassium. At the end of the experiments sugars, organic acids and amino acids in the nutrient solutions were determined. Under axenic conditions the exudates consisted of up to 65% sugars, up to 33% organic acids and only up to 2% amino acids. The same substances were detected in non-sterile nutrient solutions. In the presence of microorganisms fructose, arabinose and the predominating glucose decreased to almost one half, while sucrose was not affected. The amounts of organic acids were not changed by microbial growth. The main amino acid, glutamic acid, was nearly doubled by the presence of microorganisms, whereas other amino acids remained unchanged. The lower O₂ content of the nutrient solutions of non-sterile roots suggested microbial decomposition of monosaccharides. In another experiment with roots grown under axenic conditions and with different K treatments low K supply significantly increased the total amounts of sugars, organic acids and amino acids exuded g^?1 root dry matter. As in the previous experiment glucose, fumaric and oxalacetic acid as well as glutamic and aspartic acid dominated in the respective fractions. Again sugars and organic acids represented the major quantity of exudates, while amino acids amounted to less than 2%. In an additional experiment with another cultivar, with nitrate as N source and a 5-day longer growth period, somewhat different results were obtained. In the exudates sugars were found in lower amounts, probably due to a higher growth rate. Under these conditions organic acids were the prevailing root exudates. Unlike sugars and amino acids, their total quantity was not affected by K nutrition, but the proportion of malic acid increased with increasing K supply, while oxalacetic acid dominated at low K nutrition. Similarly the total amount of organic acids within the root was independent of K nutrition. However malic acid content increased with increasing K application, while the likewise dominating citric and oxalacetic acid decreased.     

饼肥中的有机营养物质及其在发酵过程中的变化

分析测定了在恒温、恒湿条件下 4种植物油饼发酵过程中有机物质的变化。结果表明 ,由于不同植物油饼化学组成不同在发酵过程中所产生有机营养物质的成分和数量存在较大差异。发酵过程所产生的高级脂肪酸以C18、C16酸为主 ,并且不饱和脂肪酸的含量明显高于饱和脂肪酸 ;有机酸则以乳酸、苹果酸、柠檬酸、琥珀酸、酒石酸、乙酸含量较高 ,草酸含量较低 ;氨基酸则以谷氨酸含量最高 ,天门冬氨酸、亮氨酸和精氨酸含量次之。有机酸和高级脂肪酸含量随发酵时间的增加而增加 ,腐熟到一定程度时达到最高 ;而饼肥发酵 70d时各种氨基酸含量明显降低。 4种饼肥相比 ,豆饼和花生饼在发酵过程中氨基酸产生量大于芝麻饼和菜籽饼。从速度来看 ,芝麻饼腐熟最快 ,其次是豆饼 ,再次是花生饼 ,菜籽饼腐熟最慢。     

Influence of low-molecular-weight organic acids on the solubilization of phosphates

A range of low-molecular-weight organic acids were identified in rhizosphere soil, leaf litter, and poultry manure compost. Laboratory and greenhouse experiments were carried out to examine the effects of seven low-molecular-weight organic acids on phosphate adsorption by soils, and the solubilization and plant uptake of P from soil pre-incubated with monocalcium phosphate and North Carolina phosphate rock. Acetic, formic, lactic (monocarboxylic), malic, tartaric, oxalic (dicarboxylic), and citric (tricarboxylic) acids were used in the study. The addition of organic acids decreased the adsorption of P by soils in the order tricarboxylic acid>dicarboxylic acid>monocarboxylic acid. The decreases in P adsorption with organic acid addition increased with an increase in the stability constant of the organic acid for Al (logK _Al). Organic acids extracted greater amounts of P from soils meubated with both monocalcium phosphate and phosphate rock than water did. Although more phosphate was extracted by the organic acids from monocalcium phosphate — than from phosphate rock — treated soils in absolute terms, when the results were expressed as a percentage of dissolved phosphate there was little difference between the two fertilizers. The amount of P extracted by the organic acids from both fertilizers increased with an increase in logK _Al values. The addition of oxalic and citric acids increased the dry matter yield of ryegrass and the uptake of P in soils treated with both fertilizers. The agronomic effectiveness of both fertilizers increased in the presence of organic acids and the increase was greater with the phosphate rock than with the monocalcium phosphate. The results indicated that organic acids increase the availability of P in soils mainly through both decreased adsorption of P and increased solubilization of P compounds.     

Dinitrogen fixation and growth responses to phosphorus and aluminum application in pigeon. pea (Cajanus cajan L.)

Abstract

Responses to aluminum phosphate (Al-P) and superphosphate (Sup-P) application in terms of dry matter production by pigeon pea and soybean were compared under field conditions. The effect of aluminum on growth and symbiotic dinitrogen fixation was examined under hydroponic conditions. In a field experiment, Al-P application significantly increased whole plant dry weight and P amount in pigeon pea, whereas in soybean, Al-P application had no significant effect on the two parameters. The results indicate that P absorption ability from Al-P is higher in pigeon pea compared to soybean. With Al-P application, leaf and whole plant Al concentration in pigeon pea significantly increased unlike in soybean.

Under hydroponic conditions, 50 nig L^?1 Al application increased the plant weight of pigeon pea while it decreased that of soybean. This treatment decreased the total nodule activity of soybean more appreciably than that of pigeon pea. Under the Al treatment conditions, whole plant P amount increased in pigeon pea, whereas it decreased in soybean, and the amount of Al increased more appreciably in pigeon pea, most of which was deposited in roots.

These results suggest that compared to soybean, pigeon pea displays a higher ability to absorb P from P compounds with a low solubility i.e. Al-P and that it is more tolerant to Al toxicity.     

Increase in the permeability of tonoplast of garlic (Allium sativum) by monocarboxylic acids

Immersion of intact aged garlic (Allium sativum) cloves in a series of 5% weak organic monocarboxylate solutions (pH 2.0) resulted in green color formation. No color was formed upon treatment with other weak organic acids, such as citric and malic acids, and the inorganic hydrochloric acid under the same conditions. To understand the significance of monocarboxylic acids and their differing function from that of other acids, acetic acid was compared with organic acids citric and malic and the inorganic hydrochloric acid. The effects of these acids on the permeability of plasma and intracellular membrane of garlic cells were measured by conductivity, light microscopy, and transmission electron microscopy. Except for hydrochloric acid, treatment of garlic with all three organic acids greatly increased the relative conductivity of their respective pickling solutions, indicating that all tested organic acids increased the permeability of plasma membrane. Moreover, a pickling solution containing acetic acid exhibited 1.5-fold higher relative conductivity (approximately 90%) as compared to those (approximately 60%) of both citric and malic acids, implying that exposure of garlic cloves to acetic acid not only changed the permeability of the plasma membrane but also increased the permeability of intracellular membrane. Exposure of garlic to acetic acid led to the production of precipitate along the tonoplast, but no precipitate was formed by citric and malic acids. This indicates that the structure of the tonoplast was damaged by this treatment. Further support for this conclusion comes from results showing that the concentration of thiosulfinates [which are produced only by catalytic conversion of S-alk(en)yl-l-cysteine sulfoxides in cytosol by alliinase located in the vacuole] in the acetic acid pickling solution is 1.3 mg/mL, but almost no thiosulfinates were detected in the pickling solution of citric and malic acids. Thus, all present results suggest that damage of tonoplast by treatment with monocarboxylates such as acetic acid may be the main reason for the greening of garlic.     

添加有机酸对土壤镉形态转化及苋菜镉积累的影响

植物根系分泌的低分子量有机酸能够与土壤中的镉形成镉–有机酸复合体,从而影响根际镉的移动性。本文通过添加有机酸对土壤镉形态转化的研究,阐明有机酸与镉生物积累的关系。采用盆栽试验及土壤培养等方法,研究了添加苹果酸、柠檬酸对赤红壤和黄棕壤中镉的形态转化以及超积累型苋菜天星米镉生物积累的影响。结果表明,与Cd 25 mg/kg处理比较,Cd 25 mg/kg＋苹果酸、Cd 25mg/kg＋柠檬酸处理对苋菜生物量未产生影响,但显著增加苋菜根系及地上部镉含量;添加苹果酸、柠檬酸处理显著降低土壤专性吸附态Cd含量,却显著增加了交换态Cd、碳酸盐结合态Cd和有机结合态Cd含量。说明添加苹果酸、柠檬酸还能够通过影响土壤镉形态转化而促进苋菜对镉的积累。     

Hormonal interactions in the rhizosphere of maize (Zea mays L.) and their effects on plant development

The phytohormones indole acetic acid (IAA), abscisic acid (ABA), isopentenyladenosine (iPA), dihydrozeatin riboside (DHZR), and zeatinriboside (ZR) were determined quantitatively using monoclonal antibodies by an enzyme immunoassay in the following samples: a) culture filtrates of known bacterial species, growing naturally in close, loose or without contact with higher plants; b) culture filtrates of heterogeneous populations of microorganisms, isolated from the rhizoplane and rhizosphere of maize as well as from root-free soil; c) sterile and nonsterile maize root exudates; d) in the rhizosphere of field-grown maize plants and in soil fractions distant from the roots (bulk soil). ABA was not detected in the culture filtrates of bacteria and ZR was not found in bulk soil and rhizosphere soil of field-grown maize. All phytohormones were present in the other samples analyzed. Bacterial cultures with cell concentrations roughly equal to those in the rhizosphere of field-grown maize may produce under optimal laboratory conditions phytohormone concentrations comparable to those measured in the rhizosphere of field-grown maize. During the whole vegetation period there was a steep phytohormone gradient in the first centimeter of soil around a maize root in the field. Inoculation of maize seedlings growing in nutrient solution with rhizosphere bacteria resulted in a synergistic increase in phytohormone concentration in comparison to the sum of hormone production by sterile plant roots and by bacteria cultures. Using regression analysis, the relationship between phytohormone concentration changes in the rhizosphere and different morphological characteristics of the maize plant were shown to be highly significant.