Growth, P uptake and rhizosphere properties of intercropped wheat and chickpea in soil amended with iron phosphate or phytate

Intercropping has been shown to increase total yield and nutrient uptake compared to monocropping. However, depending on crop combinations, one crop may dominate and decrease the growth of the other. Interactions in the soil, especially in the rhizosphere, may be important in the interactions between intercropped plant genotypes. To assess the role of the rhizosphere interactions, we intercropped a P-inefficient wheat genotype (Janz) with either the P-efficient wheat genotype (Goldmark) or chickpea in a soil with low P availability amended with 100 mg P kg⁻¹ as FePO₄ (FeP) or phytate. The plants were grown for 10 weeks in pots where the roots of the genotypes could intermingle (no barrier, NB), were separated by a 30 μm mesh (mesh barrier, MB), preventing direct root contact but allowing exchange of diffusible compounds and microorganisms, or were completely separated by a solid barrier (SB). When supplied with FeP, Janz intercropped with chickpea had higher shoot and grain dry weight (dw) and greater plant P uptake in NB and MB than in SB. Contact with roots of Janz increased shoot, grain and root dw, root length, shoot P concentration and shoot P uptake of chickpea compared to SB. Root contact between the two wheat genotypes, Janz and Goldmark, had no effect on growth and P uptake of Janz. Shoot and total P uptake by Goldmark were significantly increased in NB compared to MB or SB. In both crop combinations, root contact significantly increased total plant dw and P uptake per pot. Plant growth and P uptake were lower with phytate and not significantly affected by barrier treatment. Differences in microbial P, available P and phosphatase activity in the rhizosphere among genotypes and barrier treatments were generally small. Root contact changed microbial community structure (assessed by fatty acid methyl ester (FAME) analysis) and all crops had similar rhizosphere microbial community structure when their roots intermingled.     

Brassica genotypes differ in growth, phosphorus uptake and rhizosphere properties under P-limiting conditions

Compared to other crops, Brassicas are generally considered to grow well in soils with low P availability, however, little is known about genotypic differences within Brassicas in this respect. To assess the role of rhizosphere properties in growth and P uptake by Brassicas, three Brassica genotypes (mustard, Brassica juncea cv Chinese greens and canola, Brassica napus cvs Drum and Outback) were grown in an acidic soil with low P availability at two treatments of added P: 25 and 100 mg P kg⁻¹ as FePO₄ (P25 and P100). The plants were harvested at the 6-leaf stage, at flowering and at maturity. Shoot and root dry weight (dry weight) and root length increased with time and were lower in P25 than in P100. In P25, shoot dry weight was lowest in Outback and highest in Chinese greens. In the P100 treatment, Chinese greens had a higher shoot dry weight than the two canola cultivars. Chinese greens had a lower root dry weight and root length at flowering and maturity than the canola genotypes in both P treatments. Irrespective of P treatment, shoot P concentration was lower in Chinese greens than in the two canola genotypes. Specific P uptake (μg P m⁻¹ root length) decreased with time. In P25, Chinese greens had the lowest specific P uptake at the 6-leaf stage but it was higher than in the two canola genotypes at flowering and maturity. In P100, Outback had the lowest specific P uptake. Available P in the rhizosphere (resin P) decreased over time with the greatest decrease from the 6-leaf stage to flowering. In P25, resin P in the rhizosphere was greatest in Chinese greens at the 6-leaf stage and flowering and smallest in Outback at flowering. Microbial P and acid phosphatase activity changed little over time, were not affected by P treatment and there were only small differences between the genotypes. The rhizosphere microbial community composition [assessed by fatty acid methyl ester (FAME) analysis] of Outback and Chinese greens differed from that of the other two genotypes at the 6-leaf stage and flowering, respectively. At maturity, all three genotypes had distinct microbial communities. Plant traits such as production of high biomass at low shoot P concentrations as well as the capacity to maintain high P availability in the rhizosphere by P mobilisation can explain the observed differences in plant growth and P uptake among the Brassica genotypes.     

Growth,phosphorus uptake,and rhizosphere microbial‐community composition of a phosphorus‐efficient wheat cultivar in soils differing in pH

In a pot experiment, the P‐efficient wheat (Triticum aestivum L.) cultivar Goldmark was grown in ten soils from South Australia covering a wide range of pH (four acidic, two neutral, and four alkaline soils) with low to moderate P availability. Phosphorus (100 mg P kg^–1) was supplied as FePO₄ to acidic soils, CaHPO₄ to alkaline, and 1:1 mixture of FePO₄ and CaHPO₄ to neutral soils. Phosphorus uptake was correlated with P availability measured by anion‐exchange resin and microbial biomass P in the rhizosphere. Growth and P uptake were best in the neutral soils, lower in the acidic, and poorest in the alkaline soils. The good growth in the neutral soils could be explained by a combination of extensive soil exploitation by the roots and high phosphatase activity in the rhizosphere, indicating microbial facilitation of organic‐P mineralization. The plant effect (soil exploitation by roots) appeared to dominate in the acidic soils. Alkaline phosphatase and diesterase activities in acidic soils were lower than in neutral soils, but strongly increased in the rhizosphere compared with the bulk soil, suggesting that microorganisms contribute to P uptake in these acidic soils. Shoot and root growth and P uptake per unit root length were lowest in the alkaline soils. Despite high alkaline phosphatase and diesterase activities in the alkaline soils, microbial biomass P was low, suggesting that the enzymes could not mineralize sufficient organic P to meet the demands of plants and microorganisms. Microbial‐community composition, assessed by fatty acid methylester (FAME) analysis, was strongly dependent on soil pH, whereas other soil properties (organic‐C or CaCO₃ content) were less important or not important at all (soil texture).     

Effect of arbuscular mycorrhizal fungi on rhizosphere organic acid content and microbial activity of trifoliate orange under different low P conditions

ABSTRACT

Arbuscular mycorrhizal (AM) fungi can improve plant phosphorus (P) uptake; however, information about how AM fungi affect rhizosphere organic acid and microbial activity to alleviate citrus low P stress is limited. Here, a pot experiment was conducted to evaluate the effect of AM fungi (Rhizophagus intraradices, Ri) inoculation on rhizosphere organic acid content, microbial biomass (MB) and enzyme activity of trifoliate orange (Poncirus trifoliata L. Raf.) seedlings grown under three low P conditions. The results showed that mycorrhizal seedlings all recorded higher P concentrations, plant biomass and better root morphology with more lateral and fine roots, but lower root mass ratios, irrespective of P conditions. Mycorrhizal P absorption contribution did not differ significantly among three P conditions. Mycorrhizal seedling rhizosphere soil exhibited lower organic acid content, soil organic P content and ratio of MB-carbon (C)/MB-P, but higher MB and enzyme activity. Additionally, the main organic acids showed a negative relationship with mycorrhizal colonization rate and hyphal length; however, phosphatase and phytase activity had a significantly positive relationship with MB. Therefore, the results suggest that AM fungi inoculation may help citrus to efficiently utilize organic P source by improving microbial activity under low available P conditions.     

The effect of root hairs on rhizosphere phosphatase activity

Background: Phosphatases in soil are of great importance for plant P acquisition. It is hypothesized that root hairs increase rhizosphere phosphatase activity as they release enzymes into soil and stimulate microbial activity. Methods: To test the effect of root hairs on soil phosphatase activity, we grew barley (Hordeum vulgare ‘Pallas') wild type and its root‐hairless mutant in rhizoboxes and determined phosphatase activity using soil zymography. Measurements were done at three moisture levels (30, 15, and 5% VWC). Rhizosphere phosphatase activity was estimated for the two genotypes and two locations along the root [root tip region (0–4 cm behind tip) and mature roots (> 7 cm behind tip)]. Results: Rhizosphere phosphatase activity was similar in the two locations along the root (root tip region vs. mature root parts). In contrast, rhizosphere phosphatase extension was two times larger for the root tip region of the wild type than for the mutant at 30% and 15% VWC. However, as phosphatase activities at the root surface of tips and mature root parts were slightly higher for the mutant than for the wild type, average enzyme activities were unaffected by the genotype. Conclusions: We conclude that the mutant seems to compensate for the lack of root hairs by increased phosphatase activity close to the root surface. However, the increased rhizosphere phosphatase extension for the wild type may be equally efficient as it allows P mobilization and uptake from large soil volumes.     

Phosphorus Efficiency of Winter Canola Cultivars and Rhizosphere Properties in Rhizobox Technique

Production of oilseed crops requires balanced fertilization, while environmental problems of applied fertilizers must be considered. We evaluated seven winter canola cultivars (Brassica napus L.) for their relative efficiency to use or acquire phosphorus (P) under deficient and sufficient conditions. Average root mean diameter (RMD), total root length (RL) and root surface area (RA), of plants were measured as well as rhizosphere properties in rhizobox technique. Water-soluble P (WSP) and phosphatase activity of treatments containing plant were higher than control. WSP increased by 2.86 mg kg^?1 soil in Gabriela and 2.63 mg kg^?1 soil in Elvis at P deficient condition, compared to the control soil. The Olsen extractable P of the treatments decreased compared to control. Variations in total dry weights of cultivars were mainly explained by the differences in P solubilizing bacteria (PSB) population, pH, and phosphatase activity. The responsible mechanism for the P efficient (PE) cultivars can be higher P uptake through larger root and changes in rhizosphere properties.     

Rhizosphere dynamics under nitrogen‐induced root modification: The interaction of phosphorus and calcium

Optimizing root phosphorus (P) acquisition to reduce intensive fertilizer use is a crucial pathway for sustainable agriculture, particularly as P is an important plant macronutrient, often limiting in a majority of soils worldwide. Although many studies have assessed plant growth and P acquisition, few studies have investigated the interactive effects of nitrogen (N)‐induced root modification on soil P processes or the understudied effects of soil calcium (Ca) dynamics on soil P bioavailability. In this study, we investigate soil P and Ca response in the rhizosphere of durum wheat (Triticum turgidum L. spp. durum). Wheat grown under controlled conditions preloaded for 20 d with two N treatments [preloaded low N (1 mmol KNO₃ plant^?1) and preloaded high N (2 mmol KNO₃ plant^?1)] were transferred to rhizoboxes for 12 d [days after transfer (DAT)]. Shoot and root biomass, P and Ca concentration, and plant‐available P and extractable Ca were determined every three days (0, 3, 6, 9, 12 DAT). Significantly higher root mass (P = 0.7%), root length (P = 1.8%) and total biomass (P = 2.2%) were found at the end of the experiment but exclusively for high N preloaded wheat. This greater root biomass was associated with lower root P concentration, suggesting a dilution response, while little difference was observed in shoot P concentration over the 12 d. However, Ca accumulated in both roots and shoots under both preloading N levels. Concurrently, soil‐extractable Ca declined, and plant‐available P increased (r = –0.62; P = 0.03%), presumably due to a promoting effect of Ca uptake on soil P availability; lower soil Ca in turn increased the repulsive forces between P ions and the negatively charged soil surface, resulting in an increased P availability in the soil solution. This study contributes to the understanding of the complex interplay between multi‐nutrient dynamics within the rhizosphere.     

Belowground interactions between intercropped wheat and Brassicas in acidic and alkaline soils

Our previous studies showed that, under P-limiting conditions, growth and P uptake were lower in the wheat genotype Janz than in three Brassica genotypes when grown in monoculture. The present study was conducted to answer the question if P mobilised by the Brassicas is available to wheat; leading to improved growth of wheat when intercropped with Brassicas compared to monocropped wheat. To assess if the interactions between the crops depend on soil type, the wheat genotype Janz and three Brassica genotypes (two canolas and one mustard) were grown for 6 weeks in monoculture or wheat intercropped with each Brassica genotype in an acidic and an alkaline soil with low P availability (with two plants per pot). Wheat grew equally well in the two soils, but the Brassicas grew better in the acidic than in the alkaline soil. In the acidic soil, monocropped Brassicas had a 3 to 4 fold greater plant dry weight (dw) and P uptake than wheat; plant dw and P uptake in wheat were decreased or not affected by intercropping and increased in the Brassicas. In the alkaline soil, dw and P uptake of the Brassicas was twice as high as in wheat, with intercropping having no effect on these parameters. The contribution of wheat to the total shoot dw and P uptake per pot was 4-21% and 32-40% in acidic and alkaline soil, respectively. Mycorrhizal colonisation was low in all genotypes in the acidic soil (1-6%). In the alkaline soil, mycorrhizal colonisation of monocropped wheat was 62%, but only 43-47% in intercropped wheat. Intercropping decreased P availability in the rhizosphere of wheat in the acidic soil but had no effect on rhizosphere P availability in the alkaline soil. Intercropping had a variable effect on rhizosphere microbial community composition (assessed by fatty acid methylester analysis (FAME) and ribosomal intergenic spacer amplification (RISA)), ranging from intercropping having no effect on the rhizosphere communities to intercropping resulting in a new and similar rhizosphere community composition in both genotypes. The results of this study show that intercropping with Brassicas does not improve growth and P uptake of wheat; thus there is no indication that P mobilised by the Brassicas is available to wheat.     

不同磷供应水平下小麦根系形态及根际过程的变化特征

以石麦15和衡观35两个品种小麦为试验材料,应用根袋栽培方式,研究了不同施磷量对小麦根系形态和根际特征的影响。结果表明,与施磷量P2O5 0.1 g/kg相比,高量供磷（P2O5 0.3 g/kg）条件下石麦15地上部生物量和磷累积量增加幅度大于衡观35;但不施磷处理衡观35地上部生物量降低幅度小于石麦15,磷含量和累积量高于石麦15,衡观35耐低磷能力较强。土壤供磷不足时,衡观35总根长中直径0.16 mm细根所占比例高于石麦15,根系平均直径较小;而高磷供应下,石麦15根系中直径0.16 mm细根长度较长,在总根长中所占比例较高。总根长和直径0.16 mm的细根长度与植株地上部磷累积量之间呈显著正相关关系。总根长越长尤其是细根越多,有利于促进植株对磷的吸收。与非根际土壤相比,高磷供应下根际土壤有机磷含量增加,微生物量磷含量降低;而供磷不足时根际土壤碱性磷酸酶活性较高,有机磷含量较低。与施磷量P2O5 0.1 g/kg相比,高量供磷下根际土壤pH值升高、碱性磷酸酶活性下降,不施磷处理根际土壤pH值降低。本研究表明,供磷不足时,小麦根系形态和根际过程均发生适应性变化,而高量供磷条件下,小麦植株根系形态的改变因品种而异。     

Phosphatversorgung von Sommerweizen (Triticum aestivum L.) in Mischkultur mit Weißer Lupine (Lupinus albus L.)

Phosphorus nutrition of spring wheat (Triticum aestivum L.) in mixed culture with white lupin (Lupinus albus L.). Spring wheat (Triticum aestivum L. ?Schirokko”?) and white lupin (Lupinus albus L.) were grown in mixed culture in Mitscherlich pots with 20 kg of soil in a green house. The soil used was a B_t of a Parabraunerde-Pseudogley from loess low in available P and limed from pH 4.6 to pH 6.5. Phosphorus was added as phosphate rock. In half of the pots cylinders of stainless steel screen prevented intertwining of the roots of the plant species. Independent of P addition, white lupin had higher dry matter production and P uptake than wheat, even although wheat had thinner roots and higher root densities than lupin, factors which favour the utilization of soil and fertilizer P. The higher P efficiency of white lupin was due to higher P uptake rates per unit root length mainly through mobilization of P especially in the rhizosphere of the proteoid roots. When the roots of the two species were allowed to intertwine, shoot dry matter production of wheat was nearly double because of improved tillering. Higher P concentrations and a more than 2-fold higher P uptake indicated that the increase in dry matter production of wheat was due to improved P nutrition. Nitrogen concentrations, however, remained unaffected at sufficient levels. An increased P uptake rate per unit root length was responsible for the better utilization of P by wheat, rather than the increase in total root length, due to the extended root volume. White lupin was able to mobilize P in the rhizosphere in excess of its own requirements. Thus mobilized P may be available to less P-efficient plants grown in mixed culture.     

Crop species differ in root plasticity response to localised P supply

The effect of localised phosphorus (P) fertiliser placement and in particular, deep P fertiliser placement, on the comparative root growth and P uptake of fibrous vs tap‐rooted crops is not known. In this study, we examined the root growth and P uptake of wheat (Triticum aestivum L.), canola (Brassica napus L.), and narrow‐leaf lupin (Lupinus angustifolius L.) in a split‐root system and in columns with deep (19 cm) or shallow (5 cm) P fertiliser sources in glasshouse conditions. In the split‐root system, plants of all three species grown under heterogeneous soil P conditions absorbed more P and produced greater root and shoot biomass than those under homogeneous P supply. Root plasticity differed between species under heterogeneous soil P supply: canola and wheat allocated relatively more root biomass and root length to the high P zone than narrow‐leaf lupin. In the column experiment, there was no difference in the amount of P accumulated in shoots of any crops grown in the deep vs shallow P fertiliser treatments. Root proliferation occurred within the shallow and deep‐P fertiliser bands in all three species; however, root distribution above or below the bands did not differ between deep or shallow P fertiliser treatments in any species. Whilst root plasticity responses to heterogeneous soil P supply differed among species, root architecture (fibrous vs taproot) did not confer any advantage or disadvantage to the acquisition of P from deep vs shallow P fertiliser bands. Moreover, whilst roots proliferate in the vicinity of P fertiliser bands, root distribution outside of the bands appears to remain unaltered in both fibrous and tap‐rooted crops during early growth.     

两种磷效率小麦在不同土壤上根际特征的差异

以磷高效型小麦小偃54和磷低效型小麦京411为研究对象设计三室根箱试验,通过测定生物量、吸磷量、pH和酸性磷酸酶,对比两种小麦在不同磷水平(P0 mg/kg土、100 mg/kg土)及两种不同土壤上(石灰性黑垆土、酸性红壤)根际特征的差异。试验结果表明,黑垆土上,增施磷肥使小偃54的总生物量增加了14.99%,京411增加了26.53%,总吸磷量二者分别增加了99.29%和83.70%;红壤的速效磷含量高,施肥仅提高了磷低效型小麦京411的生物量。黑垆土上磷胁迫并未造成小偃54与京411各部分生物量和吸磷量的显著性差异,但小偃54的根际pH降低值和酸性磷酸酶的活性却已显著高于京411;P0处理时,红壤上小偃54的地上部和总生物量显著高于京411,虽然红壤的速效磷含量高于黑垆土,但在P0处理时两种小麦在两种土壤上的生物量和吸磷量并无显著性差异。就根际分泌物而言,石灰性黑垆土上,小偃54的根系在低磷胁迫下通过降低pH和分泌酸性磷酸酶来活化土壤中难溶态的磷,而在红壤上小偃54的pH和酸性磷酸酶的活性保持稳定。酸性红壤中两种小麦酸性磷酸酶活性显著高于石灰性黑垆土。由此可见,两种磷效率小麦在两种不同性质土壤上活化机理存在差异。     

Nitrogen and phosphorus transformations in the rhizospheres of three tree species in a nutrient-poor sandy soil

We examined acid phosphatase activity (APA), N mineralization and nitrification rates, available N and P, and microbial biomass C, N and P in rhizosphere and bulk soils of 18-year-old Siberian elm (Ulmus pumila), Simon poplar (Populus simonii) and Mongolian pine (Pinus sylvestris var. mongolica) plantations on a nutrient-poor sandy soil in Northeast China. The main objective was to compare the rhizosphere effects of different tree species on N and P cycling under nutrient-deficient conditions. All tree species had the similar pattern but considerably different magnitude of rhizosphere effects. The APA, potential net N mineralization and nitrification rates increased significantly (by 27–60%, 110–188% and 106–142% respectively across the three species) in rhizosphere soil compared to bulk soil. This led to significantly higher Olsen-P and NH₄⁺-N concentrations in rhizosphere soil, whereas NO₃⁻-N concentration was significantly lower in rhizosphere soil owing to increased microbial immobilization and root uptake. Microbial biomass C and N generally increased while microbial biomass P remained constant in rhizosphere soil relative to bulk soil, indicating the N-limited rather than P-limited microbial growth. Rhizosphere effects on P transformation were most pronounced for Siberian elm, while rhizosphere effects on N transformation were most pronounced for Mongolian pine, implying the different capacities of these species to acquire nutrients.     

Microscale distribution and function of soil microorganisms in the interface between rhizosphere and detritusphere

The rhizosphere and the detritusphere are hot spots of microbial activity, but little is known about the interface between rhizosphere and detritusphere. We used a three-compartment pot design to study microbial community structure and enzyme activity in this interface. All three compartments were filled with soil from a long-term field trial. The two outer compartments were planted with maize (root compartment) or amended with mature wheat shoot residues from a free air CO₂ enrichment experiment (residue compartment) and were separated by a 50 μm mesh from the inner compartment. Soil, residues and maize differed in ¹³C signature (δ¹³C soil −26.5‰, maize roots −14.1‰ and wheat residues −44.1‰) which allowed tracking of root- and residue-derived C into microbial phospholipid fatty acids (PLFA). The abundance of bacterial and fungal PLFAs showed clear gradients with highest abundance in the first 1–2 mm of the root and residue compartment, and generally higher values in the vicinity of the residue compartment. The δ¹³C of the PLFAs indicated that soil microorganisms incorporated more carbon from the residues than from the rhizodeposits and that the microbial use of wheat residue carbon was restricted to 1 mm from the residue compartment. Carbon incorporation into soil microorganisms in the interface was accompanied by strong microbial N immobilisation evident from the depletion of inorganic N in the rhizosphere and detritusphere. Extracellular enzyme activities involved in the degradation of organic C, N and P compounds (β-glucosidase, xylosidase, acid phosphatase and leucin peptidase) did not show distinct gradients in rhizosphere or detritusphere. Our microscale study showed that rhizosphere and detritusphere differentially influenced microbial C cycling and that the zone of influence depended on the parameter assessed. These results are highly relevant for defining the size of different microbial hot spots and understanding microbial ecology in soils.     

Phosphorus uptake and rhizosphere properties of intercropped and monocropped maize, faba bean, and white lupin in acidic soil

Little information is available on phosphorus (P) uptake and rhizosphere processes in maize (Zea mays L.), faba bean (Vicia faba L.), and white lupin (Lupinus albus L.) when intercropped or grown alone in acidic soil. We studied P uptake and soil pH, carboxylate concentration, and microbial community structure in the rhizosphere of maize, faba bean, and white lupin in an acidic soil with 0–250 mg P (kg⁻¹ soil) as KH₂PO₄ (KP) or FePO₄ (FeP) with species grown alone or intercropped. All plant species increased the pH compared to unplanted control, particularly faba bean. High KP supply (>100 mg P kg⁻¹) significantly increased carboxylate concentration in the rhizosphere of maize. The carboxylate composition of the rhizosphere soil of maize and white lupin was significantly affected by P form (KP or FeP), whereas, this was not the case for faba bean. In maize, the carboxylate composition of the rhizosphere soil differed significantly between intercropping and monocropping. Yield and P uptake were similar in monocropping and intercropping. Monocropped faba bean had a greater concentration of phospholipid fatty acids in the rhizosphere than that in intercropping. Intercropping changed the microbial community structure in faba bean but not in the other corps. The results show that P supply and P form, as well as intercropping can affect carboxylate concentration and microbial community composition in the rhizosphere, but that the effect is plant species-specific. In contrast to previous studies in alkaline soils, intercropping of maize with legumes did not result in increased maize growth suggesting that the legumes did not increase P availability to maize in this acidic soil.     

Distribution of phosphatase activity and various bacterial phyla in the rhizosphere of Hordeum vulgare L. depending on P availability

Despite the importance of the rhizosphere for nutrient turnover, little is known about the spatial patterns of organic phosphorus mineralization by plants and by microorganisms in the rhizosphere. Therefore, the distribution of acid and alkaline phosphatase activity and the abundance of bacteria belonging to various bacterial phyla were investigated in the rhizosphere of barley (Hordeum vulgare L.) as dependent on the availability of inorganic P. For this purpose, we conducted a greenhouse experiment with barley growing in inclined boxes that can be opened to the bottom side (rhizoboxes), and applied soil zymography and fluorescence-in situ-hybridization (FISH). Acid phosphatase activity was strongly associated with the root and was highest at the root tips. Due to P fertilization, acid phosphatase activity decreased in the bulk soil, and less strongly in the rhizosphere. Alkaline phosphatase activity, i.e., microbial phosphatase activity was high throughout the soil in the control treatment and was reduced due to inorganic P fertilization especially in the rhizosphere and less strongly in the bulk soil. P-fertilization slightly increased the total number of bacteria in the rhizosphere. Moreover, P-fertilization decreased the abundance of Firmicutes and increased the abundances of Beta- and Gamma-Proteobacteria. The total number of bacterial cells was significantly higher at the root surface than at the root tip and at a distance of 30 μm from the root surface. Our results show that alkaline phosphatase activity decreased more strongly in the rhizosphere than in the bulk soil due to P fertilization, which might be because of greater C deficiency in the bulk soil compared to the rhizosphere. Furthermore, the results indicate a spatial separation between hotspots of acid phosphatase activity and hotspots of bacteria in the rhizosphere of H. vulgare. Taken together, our study shows that bacteria and phosphatase activity were very heterogeneously distributed in soil, and that the effects of P fertilization on phosphatase activity differed strongly between bulk soil and rhizosphere as well as between various zones of the rhizosphere.     

Root-induced changes in the rhizosphere: Importance for the mineral nutrition of plants

Root-induced changes in the rhizosphere may affect mineral nutrition of plants in various ways. Examples for this are changes in rhizosphere pH in response to the source of nitrogen (NH₄-N versus NO₃-N), and iron and phosphorus deficiency. These pH changes can readily be demonstrated by infiltration of the soil with agar containing a pH indicator. The rhizosphere pH may be as much as 2 units higher or lower than the pH of the bulk soil. Also along the roots distinct differences in rhizosphere pH exist. In response to iron deficiency most plant species in their apical root zones increase the rate of H⁺ net excretion (acidification), the reducing capacity, the rate of Fe^III reduction and iron uptake. Also manganese reduction and uptake is increased several-fold, leading to high manganese concentrations in iron deficient plants. Low-molecular-weight root exudates may enhance mobilization of mineral nutrients in the rhizosphere. In response to iron deficiency, roots of grass species release non-proteinogenic amino acids (?phytosiderophores”?) which dissolve inorganic iron compounds by chelation of Fe^III and also mediate the plasma membrane transport of this chelated iron into the roots. A particular mechanism of mobilization of phosphorus in the rhizosphere exists in white lupin (Lupinus albus L.). In this species, phosphorus deficiency induces the formation of so-called proteoid roots. In these root zones sparingly soluble iron and aluminium phosphates are mobilized by the exudation of chelating substances (probably citrate), net excretion of H⁺ and increase in the reducing capacity. In mixed culture with white lupin, phosphorus uptake per unit root length of wheat (Triticum aestivum L.) plants from a soil low in available P is increased, indicating that wheat can take up phosphorus mobilized in the proteoid root zones of lupin. At the rhizoplane and in the root (root homogenates) of several plant species grown in different soils, of the total number of bacteria less than 1 % are N₂-fixing (diazotrophe) bacteria, mainly Enterobacter and Klebsiella. The proportion of the diazotroph bacteria is higher in the rhizosphere soil. This discrimination of diazotroph bacteria in the rhizosphere is increased with foliar application of combined nitrogen. Inoculation with the diazotroph bacteria Azospirillum increases root length and enhances formation of lateral roots and root hairs similarly as does application of auxin (IAA). Thus rhizosphere bacteria such as Azospirillum may affect mineral nutrition and plant growth indirectly rather than by supply of nitrogen.     

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.     

碳对微生物–根系介导的蔬菜作物磷吸收的影响

  【目的】  碳是微生物代谢活动的能量来源,解析碳驱动的微生物磷周转对根系/根际属性以及作物磷吸收的影响,对探索提高磷利用效率的根际调控措施具有重要的指导意义。  【方法】  以绿叶蔬菜上海青(Brassica chinensis L., Xiaqing 3)为供试作物进行盆栽试验,供试碳源为葡萄糖。设置添加葡萄糖(+G)和不添加葡萄糖(?G,对照)两个处理,在添加葡萄糖后第7天和第21天,测定土壤微生物量磷与Olsen-P含量、根际酸性磷酸酶活性以及柠檬酸和苹果酸含量、根系形态(生物量、根冠比、根长、根系直径、比根长和根系组织密度)与根际生理(酸性磷酸酶、柠檬酸和苹果酸)指标和作物磷吸收量。  【结果】  添加葡萄糖后第7天,土壤微生物量磷增加,Olsen-P含量降低;上海青根系生物量和根冠比显著高于对照,另外,与不加葡萄糖处理相比,添加葡萄糖导致上海青总根长降低33%,根系平均直径增加27%,比根长降低46%,根际柠檬酸含量增加106%。从第7天到第21天,添加葡萄糖处理土壤微生物量磷降低,Olsen-P含量增加,上海青根系生长速率显著提高。葡萄糖添加后第21天,添加葡萄糖处理土壤Olsen-P含量高于对照土壤;与不加葡萄糖的处理相比,根际酸性磷酸酶和柠檬酸的分泌降低,上海青根系总根长增加,其相对增加量为31%。添加葡萄糖对第7天和第21天上海青地上部磷吸收没有显著影响。  【结论】  添加葡萄糖提高了前期(添加葡萄糖后第7天)根际微生物量磷,降低了Olsen-P含量,促进根际柠檬酸的分泌满足作物生长对磷的需求。后期(添加葡萄糖后第21天),微生物量磷的降低促进土壤有效磷含量的增加,刺激根系快速伸长。微生物介导磷周转诱导作物调节根系形态和根际分泌物响应土壤磷环境的变化,维持地上部磷营养。     

Growth and uptake rates of P,K, Ca and Mg in wheat 1

The relation between plant age and nutrient absorption properties of red winter wheat (Triticum aestivum L.) roots were investigated. Understanding the change in ion uptake parameters with increasing plant age is helpful in devising efficient fertilization systems. Such information can be used to determine the nutrient levels needed in the soil to supply nutrients rapidly enough to the root surface to minimize deficiencies. Wheat was grown for periods up to 40 days in solution culture in a controlled climate chamber. Sequential harvest and nutrient influx measurements were made. Shoot growth was exponential with time to 30 days and linear thereafter. Root dry weight increased linearly with time at a slower rate than shoot dry weight. Root length increased linearily with time. With increasing plant age there was a reduction in average P and K uptake rate while average uptake rates for Ca and Mg remained relatively unchanged. With increasing plant age, the maximum influx, I_max. for P and Mg remained constant, but for K and Ca, there was a decrease. For the Michael is constant, Km, no change was observed for P, an increase occurred for K, and a decrease for Ca and Mg, as the wheat plant grew from 5 to 40 days.