Showing papers in "Plant and Soil in 2001"

Bioavailability of soil inorganic P in the rhizosphere as affected by root-induced chemical changes: A review

Abstract: In most soils, inorganic phosphorus occurs at fairly low concentrations in the soil solution whilst a large proportion of it is more or less strongly held by diverse soil minerals. Phosphate ions can indeed be adsorbed onto positively charged minerals such as Fe and Al oxides. Phosphate (P) ions can also form a range of minerals in combination with metals such as Ca, Fe and Al. These adsorption/desorption and precipitation/dissolution equilibria control the concentration of P in the soil solution and, thereby, both its chemical mobility and bioavailability. Apart from the concentration of P ions, the major factors that determine those equilibria as well as the speciation of soil P are (i) the pH, (ii) the concentrations of anions that compete with P ions for ligand exchange reactions and (iii) the concentrations of metals (Ca, Fe and Al) that can coprecipitate with P ions. The chemical conditions of the rhizosphere are known to considerably differ from those of the bulk soil, as a consequence of a range of processes that are induced either directly by the activity of plant roots or by the activity of rhizosphere microflora. The aim of this paper is to give an overview of those chemical processes that are directly induced by plant roots and which can affect the concentration of P in the soil solution and, ultimately, the bioavailability of soil inorganic P to plants. Amongst these, the uptake activity of plant roots should be taken into account in the first place. A second group of activities which is of major concern with respect to P bioavailability are those processes that can affect soil pH, such as proton/bicarbonate release (anion/cation balance) and gaseous (O2/CO2) exchanges. Thirdly, the release of root exudates such as organic ligands is another activity of the root that can alter the concentration of P in the soil solution. These various processes and their relative contributions to the changes in the bioavailability of soil inorganic P that can occur in the rhizosphere can considerably vary with (i) plant species, (ii) plant nutritional status and (iii) ambient soil conditions, as will be stressed in this paper. Their possible implications for the understanding and management of P nutrition of plants will be briefly addressed and discussed.

Topsoil foraging – an architectural adaptation of plants to low phosphorus availability

Abstract: Low phosphorus availability is a primary constraint to plant productivity in many natural and agricultural ecosystems. Plants display a wide array of adaptive responses to low phosphorus availability that generally serve to enhance phosphorus mobility in the soil and increase its uptake. One set of adaptive responses is the alteration of root architecture to increase phosphorus acquisition from the soil at minimum metabolic cost. In a series of studies with the common bean, work in our laboratory has shown that architectural traits that enhance topsoil foraging appear to be particularly important for genotypic adaptation to low phosphorus soils (‘phosphorus efficiency’). In particular, the gravitropic trajectory of basal roots, adventitious rooting, the dispersion of lateral roots, and the plasticity of these processes in response to phosphorus availability contribute to phosphorus efficiency in this species. These traits enhance the exploration and exploitation of shallow soil horizons, where phosphorus availability is greatest in many soils. Studies with computer models of root architecture show that root systems with enhanced topsoil foraging acquire phosphorus more efficiently than others of equivalent size. Comparisons of contrasting genotypes in controlled environments and in the field show that plants with better topsoil foraging have superior phosphorus acquisition and growth in low phosphorus soils. It appears that many architectural responses to phosphorus stress may be mediated by the plant hormone ethylene. Genetic mapping of these traits shows that they are quantitatively inherited but can be tagged with QTLs that can be used in plant breeding programs. New crop genotypes incorporating these traits have substantially improved yield in low phosphorus soils, and are being deployed in Africa and Latin America.

Effect of salt and osmotic stresses on germination in durum wheat (Triticum durum Desf.)

Abstract: In order to determine the relative importance of ionic toxicity versus the osmotic component of salt stress on germination in durum wheat (Triticum durum Desf.), seeds of three cultivars differing in their salt and drought resistance (Omrabi-5, drought-resistant; Belikh, salt-resistant and Cando, salt-sensitive) were incubated in various iso-osmotic solutions of NaCl, mannitol and polyethylene-glycol (PEG) (osmotic potential of −0.15 (control solution) −0.58, −1.05 or −1.57 MPa). Moderate stress intensities only delayed germination, whereas the highest concentration of NaCl and PEG reduced final germination percentages. PEG was the most detrimental solute, while mannitol had no effect on final germination percentages. All osmotica reduced endosperm starch and soluble sugars content as well as α-amylase activities recorded after 48 h of treatment while β-amylase activities were, in contrast, slightly stimulated in all cultivars. Deleterious effects of NaCl and PEG were higher on isolated embryos germinated onto an in vitro Linsmaier and Skoog (LS) medium comparatively to whole seeds. All PEG-treated embryos, however, recovered after the stress relief while NaCl-treated embryos exhibited a lower rate of recovery and some extent of abnormal germination after rinsing. It was concluded that stress inhibition of germination could not be attributed to an inhibition of mobilisation of reserves and that the main effect of PEG occurred via an inhibition of water uptake while detrimental effects of NaCl may be linked to long-term effects of accumulated toxic ions. The behaviour of the three cultivars during germination did not fully reflect their mean level of putative stress resistance in field conditions and germination is, therefore, not recommended as a reliable selection criterion for breeding purposes.

Large contribution of arbuscular mycorrhizal fungi to soil carbon pools in tropical forest soils

Abstract: The origins and composition of soil organic matter (SOM) are still largely uncertain. Arbuscular mycorrhizal fungi (AMF) are recognized as indirect contributors through their influence on soil aggregation, plant physiology, and plant community composition. Here we present evidence that AMF can also make large, direct contributions to SOM. Glomalin, a recently discovered glycoprotein produced by AMF hyphae, was detected in tropical soils in concentrations of over 60 mg cm−3. Along a chronosequence of soils spanning ages from 300 to 4.1 Mio years, a pattern of glomalin concentrations is consistent with the hypothesis that this protein accumulates in soil. Carbon dating of glomalin indicated turnover at time scales of several years to decades, much longer than the turnover of AMF hyphae (which is assumed to be on the order of days to weeks). This suggests that contributions of mycorrhizae to soil carbon storage based on hyphal biomass in soil and roots may be an underestimate. The amount of C and N in glomalin represented a sizeable amount (ca. 4–5%) of total soil C and N in the oldest soils. Our results thus indicate that microbial (fungal) carbon that is not derived from above- or below-ground litter can make a significant contribution to soil carbon and nitrogen pools and can far exceed the contributions of soil microbial biomass (ranging from 0.08 to 0.2% of total C for the oldest soils).

Plant and rhizosphere processes involved in phytoremediation of metal-contaminated soils

Abstract: This paper reviews the recent advances in understanding of metal removal from contaminated soils, using either hyperaccumulator plants, or high biomass crop species after soil treatment with chelating compounds. Progress has been made at the physiology and molecular level regarding Zn and Ni uptake and translocation in some hyperaccumulators. It is also known that natural hyperaccumulators do not use rhizosphere acidification to enhance their metal uptake. Recently, it has been found that some natural hyperaccumulators proliferate their roots positively in patches of high metal availability. In contrast, non-accumulators actively avoid these areas, and this is one of the mechanisms by which hyperaccumulators absorb more metals when grown in the same soil. However, there are few studies on the exudation and persistence of natural chelating compounds by these plants. It is thought that rhizosphere microorganisms are not important for the hyperaccumulation of metals from soil. Applications of chelates have been shown to induce large accumulations of metals like Pb, U and Au in the shoots of non-hyperaccumulators, by increasing metal solubility and root to shoot translocation. The efficiency of metal uptake does vary with soil properties, and a full understanding of the relative importance of mass flow and diffusion in the presence and absence of artificial chelates is not available. To successfully manipulate and optimise future phytoextraction technologies, it is argued that a fully combined understanding of soil supply and plant uptake is needed.

Effect of Trichoderma harzianum on microelement concentrations and increased growth of cucumber plants

Abstract: The potential of the biocontrol agent Trichoderma harzianum strain T-203 to induce a growth response in cucumber plants was studied in soil and under axenic hydroponic growth conditions. When soil was amended with T. harzianum propagules, a 30% increase in seedling emergence was observed up to 8 days after sowing. On day 28, these plants exhibited a 95 and 75% increase in root area and cumulative root length, respectively, and a significant increase in dry weight (80%), shoot length (45%) and leaf area (80%). Similarly, an increase of 90 and 30% in P and Fe concentration respectively, was observed in T. harzianum inoculated plants. To better characterize the effect of T. harzianum during the early stages of root colonization, experiments were carried out in a gnotobiotic hydroponic system. An increased growth response was apparent as early as 5 days post-inoculation with T. harzianum, resulting in an increase of 25 and 40% in the dry weight of roots and shoots, respectively. Similarly a significant increase in the concentration of Cu, P, Fe, Zn, Mn and Na was observed in inoculated roots. In the shoots of these plants, the concentration of Zn, P and Mn increased by 25, 30 and 70%, respectively. Using the axenic hydroponic system, we showed that the improvement of plant nutritional level may be directly related to a general beneficial growth effect of the root system following T. harzianum inoculation. This phenomenon was evident from 5 days post-inoculation throughout the rest of the growth period, resulting in biomass accumulation in both roots and shoots.

Antioxidative and growth-promoting effect of selenium on senescing lettuce

Abstract: In human and animal cells, Se plays an essential role in antioxidation and exerts an antiaging function but it is toxic at high dietary intake. To increase its intake in forage and foodstuffs, Se fertilization is adopted in some countries where soils are low in bioavailable Se, even though higher plants are regarded not to require Se. To test its ability to counteract senescence-related oxidative stress in higher plants, a pot experiment was carried out with lettuce (Lactuca sativa) cultivated with increasing amounts of H2SeO4. The yields harvested 7 or 14 weeks after sowing revealed that a low Se dosage (0.1 mg kg−1 soil) stimulated the growth of senescing seedlings (dry weight yield by 14%) despite a decreased chlorophyll concentration. The growth-promoting function was related to diminished lipid peroxidation. In young and senescing plants, the antioxidative effect of Se was associated with the increased activity of glutathione peroxidase (GSH-Px). In the senescing plants, the added Se strengthened the antioxidative capacity also by preventing the reduction of tocopherol concentration and by enhancing superoxide dismutase (SOD) activity. When no Se was added, tocopherols and SOD activity diminished during plant senescence. The higher Se dosage (1.0 mg kg−1 soil) was toxic and reduced the yield of young plants. In the senescing plants, it diminished the dry weight yield but not the fresh weight yield.

The nutritional control of root development

Abstract: Root development is remarkably sensitive to variations in the supply and distribution of inorganic nutrients in the soil. Here we review examples of the ways in which nutrients such as N, P, K and Fe can affect developmental processes such as root branching, root hair production, root diameter, root growth angle, nodulation and proteoid root formation. The nutrient supply can affect root development either directly, as a result of changes in the external concentration of the nutrient, or indirectly through changes in the internal nutrient status of the plant. The direct pathway results in developmental responses that are localized to the part of the root exposed to the nutrient supply; the indirect pathway produces systemic responses and seems to depend on long-distance signals arising in the shoot. We propose the term ‘trophomorphogenesis’ to describe the changes in plant morphology that arise from variations in the availability or distribution of nutrients in the environment. We discuss what is currently known about the mechanisms of external and internal nutrient sensing, the possible nature of the long-distance signals and the role of hormones in the trophomorphogenic response.

Phosphorus loss from land to water: integrating agricultural and environmental management

Abstract: Phosphorus (P), an essential nutrient for crop and animal production, can accelerate freshwater eutrophication, now one of the most ubiquitous forms of water quality impairment in the developed world. Repeated outbreaks of harmful algal blooms (e.g., Cyanobacteria and Pfiesteria) have increased society's awareness of eutrophication, and the need for solutions. Agriculture is regarded as an important source of P in the environment. Specifically, the concentration of specialized farming systems has led to a transfer of P from areas of grain production to animal production. This has created regional surpluses in P inputs (mineral fertilizer and feed) over outputs (crop and animal produce), built up soil P in excess of crop needs, and increased the loss of P from land to water. Recent research has shown that this loss of P in both surface runoff and subsurface flow originates primarily from small areas within watersheds during a few storms. These areas occur where high soil P, or P application in mineral fertilizer or manure, coincide with high runoff or erosion potential. We argue that the overall goal of efforts to reduce P loss to water should involve balancing P inputs and outputs at farm and watershed levels by optimizing animal feed rations and land application of P as mineral fertilizer and manure. Also, conservation practices should be targeted to relatively small but critical watershed areas for P export.

EDTA enhanced heavy metal phytoextraction: metal accumulation, leaching and toxicity

Abstract: Synthetic chelates such as ethylene diamine tetraacetic acid (EDTA) have been shown to enhance phytoextraction of some heavy metals from contaminated soil. In a soil column study, we examined the effect of EDTA on the uptake of Pb, Zn and Cd by Chinese cabbage (Brassica rapa), mobilization and leaching of heavy metals and the toxicity effects of EDTA additions on plants. The most effective was a single dose of 10 mmol EDTA kg−1 soil where we detected Pb, Zn and Cd concentrations that were 104.6, 3.2 and 2.3-times higher in the aboveground plant biomass compared to the control treatments. The same EDTA addition decreased the concentration of Pb, Zn and Cd in roots of tested plants by 41, 71 and 69%, respectively compared to concentrations in the roots of control plants. In columns treated with 10 mmol kg−1 EDTA, up to 37.9, 10.4 and 56.3% of initial total Pb, Zn and Cd in soil were leached down the soil profile, suggesting high solubility of heavy metals-EDTA complexes. EDTA treatment had a strong phytotoxic effect on the red clover (Trifolium pratense) in bioassay experiment. Moreover, the high dose EDTA additions inhibited the development of arbuscular mycorrhiza. The results of phospholipid fatty acid analyses indicated toxic effects of EDTA on soil fungi and increased environmental stress of soil microfauna.

Simulation of C and N mineralisation during crop residue decomposition: A simple dynamic model based on the C:N ratio of the residues

Abstract: C and N mineralisation kinetics obtained in laboratory incubations during decomposition of crop residues under non-limiting nitrogen conditions were simulated using a simple dynamic model. This model includes three compartments: the residues, microbial biomass and humified organic matter. Seven parameters are used to describe the C and N fluxes. The decomposed C is either mineralised as CO2 or assimilated by the soil microflora, microbial decay producing both C humification and secondary C mineralisation. The N dynamics are governed by the C rates and the C:N ratio of the compartments which remain constant in the absence of nitrogen limitation. The model was parameterised using apparent C and N mineralisation kinetics obtained for 27 different residues (organs of oilseed rape plants) that exhibited very wide variations in chemical composition and nitrogen content. Except for the C:N ratio of the residues and the soil organic matter, the other five parameters of the model were obtained by non-linear fitting and by minimising the differences between observed and simulated values of CO2 and mineral N. Three parameters, namely the decomposition rate constant of the residues, the biomass C:N ratio and humification rate, were strongly correlated with the residues C:N ratio. Hyperbolic relationships were established between these parameters and the residues C:N ratio. In contrast, the other two parameters, i.e. the decay rate of the microbial biomass and the assimilation yield of residue-C by the microbial biomass, were not correlated to the residues C:N ratio and were, therefore, fixed in the model. The model thus parameterised against the residue C:N ratio as a unique criterion, was then evaluated on a set of 48 residues. An independent validation was obtained by taking into account 21 residues which had not been used for the parameterisation. The kinetics of apparent C and N mineralisation were reasonably well simulated by the model. The model tended to over-estimate carbon mineralisation which could limit its use for C predictions, but the kinetics of N immobilisation or mineralisation due to decomposition of residues in soil were well predicted. The model indicated that the C:N ratio of decomposers increased with the residue C:N ratio. Higher humification was predicted for substrates with lower C:N ratios. This simple dynamic model effectively predicts N evolution during crop residue decomposition in soil.

Preinoculation of lettuce and onion with VA mycorrhizal fungi reduces deleterious effects of soil salinity

Abstract: The hypothesis that inoculation of transplants with vesicular-arbuscular mycorrhizal (VAM) fungi before planting into saline soils alleviates salt effects on growth and yield was tested on lettuce (Lactuca sativa L.) and onion (Allium cepa L.). A second hypothesis was that fungi isolated from saline soil are more effective in counteracting salt effects than those from nonsaline soil. VAM fungi from high- and low-salt soils were trap-cultured, their propagules quantified and adjusted to a like number, and added to a pasteurized soil mix in which seedlings were grown for 3–4 weeks. Once the seedlings were colonized by VAM fungi, they were transplanted into salinized (NaCl) soil. Preinoculated lettuce transplants grown for 11 weeks in the saline soils had greater shoot mass compared with nonVAM plants at all salt levels [2 (control), 4, 8 and 12 dS m−1] tested. Leaves of VAM lettuce at the highest salt level were significantly greener (more chlorophyll) than those of the nonVAM lettuce. NonVAM onions were stunted due to P deficiency in the soil, but inoculation with VAM fungi alleviated P deficiency and salinity effects; VAM onions were significantly larger at all salt levels than nonVAM onions. In a separate experiment, addition of P to salinized soil reduced the salt stress effect on nonVAM onions but to a lesser extent than by VAM inoculation. VAM fungi from the saline soil were not more effective in reducing growth inhibition by salt than those from the nonsaline site. Colonization of roots and length of soil hyphae produced by the VAM fungi decreased with increasing soil salt concentration. Results indicate that preinoculation of transplants with VAM fungi can help alleviate deleterious effects of saline soils on crop yield.

Phosphorus Transformations in an Oxisol under contrasting land-use systems: The role of the soil microbial biomass

Abstract: It is generally assumed that phosphorus (P) availability for plant growth on highly weathered and P-deficient tropical soils may depend more on biologically mediated organic P (Po) turnover processes than on the release of adsorbed inorganic P (Pi). However, experimental evidence showing the linkages between Po, microbial activity, P cycling and soil P availability is scarce. To test whether land-use systems with higher soil Po are characterized by greater soil biological activity and increased P mineralization, we analyzed the partitioning of P among various organic and inorganic P fractions in soils of contrasting agricultural land-use systems and related it to biological soil properties. Isotopic labeling was used to obtain information on the turnover of P held in the microbial biomass. Soil samples were taken from grass–legume pasture (GL), continuous rice (CR) and native savanna (SAV) which served as reference. In agreement with estimated P budgets (+277, +70 and 0 kg P ha−1 for CR, GL and SAV, respectively), available P estimated using Bray-2 and resin extraction declined in the order CR > GL > SAV. Increases in Bray-2 and resin Pi were greater in CR than GL relative to total soil P increase. Organic P fractions were significantly less affected by P inputs than inorganic fractions, but were a more important sink in GL than CR soils. Extractable microbial P (Pchl) was slightly higher in GL (6.6 mg P kg−1) than SAV soils (5.4 mg P kg−1), and significantly lowest in CR (2.6 mg P kg−1). Two days after labeling the soil with carrier free 33P, 25, 10 and 2% of the added 33P were found in Pchl in GL, SAV and CR soils, respectively, suggesting a high and rapid microbial P turnover that was highest in GL soils. Indicators of P mineralization were higher in GL than CR soils, suggesting a greater transformation potential to render Po available. Legume-based pastures (GL) can be considered as an important land-use option as they stimulate P cycling. However, it remains to be investigated whether crops planted in pasture–crop rotations could benefit from the enhanced Po cycling in grass–legume soils. Furthermore, there is need to develop and test a direct method to quantify Po mineralization in these systems.

Nitrogen cycling in low input legume-based agriculture, with emphasis on legume/grass pastures

Abstract: Low input legume-based agriculture exists in a continuum between subsistence farming and intensive arable and pastoral systems. This review covers this range, but with most emphasis on temperate legume/grass pastures under grazing by livestock. Key determinants of nitrogen (N) flows in grazed legume/grass pastures are: inputs of N from symbiotic N2 fixation which are constrained through self-regulation via grass/legume interactions; large quantities of N cycling through grazing animals with localised return in excreta; low direct conversion of pasture N into produce (typically 5–20%) but with N recycling under intensive grazing the farm efficiency of product N: fixed N can be up to 50%; and regulation of N flows by mineralisation/immobilisation reactions. Pastoral systems reliant solely on fixed N are capable of moderate-high production with modest N losses e.g. average denitrification and leaching losses from grazed pastures of 6 and 23 kg N ha−1 yr−1. Methods for improving efficiency of N cycling in legume-based cropping and legume/grass pasture systems are discussed. In legume/arable rotations, the utilisation of fixed N by crops is influenced greatly by the timing of management practices for synchrony of N supply via mineralisation and crop N uptake. In legume/grass pastures, the spatial return of excreta and the uptake of excreta N by pastures can potentially be improved through dietary manipulation and management strategies. Plant species selection and plant constituent modification also offer the potential to increase N efficiency through greater conversion into animal produce, improved N uptake from soil and manipulation of mineralisation/immobilisation/nitrification reactions.

Are differences in root growth of nitrogen catch crops important for their ability to reduce soil nitrate-N content, and how can this be measured?

Abstract: An experiment was made to measure root growth of nitrogen catch crops, to investigate whether differences in root growth among plant species are related to their ability to deplete the soil nitrate-N pool. Large differences were observed in root growth parameters. Monocot species had rooting depth penetration rates in the range of 1.0 to 1.2 mm d−1 °C−1, whereas the non-legume dicot species had rates between 1.5 and 2.3 mm d−1 °C−1. Substantial differences were also found in the lag time from sowing until significant root growth was observed. The estimated temperature sum needed for the crops to reach a rooting depth of 1.0 m varied from 750 d °C for fodder radish to 1375 d °C for Italian ryegrass. The depth distribution of the root system varied strongly, and at a depth of 1.0 m the non-legume dicot species generally had root intensities (number of root intersections m−1 line on the minirhizotrons) 12 times as high as the monocot species. The amount of nitrate left in the topsoil (0–0.5 m) was only weakly correlated to a few of the measured plant and root parameters, whereas nitrate left in the subsoil (0.5–1.0 m) was clearly correlated to several root parameters. Subsoil nitrate residues were well correlated to root intensity, but showed even stronger correlations to more simple estimates of rooting depth. In the deepest soil layer measured (1.0–1.5 m), the soil water nitrate concentration was reduced from 119 μg L−1 without a catch crop to 61 μg L−1 under Italian ryegrass and to only 1.5 μg L−1 under fodder radish. The results show that to identify the important differences in root growth among catch crops, root growth must be measured in deep soil layers. In this study, none of the measurements made aboveground or in the upper soil layers were well related to subsoil nitrate depletion.

Isolation, partial characterization, and the effect of plant growth-promoting bacteria (PGPB) on micro-propagated sugarcane in vitro

Abstract: We report the isolation of nitrogen fixing, phytohormone producing bacteria from sugarcane and their beneficial effects on the growth of micropropagated sugarcane plantlets. Detection of the nitrogen fixing bacteria by ARA-based MPN (acetylene reduction assay-based most probable number) method indicated the presence of up to 106 bacteria per gram dry weight of stem and 107 bacteria per gram dry weight of root of field-grown sugarcane. Two nitrogen fixing bacterial isolates were obtained from stem (SC11, SC20) and two from the roots (SR12, SR13) of field-grown plants. These isolates were identified as Enterobacter sp. strains on the basis of their morphological characteristics and biochemical tests. The isolate SC20 was further characterized by 16S rRNA sequence analysis, which showed high sequence similarity to the sequence of Enterobacter cloacae and Klebsiella oxytoca. All the isolates produced the phytohormone indoleacetic acid (IAA) in pure culture and this IAA production was enhanced in growth medium containing tryptophan. The bacterial isolates were used to inoculate micro-propagated sugarcane in vitro where maximum increase in the root and shoot weight over control was observed in the plantlets inoculated with strain SC20. By using the15N isotope dilution technique, maximum nitrogen fixation contribution (28% of total plant nitrogen) was detected in plantlets inoculated with isolate SC20.

Effect of phosphorus availability on basal root shallowness in common bean.

Abstract: Root gravitropism may be an important element of plant response to phosphorus availability because it determines root foraging in fertile topsoil horizons, and thereby phosphorus acquisition. In this study we seek to test this hypothesis in both two dimensional paper growth pouch and three-dimensional solid media of sand and soil cultures. Five common bean (Phaseolus vulgaris L.) genotypes with contrasting adaptation to low phosphorus availability were evaluated in growth pouches over 6 days of growth, and in sand culture and soil culture over 4 weeks of growth. In all three media, phosphorus availability regulated the gravitropic response of basal roots in a genotype-dependent manner. In pouches, sand, and soil, the phosphorus-inefficient genotype DOR 364 had deeper roots with phosphorus stress, whereas the phosphorus-efficient genotype G19833 responded to phosphorus stress by producing shallower roots. Genotypes were most responsive to phosphorus stress in sand culture, where relative root allocation to the 0–3- and 3–6-cm horizons increased 50% with phosphorus stress, and varied 300% (3–6 cm) to 500% (0–3 cm) among genotypes. Our results indicate that (1) phosphorus availability regulates root gravitropic growth in both paper and solid media, (2) responses observed in young seedlings continue throughout vegetative growth, (3) the response of root gravitropism to phosphorus availability varies among genotypes, and (4) genotypic adaptation to low phosphorus availability is correlated with the ability to allocate roots to shallow soil horizons under phosphorus stress.

Trends in rhizobial inoculant production and use

Abstract: Rhizobia inoculants have contributed to increase N2 fixation and yield in legumes crops. However, most of the inoculants produced world-wide are of poor or suboptimal quality. We discuss here why some of them are poor products and how to improve their quality and efficacy. Reported data on the inoculation rate effect can be used to design good inoculants. Technologies are now available to produce inoculants with a shelf-life of more than 1 year. Available quality control methods can help to improve the quality of inoculants although they do not take into account the physiological satus of the rhizobia. Unfortunately quality control is not commonly used except in major inoculant companies and the quality of inoculants sold on the market is low. The need for an increase in quality standards is discussed especially for the number of rhizobia delivered per seed and for the presence of contaminants. Some new technologies which able to increase efficacy and reliability of inoculation are discussed.

Plants and fertilisers as drivers of change in microbial community structure and function in soils

Abstract: Here we report on a range of studies designed to understand the link between diversity and function in soils and in particular how plants and fertilisers might interact with microbial community dynamics in soils. The data presented indicate that although plants and fertilisers do impact on microbial community structure, the relationship between diversity, community structure and function remains complex and difficult to interpret using currently available chemical and molecular fingerprinting techniques. The paper assesses plants and management practices as drivers of change in soil and argues that whilst understanding diversity per se is unlikely to contribute to our understanding of function, an appreciation of what causes communities to change and also the relative importance of such drivers, could lead to new insights into the sustainable management and conservation of soils and natural resources.

Root hairs confer a competitive advantage under low phosphorus availability

Abstract: Root hairs are presumably important in the acquisition of immobile soil resources such as phosphorus. The density and length of root hairs vary substantially within and between species, and are highly regulated by soil phosphorus availability, which suggests that at high nutrient availability, root hairs may have a neutral or negative impact on fitness. We used a root-hairless mutant of the small herbaceous dicot Arabidopsis thaliana to assess the effect of root hairs on plant competition under contrasting phosphorus regimes. Wildtype plants were grown with hairless plants in a replacement series design at high (60 μm phosphate in soil solution) and low (1 μm phosphate in soil solution) phosphorus availability. At high phosphorus availability, wildtype and mutant plants were equal in growth, phosphorus acquisition, fecundity and relative crowding coefficient (RCC). At low phosphorus availability, hairless plants accumulated less biomass and phosphorus, and produced less seed when planted with wildtype plants. Wildtype plants were unaffected by the presence of hairless plants in mixed genotype plantings. Wildtype plants had RCC values greater than one while hairless plants had RCC values less than one. We conclude that root hairs increase the competitiveness of plants under low phosphorus availability but do not reduce growth or competitiveness under high phosphorus availability.

A comparison of soil quality in adjacent cultivated, forest and native grassland soils

Abstract: Changes in soil quality after 45 years of continuous production of corn (Zea mays L) by the conventional tillage method (C) compared with adjacent poplar forest (F) and native grassland (G) sites were examined The investigated parameters were: total and humified organic C, total N, light fraction content and composition, water-soluble organic C (WSOC), water-soluble carbohydrates (WSC), phenolic substances, biomass C, cumulative CO2-C (soil respiration) (Cm), enzyme activities (alkaline phosphatase, protease, β-glucosidase, urease, catalase and dehydrogenase) Empirical indexes of soil quality were also calculated: biomass C/organic C, specific respiration of biomass C (qCO2), death rate quotient (qD), metabolic potential (MP), biological index of fertility (BIF), enzyme activity number (EAN) and hydrolysing coefficient (HC) Results indicate that long-term corn production at an intensive level caused a marked decline in all examined parameters Between the undisturbed systems, native grassland showed higher values of soil quality parameters than forest site The indexes most responsive to management practices that may provide indications of the effects of soil cultivation, as well as of the differently undisturbed ecosystems were: organic C, WSC, Cm, protease, β-glucosidase, urease and HC Soil enzyme activities were well related with, and not more sensitive than organic carbon

Agronomic measures for increasing P availability to crops

Abstract: Acquisition of soil and fertiliser phosphorus (P) by crops depends on soil and plant properties Soil processes determining P availability to plants are P solubility/sorption, P transport, root/soil contact and mineralisation/immobilisation Plants have evolved properties contributing to a more efficient use of plant-available soil P and to mobilise P from less available soil P fractions Agronomic measures may affect P availability to crops through the modification of soil properties or through direct quantitative and qualitative crop impact on soil P dynamics Among the agronomic measures, the application of organic matter such as green manure and crop residues to maintain or increase soil organic matter content and to enhance soil biological activity, and the incorporation into the cropping system of P-mobilising plant species are particularly beneficial Our experimental activities have concentrated on the characterisation of the P mobilising capacity of different leguminous grain and cover crops, and their effect on P availability to less P-efficient cereals grown in mixed culture and in rotation Fractionation of P in the rhizosphere soil revealed the capacity of some legumes to better use P from sparingly soluble soil P fractions than maize Field experiments conducted on 2 sites in the Northern Guinea Savannah of Nigeria and accompanying green-house pot experiments revealed a positive rotational effect of P-efficient cover crops on maize growth and grain yield with and without the return of crop residues This could unequivocally be attributed to a better P supply to maize, especially on strongly P-fixing soil However, the residual effect was small compared to the application of water-soluble P fertiliser This clearly indicates the need for a maintenance application of fertiliser P in addition to the agronomic measures for sustainable crop production

Interactive effects of boron and salinity stress on the growth, membrane permeability and mineral composition of tomato and cucumber plants

Abstract: A greenhouse study was conducted in order to determine interactive effects of NaCl salinity and B on the growth, sodium (Na), chloride (Cl), boron (B), potassium (K) concentrations and membrane permeability of salt resistant Tomato (Lycopersicon esculentum L cv Lale F1) and salt sensitive cucumber (Cucumis sativus L cv Santana F1) plants Plants were grown in a factorial combination of NaCl (0 and 30 mM for cucumber and 0 and 40 mM for tomato) and B (0, 5, 10 and 20 mg kg−1 soil) Boron toxicity symptoms appeared at 5 mg kg−1 B treatments in both plants Salinity caused an increase in leaf injury due to B toxicity, but it was more severe in cucumber Dry weights of the plants decreased with the increasing levels of applied B in nonsaline conditions, but the decrease in dry weights due to B toxicity was more pronounced in saline conditions especially in cucumber Salinity × B interaction on the concentration of B in both plants was found significant However, increase in B concentrations of tomato decreased under saline conditions when compared to nonsaline conditions Contrary to this, B concentration of cucumber increased as a result of increasing levels of applied B and salinity Salinity increased Na and Cl concentrations of both plants Potassium concentration of tomato was not affected by salinity and B treatments, but K concentration of cucumber was decreased by salinity Membrane permeability of the plants was increased by salinity while toxic levels of B had no effect on membrane permeability in nonsaline conditions Membrane permeability was significantly increased in the presence of salinity by the increasing levels of applied B

Salinity tolerance in Hordeum vulgare : ion concentrations in root cells of cultivars differing in salt tolerance

Abstract: The tolerance of 24 genotypes of barley was assessed by estimating their survival in saline conditions either in a glasshouse or in a controlled environment cabinet. Two cultivars, sensitive Triumph and resistant Gerbel, were picked for further study, which involved sequential harvesting of plants grown in a range of salinities. After about one month in 200 mol m_3 sodium chloride, the sodium concentration in the roots and shoots of the sensitive Triumph was about 1.5 times that in the roots of resistant Gerbel. The addition of Na to the root medium reduced the potassium transport to the shoot in Triumph to a much greater extent than in Gerbel, so the K:Na ratio of Gerbel was twice that for Triumph, when averaged over all treatments and harvests. The sodium, potassium and chloride concentrations within the major subcellular compartments of the cortical cells of roots of Triumph and Gerbel were determined by X-ray microanalysis following freeze-substitution and dry-sectioning. The mean cytoplasmic sodium concentration (245 molm_3 analysed volume) in Triumph grown in 200 molm_3 NaCl for 15 d was almost 1.4 times greater than that in the resistant Gerbel: the potassium concentration in Gerbel showed a lower reduction than did that of Triumph. Another major difference between the two cultivars was the higher concentrations of sodium and chloride in the cell walls of Triumph than Gerbel: the sodium concentration in the cortical cell walls of the salt-sensitive cultivar was about 1.75 times that in the more salt-resistant cultivar. The exchange capacity of the cell walls of Gerbel was greater than that of Triumph. We hypothesise that ion transport to the shoot reflects cytosolic ion concentrations, with a more sensitive cultivar having a higher sodium concentration in its cytoplasm than a more resistant variety. It is noteworthy that the difference in the K:Na ratio between the shoots of Gerbel and Triumph after 15 days of exposure to 200 mol m_3 NaCl was similar to the difference in their symplastic K:Na ratios.

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

Abstract: 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.

Moisture retention properties of a mycorrhizal soil

Abstract: The water relations of arbuscular mycorrhizal plants have been compared often, but virtually nothing is known about the comparative water relations of mycorrhizal and nonmycorrhizal soils. Mycorrhizal symbiosis typically affects soil structure, and soil structure affects water retention properties; therefore, it seems likely that mycorrhizal symbiosis may affect soil water relations. We examined the water retention properties of a Sequatchie fine sandy loam subjected to three treatments: seven months of root growth by (1) nonmycorrhizal Vigna unguiculata given low phosphorus fertilization, (2) nonmycorrhizal Vigna unguiculata given high phosphorus fertilization, (3) Vigna unguiculata colonized by Glomus intraradices and given low phosphorus fertilization. Mycorrhization of soil had a slight but significant effect on the soil moisture characteristic curve. Once soil matric potential (Ψm) began to decline, changes in Ψm per unit change in soil water content were smaller in mycorrhizal than in the two nonmycorrhizal soils. Within the range of about −1 to −5 MPa, the mycorrhizal soil had to dry more than the nonmycorrhizal soils to reach the same Ψm. Soil characteristic curves of nonmycorrhizal soils were similar, whether they contained roots of plants fed high or low phosphorus. The mycorrhizal soil had significantly more water stable aggregates and substantially higher extraradical hyphal densities than the nonmycorrhizal soils. Importantly, we were able to factor out the possibly confounding influence of differential root growth among mycorrhizal and nonmycorrhizal soils. Mycorrhizal symbiosis affected the soil moisture characteristic and soil structure, even though root mass, root length, root surface area and root volume densities were similar in mycorrhizal and nonmycorrhizal soils.

Utilization of phosphorus by pasture plants supplied with myo-inositol hexaphosphate is enhanced by the presence of soil micro-organisms

Abstract: A range of pasture grass (Danthonia richardsonii and Phalaris aquatica) and legume (Medicago polymorpha, M. sativa, Trifolium repens and T. subterraneum) species showed limited capacity to obtain phosphorus (P) from inositol hexaphosphate (IHP), when grown in either sterile agar (pH 5.0 or 5.5) or sand-vermiculite media (pH 5.0). The total P content of shoots from IHP-supplied plants grown in agar was between 20% and 34% of that for seedlings supplied with an equivalent amount of P as inorganic phosphate (Pi), while in sand-vermiculite, the total P content of IHP-grown plants was between 5 and 10% of control plants. The poor ability of plants to utilize P from IHP resulted in significantly lower tissue P concentrations and, in general, reduced plant dry weight accumulation. In contrast, the P nutrition of plants supplied with IHP was significantly improved by inoculating media with either a cultured population of total soil micro-organisms or with a specific isolate of Pseudomonas sp., selected for its ability to release phosphate from IHP (strain CCAR59; Richardson and Hadobas, 1997 Can. J. Micro. 43, 509-516). In agar and sand-vermiculite media, respectively, the P content of IHP-grown plants increased with inoculation by up to 3.9- and 6.8-fold, such that the dry weight and P content of the plant material were equivalent to those observed for control plants supplied with Pi. However, the response to inoculation was dependent on the growth medium and the source of micro-organisms used. In sand-vermiculite, the cultured population of soil micro-organisms was effective when IHP was supplied at an equivalent level of Pi required for maximum plant growth. By comparison, inoculation of plants with the Pseudomonas strain was only effective at very high levels of IHP supply (×36), whereas in agar a response to inoculation occurred at all levels of IHP. The ability of pasture plants to acquire P from phytate was, therefore, influenced by the availability of IHP substrate, which was further affected by the presence of soil micro-organisms. Our results show that in addition to having an effect on the sorption characteristics of the growth media, soil micro-organisms also provided a source of phytase for the dephosphorylation of phytate for subsequent utilization of Pi by plants.

Combined effects of drought and high temperature on water relations of wheat and sorghum

Abstract: Drought and high temperature are major factors limiting crop production The two stresses occur together in many regions, but they usually are investigated separately This study tested the hypothesis that high temperature interacts with drought to affect water relations, and the effect is greater in heat-sensitive wheat (Triticum aestivum L) than in sorghum (Sorghum bicolor L Moench) Wheat and sorghum were grown in soil that was well watered or not watered in controlled chambers at 15/10, 25/20, 35/30 and 40/35 °C day/night Soil water content (SWC), leaf relative water content (RWC), leaf water potential (Ψ), leaf osmotic potential (π), leaf turgor potential (P) and osmotic adjustment (OA) were determined at 2-d intervals All values held nearly constant at all temperatures when soil was well watered but were affected strongly by high temperature when water was withheld The combined stresses reduced SWC, RWC, Psi and π, and unevenly raised P over time, particularly in sorghum Sorghum also exhibited marked OA at high temperature, which was usually lethal to wheat High temperature appeared to interact with drought to affect water relations by altering SWC and not by influencing OA The results demonstrated that crops maintain nearly stable water relations regardless of temperature when moisture is ample, but high temperature strongly affects water relations when water is limiting Increasing the thermotolerance of wheat might improve its potential to acclimate to both high temperature and drought

Temporal and spatial distribution of roots and competition for nitrogen in pea-barley intercrops – a field study employing 32P technique

Abstract: Root system dynamics, productivity and N use were studied in inter- and sole crops of field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L.) on a temperate sandy loam. A 32P tracer placed at a depth of 12.5, 37.5, 62.5 or 87.5 cm was employed to determine root system dynamics by sampling crop leaves at 0, 15, 30 and 45 cm lateral distance. 15N addition was used to estimate N2 fixation by pea, using sole cropped barley as reference crop. The Land Equivalent Ratio (LER), which is defined as the relative land area under sole crops that is required to produce the yields achieved in intercropping, were used to compare the crop growth in intercrops relative to the respective sole crops. The 32P appearance in leaves revealed that the barley root system grows faster than that of pea. P uptake by the barley root system during early growth stages was approximately 10 days ahead of that of the pea root system in root depth and lateral root distribution. More than 90% of the P uptake by the pea root system was confined to the top 12.5 cm of soil, whereas barley had about 25–30% of tracer P uptake in the 12.5 – 62.5 cm soil layer. Judging from this P uptake, intercropping caused the barley root system to grow deeper and faster lateral root development of both species was observed. Barley accumulated similar amounts of aboveground N when grown as inter- and sole crop, whereas the total aboveground N acquired by pea in the intercrop was only 16% of that acquired in the pea sole crop. The percentage of total aboveground N derived from N2 fixation in sole cropped pea increased from 40% to 80% during the growth period, whereas it was almost constant at 85% in intercropped pea. The total amounts of N2 fixed were 95 and 15 kg N ha−1 in sole cropped and intercropped pea, respectively. Barley was the dominant component of the pea-barley intercrop, obtaining 90% of its sole crop yield, while pea produced only 15% of the grains of a sole crop pea. Intercropping of pea and barley improved the utilization of plant growth resources (LER > 1) as compared to sole crops. Root system distribution in time and space can partly explain interspecific competition. The 32P methodology proved to be a valuable tool for determining root dynamics in intercropping systems.

Cadmium mobilisation and plant availability – the impact of organic acids commonly exuded from roots

Abstract: The present work highlights metal-organic acid interactions with special reference to their plant availability Pot experiments were conducted to investigate the effect of various organic (carboxylic and amino) acids on the uptake and translocation of root-absorbed Cd by maize (Zea mays) plants grown in sand and soil culture Statistically significant increases in Cd accumulation from Cd-treated plants in the presence of increasing concentrations of organic acids, suggest the existence of Cd-organic acid interactions in the soil-plant system In order to support the above hypothesis of formation of organically bound Cd, separate experiments were performed to synthesize and estimate its various forms viz cationic, anionic and neutral The chemical nature of the organically bound forms was ascertained by electrophoretic experiments Amino acids have been found to be less effective in the mobilisation of cadmium compared to carboxylic acids The results are discussed on the basis of the potential of organic acids to form complexes with Cd