Neighbouring plants modify maize root foraging for phosphorus: coupling nutrients and neighbours for improved nutrient-use efficiency.

Nutrient distribution and neighbours can impact plant growth, but how neighbours shape root-foraging strategy for nutrients is unclear. Here, we explore new patterns of plant foraging for nutrients as affected by neighbours to improve nutrient acquisition. Maize (Zea mays) was grown alone (maize), or with maize (maize/maize) or faba bean (Vicia faba) (maize/faba bean) as a neighbour on one side and with or without a phosphorus (P)-rich zone on the other in a rhizo-box experiment. Maize demonstrated root avoidance in maize/maize, with reduced root growth in 'shared' soil, and increased growth away from its neighbours. Conversely, maize proliferated roots in the proximity of neighbouring faba bean roots that had greater P availability in the rhizosphere (as a result of citrate and acid phosphatase exudation) compared with maize roots. Maize proliferated more roots, but spent less time to reach, and grow out of, the P patches away from neighbours in the maize/maize than in the maize/faba bean experiment. Maize shoot biomass and P uptake were greater in the heterogeneous P treatment with maize/faba bean than with maize/maize system. The foraging strategy of maize roots is an integrated function of heterogeneous distribution of nutrients and neighbouring plants, thus improving nutrient acquisition and maize growth. Understanding the foraging patterns is critical for optimizing nutrient management in crops.

Elevated [CO2 ] alleviates the impacts of water deficit on xylem anatomy and hydraulic properties of maize stems.

Plants can modify xylem anatomy and hydraulic properties to adjust to water status. Elevated [CO2 ] can increase plant water potential via reduced stomatal conductance and water loss. This raises the question of whether elevated [CO2 ], which thus improves plant water status, will reduce the impacts of soil water deficit on xylem anatomy and hydraulic properties of plants. To analyse the impacts of water and [CO2 ] on maize stem xylem anatomy and hydraulic properties, we exposed potted maize plants to varying [CO2 ] levels (400, 700, 900, and 1,200 ppm) and water levels (full irrigation and deficit irrigation). Results showed that at current [CO2 ], vessel diameter, vessel roundness, stem cross-section area, specific hydraulic conductivity, and vulnerability to embolism decreased under deficit irrigation; yet, these impacts of deficit irrigation were reduced at elevated [CO2 ]. Across all treatments, midday stem water potential was tightly correlated with xylem traits and displayed similar responses. A distinct trade-off between efficiency and safety in stem xylem water transportation in response to water deficit was observed at current [CO2 ] but not observed at elevated [CO2 ]. The results of this study enhance our knowledge of plant hydraulic acclimation under future climate environments and provide insights into trade-offs in xylem structure and function.

Responses of water accumulation and solute metabolism in tomato fruit to water scarcity and implications for main fruit quality variables.

Fruit is important for human health, and applying deficit irrigation in fruit production is a strategy to regulate fruit quality and support environmental sustainability. Responses of different fruit quality variables to deficit irrigation have been widely documented, and much progress has been made in understanding the mechanisms of these responses. We review the effects of water shortage on fruit water accumulation considering water transport from the parent plant into the fruit determined by hydraulic properties of the pathway (including xylem water transport and transmembrane water transport regulated by aquaporins) and the driving force for water movement. We discuss water relations and solute metabolism that affect the main fruit quality variables (e.g. size, flavour, nutrition, and firmness) at the cellular level under water shortage. We also summarize the most recent advances in the understanding of responses of the main fruit quality variables to water shortage, considering the effects of variety, the severity of water deficit imposed, and the developmental stage of the fruit. We finally identify knowledge gaps and suggest avenues for future research. This review provides new insights into the stress physiology of fleshy fruit, which will be beneficial for the sustainable production of high-quality fruit under deficit irrigation.

Ozone pollution will compromise efforts to increase global wheat production.

Introduction of high-performing crop cultivars and crop/soil water management practices that increase the stomatal uptake of carbon dioxide and photosynthesis will be instrumental in realizing the United Nations Sustainable Development Goal (SDG) of achieving food security. To date, however, global assessments of how to increase crop yield have failed to consider the negative effects of tropospheric ozone, a gaseous pollutant that enters the leaf stomatal pores of plants along with carbon dioxide, and is increasing in concentration globally, particularly in rapidly developing countries. Earlier studies have simply estimated that the largest effects are in the areas with the highest ozone concentrations. Using a modelling method that accounts for the effects of soil moisture deficit and meteorological factors on the stomatal uptake of ozone, we show for the first time that ozone impacts on wheat yield are particularly large in humid rain-fed and irrigated areas of major wheat-producing countries (e.g. United States, France, India, China and Russia). Averaged over 2010-2012, we estimate that ozone reduces wheat yields by a mean 9.9% in the northern hemisphere and 6.2% in the southern hemisphere, corresponding to some 85 Tg (million tonnes) of lost grain. Total production losses in developing countries receiving Official Development Assistance are 50% higher than those in developed countries, potentially reducing the possibility of achieving UN SDG2. Crucially, our analysis shows that ozone could reduce the potential yield benefits of increasing irrigation usage in response to climate change because added irrigation increases the uptake and subsequent negative effects of the pollutant. We show that mitigation of air pollution in a changing climate could play a vital role in achieving the above-mentioned UN SDG, while also contributing to other SDGs related to human health and well-being, ecosystems and climate change.

Phenotypic and genome-wide association analysis of spike ethylene in diverse wheat genotypes under heat stress.

The gaseous phytohormone ethylene plays an important role in spike development in wheat (Triticum aestivum). However, the genotypic variation and the genomic regions governing spike ethylene (SET) production in wheat under long-term heat stress remain unexplored. We investigated genotypic variation in the production of SET and its relationship with spike dry weight (SDW) in 130 diverse wheat elite lines and landraces under heat-stressed field conditions. We employed an Illumina iSelect 90K single nucleotide polymorphism (SNP) genotyping array to identify the genetic loci for SET and SDW through a genome-wide association study (GWAS) in a subset of the Wheat Association Mapping Initiative (WAMI) panel. The SET and SDW exhibited appreciable genotypic variation among wheat genotypes at the anthesis stage. There was a strong negative correlation between SET and SDW. The GWAS uncovered five and 32 significant SNPs for SET, and 22 and 142 significant SNPs for SDW, in glasshouse and field conditions, respectively. Some of these SNPs closely localized to the SNPs for plant height, suggesting close associations between plant height and spike-related traits. The phenotypic and genetic elucidation of SET and its relationship with SDW supports future efforts toward gene discovery and breeding wheat cultivars with reduced ethylene effects on yield under heat stress.

Applying 'drought' to potted plants by maintaining suboptimal soil moisture improves plant water relations.

Pot-based phenotyping of drought response sometimes maintains suboptimal soil water content by applying high-frequency deficit irrigation (HFDI). We examined the effect of this treatment on water and abscisic acid (ABA) relations of two species (Helianthus annuus and Populus nigra). Suboptimal soil water content was maintained by frequent irrigation, and compared with the effects of withholding water and with adequate irrigation. At the same average whole-pot soil moisture, frequent irrigation resulted in larger soil water content gradients, lower root and xylem ABA concentrations ([X-ABA]), along with higher transpiration rates or stomatal conductance, compared with plants from which water was withheld. [X-ABA] was not uniquely related to transpiration rate or stomatal conductance, as frequently irrigated plants showed partial stomatal closure compared with well-watered controls, without differing in [X-ABA] and, in H. annuus, [ABA]leaf. In two P. nigra genotypes differing in leaf area, the ratio between leaf area and root weight in the upper soil layer influenced the soil water content of this layer. Maintaining suboptimal soil water content alters water relations, which might become dependent on root distribution and leaf area, which influences soil water content gradients. Thus genotypic variation in 'drought tolerance' derived from phenotyping platforms must be carefully interpreted.

Stomatal and growth responses to hydraulic and chemical changes induced by progressive soil drying.

A better understanding of physiological responses of crops to drought stress is important for ensuring sustained crop productivity under climate change. Here, we studied the effect on 15-day-old maize (Zea mays L.) plants of a 6 d non-lethal period of soil drying [soil water potential (SWP) decreased from -0.20 MPa to -0.81 MPa]. Root growth was initially stimulated during drying (when SWP decreased from -0.31 MPa to -0.38 MPa, compared with -0.29 MPa in well-watered pots), followed by inhibition during Days 5-6 (SWP from -0.63 MPa to -0.81 MPa). Abscisic acid (ABA) in the root began to accumulate as the root water potential declined during Days 2-3. Leaf elongation was inhibited from Day 4 (SWP less than -0.51 MPa), just after leaf ABA content began to increase, but coinciding with a decline in leaf water potential. The stomatal conductance was restricted earlier in the younger leaf (fourth) (on Day 3) than in the older leaf (third). The ethylene content of leaves and roots decreased during drying, but after the respective increase in ABA contents. This work identified critical timing of hydraulic and chemical changes at the onset of soil drying, which can be important in initiating early stomatal and growth responses to drought.

Increased soil phosphorus availability induced by faba bean root exudation stimulates root growth and phosphorus uptake in neighbouring maize.

Root growth is influenced by soil nutrients and neighbouring plants, but how these two drivers affect root interactions and regulate plant growth dynamics is poorly understood. Here, interactions between the roots of maize (Zea mays) and faba bean (Vicia faba) are characterized. Maize was grown alone (maize) or with maize (maize/maize) or faba bean (maize/faba bean) as competitors under five levels of phosphorus (P) supply, and with homogeneous or heterogeneous P distribution. Maize had longer root length and greater shoot biomass and P content when grown with faba bean than with maize. At each P supply rate, faba bean had a smaller root system than maize but greater exudation of citrate and acid phosphatase, suggesting a greater capacity to mobilize P in the rhizosphere. Heterogeneous P availability enhanced the root-length density of maize but not faba bean. Maize root proliferation in the P-rich patches was associated with increased shoot P uptake. Increased P availability by localized P application or by the presence of faba bean exudation stimulated root morphological plasticity and increased shoot growth in maize in the maize/faba bean mixture, suggesting that root interactions of neighbouring plants can be modified by increased P availability.

How can we improve crop genotypes to increase stress resilience and productivity in a future climate? A new crop screening method based on productivity and resistance to abiotic stress.

The need to accelerate the selection of crop genotypes that are both resistant to and productive under abiotic stress is enhanced by global warming and the increase in demand for food by a growing world population. In this paper, we propose a new method for evaluation of wheat genotypes in terms of their resilience to stress and their production capacity. The method quantifies the components of a new index related to yield under abiotic stress based on previously developed stress indices, namely the stress susceptibility index, the stress tolerance index, the mean production index, the geometric mean production index, and the tolerance index, which were created originally to evaluate drought adaptation. The method, based on a scoring scale, offers simple and easy visualization and identification of resilient, productive and/or contrasting genotypes according to grain yield. This new selection method could help breeders and researchers by defining clear and strong criteria to identify genotypes with high resilience and high productivity and provide a clear visualization of contrasts in terms of grain yield production under stress. It is also expected that this methodology will reduce the time required for first selection and the number of first-selected genotypes for further evaluation by breeders and provide a basis for appropriate comparisons of genotypes that would help reveal the biology behind high stress productivity of crops.

Grain production versus resource and environmental costs: towards increasing sustainability of nutrient use in China.

Over the past five decades, Chinese grain production has increased 4-fold, from 110 Mt in 1961 to 557 Mt in 2014, with less than 9% of the world's arable land feeding 22% of the world's population, indicating a substantial contribution to global food security. However, compared with developed economies, such as the USA and the European Union, more than half of the increased crop production in China can be attributed to a rapid increase in the consumption of chemicals, particularly fertilizers. Excessive fertilization has caused low nutrient use efficiency and high environmental costs in grain production. We analysed the key requirements underpinning increased sustainability of crop production in China, as follows: (i) enhance nutrient use efficiency and reduce nutrient losses by fertilizing roots not soil to maximize root/rhizosphere efficiency with innovative root zone nutrient management; (ii) improve crop productivity and resource use efficiency by matching the best agronomic management practices with crop improvement; and (iii) promote technology transfer of the root zone nutrient management to achieve the target of high yields and high efficiency with low environmental risks on a broad scale. Coordinating grain production and environmental protection by increasing the sustainability of nutrient use will be a key step in achieving sustainable crop production in Chinese agriculture.

Deficit irrigation and sustainable water-resource strategies in agriculture for China's food security.

More than 70% of fresh water is used in agriculture in many parts of the world, but competition for domestic and industrial water use is intense. For future global food security, water use in agriculture must become sustainable. Agricultural water-use efficiency and water productivity can be improved at different points from the stomatal to the regional scale. A promising approach is the use of deficit irrigation, which can both save water and induce plant physiological regulations such as stomatal opening and reproductive and vegetative growth. At the scales of the irrigation district, the catchment, and the region, there can be many other components to a sustainable water-resources strategy. There is much interest in whether crop water use can be regulated as a function of understanding of physiological responses. If this is the case, then agricultural water resources can be reallocated to the benefit of the broader community. We summarize the extent of use and impact of deficit irrigation within China. A sustainable strategy for allocation of agricultural water resources for food security is proposed. Our intention is to build an integrative system to control crop water use during different cropping stages and actively regulate the plant's growth, productivity, and development based on physiological responses. This is done with a view to improving the allocation of limited agricultural water resources.

The cadmium-tolerant pea (Pisum sativum L.) mutant SGECdt is more sensitive to mercury: assessing plant water relations.

Heavy metals have multiple effects on plant growth and physiology, including perturbation of plant water status. These effects were assessed by exposing the unique Cd-tolerant and Cd-accumulating pea (Pisum sativum L.) mutant SGECd(t) and its wild-type (WT) line SGE to either cadmium (1, 4 µM CdCl2) or mercury (0.5, 1, 2 µM HgCl2) in hydroponic culture for 12 days. When exposed to Cd, SGECd(t) accumulated more Cd in roots, xylem sap, and shoot, and had considerably more biomass than WT plants. WT plants lost circa 0.2 MPa turgor when grown in 4 µM CdCl2, despite massive decreases in whole-plant transpiration rate and stomatal conductance. In contrast, root Hg accumulation was similar in both genotypes, but WT plants accumulated more Hg in leaves and had a higher stomatal conductance, and root and shoot biomass compared with SGECd(t). Shoot excision resulted in greater root-pressure induced xylem exudation of SGECd(t) in the absence of Cd or Hg and following Cd exposure, whereas the opposite response or no genotypic differences occurred following Hg exposure. Exposing plants that had not been treated with metal to 50 µM CdCl2 for 1h increased root xylem exudation of WT, whereas 50 µM HgCl2 inhibited and eliminated genotypic differences in root xylem exudation, suggesting differences between WT and SGECd(t) plants in aquaporin function. Thus, root water transport might be involved in mechanisms of increased tolerance and accumulation of Cd in the SGECd(t) mutant. However, the lack of cross-tolerance to Cd and Hg stress in the mutant indicates metal-specific mechanisms related to plant adaptation.

Isoprene emission protects photosynthesis but reduces plant productivity during drought in transgenic tobacco (Nicotiana tabacum) plants.

Isoprene protects the photosynthetic apparatus of isoprene-emitting plants from oxidative stress. The role of isoprene in the response of plants to drought is less clear. Water was withheld from transgenic isoprene-emitting and non-emitting tobacco (Nicotiana tabacum) plants, to examine: the response of isoprene emission to plant water deficit; a possible relationship between concentrations of the drought-induced phytohormone abscisic acid (ABA) and isoprene; and whether isoprene affected foliar reactive oxygen species (ROS) and lipid peroxidation levels. Isoprene emission did not affect whole-plant water use, foliar ABA concentration or leaf water potential under water deficit. Compared with well-watered controls, droughted non-emitting plants significantly increased ROS content (31-46%) and lipid peroxidation (30-47%), concomitant with decreased operating and maximum efficiencies of photosystem II photochemistry and lower leaf and whole-plant water use efficiency (WUE). Droughted isoprene-emitting plants showed no increase in ROS content or lipid peroxidation relative to well-watered controls, despite isoprene emission decreasing before leaf wilting. Although isoprene emission protected the photosynthetic apparatus and enhanced leaf and whole-plant WUE, non-emitting plants had 8-24% more biomass under drought, implying that isoprene emission incurred a yield penalty.

Ethylene limits abscisic acid- or soil drying-induced stomatal closure in aged wheat leaves.

The mechanism of age-induced decreased stomatal sensitivity to abscisic acid (ABA) and soil drying has been explored here. Older, fully expanded leaves partly lost their ability to close stomata in response to foliar ABA sprays, and soil drying which stimulated endogenous ABA production, while young fully expanded leaves closed their stomata more fully. However, ABA- or soil drying-induced stomatal closure of older leaves was partly restored by pretreating plants with 1-methylcyclopropene (1-MCP), which can antagonize ethylene receptors, or by inoculating soil around the roots with the rhizobacterium Variovorax paradoxus 5C-2, which contains 1-aminocyclopropane-1-carboxylic acid (ACC)-deaminase. ACC (the immediate biosynthetic precursor of ethylene) sprays revealed higher sensitivity of stomata to ethylene in older leaves than younger leaves, despite no differences in endogenous ACC concentrations or ethylene emission. Taken together, these results indicate that the relative insensitivity of stomatal closure to ABA and soil drying in older leaves is likely due to altered stomatal sensitivity to ethylene, rather than ethylene production. To our knowledge, this is the first study to mechanistically explain diminished stomatal responses to soil moisture deficit in older leaves, and the associated reduction in leaf water-use efficiency.

The rhizobacterium Variovorax paradoxus 5C-2, containing ACC deaminase, promotes growth and development of Arabidopsis thaliana via an ethylene-dependent pathway.

Many plant-growth-promoting rhizobacteria (PGPR) associated with plant roots contain the enzyme 1-aminocyclopropane-1-carboxylate (ACC) deaminase and can metabolize ACC, the immediate precursor of the plant hormone ethylene, thereby decreasing plant ethylene production and increasing plant growth. However, relatively few studies have explicitly linked ethylene emission and/or action to growth promotion in these plant-microbe interactions. This study examined effects of the PGPR Variovorax paradoxus 5C-2 containing ACC deaminase on the growth and development of Arabidopsis thaliana using wild-type (WT) plants and several ethylene-related mutants (etr1-1, ein2-1, and eto1-1). Soil inoculation with V. paradoxus 5C-2 promoted growth (leaf area and shoot biomass) of WT plants and the ethylene-overproducing mutant eto1-1, and also enhanced floral initiation of WT plants by 2.5 days. However, these effects were not seen in ethylene-insensitive mutants (etr1-1 and ein2-1) even though bacterial colonization of the root system was similar. Furthermore, V. paradoxus 5C-2 decreased ACC concentrations of rosette leaves of WT plants by 59% and foliar ethylene emission of both WT plants and eto1-1 mutants by 42 and 37%, respectively. Taken together, these results demonstrate that a fully functional ethylene signal transduction pathway is required for V. paradoxus 5C-2 to stimulate leaf growth and flowering of A. thaliana.

Eco-physiology and environmental impacts of newly developed rice genotypes for improved yield and nitrogen use efficiency coordinately.

Significant advancements have been made in understanding the genetic regulation of nitrogen use efficiency (NUE) and identifying crucial NUE genes in rice. However, the development of rice genotypes that simultaneously exhibit high yield and NUE has lagged behind these theoretical advancements. The grain yield, NUE, and greenhouse gas (GHG) emissions of newly-bred rice genotypes under reduced nitrogen application remain largely unknown. To address this knowledge gap, field experiments were conducted, involving 80 indica (14 to 19 rice genotypes each year in Wuxue, Hubei) and 12 japonica (8 to 12 rice genotypes each year in Yangzhou, Jiangsu). Yield, NUE, agronomy, and soil parameters were assessed, and climate data were recorded. The experiments aimed to assess genotypic variations in yield and NUE among these genotypes and to investigate the eco-physiological basis and environmental impacts of coordinating high yield and high NUE. The results showed significant variations in yield and NUE among the genotypes, with 47 genotypes classified as moderate-high yield with high NUE (MHY_HNUE). These genotypes demonstrated the higher yields and NUE levels, with 9.6 t ha-1, 54.4 kg kg-1, 108.1 kg kg-1, and 64 % for yield, NUE for grain and biomass production, and N harvest index, respectively. Nitrogen uptake and tissue concentration were key drivers of the relationship between yield and NUE, particularly N uptake at heading and N concentrations in both straw and grain at maturity. Increase in pre-anthesis temperature consistently lowered yield and NUE. Genotypes within the MHY_HNUE group exhibited higher methane emissions but lower nitrous oxide emissions compared to those in the low to middle yield and NUE group, resulting in a 12.8 % reduction in the yield-scaled greenhouse gas balance. In conclusion, prioritizing crop breeding efforts on yield and resource use efficiency, as well as developing genotypes resilient to high temperatures with lower GHGs, can mitigate planetary warming.

How is ozone pollution reducing our food supply?

Ground-level ozone pollution is already decreasing global crop yields (from ∼2.2-5.5% for maize to 3.9-15% and 8.5-14% for wheat and soybean, respectively), to differing extents depending on genotype and environmental conditions, and this problem is predicted to escalate given climate change and increasing ozone precursor emissions in many areas. Here a summary is provided of how ozone pollution affects yield in a variety of crops, thus impacting global food security. Ozone causes visible injury symptoms to foliage; it induces early senescence and abscission of leaves; it can reduce stomatal aperture and thereby carbon uptake, and/or directly reduce photosynthetic carbon fixation; it can moderate biomass growth via carbon availability or more directly; it can decrease translocation of fixed carbon to edible plant parts (grains, fruits, pods, roots) due either to reduced availability at source, redirection to synthesis of chemical protectants, or reduced transport capabilities via phloem; decreased carbon transport to roots reduces nutrient and water uptake and affects anchorage; ozone can moderate or bring forward flowering and induce pollen sterility; it induces ovule and/or grain abortion; and finally it reduces the ability of some genotypes to withstand other stresses such as drought, high vapour pressure deficit, and high photon flux density via effects on stomatal control. This latter point is emphasized here, given predictions that atmospheric conditions conducive to drought formation that also give rise to intense precursor emission events will become more severe over the coming decades.

Improving crop productivity and resource use efficiency to ensure food security and environmental quality in China.

In recent years, agricultural growth in China has accelerated remarkably, but most of this growth has been driven by increased yield per unit area rather than by expansion of the cultivated area. Looking towards 2030, to meet the demand for grain and to feed a growing population on the available arable land, it is suggested that annual crop production should be increased to around 580 Mt and that yield should increase by at least 2% annually. Crop production will become more difficult with climate change, resource scarcity (e.g. land, water, energy, and nutrients) and environmental degradation (e.g. declining soil quality, increased greenhouse gas emissions, and surface water eutrophication). To pursue the fastest and most practical route to improved yield, the near-term strategy is application and extension of existing agricultural technologies. This would lead to substantial improvement in crop and soil management practices, which are currently suboptimal. Two pivotal components are required if we are to follow new trajectories. First, the disciplines of soil management and agronomy need to be given increased emphasis in research and teaching, as part of a grand food security challenge. Second, continued genetic improvement in crop varieties will be vital. However, our view is that the biggest gains from improved technology will come most immediately from combinations of improved crops and improved agronomical practices. The objectives of this paper are to summarize the historical trend of crop production in China and to examine the main constraints to the further increase of crop productivity. The paper provides a perspective on the challenge faced by science and technology in agriculture which must be met both in terms of increased crop productivity but also in increased resource use efficiency and the protection of environmental quality.

Plant hormone interactions: innovative targets for crop breeding and management.

Here we highlight how both the root and shoot environment impact on whole plant hormone balance, particularly under stresses such as soil drying, and relate hormone ratios and relative abundances to processes influencing plant performance and yield under both mild and more severe stress. We discuss evidence (i) that abscisic acid (ABA) and ethylene act antagonistically on grain-filling rate amongst other yield-impacting processes; (ii) that ABA's effectiveness as an agent of stomatal closure can be modulated by coincident ethylene or cytokinin accumulation; and (iii) that enhanced cytokinin production can increase growth and yield by improving foliar stay-green indices under stress, and by improving processes that impact grain-filling and number, and that this can be the result of altered relative abundances of cytokinin and ABA (and other hormones). We describe evidence and novel processes whereby these phenomena are/could be amenable to manipulation through genetic and management routes, such that plant performance and yield can be improved. We explore the possibility that a range of ABA-ethylene and ABA-cytokinin relative abundances could represent targets for breeding/managing for yield resilience under a spectrum of stress levels between severe and mild, and could circumvent some of the pitfalls so far encountered in the massive research effort towards breeding for increases in the complex trait of yield.

Multiple impacts of the plant growth-promoting rhizobacterium Variovorax paradoxus 5C-2 on nutrient and ABA relations of Pisum sativum.

Resolving the physiological mechanisms by which rhizobacteria enhance plant growth is difficult, since many such bacteria contain multiple plant growth-promoting properties. To understand further how the 1-aminocyclopropane-1-carboxylate (ACC) deaminase (ACCd)-containing rhizobacterium Variovorax paradoxus 5C-2 affects plant growth, the flows and partitioning of mineral nutrients and abscisic acid (ABA) and ABA metabolism were studied in pea (Pisum sativum) plants following rhizosphere bacterial inoculation. Although root architecture was not affected, inoculation increased root and shoot biomass, and stomatal conductance, by 20, 15, and 24%, respectively, and increased N, P, K, Ca, and Mg uptake by 16, 81, 50, 46, and 58%, respectively. P deposition in inoculated plant roots was 4.9 times higher than that in uninoculated controls. Rhizobacterial inoculation increased root to shoot xylem flows and shoot to root phloem flows of K by 1.8- and 2.1-fold, respectively. In control plants, major sinks for K deposition were the roots and upper shoot (43% and 49% of total uptake, respectively), while rhizobacterial inoculation increased K distribution to the lower shoot at the expense of other compartments (xylem, phloem, and upper shoot). Despite being unable to metabolize ABA in vitro, V. paradoxus 5C-2 decreased root ABA concentrations and accumulation by 40-60%. Although inoculation decreased xylem ABA flows, phloem ABA flows increased. Whether bacterial ACCd attenuates root to shoot ABA signalling requires further investigation, since ABA is critical to maintain growth of droughted plants, and ACCd-containing organisms have been advocated as a means of minimizing growth inhibition of plants in drying soil.