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BACKGROUND: Nitrate accumulates in plants in response to N supply, aerial environment (predominantly light), and genotype. This paper characterises the effects of genotype, environment, and their interactions on nitrate accumulation by 24 cultivated and wild lettuce accessions grown hydroponically in winter and summer. The results will inform future strategies for selecting for low-nitrate varieties. RESULTS: A preliminary study in which two accessions were sampled for nitrate over time showed largest differences between cultivars in the early-middle period of growth. Sampling the whole population of lettuce at this stage revealed significant effects of genotype, environment (with nitrate concentrations generally higher in winter), and genotype x environment interactions (largely due to a wider range of concentrations in summer). Changes in the ranking of accessions for nitrate accumulation between the two growing seasons were generally small for cultivated morphotypes. Shoot nitrate concentrations and water contents were positively associated, particularly in summer when separate relationships for different cultivated morphotypes (butterhead, cos/Romaine, crisp, leaf, and stem lettuce) were detected. Expressing nitrate concentration on either a shoot fresh or dry matter basis had relatively little effect on the ranking of most cultivated accessions, but not for the wild types. CONCLUSION: There is a well-defined sampling window when differences in nitrate accumulation between lettuce genotypes are at a maximum. Delaying sampling may allow morphological changes in head form to mask earlier genotypic differences. Genotype x environment interactions are predominantly of the non-crossover type and have only a small effect on changes in the ranking of accessions between seasons, allowing selections to be made at any time of year. At least part of the genotypic variation in nitrate accumulation is associated with differences in shoot water content. (C) 2010 Society of Chemical Industry

Human individuals require at least 20 inorganic elements (‘minerals’) for normal functioning. However, much of the world's population is probably deficient in one or more essential minerals and at increased risk of physiological disorders. Addressing these ‘hidden hungers’ is a challenge for the nutrition and agriculture sectors. Mineral deficiencies among populations are typically identified from dietary surveys because (1) minerals are acquired primarily from dietary sources and (2) (bio)assays of mineral status can be unreliable. While dietary surveys are likely to under-report energy intakes, surveys show that 9% of all UK and US adults consume Ca and Mg, and 14% of adults consume K, at quantities below the UK lower reference nutrient intake, and are therefore at risk of deficiency. Low dietary Ca, Mg and K intakes can be caused by energy-malnourishment and by cultural and economic factors driving dietary conservatism. For example, cereal grains routinely displace vegetables and fruits in the diet. Cereal grains have low concentrations of several minerals, notably Ca, as a consequence of their physiology. Low grain mineral concentrations are compounded when cereal crops are grown in soils of low mineral phytoavailability and when grain is processed. In this paper, the impact of increased vegetable consumption and horticultural biofortification, i.e. enhancing crop mineral content through breeding and agronomy, on intakes of the major minerals Ca, Mg and K is assessed. Despite low energy intake from horticultural crops generally, increased vegetable consumption and biofortification would significantly improve dietary intakes of Ca, Mg and K.

Background: Dietary recommendations for selenium differ between countries, mainly because of uncertainties over the definition of optimal selenium status. Objective: The objective was to examine the dose-response relations for different forms of selenium. Design: A randomized, double-blind, placebo-controlled dietary intervention was carried out in 119 healthy men and women aged 50‐ 64 y living in the United Kingdom. Daily placebo or seleniumenriched yeast tablets containing 50, 100, or 200 l gS e (’60% selenomethionine), selenium-enriched onion meals (’66% cglutamyl-methylselenocysteine, providing the equivalent of 50 lg Se/d), or unenriched onion meals were consumed for 12 wk. Changes in platelet glutathione peroxidase activity and in plasma selenium and selenoprotein P concentrations were measured. Results: The mean baseline plasma selenium concentration for all subjects was 95.7 6 11.5 ng/mL, which increased significantly by 10 wk to steady state concentrations of 118.3 6 13.1, 152.0 6 24.3, and 177.4 6 26.3 ng/mL in those who consumed 50, 100, or 200 lg Se-yeast/d, respectively. Platelet glutathione peroxidase activity did not change significantly in response to either dose or form of selenium. Selenoprotein P increased significantly in all selenium intervention groups from an overall baseline mean of 4.99 6 0.80 lg/mL to 6.17 6 0.85, 6.73 6 1.01, 6.59 6 0.64, and 5.72 6 0.75 lg/mL in those who consumed 50, 100, or 200 lg Se-yeast/d and 50 lg Seenriched onions/d, respectively. Conclusions: Plasma selenoprotein P is a useful biomarker of status in populations with relatively low selenium intakes because it responds to different dietary forms of selenium. To optimize the plasma selenoprotein P concentration in this study, 50 lg Se/d was required in addition to the habitual intake of ’55 lg/d. In the context of established relations between plasma selenium and risk of cancer and mortality, and recognizing the important functions of selenoprotein P, these results provide important evidence for deriving estimated average requirements for selenium in adults. This trial was registered at clinicaltrials.gov as NCT00279812. Am J Clin Nutr 2010;91:923‐31.

There is concern that modern cultivars and/or agronomic practices have resulted in reduced concentrations of mineral elements essential to human nutrition in edible crops. Increased yields are often associated with reduced concentrations of mineral elements in produce, and a number of recent studies have indicated that, when grown under identical conditions, the concentrations of several mineral elements are lower in genotypes yielding more grain or shoot biomass than in older, lower-yielding genotypes. Potato is a significant crop, grown worldwide, yet few studies have investigated whether increasing yields, through agronomy or breeding, affects the concentrations of mineral elements in tubers. This article examines the hypothesis that increasing yields, either by the application of mineral fertilizers and/or by growing higher-yielding varieties, leads to decreased concentrations of mineral elements in tubers. It reports that the application of fertilizers influences tuber elemental composition in a complex manner, presumably as a consequence of soil chemistry and interactions between mineral elements within the plant, that considerable variation exists between potato genotypes in the concentrations of mineral elements in their tubers, and that, like in other crops, higher-yielding genotypes occasionally have lower concentrations of some mineral elements in their edible tissues than lower-yielding genotypes.

Summary The diets of over two-thirds of the world's population lack one or more essential mineral elements. This can be remedied through dietary diversification, mineral supplementation, food fortification, or increasing the concentrations and/or bioavailability of mineral elements in produce (biofortification). This article reviews aspects of soil science, plant physiology and genetics underpinning crop biofortification strategies, as well as agronomic and genetic approaches currently taken to biofortify food crops with the mineral elements most commonly lacking in human diets: iron (Fe), zinc (Zn), copper (Cu), calcium (Ca), magnesium (Mg), iodine (I) and selenium (Se). Two complementary approaches have been successfully adopted to increase the concentrations of bioavailable mineral elements in food crops. First, agronomic approaches optimizing the application of mineral fertilizers and/or improving the solubilization and mobilization of mineral elements in the soil have been implemented. Secondly, crops have been developed with: increased abilities to acquire mineral elements and accumulate them in edible tissues; increased concentrations of ‘promoter’ substances, such as ascorbate, β-carotene and cysteine-rich polypeptides which stimulate the absorption of essential mineral elements by the gut; and reduced concentrations of ‘antinutrients’, such as oxalate, polyphenolics or phytate, which interfere with their absorption. These approaches are addressing mineral malnutrition in humans globally.

Summary • The transcriptome of an organism is its set of gene transcripts (mRNAs) at a defined spatial and temporal locus. Because gene expression is affected markedly by environmental and developmental perturbations, it is widely assumed that transcriptome divergence among taxa represents adaptive phenotypic selection. This assumption has been challenged by neutral theories which propose that stochastic processes drive transcriptome evolution. • To test for evidence of neutral transcriptome evolution in plants, we quantified 18 494 gene transcripts in nonsenescent leaves of 14 taxa of Brassicaceae using robust cross-species transcriptomics which includes a two-step physical and in silico-based normalization procedure based on DNA similarity among taxa. • Transcriptome divergence correlates positively with evolutionary distance between taxa and with variation in gene expression among samples. Results are similar for pseudogenes and chloroplast genes evolving at different rates. Remarkably, variation in transcript abundance among root-cell samples correlates positively with transcriptome divergence among root tissues and among taxa. • Because neutral processes affect transcriptome evolution in plants, many differences in gene expression among or within taxa may be nonfunctional, reflecting ancestral plasticity and founder effects. Appropriate null models are required when comparing transcriptomes in space and time.

Major research efforts are targeting the improved performance of root systems for more efficient use of water and nutrients by crops. However, characterizing root system architecture (RSA) is challenging, because roots are difficult objects to observe and analyse. A model-based analysis of RSA traits from phenotyping image data is presented. The model can successfully back-calculate growth parameters without the need to measure individual roots. The mathematical model uses partial differential equations to describe root system development. Methods based on kernel estimators were used to quantify root density distributions from experimental image data, and different optimization approaches to parameterize the model were tested. The model was tested on root images of a set of 89 Brassica rapa L. individuals of the same genotype grown for 14 d after sowing on blue filter paper. Optimized root growth parameters enabled the final (modelled) length of the main root axes to be matched within 1% of their mean values observed in experiments. Parameterized values for elongation rates were within ±4% of the values measured directly on images. Future work should investigate the time dependency of growth parameters using time-lapse image data. The approach is a potentially powerful quantitative technique for identifying crop genotypes with more efficient root systems, using (even incomplete) data from high-throughput phenotyping systems.

Rooting media used in current root phenotyping studies can have substantial effect. In this study, the effects of three different nutrient conducting papers (Black construction paper, Anchor blue germination paper and Kimpak paper) and soil-filled boxes on root growth and root system architecture (RSA) of Brassica rapa (cultivars ‘R500’ and ‘IMB211’) were investigated. Seedlings of the two B. rapa genotypes were supplied with nutrients on the nutrient conducting papers and in the soil-filled boxes. The papers and soil-filled boxes were fixed to flatbed scanners and two-dimensional images of roots were periodically taken and analysed. Root media effects on shoot and root biomass and on topological indices (TI) were observed. For example, root branching was more pronounced on the construction paper. Mean TI of 0.82 and 0.93, recorded for R500 and IMB211, respectively, on the construction paper indicated that substrates affect the herringbone pattern of brassica roots. Whilst it was indicated that different results could be obtained for the same RSA when different germination papers are used, the results showed that Anchor blue germination paper is an ideal proxy for soil in phenotyping seedlings for RSA traits and root growth.

Zinc (Zn) is an essential component of thousands of proteins in plants, although it is toxic in excess. In this review, the dominant fluxes of Zn in the soil-root-shoot continuum are described, including Zn inputs to soils, the plant availability of soluble Zn(2+) at the root surface, and plant uptake and accumulation of Zn. Knowledge of these fluxes can inform agronomic and genetic strategies to address the widespread problem of Zn-limited crop growth. Substantial within-species genetic variation in Zn composition is being used to alleviate human dietary Zn deficiencies through biofortification. Intriguingly, a meta-analysis of data from an extensive literature survey indicates that a small proportion of the genetic variation in shoot Zn concentration can be attributed to evolutionary processes whose effects manifest above the family level. Remarkable insights into the evolutionary potential of plants to respond to elevated soil Zn have recently been made through detailed anatomical, physiological, chemical, genetic and molecular characterizations of the brassicaceous Zn hyperaccumulators Thlaspi caerulescens and Arabidopsis halleri.

The aim of this study was to characterise nutritional-I status in Malawi. Dietary-I intakes were assessed using new datasets of crop, fish, salt and water-I concentrations, while I status was assessed for 60 women living on each of calcareous and non-calcareous soils as defined by urinary iodine concentration (UIC). Iodine concentration in staple foods was low, with median concentrations of 0.01 mg kg−1 in maize grain, 0.008 mg kg−1 in roots and tubers, but 0.155 mg kg−1 in leafy vegetables. Freshwater fish is a good source of dietary-I with a median concentration of 0.51 mg kg−1. Mean Malawian dietary-Iodine intake from food, excluding salt, was just 7.8 μg d−1 compared to an adult requirement of 150 μg d−1. Despite low dietary-I intake from food, median UICs were 203 μg L−1 with only 12% defined as I deficient whilst 21% exhibited excessive I intake. Iodised salt is likely to be the main source of dietary I intake in Malawi; thus, I nutrition mainly depends on the usage and concentration of I in iodised salt. Drinking water could be a significant source of I in some areas, providing up to 108 μg d−1 based on consumption of 2 L d−1.

Background In this review, we examine the potential of Zn-enriched fertilisers to alleviate human dietary Zn deficiency. The focus is on ten African countries where dietary Zn supply is low and where fertiliser subsidies are routinely deployed on cereal crops. Scope Dietary Zn supply and deficiency prevalence were quantified from food supply and composition data. Typical effects of soil (granular) and foliar Zn applications on Zn concentrations in maize (Zea mays L.), rice (Oryza sativa L.) and wheat (Triticum aestivum L.) grains were based on a systematic literature review. Reductions in disease burdens attributable to Zn deficiency and cost-effectiveness were estimated using a disability-adjusted life years (DALYs) approach. Conclusions Baseline Zn supply in 2009 ranged from 7.1 (Zambia) to 11.9 (Mali) mg capita−1 day−1; prevalence of Zn deficiency ranged from 24 (Nigeria) to 66 % (Zambia). In reviewed studies, soil Zn application led to an increase in median Zn concentration in maize, rice and wheat grains of 23, 7 and 19 %; foliar application led to increases of 30, 25 and 63 %. Enriching granular fertilisers within current subsidy schemes would be most effective in Malawi, reducing DALYs lost due to Zn deficiency by 10 %. The cost per DALY saved ranged from US$ 624 to 5893 via granular fertilisers and from US$ 46 to 347 via foliar fertilisers. Foliar applications are likely to be more cost effective than soil applications due to fixation of Zn in the soil but may be more difficult to deploy. Zinc fertilisation is likely to be less cost-effective than breeding in the longer term although other micronutrients such as selenium could be incorporated.

Improved methods for selecting cultivars for their ability to grow well on low P soils will improve productivity and minimise pollution. Estimates are required of the relative values for different genotypes of physiologically meaningful parameters that define yield response to increasing levels of plant available soil P. The object of this study was to devise ways of obtaining these estimates from conventional trials that include numerous genotypes but only two P treatments. The parameters were estimated as coefficients of a modified Michaelis–Menten equation, namely A, the maximum yield that can be obtained with ample P, Km, the concentration of plant available P in the rooting medium at which yield is half the maximum, and B, the gradient of yield against available P as it approaches the origin. Two novel methods of estimation were devised. They both require inputs of yield and its variance at each of two concentrations of available P, but these must be within certain limits. Estimates of parameter values for 12 Brassica oleracea genotypes made by these methods from two P-levels in each of two experiments with six P levels were compared with the values obtained by conventional fitting to the data from all six P levels. According to regression and ranking analyses, the relative values of A obtained for the different genotypes by both two P-level methods were similar to those from the six P-level method and both were reproducible. None of the methods however detected any reproducible differences in Km. Application of one of the two P-level methods to published data for genotypes of other species indicated that there was little intra-species variation in Km within some data sets on maize, beans and leguminous cover crops but much within other data sets. Sensitivity and algebraic analyses are presented to define the experimental conditions required for successful use of two P-level methods. It is concluded that, provided these experimental conditions are met, measurements at two P levels can give almost as much information about intra-species differences in P response parameters as measurements at six P levels. The procedure may also be useful for interpreting other responses of a “diminishing returns” type such as those to other nutrients or plant spacing.

Background and Aims Silicon (Si) in plants provides structural support and improves tolerance to diseases, drought and metal toxicity. Shoot Si concentrations are generally considered to be greater in monocotyledonous than in non-monocot plant species. The phylogenetic variation in the shoot Si concentration of plants reported in the primary literature has been quantified. Methods Studies were identified which reported Si concentrations in leaf or non-woody shoot tissues from at least two plant species growing in the same environment. Each study contained at least one species in common with another study. Key Results Meta-analysis of the data revealed that, in general, ferns, gymnosperms and angiosperms accumulated less Si in their shoots than non-vascular plant species and horsetails. Within angiosperms and ferns, differences in shoot Si concentration between species grouped by their higher-level phylogenetic position were identified. Within the angiosperms, species from the commelinoid monocot orders Poales and Arecales accumulated substantially more Si in their shoots than species from other monocot clades. Conclusions A high shoot Si concentration is not a general feature of monocot species. Information on the phylogenetic variation in shoot Si concentration may provide useful palaeoecological and archaeological information, and inform studies of the biogeochemical cycling of Si and those of the molecular genetics of Si uptake and transport in plants.

Better criteria for selecting cultivars for their ability to grow well on low-P soils could reduce waste of fertilizer P. The objective of this study was to quantify and determine the reproducibility of two crop parameters (A and K m ) responsible for differences in the response of 12 Brassica oleracea L. genotypes to P supply. Modified Michaelis-Menten equations were fitted to shoot dry weight responses to P supply for each genotype in each of three glasshouse and three field experiments. One of the fitted parameters (A) defined the maximum shoot dry weight that could be obtained with ample P, and the other (K m ) defined the efficiency of root P-uptake as the concentration of extractable soil P in the rooting medium at which growth was half the maximum. The equations fitted the data well. Ranking and regression analyses showed that intergenotypic differences in A were considerable, and reproducible, but that differences in K m were small. Thus, B. oleracea genotypes yielding most on a P-sufficient soil will also yield most on a P-deficient soil. However, both the maximum yield (A) and the efficiency of root-P uptake (K m ) vary considerably between different vegetable crops. Thus, a graphical procedure to select crops for highest yields on low-P soils, on the basis of a comparison of A and K m values, is described.

SummaryHistorical variation in the mineral composition of edible horticultural products was determined from UK and USA food survey data. From these data, it was possible to measure the variation in the mineral composition of edible horticultural products in general, and in edible horticultural products grouped as vegetables, fruits or nuts, in the 1930s and in the 1980s (or later) for both countries. Thus, the hypothesis that the mineral composition of edible horticultural products had altered since the 1930s was tested. The average concentrations of Cu, Mg and Na in the dry matter of vegetables, and the average concentrations of Cu, Fe and K in fruits decreased significantly between the 1930s and the 1980s in the UK. The same hypothesis was tested with comparable data from the USA, whose historical horticultural and consumer practices have paralleled those of the UK. Data from the USA showed that the average Ca, Cu and Fe concentrations in the dry matter of vegetables, and the average concentrations of C...

Assessment of the transfer of radionuclides to wild species is an important component in the estimation of predicted doses to biota. Reviews of available data for the many potential radionuclide-biota combinations which may be required for environmental assessments highlight many data gaps for terrestrial species. Here, we discuss different approaches which have been suggested to compensate for these data gaps. All of the reviewed approaches have merit; however, there is a requirement for transparency in methodology and data provenance which in some instances is currently missing. Furthermore, there is a need to validate the various methodologies to enable their use with confidence. The requirements of improving our ability to predict radionuclide transfer to wild species are discussed and recommendations made.

Cesium (Cs) is chemically similar to potassium (K). However, although K is an essential element, Cs is toxic to plants. Two contrasting hypotheses to explain Cs toxicity have been proposed: (1) extracellular Cs+ prevents K+ uptake and, thereby, induces K starvation; and (2) intracellular Cs+ interacts with vital K+-binding sites in proteins, either competitively or noncompetitively, impairing their activities. We tested these hypotheses with Arabidopsis (Arabidopsis thaliana). Increasing the Cs concentration in the agar ([Cs]agar) on which Arabidopsis were grown reduced shoot growth. Increasing the K concentration in the agar ([K]agar) increased the [Cs]agar at which Cs toxicity was observed. However, although increasing [Cs]agar reduced shoot K concentration ([K]shoot), the decrease in shoot growth appeared unrelated to [K]shoot per se. Furthermore, the changes in gene expression in Cs-intoxicated plants differed from those of K-starved plants, suggesting that Cs intoxication was not perceived genetically solely as K starvation. In addition to reducing [K]shoot, increasing [Cs]agar also increased shoot Cs concentration ([Cs]shoot), but shoot growth appeared unrelated to [Cs]shoot per se. The relationship between shoot growth and [Cs]shoot/[K]shoot suggested that, at a nontoxic [Cs]shoot, growth was determined by [K]shoot but that the growth of Cs-intoxicated plants was related to the [Cs]shoot/[K]shoot quotient. This is consistent with Cs intoxication resulting from competition between K+ and Cs+ for K+-binding sites on essential proteins.

BackgroundPhosphorus (P) is an essential macronutrient for plants. Plants take up P as phosphate (Pi) from the soil solution. Since little Pi is available in most soils, P fertilizers are applied to crops. However, the use of P fertilizers is unsustainable and may cause pollution. Consequently, there is a need to develop more P-use-efficient (PUE) crops and precise methods to monitor crop P-status. Scope Manipulating the expression of genes to improve the PUE of crops could reduce their P fertilizer requirement. This has stimulated research towards the identification of genes and signalling cascades involved in plant responses to P deficiency. Genes that respond to P deficiency can be grouped into ‘early’ genes that respond rapidly and often non-specifically to P deficiency, or ‘late’ genes that impact on the morphology, physiology or metabolism of plants upon prolonged P deficiency. SummaryThe use of micro-array technology has allowed researchers to catalogue the genetic responses of plants to P deficiency. Genes whose expression is altered by P deficiency include various transcription factors, which are thought to coordinate plant responses to P deficiency, and other genes involved in P acquisition and tissue P economy. Several common cis-regulatory elements have been identified in the promoters of these genes, suggesting that their expression might be coordinated. It is suggested that knowledge of the genes whose expression changes in response to P deficiency might allow the development of crops with improved PUE, and could be used in diagnostic techniques to monitor P deficiency in crops either directly using ‘smart’ indicator plants or indirectly through transcript profiling. The development of crops with improved PUE and the adoption of diagnostic technology could reduce production costs, minimize the use of a non-renewable resource, reduce pollution and enhance biodiversity. a 2004 Annals of Botany Company

Selenium (Se) is an essential plant micronutrient, but is toxic at high tissue concentrations. It is chemically similar to sulphur (S), an essential plant macronutrient. The interactions between Se and S nutrition were investigated in the model plant Arabidopsis thaliana (L.) Heynh. Arabidopsis plants were grown on agar containing a complete mineral complement and various concentrations of selenate and sulphate. The Se/S concentration ratio in the shoot ([Se](shoot)/[S](shoot)) showed a complex dependence on the ratio of selenate to sulphate concentration in the agar ([Se](agar)/[S](agar)). Increasing [S](agar) increased shoot fresh weight (FW) and [S](shoot), but decreased [Se](shoot). Increasing [Se](agar) increased both [Se](shoot) and [S](shoot), but reduced shoot FW. The reduction in shoot FW in the presence of Se was linearly related to the shoot Se/S concentration ratio. These data suggest (i) that Se and S enter Arabidopsis through multiple transport pathways with contrasting sulphate/selenate selectivities, whose activities vary between plants of contrasting nutritional status, (ii) that rhizosphere sulphate inhibits selenate uptake, (iii) that rhizosphere selenate promotes sulphate uptake, possibly by preventing the reduction in the abundance and/or activity of sulphate transporters by sulphate and/or its metabolites, and (iv) that Se toxicity occurs because Se and S compete for a biochemical process, such as assimilation into amino acids of essential proteins.

Summary • Ingestion of caesium (Cs) radioisotopes poses a health risk to humans. Crop varieties that accumulate less Cs in their edible tissues may provide a useful countermeasure. This study was performed to determine whether quantitative genetics on a model plant (Arabidopsis thaliana) might inform such ‘safe’-crop strategies. • Arabidopsis accessions and recombinant inbred lines (RILs), from Landsberg erecta (Ler) × Cape Verdi Island (Cvi), Ler × Columbia (Col), and Niederzenz (Nd) × Col mapping populations, were grown on agar supplemented with subtoxic levels of Cs. • Shoot Cs concentration varied up to three-fold, and shoot f. wt varied up to 25-fold within populations. The heritability of growth and Cs accumulation traits ranged from 0.06 to 0.28. Four quantitative trait loci (QTL) accounted for > 80% of the genetic contribution to the total phenotypic variation in shoot Cs concentration in the Ler × Col population. • QTL identified in this study, in particular, QTL co-localizing to the top and bottom regions of Chromosomes I and V in two different mapping populations, are amenable to positional cloning and, through collinearity, may inform selection or breeding strategies for the development of ‘safe’ crops.

The calcium (Ca) concentration of plant shoot tissues varies systematically between angiosperm orders. The phylogenetic variation in the shoot concentration of other mineral nutrients has not yet been described at an ordinal level. The aims of this study were (1) to quantify the shoot mineral concentration of different angiosperm orders, (2) to partition the phylogenetic variation in shoot mineral concentration between and within orders, (3) to determine if the shoot concentration of different minerals are correlated across angiosperm species, and (4) to compare experimental data with published ecological survey data on 81 species sampled from their natural habitats. Species, selected pro rata from different angiosperm orders, were grown in a hydroponic system under a constant external nutrient regime. Shoots of 117 species were sampled during vegetative growth. Significant variation in shoot carbon (C), calcium (Ca), potassium (K), and magnesium (Mg) concentration occurred between angiosperm orders. There was no evidence for systematic differences in shoot phosphorus (P) or organic-nitrogen (N) concentration between orders. At a species level, there were strong positive correlations between shoot Ca and Mg concentration, between shoot P and organic-N concentration, and between shoot K concentration and shoot fresh weight:dry weight ratio. Shoot C and cation concentration correlated negatively at a species level. Species within the Poales and the Caryophyllales had distinct shoot mineralogies in both the designed experiment and in the ecological survey.

Our aim was to generate and prove the concept of “smart” plants to monitor plant phosphorus (P) status in Arabidopsis. Smart plants can be genetically engineered by transformation with a construct containing the promoter of a gene up-regulated specifically by P starvation in an accessible tissue upstream of a marker gene such as β-glucuronidase (GUS). First, using microarrays, we identified genes whose expression changed more than 2.5-fold in shoots of plants growing hydroponically when P, but not N or K, was withheld from the nutrient solution. The transient changes in gene expression occurring immediately (4 h) after P withdrawal were highly variable, and many nonspecific, shock-induced genes were up-regulated during this period. However, two common putative cis-regulatory elements (a PHO-like element and a TATA box-like element) were present significantly more often in the promoters of genes whose expression increased 4 h after the withdrawal of P compared with their general occurrence in the promoters of all genes represented on the microarray. Surprisingly, the expression of only four genes differed between shoots of P-starved and -replete plants 28 h after P was withdrawn. This lull in differential gene expression preceded the differential expression of a new group of 61 genes 100 h after withdrawing P. A literature survey indicated that the expression of many of these “late” genes responded specifically to P starvation. Shoots had reduced P after 100 h, but growth was unaffected. The expression of SQD1, a gene involved in the synthesis of sulfolipids, responded specifically to P starvation and was increased 100 h after withdrawing P. Leaves of Arabidopsis bearing a SQD1::GUS construct showed increased GUS activity after P withdrawal, which was detectable before P starvation limited growth. Hence, smart plants can monitor plant P status. Transferring this technology to crops would allow precision management of P fertilization, thereby maintaining yields while reducing costs, conserving natural resources, and preventing pollution.

Phytoextraction is the removal of metals from contaminated soils into harvested plant tissues. The rate of phytoextraction is governed by both soil and plant characteristics. Most effort has focused on identifying appropriate plants for phytoextraction, but the benefits from this effort will be marginal unless the metals are in phytoavailable forms in the rhizosphere. The concentration of a metal in the rhizosphere can be estimated using solute transfer models that incorporate: the metal concentration in the bulk soil solution, the buffer power of the soil, diffusion coefficient for the metal, water movement, root size and morphology, and the rate of entry of metal into the roots. Here a solute transfer model is developed to predict the concentration of Zn in the rhizosphere solution ([Zn]ext) of Thlaspi caerulescens, a hyperaccumulator species that could be exploited for Zn phytoextraction. The model predicts that Zn accumulation by T. caerulescens is sub-optimal when the Zn concentration in the bulk soil solution is

Caesium (Cs) is an alkali metal with chemical properties similar to potassium (K). It has no known role in plant nutrition and it is not toxic to plants at the micromolar concentrations occurring naturally in soil solutions. However, two radioisotopes of Cs (134Cs and 137Cs) are of environmental concern due to their relatively long half-lives, emissions of β and γ radiation during decay, and rapid incorporation into biological systems. There is considerable interest in remediating sites contaminated by these isotopes using phytoextraction and, since the produce from radiocaesium-contaminated areas may enter the food chain, the introduction of `safe' crops that do not accumulate Cs. This article reviews the molecular mechanisms of Cs uptake by plants, and provides a perspective on strategies to develop: (1) plants that extract Cs efficiently from soils (for the phytoremediation of land), or (2) `safe' crops that minimise the entry of radiocaesium directly into the human food chain.

1 Department of Ecological Sciences, James Hutton Institute, Invergowrie, Dundee, UK; 2 Plant and Crop Sciences Division, School of Biosciences, University of Nottingham, Sutton Bonington Campus, Leicestershire, UK; 3 Centre for Plant Integrative Biology, University of Nottingham, Sutton Bonington Campus, Leicestershire, UK; 4 College of Science, King Saud University, Riyadh, Kingdom of Saudi Arabia; 5 Current address: Department of Crop Science, School of Agriculture, College of Agriculture and Natural Sciences, University of Cape Coast, Central Region, Ghana * For correspondence: michael.adu@ucc.edu.gh

Globally, more than 800 million people are undernourished while >2 billion people have one or more chronic micronutrient deficiencies (MNDs). More than 6% of global mortality and morbidity burdens are associated with undernourishment and MNDs. Here we show that, in 2011, 3.5 and 1.1 billion people were at risk of calcium (Ca) and zinc (Zn) deficiency respectively due to inadequate dietary supply. The global mean dietary supply of Ca and Zn in 2011 was 684 ± 211 and 16 ± 3 mg capita−1 d−1 (±SD) respectively. Between 1992 and 2011, global risk of deficiency of Ca and Zn decreased from 76 to 51% and 22 to 16%, respectively. Approximately 90% of those at risk of Ca and Zn deficiency in 2011 were in Africa and Asia. To our knowledge, these are the first global estimates of dietary Ca deficiency risks based on food supply. We conclude that continuing to reduce Ca and Zn deficiency risks through dietary diversification and food and agricultural interventions including fortification, crop breeding and use of micronutrient fertilisers will remain a significant challenge.

To minimize the financial and environmental costs of fertilizer phosphorus (P) applications, it is important to maximize whole-plant P-use efficiencies. Nutrient efficiencies are influenced by both the uptake and partitioning of P between pools within the plant. This study investigates the partitioning of total plant P between storage/metabolic P and structural P pools, and their relationships with plant growth. Lettuce (Lactuca sativa L. cv. Saladin) plants were grown hydroponically under contrasting P-supply regimes. A rapid drying step was employed to prevent the breakdown of structural P molecules during conventional oven drying. Richards' growth functions accounted for 96.4 and 94.4% of the variation in plant dry weight for well-nourished and P-limited plants, respectively. There were sub-linear (or bi-linear) relationships between different P pools and relative growth rates for plants grown under restricted P supply conditions. Reanalysis of published data for tobacco (Nicotiana tabacum L.) revealed...

There is increasing interest in enhancing the micronutrient composition of cereals through fertilization. The aims of this study were (1) to determine the Se concentration of commercial beers retailing in the UK, and (2) to test if the transfer of Se, from biofortified grain to final beer product, is

In the current study, potential involvement of anthocyanins in the defence of the Zn hyperaccumulator plant, Noccaea caerulescens, was investigated in two independent glasshouse experiments. Plants were grown in compost amended with a range of Zn concentrations. In the first experiment, foliar herbivory was simulated by clipping the shoots with scissors, whilst in second experiment, plants were subjected to natural herbivory by thrips (Frankliniella occidentalis). In both experiments, foliar Zn concentrations were approximately twice as high as that applied to the soil, indicating accumulation occurred. In the clipping experiment, foliar anthocyanin concentrations decreased by 22–45 % with increasing soil Zn amendments and clipping increased anthocyanin concentrations by 17–24 % relative to unclipped plants. In a thrips feeding experiment, Zn treatment had little effect on anthocyanin content although there was a trend towards increased anthocyanin production in plants grown at the lower concentrations of Zn. Anthocyanin concentration increased by 49–52 % in N. caerulescens leaves subjected to thrip attack compared to that of thrip-free shoots. Thus, anthocyanins increased as a result of foliar damage (manual clipping and natural herbivory) and generally decreased with enhanced Zn concentrations. Foliar glucosinolate concentrations were higher in ungrazed leaves than in grazed leaves of the same thrip-infested plants, whilst anthocyanin production was higher in the grazed leaves. The possibility of an interaction between glucosinolate and anthocyanin production resulting from herbivory is discussed.

Although Ca transport in plants is highly complex, the overexpression of vacuolar Ca2+ transporters in crops is a promising new technology to improve dietary Ca supplies through biofortification. Here, we sought to identify novel targets for increasing plant Ca accumulation using genetical and comparative genomics. Expression quantitative trait locus (eQTL) mapping to 1895 cis- and 8015 trans-loci were identified in shoots of an inbred mapping population of Brassica rapa (IMB211 × R500); 23 cis- and 948 trans- eQTLs responded specifically to altered Ca supply. eQTLs were screened for functional significance using a large database of shoot Ca concentration phenotypes of Arabidopsis thaliana. From 31 Arabidopsis gene identifiers tagged to robust shoot Ca concentration phenotypes, 21 mapped to 27 B. rapa eQTLs, including orthologs of the Ca2+ transporters At-CAX1 and At-ACA8. Two of three independent missense mutants of BraA.cax1a, isolated previously by targeting induced local lesions in genomes, have allele-specific shoot Ca concentration phenotypes compared with their segregating wild types. BraA.CAX1a is a promising target for altering the Ca composition of Brassica, consistent with prior knowledge from Arabidopsis. We conclude that multiple-environment eQTL analysis of complex crop genomes combined with comparative genomics is a powerful technique for novel gene identification/prioritization.

Contamination of soils with radioisotopes of caesium (Cs) is of concern because of their emissions of harmful β and γ radiation. Radiocaesium enters the food chain through vegetation and the intake of Cs can affect the health of organisms. Arbuscular mycorrhizal (AM) fungi form mutualistic symbioses with plants through colonization of the roots and previous studies on the influence of AM on Cs concentrations in plants have given inconsistent results. These studies did not investigate the influence of Cs on AM fungi and it is therefore not known if Cs has a direct effect on AM colonization. Here, we investigated whether Cs influences AM colonization and if this effect impacts on the influence of Rhizophagus intraradices on Cs accumulation by Medicago truncatula. M. truncatula was grown with or without R. intraradices in pots containing different concentrations of Cs. Here, we present the first evidence that colonization of plants by AM fungi can be negatively affected by increasing Cs concentrations in the soil. Mycorrhizal colonization had little effect on root or shoot Cs concentrations. In conclusion, the colonization by AM fungi is impaired by high Cs concentrations and this direct effect of soil Cs on AM colonization might explain the inconsistent results reported in literature that have shown increased, decreased or unaffected Cs concentrations in AM plants.

Natural inputs of chlorine (Cl) to soils come mainly from rainwater, sea spray, dust and air pollution. In addition, human practices, such as irrigation and fertilization, contribute significantly to Cl deposition. In the soil solution, Cl occurs predominantly as the chloride anion (Cl−). The Cl−anion does not form complexes readily, and shows little affinity (or specificity) in its adsorption to soil components. Thus, Cl−movement within the soil is largely determined by water flows. Chlorine is an essential micronutrient for higher plants. It is present mainly as Cl−. Chloride is a major osmotically active solute in the vacuole and is involved in both turgor- and osmoregulation. In the cytoplasm it may regulate the activities of key enzymes. In addition, Cl−also acts as a counter anion, and Cl−fluxes are implicated in the stabilization of membrane potential, regulation of intracellular pH gradients and electrical excitability. Chloride enters plants through the roots, and there is some concern over the uptake of the long-lived radionuclide36Cl, which enters into the food chain through plants. Chloride is thought to traverse the root by a symplastic pathway, and Cl−fluxes across the plasma membrane and tonoplast of root cells have been estimated. These fluxes are regulated by the Cl−content of the root. Chloride is mobile within the plant. The Cl−concentrations of xylem and phloem saps have been determined and Cl−fluxes through the xylem and phloem have been modelled. Measurements of transmembrane voltages and Cl−activities in cellular compartments suggest (1) that active Cl−transport across the plasma membrane dominates Cl−influx to root cells at low Cl−concentrations in the soil solution and that passive Cl−influx to root cells occurs under more saline conditions, and (2) that both active and passive Cl−transport occurs at the tonoplast. Electrophysiological studies have demonstrated the presence of an electrogenic Cl−/2H+symporter in the plasma membrane of root-hair cells and Cl−channels mediating either Cl−influx or Cl−efflux across the plasma membrane. Similarly, there is both biochemical and electrophysiological evidence that Cl−channels mediate Cl−fluxes in either direction across the tonoplast and that a Cl−/nH+antiport mediates Cl−influx to the vacuole. This article reviews the availability of Cl−in the soil, the roles and distribution of Cl−within the plant, the magnitude of Cl−fluxes across membranes and between tissues, the mechanisms of Cl−transport across membranes and the electrical characteristics and molecular biology of Cl−channels.

Summary 241 I. INTRODUCTION: CAESIUM IN THE ENVIRONMENT 242 II. UPTAKE OF CAESIUM BY PLANT ROOTS 243 1. Evidence for multiple mechanisms of Cs+uptake by plant roots 243 2. Caesium uptake is affected by the presence of other cations 244 3. Caesium inhibits the uptake of other cations 244 III. MOLECULAR MECHANISMS CATALYSING CAESIUM UPTAKE 245 1. ‘High-affinity’transport mechanisms 245 2. Inward-rectifying potassium (KIR) channels 245 3. Outward-rectifying potassium (KOR) channels 248 4. Voltage-insensitive cation (VIC) channels 249 5. Ca2+-permeable channels 249 IV. MODELLING CAESIUM INFLUX TO ROOT CELLS 249 1. Predicted Cs+influx through high-affinity mechanisms 250 2. Predicted Cs+influx through cation channels 250 3. Predicted dependence of Cs+influx on[Cs+]ext 252 V. PERSPECTIVE 253 Acknowledgements 254 References 254 Caesium (Cs) is a Group I alkali metal with chemical properties similar to potassium (K). It is present in solution as the monovalent cation Cs+. Concentrations of the stable caesium isotope 133Cs in soils occur up to 25 μg g−1 dry soil. This corresponds to low micromolar Cs+ concentrations in soil solutions. There is no known role for Cs in plant nutrition, but excessive Cs can be toxic to plants. Studies of the mechanism of Cs+ uptake are important for understanding the implications arising from releases of radioisotopes of Cs, which are produced in nuclear reactors and thermonuclear explosions. Two radioisotopes of Cs (134Cs and 137Cs) are of environmental concern owing to their relatively long half-lives, emissions of β and γ radiation during decay and rapid incorporation into biological systems. The soil concentrations of these radioisotopes are six orders of magnitude lower than those of 133Cs. Early physiological studies demonstrated that K+ and Cs+ competed for influx to excised roots, suggesting that the influx of these cations to root cells is mediated by the same molecular mechanism(s). The molecular identity and/or electrophysiological signature of many K+ transporters expressed in the plasma membrane of root cells have been described. The inward-rectifying K+ (KIR), outward-rectifying K+ (KOR) and voltage-insensitive cation (VIC) channels are all permeable to Cs+ and, by analogy with their bacterial counterparts, it is likely that ‘high-affinity’ K+/H+ symporters (tentatively ascribed here to KUP genes) also transport Cs+. By modelling cation fluxes through these transporters into a stereotypical root cell, it can be predicted that VIC channels mediate most (30–90%) of the Cs+ influx under physiological conditions and that the KUP transporters mediate the bulk of the remainder. Cation influx through KIR channels is likely to be blocked by extracellular Cs+ under typical ionic conditions in the soil. Further simulations suggest that the combined Cs+ influxes through VIC channels and KUP transporters can produce the characteristic ‘dual isotherm’ relationship between Cs+ influx to excised roots and external Cs+ concentrations below 200 μM. Thus, molecular targets for modulating Cs+ influx to root cells have been identified. This information can be used to direct future genetic modification of plants, allowing them to accumulate more, or less, Cs and thereby to remediate contaminated sites.

The Michaelis–Menten kinetic parameters ( I max , K m and Cmin) of Cs uptake were estimated using a solution-depletion method, after 0, 1 and 4 d K-starvation in eight mature grass species of different relative growth rates (Bromus sterilis L., Elymus repens (L.) Gould, Poa annua L., Lolium perenne L. var. ‘Taya’, Anthoxanthum odoratum L., Festuca gigantea (L.) Vill., Festuca ovina L. and Nardus stricta L.). There were species differences in Imax and Cmin, a positive relationship between Km and RGR (r2=0.53; p=0.03), and a negative relationship between Cmin and RGR (r2=0.89; p=0.001) in 0 d K starved plants. In general, Imax increased with K starvation, as previously reported for K uptake kinetics. Km increased with K starvation, contrasting with reported studies on K, and Cmin decreased, although all species did not respond similarly.

Dietary micronutrient deficiencies (MNDs) are widespread, yet their prevalence can be difficult to assess. Here, we estimate MND risks due to inadequate intakes for seven minerals in Africa using food supply and composition data, and consider the potential of food-based and agricultural interventions. Food Balance Sheets (FBSs) for 46 countries were integrated with food composition data to estimate per capita supply of calcium (Ca), copper (Cu), iron (Fe), iodine (I), magnesium (Mg), selenium (Se) and zinc (Zn), and also phytate. Deficiency risks were quantified using an estimated average requirement (EAR) 'cut-point' approach. Deficiency risks are highest for Ca (54% of the population), followed by Zn (40%), Se (28%) and I (19%, after accounting for iodized salt consumption). The risk of Cu (1%) and Mg (1%) deficiency are low. Deficiency risks are generally lower in the north and west of Africa. Multiple MND risks are high in many countries. The population-weighted mean phytate supply is 2770 mg capita(-1) day(-1). Deficiency risks for Fe are lower than expected (5%). However, 'cut-point' approaches for Fe are sensitive to assumptions regarding requirements; e.g. estimates of Fe deficiency risks are 43% under very low bioavailability scenarios consistent with high-phytate, low-animal protein diets. Fertilization and breeding strategies could greatly reduce certain MNDs. For example, meeting HarvestPlus breeding targets for Zn would reduce dietary Zn deficiency risk by 90% based on supply data. Dietary diversification or direct fortification is likely to be needed to address Ca deficiency risks.

Mineral malnutrition is widespread in sub-Saharan Africa but its extent is difficult to quantify. Using Malawi as a case study, the aim of this work was to investigate the adequacy of calcium (Ca) and magnesium (Mg) nutrition by combining national food supply and food composition data with a new spatial survey of maize grain. Non-maize dietary sources of Ca and Mg were estimated using existing food supply and composition data. Calcium and Mg concentrations in maize grain were determined at 88 field sites, representing > 75 % of Malawi’s land area in terms of soil classification. Median maize grain concentrations from the survey were 34 and 845 mg kg-1, representing a per capita supply of 12 and 299 mg d-1 of Ca and Mg, respectively. Combining these data with food supply and composition data reveals that average Ca nutrition is likely to be inadequate for many individuals, whereas average Mg nutrition appears adequate. Optimal supply of Ca per capita depends critically on balanced food availability and choice. Since maize grain sourced from highly calcareous soils is still unlikely to deliver > 5 % of estimated average requirements, agronomic solutions to rectify Ca malnutrition via maize are limited, in comparison with strategies for dietary diversification.

Selenium (Se) is an essential human micronutrient with critical roles in immune functioning and antioxidant defence. Estimates of dietary Se intakes and status are scarce for Africa although crop surveys indicate deficiency is probably widespread in Malawi. Here we show that Se deficiency is likely endemic in Malawi based on the Se status of adults consuming food from contrasting soil types. These data are consistent with food balance sheets and composition tables revealing that >80% of the Malawi population is at risk of dietary Se inadequacy. Risk of dietary Se inadequacy is >60% in seven other countries in Southern Africa, and 22% across Africa as a whole. Given that most Malawi soils cannot supply sufficient Se to crops for adequate human nutrition, the cost and benefits of interventions to alleviate Se deficiency should be determined; for example, Se-enriched nitrogen fertilisers could be adopted as in Finland.

• The ionome is the elemental composition of a tissue or organism. Phylogenetic variation in the ionomes of plant shoots has been widely reported based on controlled experiments, vegetation surveys and literature meta-analyses. However, environmental effects on phylogenetic variation in shoot ionomes have not been quantified. This study tests the hypothesis that phylogenetic variation in shoot ionomes is robust to environmental perturbation and that plant families can be distinguished by their shoot ionomes. • Herbage was sampled from six subplots of the Rothamsted Park Grass Experiment. Subplots had received contrasting fertilizer treatments since 1856. Herbage was separated into its constituent species (n = 21) and concentrations of eleven mineral elements were determined in dried shoot material. • Shoot concentrations of calcium (Ca), zinc (Zn), manganese (Mn), magnesium (Mg) and sodium (Na) showed significant variation associated with plant species, and responded similarly to fertilizer treatments in diverse plant species. Species × treatment interactions were indicated for phosphorus (P), potassium (K), nickel (Ni), copper (Cu) and iron (Fe). Plant families could be distinguished by their shoot ionomes. The most informative elements for discriminant analysis were Ca > Mg > Ni > S > Na > Zn > K > Cu > Fe > Mn > P. • Whilst shoot ionomes were sensitive to fertilizer treatment, phylogenetic variation in a subset of the shoot ionome (Ca, Zn, Mn, Mg) was robust to this environmental perturbation.

Publisher Summary This chapter focuses on the functions of iron, manganese, copper, zinc, nickel, molybdenum, boron, and chlorine in plants and describes the effects of their deficiency and toxicity. Iron (Fe) plays a crucial role in redox systems in cells and in various enzymes. In dicotyledonous and monocotyledonous plant species, Fe deficiency is associated with the formation of rhizodermal transfer cells, which is a part of a their strategy to enhance iron uptake. Manganese (Mn) and copper (Cu) are important for redox systems, as activators of various enzymes including those involved in the detoxification of superoxide radicals, and for the synthesis of lignin. In dicotyledonous plants, intercostal chlorosis of the younger leaves is the most distinct symptom of Mn deficiency, whereas in cereals, greenish grey spots on the older leaves are the major symptoms. Stunted growth, distortion of young leaves, chlorosis/ necrosis starting at the apical meristem extending down to the leaf margins, bleaching of young leaves, and/or “summer dieback” in trees are typical visible symptoms of Cu deficiency. Zinc (Zn) plays a role in the detoxification of superoxide radicals, membrane integrity, as well as the synthesis of proteins and the phytohormone IAA. Nickel (Ni) is involved in N metabolism as a metal component of the enzyme urease, whereas molybdenum (Mo) helps in N metabolism by acting as a metal component of the nitrogenase (N2 fixation) and nitrate reductase enzymes. Boron (B) is crucial for cell wall and membrane integrity, whereas chlorine plays a role in osmoregulation and stomata movement.

The plant ionome varies both inter- and intraspecifically despite the highly conserved roles for particular elements across the plant kingdom. Element storage requires transport across the plasma membrane and commonly deposition within the central vacuole. Therefore, tonoplast transport characteristics can be highly influential in controlling the plant ionome. As a result, individual cell types of the same plant, each with unique transcriptomes and vacuolar proteomes, can display very different elemental profiles. Here we address the use of natural variation in Arabidopsis thaliana for identifying genes involved in elemental accumulation. We present a conceptual framework, exploiting publicly available leaf ionomic and transcriptomic data across 31 Arabidopsis accessions, that promises to accelerate conventional forward genetics approaches for candidate gene discovery. Utilizing this framework, we identify numerous genes with documented roles in accumulation of calcium, magnesium and zinc and implicate additional candidate genes. Where appropriate, we discuss their role in cell-specific elemental accumulation. Currently, this framework could represent an alternate approach for identifying genes suitable for element biofortification of plants. Integration of additional cell-specific and whole-plant 'omics' datasets across Arabidopsis accessions under diverse environmental conditions should enable this concept to be developed into a scalable and robust tool for linking genotype and phenotype. Refereed/Peer-reviewed

L'article original est publié par The American Society of Plant Biologists; International audience; Roots are important to plants for a wide variety of processes, including nutrient and water uptake, anchoring and mechanical support, storage functions, and as the major interface between the plant and various biotic and abiotic factors in the soil environment. Therefore, understanding the development and architecture of roots holds potential for the manipulation of root traits to improve the productivity and sustainability of agricultural systems and to better understand and manage natural ecosystems. While lateral root development is a traceable process along the primary root and different stages can be found along this longitudinal axis of time and development, root system architecture is complex and difficult to quantify. Here, we comment on assays to describe lateral root phenotypes and propose ways to move forward regarding the description of root system architecture, also considering crops and the environment.