Superior Root Hair Formation Confers Root Efficiency in Some, But Not All, Rice Genotypes upon P Deficiency

Root hairs are a low-cost way to extend root surface area (RSA), water and nutrient acquisition. This study investigated to what extend variation exists for root hair formation in rice in dependence of genotype, phosphorus (P) supply, growth medium, and root type. In general, genotypic variation was found for three root hair properties: root hair length, density, and longevity. In low P nutrient solution more than twofold genotypic difference was detected for root hair length while only onefold variation was found in low P soil. These differences were mostly due to the ability of some genotypes to increase root hair length in response to P deficiency. In addition, we were able to show that a higher proportion of root hairs remain viable even in mature, field-grown plants under low P conditions. All investigated root hair parameters exhibited high correlations across root types which were always higher in the low P conditions compared to the high P controls. Therefore we hypothesize that a low P response leads to a systemic signal in the entire root system. The genotype DJ123 consistently had the longest root hairs under low P conditions and we estimated that, across the field-grown root system, root hairs increased the total RSA by 31% in this genotype. This would explain why DJ123 is considered to be very root efficient in P uptake and suggests that DJ123 should be utilized as a donor in breeding for enhanced P uptake. Surprisingly, another root and P efficient genotype seemed not to rely on root hair growth upon P deficiency and therefore must contain different methods of low P adaptation. Genotypic ranking of root hair properties did change substantially with growth condition highlighting the need to phenotype plants in soil-based conditions or at least to validate results obtained in solution-based growth conditions

Stability of grain zinc concentrations across lowland rice environments favors zinc biofortification breeding

IntroductionOne-third of the human population consumes insufficient zinc (Zn) to sustain a healthy life. Zn deficiency can be relieved by increasing the Zn concentration ([Zn]) in staple food crops through biofortification breeding. Rice is a poor source of Zn, and in countries predominantly relying on rice without sufficient dietary diversification, such as Madagascar, Zn biofortification is a priority.MethodsMulti-environmental trials were performed in Madagascar over two years, 2019 and 2020, to screen a total of 28 genotypes including local and imported germplasm. The trials were conducted in the highlands of Ankazomiriotra, Anjiro, and Behenji and in Morovoay, a location representative of the coastal ecosystem. Contributions of genotype (G), environment (E), and G by E interactions (GEIs) were investigated.ResultThe grain [Zn] of local Malagasy rice varieties was similar to the internationally established grain [Zn] baseline of 18–20 μg/g for brown rice. While several imported breeding lines reached 50% of our breeding target set at +12 μg/g, only few met farmers’ appreciation criteria. Levels of grain [Zn] were stable across E. The G effects accounted for a main fraction of the variation, 76% to 83% of the variation for year 1 and year 2 trials, respectively, while GEI effects were comparatively small, contributing 23% to 9%. This contrasted with dominant E and GEI effects for grain yield. Our results indicate that local varieties tested contained insufficient Zn to alleviate Zn malnutrition, and developing new Zn-biofortified varieties should therefore be a priority. GGE analysis did not distinguish mega-environments for grain [Zn], whereas at least three mega-environments existed for grain yield, differentiated by the presence of limiting environmental conditions and responsiveness to improved soil fertility.DiscussionOur main conclusion reveals that grain [Zn] seems to be under strong genetic control in the agro-climatic conditions of Madagascar. We could identify several interesting genotypes as potential donors for the breeding program, among those BF156, with a relatively stable grain [Zn] (AMMI stability value (ASV) = 0.89) reaching our target (>26 μg/g). While selection for grain yield, general adaptation, and farmers’ appreciation would have to rely on multi-environment testing, selection for grain [Zn] could be centralized in earlier generations

Rice genotype differences in tolerance of zinc-deficient soils: evidence for the importance of root-induced changes in the rhizosphere

The Supplementary Material for this article can be found online at: http://journal.frontiersin.org/article/10.3389/fpls.2015.01160Zinc (Zn) deficiency is a major constraint to rice production and Zn is also often deficient in humans with rice-based diets. Efforts to breed more Zn-efficient rice are constrained by poor understanding of the mechanisms of tolerance to deficiency. Here we assess the contributions of root growth and root Zn uptake efficiency, and we seek to explain the results in terms of specific mechanisms. We made a field experiment in a highly Zn-deficient rice soil in the Philippines with deficiency-tolerant and -sensitive genotypes, and measured growth, Zn uptake and root development. We also measured the effect of planting density. Tolerant genotypes produced more crown roots per plant and had greater uptake rates per unit root surface area; the latter was at least as important as root number to overall tolerance. Tolerant and sensitive genotypes took up more Zn per plant at greater planting densities. The greater uptake per unit root surface area, and the planting density effect can only be explained by root-induced changes in the rhizosphere, either solubilizing Zn, or neutralizing a toxin that impedes Zn uptake (possibly HCO − 3 HCO3− or Fe2+), or both. Traits for these and crown root number are potential breeding targets.This research was funded by a grant from the UK's Biotechnology and Biological Sciences Research Council (BBSRC, Grant Ref. BB/J011584/1) under the Sustainable Crop Production Research for International Development (SCPRID) programme, a joint multi-national initiative of BBSRC, the UK Government's Department for International Development (DFID) and (through a grant awarded to BBSRC) the Bill & Melinda Gates Foundation. Support to AKN in the form of a fellowship awarded by the Japan Society for the Promotion of Science (JSPS) is gratefully acknowledged

Phosphorus Efficient Phenotype of Rice

The ideal phenotype to cope with P deficiency is suggested to be a larger root system, both in terms of length and foraging area, coupled with a high capacity for P solubilization from compounds exuded from roots. Greater soil exploration results in a large number of roots in the top soil, longer roots in general with more cortical aerenchyma, more and longer root hairs, and a shift in mycorrhizal and bacterial colonization. However, these assumptions often result from experiments in highly controlled, sterile and soil-free conditions using model plants or single ecotypes where results are then extrapolated to all genotypes and plant species. In recent years this generalization has been questioned. Here, we summarize recent rice research analyzing the natural diversity of rice root systems under P deficiency. Interestingly, while some of the high yielding genotypes do show the expected, large root system phenotype, some have a surprisingly small root system—as little as a quarter of that of the large root system varieties—but achieve similar yield and P uptake under P deficiency. This effect has recently been termed root efficiency, which we discuss in this chapter in conjunction with root foraging traits

The Frustration with Utilization: Why Have Improvements in Internal Phosphorus Utilization Efficiency in Crops Remained so Elusive?

Despite the attention internal phosphorus utilization efficiency (PUE) of crops has received in the literature, little progress in breeding crop cultivars with high PUE has been made. Surprisingly few studies have specifically investigated PUE; instead, genotypic variation for PUE has been investigated in studies that concurrently assess phosphorus acquisition efficiency (PAE). We hypothesized that genotypic differences in PAE confound PUE rankings because genotypes with higher PAE suffer a lower degree of P stress, resulting in lower PUE. The hypothesis was tested by comparing soil-based screening to a modified technique whereby rice genotypes were grown in individual containers with a single dose of solution P, to eliminate differences in P uptake among genotypes. Genotypic differences in PUE were apparent in root and shoot tissue using the modified nutrient solution technique, but PUE rankings showed no correlation with those from traditional soil-based screening. We conclude that PUE in soil-based screening systems is unavoidably linked with genotypic PAE, resulting in PUE rankings confounded by differences in P uptake. Only screening techniques assuring equal P uptake are suitable for the exploitation of genotypic variation for PUE

Soil CO2 venting as one of the mechanisms for tolerance of Zn deficiency by rice in flooded soils

We sought to explain rice (Oryza sativa) genotype differences in tolerance of zinc (Zn) deficiency in flooded paddy soils and the counter-intuitive observation, made in earlier field experiments, that Zn uptake per plant increases with increasing planting density. We grew tolerant and intolerant genotypes in a Zn-deficient flooded soil at high and low planting densities, and found (a) plant Zn concentrations and growth increased with planting density and more so in the tolerant genotype, whereas the concentrations of other nutrients decreased, indicating a specific effect on Zn uptake; (b) the effects of planting density and genotype on Zn uptake could only be explained if the plants induced changes in the soil to make Zn more soluble; and (c) the genotype and planting density effects were both associated with decreases in dissolved CO2 in the rhizosphere soil solution and resulting increases in pH. We suggest the increases in pH caused solubilisation of soil Zn by dissolution of alkali-soluble, Zn-complexing organic ligands from soil organic matter. We conclude that differences in venting of soil CO2 through root aerenchyma were responsible for the genotype and planting density effects

Rice increases phosphorus uptake in strongly sorbing soils by intra-root facilitation

Upland rice (Oryza sativa) is adapted to strongly phosphorus (P) sorbing soils. The mechanisms underlying P acquisition, however, are not well understood, and models typically underestimate uptake. This complicates root ideotype development and trait-based selection for further improvement. We present a novel model, which correctly simulates the P uptake by a P-efficient rice genotype measured over 48 days of growth. The model represents root morphology at the local rhizosphere scale, including root hairs and fine S-type laterals. It simulates fast- and slowly reacting soil P and the P-solubilizing effect of root-induced pH changes in the soil. Simulations predict that the zone of pH changes and P solubilization around a root spreads further into the soil than the zone of P depletion. A root needs to place laterals outside its depletion- but inside its solubilization zone to maximize P uptake. S-type laterals, which are short but hairy, appear to be the key root structures to achieve that. Thus, thicker roots facilitate the P uptake by fine lateral roots. Uptake can be enhanced through longer root hairs and greater root length density but was less sensitive to total root length and root class proportions.Biotechnology and Biological Sciences Research Council (BBSRC): BB/R020388/

Business Rules: Die Wissensverarbeitung erreicht die Betriebswirtschaft ; Einsatzmöglichkeiten und Marktübersicht

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