With increasing demand of agricultural production and as the peak in global production will occur in the next decades, phosphorus (P) is receiving more attention as a nonrenewable resource (Cordell et al., 2009;Gilbert, 2009). One unique characteristic of P is its low availability due to slow diffusion and high fixation in soils. All of this means that P can be a major limiting factor for plant growth. Applications of chemical P fertilizers and animal manure to agricultural land have improved soil P fertility and crop production, but caused environmental damage in the past decades. Maintaining a proper P-supplying level at the root zone can maximize the efficiency of plant roots to mobilize and acquire P from the rhizosphere by an integration of root morphological and physiological adaptive strategies. Furthermore, P uptake and utilization by plants plays a vital role in the determination of final crop yield. A holistic understanding of P dynamics from soil to plant is necessary for optimizing P management and improving P-use efficiency, aiming at reducing consumption of chemical P fertilizer, maximizing exploitation of the biological potential of root/rhizosphere processes for efficient mobilization, and acquisition of soil P by plants as well as recycling P from manure and waste. Taken together, overall P dynamics in the soilplant system is a function of the integrative effects of P transformation, availability, and utilization caused by soil, rhizosphere, and plant processes. This Update focuses on the dynamic processes determining P availability in the soil and in the rhizosphere, P mobilization, uptake, and utilization by plants. It highlights recent advances in the understanding of the P dynamics in the soil/rhizosphere-plant continuum.
Summary Plant roots exhibit diverse root functional traits to enable soil phosphorus (P) acquisition, including changes in root morphology, root exudation and mycorrhizal symbioses. Yet, whether these traits are differently coordinated among crop species to enhance P acquisition is unclear. Here, eight root functional traits for P acquisition were characterized in 16 major herbaceous crop species grown in a glasshouse under limiting and adequate soil P availability. We found substantial interspecific variation in root functional traits among species. Those with thinner roots showed more root branching and less first‐order root length, and had consistently lower colonization by arbuscular mycorrhizal fungi (AMF), fewer rhizosheath carboxylates and reduced acid phosphatase activity. In response to limiting soil P, species with thinner roots showed a stronger response in root branching, first‐order root length and specific root length of the whole root system, Conversely, species with thicker roots exhibited higher colonization by AMF and/or more P‐mobilizing exudates in the rhizosheath. We conclude that, at the species level, tradeoffs occur among the three groups of root functional traits we examined. Root diameter is a good predictor of the relative expression of these traits and how they change when P is limiting.
Legacy phosphorus (P) that has accumulated in soils from past inputs of fertilizers and manures is a large secondary global source of P that could substitute manufactured fertilizers, help preserve critical reserves of finite phosphate rock to ensure future food and bioenergy supply, and gradually improve water quality. We explore the issues and management options to better utilize legacy soil P and conclude that it represents a valuable and largely accessible P resource. The future value and period over which legacy soil P can be accessed depends on the amount present and its distribution, its availability to crops and rates of drawdown determined by the cropping system. Full exploitation of legacy P requires a transition to a more holistic system approach to nutrient management based on technological advances in precision farming, plant breeding and microbial engineering together with a greater reliance on recovered and recycled P. We propose the term 'agro-engineering' to encompass this integrated approach. Smaller targeted applications of fertilizer P may still be needed to optimize crop yields where legacy soil P cannot fully meet crop demands. Farm profitability margins, the need to recycle animal manures and the extent of local eutrophication problems will dictate when, where and how quickly legacy P is best exploited. Based on our analysis, we outline the stages and drivers in a transition to the full utilization of legacy soil P as part of more sustainable regional and global nutrient management.
BackgroundThe human genome encodes many long non-coding RNAs (lncRNAs). However, their biological functions, molecular mechanisms, and the prognostic value associated with pancreatic ductal adenocarcinoma (PDAC) remain to be elucidated. Here, we identify a fundamental role for the lncRNA HOXA transcript at the distal tip (HOTTIP) in the progression and chemoresistance of PDAC.MethodsHigh-throughput microarrays were performed to detect the expression profiles of lncRNAs and messenger RNAs in eight human PDAC tissues and four pancreatic tissues. Quantitative real-time PCR was used to determine the levels of HOTTIP and HOXA13 transcripts in PDAC cell lines and 90 PDAC samples from patients. HPDE6 cells (immortalized human pancreatic ductal epithelial cells) and corresponding adjacent non-neoplastic tissues were used as controls, respectively. The functions of HOTTIP and HOXA13 in cell proliferation, invasion, and epithelial-mesenchymal transition were evaluated by targeted knockdown in vitro. CCK-8 assays, colony formation assays, and xenografts in nude mice were used to investigate whether targeted silencing of HOTTIP could sensitize pancreatic cancer cells to gemcitabine. Immunohistochemistry was performed to investigate the relationship between HOXA13 expression and patient outcome.ResultsMicroarray analyses revealed that HOTTIP was one of the most significantly upregulated lncRNAs in PDAC tissues compared with pancreatic tissues. Quantitative PCR further verified that HOTTIP levels were increased in PDAC cell lines and patient samples compared with controls. Functionally, HOTTIP silencing resulted in proliferation arrest by altering cell-cycle progression, and impaired cell invasion by inhibiting epithelial-mesenchymal transition in pancreatic cancer. Additionally, inhibition of HOTTIP potentiated the antitumor effects of gemcitabine in vitro and in vivo. Furthermore, knockdown of HOXA13 by RNA interference (siHOXA13) revealed that HOTTIP promoted PDAC cell proliferation, invasion, and chemoresistance, at least partly through regulating HOXA13. Immunohistochemistry results revealed that higher HOXA13 expression was correlated with lymph node metastasis, poor histological differentiation, and decreased overall survival in PDAC patients.ConclusionsAs a crucial tumor promoter, HOTTIP promotes cell proliferation, invasion, and chemoresistance by modulating HOXA13. Therefore, the HOTTIP/HOXA13 axis is a potential therapeutic target and molecular biomarker for PDAC.Electronic supplementary materialThe online version of this article (doi:10.1186/s12967-015-0442-z) contains supplementary material, which is available to authorized users.
This publication is made publicly available in the institutional repository of Wageningen University and Research, under the terms of article 25fa of the Dutch Copyright Act, also known as the Amendment Taverne. This has been done with explicit consent by the author.Article 25fa states that the author of a short scientific work funded either wholly or partially by Dutch public funds is entitled to make that work publicly available for no consideration following a reasonable period of time after the work was first published, provided that clear reference is made to the source of the first publication of the work.This publication is distributed under The Association of Universities in the Netherlands (VSNU) 'Article 25fa implementation' project. In this project research outputs of researchers employed by Dutch Universities that comply with the legal requirements of Article 25fa of the Dutch Copyright Act are distributed online and free of cost or other barriers in institutional repositories. Research outputs are distributed six months after their first online publication in the original published version and with proper attribution to the source of the original publication.
The relationship between root morphological and physiological responses to variable P supply in different plant species is poorly understood. We compared root morphological and physiological responses to P supply in seven crop species (Zea mays, Triticum aestivum, Brassica napus, Lupinus albus, Glycine max, Vicia faba, Cicer arietinum) treated with or without 100 mg P kg-1 in two soils (acidic and calcareous). Phosphorus deficiency decreased root length more in fibrous root species (Zea mays, Triticum aestivum, Brassica napus) than legumes. Zea mays and Triticum aestivum had higher root/shoot biomass ratio and Brassica napus had higher specific root length compared to legumes, whereas legumes (except soybean) had higher carboxylate exudation than fibrous root species. Lupinus albus exhibited the highest P-acquisition efficiency due to high exudation of carboxylates and acid phosphatases. Lupinus albus and Cicer arietinum depended mostly on root exudation (i.e., physiological response) to enhance P acquisition, whereas Zea mays, Triticum aestivum and Brassica napus had higher root morphology dependence, with Glycine max and Vicia faba in between. Principal component analysis using six morphological and six physiological responses identified root size and diameter as the most important morphological traits, whereas important physiological responses included carboxylate exudation, and P-acquisition and P-utilization efficiency followed by rhizosphere soil pH and acid phosphatase activity. In conclusion, plant species can be grouped on the basis of their response to soil P being primarily via root architectural or exudation plasticity, suggesting a potential benefit of crop-specific root-trait-based management to cope with variable soil P supply in sustainable grain production.
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