Synchronization of developmental, molecular and metabolic aspects of source–sink interactions

Plants have evolved a multitude of strategies to adjust their growth according to external and internal signals. Interconnected metabolic and phytohormonal signalling networks allow adaption to changing environmental and developmental conditions and ensure the survival of species in fluctuating environments. In agricultural ecosystems, many of these adaptive responses are not required or may even limit crop yield, as they prevent plants from realizing their fullest potential. By lifting source and sink activities to their maximum, massive yield increases can be foreseen, potentially closing the future yield gap resulting from an increasing world population and the transition to a carbon-neutral economy. To do so, a better understanding of the interplay between metabolic and developmental processes is required. In the past, these processes have been tackled independently from each other, but coordinated efforts are required to understand the fine mechanics of source–sink relations and thus optimize crop yield. Here, we describe approaches to design high-yielding crop plants utilizing strategies derived from current metabolic concepts and our understanding of the molecular processes determining sink development.Research in the authors’ laboratories was supported by the following grants: the cassava source–sink (CASS) project of the Bill and Melinda Gates Foundation (to A.R.F., H.E.N., M.S. and U.S.); the ERA-CAPs project SourSi (to A.R.F. and L.J.S.); the BIO2015-3019-EXP grant from the Spanish Ministry of Economy, Industry and Competitiveness and the PCIN-2017-032 CONCERT-JAPAN project financed by the Ministry of Science, Innovation and Universities (to S.P.); Australian Research Council DP180103834 (to Y.L.R.); the US National Science Foundation (grant no. IOS-1457183); the Agriculture and Food Research Initiative (AFRI; grant no. 2017-67013-26158) from the USDA National Institute of Food and Agriculture (to M.T.); the Finnish Centre of Excellence in Molecular Biology of Primary Producers (Academy of Finland CoE program 2014–2019; grant no. 271832); the Gatsby Foundation (grant no. GAT3395/PR3); the University of Helsinki (grant no. 799992091); the European Research Council Advanced Investigator Grant SYMDEV (grant no. 323052; to Y.H.); the BMBF (grant no. 031B0191); the DFG (SPP1530: WA3639/1-2, 2-1); and the Max-Planck-Society (to V.W.). We additionally thank D. Ko and R. Ruonala for their comments on the manuscript

A Research Road Map for Responsible Use of Agricultural Nitrogen

Nitrogen (N) is an essential but generally limiting nutrient for biological systems. Development of the Haber-Bosch industrial process for ammonia synthesis helped to relieve N limitation of agricultural production, fueling the Green Revolution and reducing hunger. However, the massive use of industrial N fertilizer has doubled the N moving through the global N cycle with dramatic environmental consequences that threaten planetary health. Thus, there is an urgent need to reduce losses of reactive N from agriculture, while ensuring sufficient N inputs for food security. Here we review current knowledge related to N use efficiency (NUE) in agriculture and identify research opportunities in the areas of agronomy, plant breeding, biological N fixation (BNF), soil N cycling, and modeling to achieve responsible, sustainable use of N in agriculture. Amongst these opportunities, improved agricultural practices that synchronize crop N demand with soil N availability are low-hanging fruit. Crop breeding that targets root and shoot physiological processes will likely increase N uptake and utilization of soil N, while breeding for BNF effectiveness in legumes will enhance overall system NUE. Likewise, engineering of novel N-fixing symbioses in non-legumes could reduce the need for chemical fertilizers in agroecosystems but is a much longer-term goal. The use of simulation modeling to conceptualize the complex, interwoven processes that affect agroecosystem NUE, along with multi-objective optimization, will also accelerate NUE gains

Uptake and partitioning of amino acids and peptides

Plant growth, productivity, and seed yield depend on the efficient uptake, metabolism, and allocation of nutrients. Nitrogen is an essential macronutrient needed in high amounts. Plants have evolved efficient and selective transport systems for nitrogen uptake and transport within the plant to sustain development, growth, and finally reproduction. This review summarizes current knowledge on membrane proteins involved in transport of amino acids and peptides. A special emphasis was put on their function in planta. We focus on uptake of the organic nitrogen by the root, source-sink partitioning, and import into floral tissues and seeds

Source and sink mechanisms of nitrogen transport and use

Contents Summary 35 I. Introduction 35 II. Nitrogen acquisition and assimilation 36 III. Root‐to‐shoot transport of nitrogen 38 IV. Nitrogen storage pools in vegetative tissues 39 V. Nitrogen transport from source leaf to sink 40 VI. Nitrogen import into sinks 42 VII. Relationship between source and sink nitrogen transport processes and metabolism 43 VIII. Regulation of nitrogen transport 43 IX. Strategies for crop improvement 44 X. Conclusions 46 Acknowledgements 47 References 47 Summary Nitrogen is an essential nutrient for plant growth. World‐wide, large quantities of nitrogenous fertilizer are applied to ensure maximum crop productivity. However, nitrogen fertilizer application is expensive and negatively affects the environment, and subsequently human health. A strategy to address this problem is the development of crops that are efficient in acquiring and using nitrogen and that can achieve high seed yields with reduced nitrogen input. This review integrates the current knowledge regarding inorganic and organic nitrogen management at the whole‐plant level, spanning from nitrogen uptake to remobilization and utilization in source and sink organs. Plant partitioning and transient storage of inorganic and organic nitrogen forms are evaluated, as is how they affect nitrogen availability, metabolism and mobilization. Essential functions of nitrogen transporters in source and sink organs and their importance in regulating nitrogen movement in support of metabolism, and vegetative and reproductive growth are assessed. Finally, we discuss recent advances in plant engineering, demonstrating that nitrogen transporters are effective targets to improve crop productivity and nitrogen use efficiency. While inorganic and organic nitrogen transporters were examined separately in these studies, they provide valuable clues about how to successfully combine approaches for future crop engineering

Molecular Evolution of Plant AAP and LHT Amino Acid Transporters

Nitrogen is an essential mineral nutrient and it is often transported within living organisms in its reduced form, as amino acids. Transport of amino acids across cellular membranes requires proteins, and here we report the phylogenetic analysis across taxa of two amino acid transporter families, the amino acid permeases (AAPs) and the lysine–histidine-like transporters (LHTs). We found that the two transporter families form two distinct groups in plants supporting the concept that both are essential. AAP transporters seem to be restricted to land plants. They were found in Selaginella moellendorffii and Physcomitrella patens but not in Chlorophyte, Charophyte, or Rhodophyte algae. AAPs were strongly represented in vascular plants, consistent with their major function in phloem (vascular tissue) loading of amino acids for sink nitrogen supply. LHTs on the other hand appeared prior to land plants. LHTs were not found in chlorophyte algae Chlamydomonas reinhardtii and Volvox carterii . However, the characean alga Klebsormidium flaccidum encodes KfLHT13 and phylogenetic analysis indicates that it is basal to land plant LHTs. This is consistent with the hypothesis that characean algae are ancestral to land plants. LHTs were also found in both S. moellendorffii and P. patens as well as in monocots and eudicots. To date, AAPs and LHTs have mainly been characterized in Arabidopsis (eudicots) and these studies provide clues to the functions of the newly identified homologs

Transporters for uptake and allocation of organic nitrogen compounds in plants

Nitrogen is an essential macronutrient for plant growth. Following uptake from the soil or assimilation within the plant, organic nitrogen compounds are transported between organelles, from cell to cell and over long distances in support of plant metabolism and development. These translocation processes require the function of integral membrane transporters. The review summarizes our current understanding of the molecular mechanisms of organic nitrogen transport processes, with a focus on amino acid, ureide and peptide transporters

Abstract

ii ACKNOWLEDGMENTS I would like to express my gratitude to my advisor, Gerry Edwards, for his continual support and patience. He gave me the opportunity to develop my own skills in scientific inquiry and discover my passion for botanical science. I have learned the most about myself and scientific investigation from this experience. I would also like to extend thanks to my committee members, Mechthild Tegeder and Asaph Cousins, for their guidance and valuable insight. I would also like to thank Valerie Lynch-Holm and Chris Davitt for their help with my microscopy work in the Franceschi Microscopy and Imaging Center, and their guidance during my time as a teaching assistant there. I would also like to thank Chuck Cody, Elena Voznesenskaya and Nouria Koteeva for their extensive knowledge and assistance in growing the plants needed for my project. I would also like to thank Ray Lee and the Cousin’s lab for their assistance in completing various analyses needed in my project. I would like to thank Marc Evans for his extensive help with all of my statistical questions related to this project. I have had the privilege of working in a lab with a group of helpful and inventive students and scientists. I would like to thank Monica Smith, JoonHo Park, Yasuko Nagai, Olavi Kiirats, Jos

Organic Carbon and Nitrogen Transporters

During growth and storage phases, plant cells import large amounts of carbon (C) and nitrogen (N) assimilates to drive their metabolism and to facilitate synthesis of storage products. Following assimilation, organic C and N metabolites are transported via the phloem (C) or via the xylem followed by phloem (N) to sink tissues. These translocation processes involve participation of metabolite transporters located in source and sink cells. While information on transporters mediating cellular efflux of C and N metabolites is scarce, cell import systems have been identified and characterized. An overview is provided on transporters importing amino acids, peptides, and sugars into plant cells with particular emphasis on their substrate selectivity, expression, and function

Roles of plasma membrane transporters in phloem functions

Phloem functions as a supracellular highway for long-distance transport of resources (nutrients and water), toxic elements/compounds and signals (macromolecules, phytohormones, redox reagents and electrical potential waves) coordinating growth, development, plant homeostasis, and defense responses. Apart from macromolecules, phloem transport function includes membrane passage of the above constituents at some point(s) along the phloem transport pathway. These membrane transport events are mediated by transporters localized to plasma membrane (primarily) or tonoplast with the latter functioning in buffering transport fluxes by exchange to, and from, cell vacuoles. Transporter proteins function as carriers (± energy coupled) or channels. For each transporter, dependent upon published information, consideration will be given to their intra- and intercellular localization, transport properties (biochemical and biophysical) related to perceived cellular function, and physiological significance (deduced from knockout/knockdown approaches) to achieve an overall phloem transport flux