► Starch is a key substance in plant metabolism and a vital product for our society. ► Plants remobilise leaf starch at night to support respiration and growth. ► Premature or incomplete exhaustion ...of starch lowers plant growth rates. ► Diurnal starch turnover is tightly regulated, in part by the circadian clock. ► We provide an update on starch metabolism, its control, and carbon use for growth.
Many plants store part of their photosynthate as starch during the day and remobilise it to support metabolism and growth at night. Mutants unable to synthesize or degrade starch show strongly impaired growth except in long day conditions. In rapidly growing plants, starch turnover is regulated such that it is almost, but not completely, exhausted at dawn. There is increasing evidence that premature or incomplete exhaustion of starch turnover results in lower rates of plant growth. This review provides an update on the pathways for starch synthesis and degradation. We discuss recent advances in understanding how starch turnover and the use of carbon for growth is regulated during diurnal cycles, with special emphasis on the role of the biological clock. Much of the molecular and genetic research on starch turnover has been performed in the reference system Arabidopsis. This review considers to what extent information gained in this weed species maybe applicable to annual crops and perennial species.
In addition to light and water, CO2 and mineral elements are required for plant growth and development. Among these factors, nitrogen is critical, since it is needed to synthesize amino acids, which ...are the building elements of protein, nucleotides, chlorophyll, and numerous other metabolites and cellular components. Therefore, nitrogen is required by plants in higher quantities and this investment in nitrogen supports the use of CO2, water, and inorganic nitrogen to produce sugars, organic acids, and amino acids, the basic building blocks of biomass accumulation. This system is maintained by complex metabolic machinery, which is regulated at different levels according to environmental factors such as light, CO2, and nutrient availability. Plants integrate these signals via a signaling network, which involves metabolites as well as nutrient-sensing proteins. Due to its importance, much research effort has been expended to understand how carbon and nitrogen metabolism are integrated and regulated according to the rates of photosynthesis, photorespiration, and respiration. Thus, in this article, we both discuss recent advances in carbon/nitrogen metabolisms as well as sensing and signaling systems in illuminated leaves of C3-plants and provide a perspective of the type of experiments that are now required in order to take our understanding to a higher level.
All plant cells are surrounded by a cell wall that determines the directionality of cell growth and protects the cell against its environment. Plant cell walls are comprised primarily of ...polysaccharides and represent the largest sink for photosynthetically fixed carbon, both for individual plants and in the terrestrial biosphere as a whole. Cell wall synthesis is a highly sophisticated process, involving multiple enzymes and metabolic intermediates, intracellular trafficking of proteins and cell wall precursors, assembly of cell wall polymers into the extracellular matrix, remodeling of polymers and their interactions, and recycling of cell wall sugars. In this review we discuss how newly fixed carbon, in the form of UDP-glucose and other nucleotide sugars, contributes to the synthesis of cell wall polysaccharides, and how cell wall synthesis is influenced by the carbon status of the plant, with a focus on the model species Arabidopsis (Arabidopsis thaliana).
In this review we discuss how newly fixed carbon, in the form of UDP-glucose and other nucleotide sugars, contributes to the synthesis of cell wall polysaccharides, and how cell wall synthesis is influenced by the carbon status of the plant, with a focus on Arabidopsis thaliana.
Plants must achieve a balance between carbon assimilation, storage and growth, but little is known about how this is achieved. We describe evidence for the existence of regulatory mechanisms that ...coordinate carbon supply and use, and the likely central role of sugar signalling. We propose the existence of both 'acute' and 'acclimatory' responses to alterations in carbon supply, the latter tuning the balance between carbon supply and demand to optimise the capacity for sustained growth. A full understanding of these responses requires new, systems-level approaches that integrate information from transcriptomic, enzyme activity, metabolomic and growth analyses. We illustrate the complexity of acute and acclimatory responses by consideration of the control of starch synthesis and degradation in leaves. Finally, we consider how carbon balance may be linked to growth, and the importance of these linkages for sustained plant growth in a changing environment.
Historically speaking, Arabidopsis was not the plant of choice for investigating photosynthesis, with physiologists and biochemists favouring other species such as Chlorella, spinach and pea. ...However, its inherent advantages for forward genetics rapidly led to its adoption for photosynthesis research. In the last ten years, the availability of the Arabidopsis genome sequence - still the gold-standard for plant genomes - and the rapid expansion of genetic and genomic resources have further increased its importance. Research in Arabidopsis has not only provided comprehensive information about the enzymes and other proteins involved in photosynthesis, but has also allowed transcriptional responses, protein levels and compartmentation to be analysed at a global level for the first time. Emerging technical and theoretical advances offer another leap forward in our understanding of post-translational regulation and the control of metabolism. To illustrate the impact of Arabidopsis, we provide a historical review of research in primary photosynthetic metabolism, highlighting the role of Arabidopsis in elucidation of the pathway of photorespiration and the regulation of RubisCO, as well as elucidation of the pathways of starch turnover and studies of the significance of starch for plant growth.
Trehalose metabolism in plants Lunn, John Edward; Delorge, Ines; Figueroa, Carlos María ...
The Plant journal : for cell and molecular biology,
August 2014, Volume:
79, Issue:
4
Journal Article
Peer reviewed
Open access
Summary
Trehalose is a quantitatively important compatible solute and stress protectant in many organisms, including green algae and primitive plants. These functions have largely been replaced by ...sucrose in vascular plants, and trehalose metabolism has taken on new roles. Trehalose is a potential signal metabolite in plant interactions with pathogenic or symbiotic micro‐organisms and herbivorous insects. It is also implicated in responses to cold and salinity, and in regulation of stomatal conductance and water‐use efficiency. In plants, as in other eukaryotes and many prokaryotes, trehalose is synthesized via a phosphorylated intermediate, trehalose 6‐phosphate (Tre6P). A meta‐analysis revealed that the levels of Tre6P change in parallel with sucrose, which is the major product of photosynthesis and the main transport sugar in plants. We propose the existence of a bi‐directional network, in which Tre6P is a signal of sucrose availability and acts to maintain sucrose concentrations within an appropriate range. Tre6P influences the relative amounts of sucrose and starch that accumulate in leaves during the day, and regulates the rate of starch degradation at night to match the demand for sucrose. Mutants in Tre6P metabolism have highly pleiotropic phenotypes, showing defects in embryogenesis, leaf growth, flowering, inflorescence branching and seed set. It has been proposed that Tre6P influences plant growth and development via inhibition of the SNF1‐related protein kinase (SnRK1). However, current models conflict with some experimental data, and do not completely explain the pleiotropic phenotypes exhibited by mutants in Tre6P metabolism. Additional explanations for the diverse effects of alterations in Tre6P metabolism are discussed.
Plant growth is driven by photosynthetic carbon fixation during the day. Some photosynthate is accumulated, often as starch, to support nocturnal metabolism and growth at night. The rate of starch ...degradation in Arabidopsis leaves at night is essentially linear, and is such that almost all of the starch is used by dawn. We have investigated the timer that matches starch utilization to the duration of the night. The rate of degradation adjusted immediately and appropriately to an unexpected early onset of night. Starch was still degraded in an appropriate manner when the preceding light period was interrupted by a period of darkness. However, when Arabidopsis was grown in abnormal day lengths (28 h or 17 h) starch was exhausted ∼24 h after the last dawn, irrespective of the actual dawn. A mutant lacking the LHY and CCA1 clock components exhausted its starch at the dawn anticipated by its fast-running circadian clock, rather than the actual dawn. Reduced growth of wild-type plants in 28-h days and lhy/cca1 mutants in 24-h days was attributable to the inappropriate rate of starch degradation and the consequent carbon starvation at the end of night. Thus, starch degradation is under circadian control to ensure that carbohydrate availability is maintained until the next anticipated dawn, and this control is necessary for maintaining plant productivity.
Genome sequences from over 200 plant species have already been published, with this number expected to increase rapidly due to advances in sequencing technologies. Once a new genome has been ...assembled and the genes identified, the functional annotation of their putative translational products, proteins, using ontologies is of key importance as it places the sequencing data in a biological context. Furthermore, to keep pace with rapid production of genome sequences, this functional annotation process must be fully automated. Here we present a redesigned and significantly enhanced MapMan4 framework, together with a revised version of the associated online Mercator annotation tool. Compared with the original MapMan, the new ontology has been expanded almost threefold and enforces stricter assignment rules. This framework was then incorporated into Mercator4, which has been upgraded to reflect current knowledge across the land plant group, providing protein annotations for all embryophytes with a comparably high quality. The annotation process has been optimized to allow a plant genome to be annotated in a matter of minutes. The output results continue to be compatible with the established MapMan desktop application.
MapMan4 is a substantial redesign of the MapMan framework incorporating the latest literature knowledge to provide greatly enhanced protein family granularity. The online Mercator4 tool uses this framework to rapidly functionally annotate protein sequences from any land plant species.
Starch is a polysaccharide that is stored to be used in different timescales. Transitory starch is used during nighttime when photosynthesis is unavailable. Long-term starch is stored to support ...vegetative or reproductive growth, reproduction, or stress responses. Starch is not just a reserve of energy for most plants but also has many other roles, such as promoting rapid stomatal opening, making osmoprotectants, cryoprotectants, scavengers of free radicals and signals, and reverting embolised vessels. Biotic and abiotic stress vary according to their nature, strength, duration, developmental stage of the plant, time of the day, and how gradually they develop. The impact of stress on starch metabolism depends on many factors: how the stress impacts the rate of photosynthesis, the affected organs, how the stress impacts carbon allocation, and the energy requirements involved in response to stress. Under abiotic stresses, starch degradation is usually activated, but starch accumulation may also be observed when growth is inhibited more than photosynthesis. Under biotic stresses, starch is usually accumulated, but the molecular mechanisms involved are largely unknown. In this mini-review, we explore what has been learned about starch metabolism and plant stress responses and discuss the current obstacles to fully understanding their interactions.
Next‐generation technologies generate an overwhelming amount of gene sequence data. Efficient annotation tools are required to make these data amenable to functional genomics analyses. The Mercator ...pipeline automatically assigns functional terms to protein or nucleotide sequences. It uses the MapMan ‘BIN’ ontology, which is tailored for functional annotation of plant ‘omics’ data. The classification procedure performs parallel sequence searches against reference databases, compiles the results and computes the most likely MapMan BINs for each query. In the current version, the pipeline relies on manually curated reference classifications originating from the three reference organisms (Arabidopsis, Chlamydomonas, rice), various other plant species that have a reviewed SwissProt annotation, and more than 2000 protein domain and family profiles at InterPro, CDD and KOG. Functional annotations predicted by Mercator achieve accuracies above 90% when benchmarked against manual annotation. In addition to mapping files for direct use in the visualization software MapMan, Mercator provides graphical overview charts, detailed annotation information in a convenient web browser interface and a MapMan‐to‐GO translation table to export results as GO terms. Mercator is available free of charge via http://mapman.gabipd.org/web/guest/app/Mercator.
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