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      Light quality modulates metabolic synchronization over the diel phases of crassulacean acid metabolism

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          Summary

          Besides the acknowledged roles of red light, blue light is a key determinant for synchronizing the metabolic and physiological components of CAM over the day/night cycle.

          Abstract

          Temporal compartmentation of carboxylation processes is a defining feature of crassulacean acid metabolism and involves circadian control of key metabolic and transport steps that regulate the supply and demand for carbon over a 24h cycle. Recent insights on the molecular workings of the circadian clock and its connection with environmental inputs raise new questions on the importance of light quality and, by analogy, certain photoreceptors for synchronizing the metabolic components of CAM. The present work tested the hypothesis that optimal coupling of stomatal conductance, net CO 2 uptake, and the reciprocal turnover of carbohydrates and organic acids over the diel CAM cycle requires both blue and red light input signals. Contrasting monochromatic wavelengths of blue, green, and red light (i.e. 475, 530, 630nm) with low fluence rates (10 μmol m –2 s –1) were administered for 16 hours each diel cycle for a total treatment time of 48 hours to the obligate CAM bromeliad, Aechmea ‘Maya’. Of the light treatments imposed, low-fluence blue light was a key determinant in regulating stomatal responses, organic acid mobilization from the vacuole, and daytime decarboxylation. However, the reciprocal relationship between starch and organic acid turnover that is typical for CAM was uncoupled under low-fluence blue light. Under low-fluence red or green light, the diel turnover of storage carbohydrates was orchestrated in line with the requirements of CAM, but a consistent delay in acid consumption at dawn compared with plants under white or low-fluence blue light was noted. Consistent with the acknowledged influences of both red and blue light as input signals for the circadian clock, the data stress the importance of both red and blue-light signalling pathways for synchronizing the metabolic and physiological components of CAM over the day/night cycle.

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          Ecophysiology of Crassulacean Acid Metabolism (CAM).

          Crassulacean Acid Metabolism (CAM) as an ecophysiological modification of photosynthetic carbon acquisition has been reviewed extensively before. Cell biology, enzymology and the flow of carbon along various pathways and through various cellular compartments have been well documented and discussed. The present attempt at reviewing CAM once again tries to use a different approach, considering a wide range of inputs, receivers and outputs. Input is given by a network of environmental parameters. Six major ones, CO(2), H(2)O, light, temperature, nutrients and salinity, are considered in detail, which allows discussion of the effects of these factors, and combinations thereof, at the individual plant level ('physiological aut-ecology'). Receivers of the environmental cues are the plant types genotypes and phenotypes, the latter including morphotypes and physiotypes. CAM genotypes largely remain 'black boxes', and research endeavours of genomics, producing mutants and following molecular phylogeny, are just beginning. There is no special development of CAM morphotypes except for a strong tendency for leaf or stem succulence with large cells with big vacuoles and often, but not always, special water storage tissues. Various CAM physiotypes with differing degrees of CAM expression are well characterized. Output is the shaping of habitats, ecosystems and communities by CAM. A number of systems are briefly surveyed, namely aquatic systems, deserts, salinas, savannas, restingas, various types of forests, inselbergs and paramós. While quantitative census data for CAM diversity and biomass are largely missing, intuition suggests that the larger CAM domains are those systems which are governed by a network of interacting stress factors requiring versatile responses and not systems where a single stress factor strongly prevails. CAM is noted to be a strategy for variable, flexible and plastic niche occupation rather than lush productivity. 'Physiological syn-ecology' reveals that phenotypic plasticity constitutes the ecophysiological advantage of CAM.
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            Exploiting the potential of plants with crassulacean acid metabolism for bioenergy production on marginal lands.

            Crassulacean acid metabolism (CAM) is a photosynthetic adaptation that facilitates the uptake of CO(2) at night and thereby optimizes the water-use efficiency of carbon assimilation in plants growing in arid habitats. A number of CAM species have been exploited agronomically in marginal habitats, displaying annual above-ground productivities comparable with those of the most water-use efficient C(3) or C(4) crops but with only 20% of the water required for cultivation. Such attributes highlight the potential of CAM plants for carbon sequestration and as feed stocks for bioenergy production on marginal and degraded lands. This review highlights the metabolic and morphological features of CAM that contribute towards high biomass production in water-limited environments. The temporal separation of carboxylation processes that underpins CAM provides flexibility for modulating carbon gain over the day and night, and poses fundamental questions in terms of circadian control of metabolism, growth, and productivity. The advantages conferred by a high water-storage capacitance, which translate into an ability to buffer fluctuations in environmental water availability, must be traded against diffusive (stomatal plus internal) constraints imposed by succulent CAM tissues on CO(2) supply to the cellular sites of carbon assimilation. The practicalities for maximizing CAM biomass and carbon sequestration need to be informed by underlying molecular, physiological, and ecological processes. Recent progress in developing genetic models for CAM are outlined and discussed in light of the need to achieve a systems-level understanding that spans the molecular controls over the pathway through to the agronomic performance of CAM and provision of ecosystem services on marginal lands.
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              Guard cell photosynthesis and stomatal function.

              Chloroplasts are a key feature of most guard cells; however, the function of these organelles in stomatal responses has been a subject of debate. This review examines evidence for and against a role of guard cell chloroplasts in stimulating stomatal opening. Controversy remains over the extent to which guard cell Calvin cycle activity contributes to stomatal regulation. However, this is only one of four possible functions of guard cell chloroplasts; other roles include supply of ATP, blue-light signalling and starch storage. Evidence exists for all these mechanisms, but is highly dependent upon species and growth/measurement conditions, with inconsistencies between different laboratories reported. Significant plasticity and extreme flexibility in guard cell osmoregulatory, signalling and sensory pathways may be one explanation. The use of chlorophyll a fluorescence analysis of individual guard cells is discussed in assessing guard and mesophyll cell physiology in relation to stomatal function. Developments in transgenic and molecular techniques have recently provided interesting, albeit contrasting, data regarding the role of these highly conserved organelles in stomatal function. Recent studies examining the link between mesophyll photosynthesis and stomatal conductance are discussed. An enhanced understanding of these processes may be fundamental in generating crop plants with greater water use efficiencies, capable of combating future climatic changes.
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                Author and article information

                Journal
                J Exp Bot
                J. Exp. Bot
                jexbot
                exbotj
                Journal of Experimental Botany
                Oxford University Press (UK )
                0022-0957
                1460-2431
                July 2014
                6 May 2014
                6 May 2014
                : 65
                : 13 , Special Issue: C4 and CAM Photosynthesis in the New Millenium
                : 3705-3714
                Affiliations
                1Faculty of Engineering Technology, Department of Microbial and Molecular systems, Bioengineering Technology TC , KU Leuven Campus Geel, Kleinhoefstraat 4, B-2440 Geel, Belgium
                2School of Biology, Newcastle Institute for Research on Sustainability, Devonshire Building, Newcastle University , Newcastle upon Tyne, NE1 7RU, UK
                3Biosciences Division, Oak Ridge National Laboratory , Oak Ridge, TN 37831-6407, USA
                4Faculty of Bioscience Engineering, Department of Biosystems, Division of Crop Biotechnics , KU Leuven, Willem De Croylaan 42, B-3001 Heverlee, Belgium
                Author notes
                * To whom correspondence should be addressed. E-mail: johan.ceusters@ 123456kuleuven.be
                Article
                10.1093/jxb/eru185
                4085966
                24803500
                cce5de1d-f7e2-461d-98b5-f4248d243762
                © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/3.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                Page count
                Pages: 10
                Categories
                Research Paper

                Plant science & Botany
                cam,carbohydrate,gas exchange,light quality,pepc,pepck,titratable acidity.
                Plant science & Botany
                cam, carbohydrate, gas exchange, light quality, pepc, pepck, titratable acidity.

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