[Show abstract][Hide abstract] ABSTRACT: All plant productivity, including the food that we eat, arises from the capture of solar energy by plants. At most latitudes sunlight is available for only part of the 24 h day due to the rotation of the planet. This rhythmic and predictable alteration in the environment has driven the evolution of the circadian clock, which has an extremely pervasive influence upon plant molecular biology, physiology and phenology. A number of recent studies have demonstrated that the circadian clock is integrated very closely with photosynthesis and its metabolic products. We consider the coupling of the circadian oscillator with carbohydrate biochemistry and the connections between the nuclear-encoded circadian clock and processes within chloroplasts. We describe how this might provide adaptations to optimize plant performance in an environment that varies both predictably upon a daily and seasonal basis, and unpredictably due to the weather.
[Show abstract][Hide abstract] ABSTRACT: Circadian rhythms produce a biological measure of time that increases plant performance. The mechanisms that underlie this increase in productivity require investigation to provide information that will underpin future crop improvement. There is a growing body of evidence that a sophisticated signalling network interconnects the circadian oscillator and chloroplasts. We consider this in the context of circadian signalling to chloroplasts and the relationship between retrograde signalling and circadian regulation. We place circadian signalling to chloroplasts by sigma factors within an evolutionary context. We describe selected recent developments in the integration of light and circadian signals that control chloroplast gene expression.
Current Opinion in Plant Biology 10/2014; 21:43–50. DOI:10.1016/j.pbi.2014.06.008 · 7.85 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: A circadian rhythm matched to the phase and period of the day-night cycle has measurable benefits for land plants. We assessed the contribution of circadian period to the phasing of cellular events with the light : dark cycle. We also investigated the plasticity of circadian period within the Arabidopsis circadian oscillator. We monitored the circadian oscillator in wild-type and circadian period mutants under light : dark cycles of varying total duration. We also investigated changes in oscillator dynamics during and after the transition from light : dark cycles to free running conditions. Under light : dark cycles, dawn and dusk were anticipated differently when the circadian period was not resonant with the environmental period ('T cycle'). Entrainment to T cycles differing from the free-running period caused a short-term alteration in oscillator period. The transient plasticity of period was described by existing mathematical models of the Arabidopsis circadian network. We conclude that a circadian period resonant with the period of the environment is particularly important for anticipation of dawn and the timing of nocturnal events; and there is short-term and transient plasticity of period of the Arabidopsis circadian network.
New Phytologist 09/2013; 201(1). DOI:10.1111/nph.12489 · 7.67 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Correct circadian regulation increases plant productivity, and photosynthesis is circadian-regulated. Here, we discuss the regulatory basis for the circadian control of photosynthesis. We discuss candidate mechanisms underpinning circadian oscillations of light harvesting and consider how the circadian clock modulates CO2 fixation by Rubisco. We show that new techniques may provide a platform to better understand the signalling pathways that couple the circadian clock with the photosynthetic apparatus. Finally, we discuss how understanding circadian regulation in model systems is underpinning research into the impact of circadian regulation in crop species.
Photosynthesis Research 03/2013; 119(1-2). DOI:10.1007/s11120-013-9811-8 · 3.50 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Circadian timekeeping in plants increases photosynthesis and productivity. There are circadian oscillations in the abundance of many chloroplast-encoded transcripts, but it is not known how the circadian clock regulates chloroplast transcription or the photosynthetic apparatus. We show that, in Arabidopsis, nuclear-encoded SIGMA FACTOR5 (SIG5) controls circadian rhythms of transcription of several chloroplast genes, revealing one pathway by which the nuclear-encoded circadian oscillator controls rhythms of chloroplast gene expression. We also show that SIG5 mediates the circadian gating of light input to a chloroplast-encoded gene. We have identified an evolutionarily conserved mechanism that communicates circadian timing information between organelles with distinct genetic systems and have established a new level of integration between eukaryotic circadian clocks and organelles of endosymbiotic origin.
[Show abstract][Hide abstract] ABSTRACT: Circadian regulation is essential for optimum plant performance. In addition to loops and cascades of transcription and translation, the plant circadian clock and its associated signal transduction networks incorporate many post-translational mechanisms. Phosphorylation is a common feature of signal transduction and gene regulation. In this opinion article, we illustrate how phosphorylation events are positioned within the entrainment, functioning, and regulation of the circadian timing system. Phosphorylation regulates protein stability, protein-protein interactions and protein-DNA interactions within the core oscillator. We suggest that phosphorylation provides a potential mechanism for the distribution of circadian timing information within the cell and for the integration of circadian timing information with other signaling pathways.
[Show abstract][Hide abstract] ABSTRACT: Circadian clocks are 24-h timing devices that phase cellular responses; coordinate growth, physiology, and metabolism; and anticipate the day-night cycle. Here we report sensitivity of the Arabidopsis thaliana circadian oscillator to sucrose, providing evidence that plant metabolism can regulate circadian function. We found that the Arabidopsis circadian system is particularly sensitive to sucrose in the dark. These data suggest that there is a feedback between the molecular components that comprise the circadian oscillator and plant metabolism, with the circadian clock both regulating and being regulated by metabolism. We used also simulations within a three-loop mathematical model of the Arabidopsis circadian oscillator to identify components of the circadian clock sensitive to sucrose. The mathematical studies identified GIGANTEA (GI) as being associated with sucrose sensing. Experimental validation of this prediction demonstrated that GI is required for the full response of the circadian clock to sucrose. We demonstrate that GI acts as part of the sucrose-signaling network and propose this role permits metabolic input into circadian timing in Arabidopsis.
Proceedings of the National Academy of Sciences 03/2011; 108(12):5104-9. DOI:10.1073/pnas.1015452108 · 9.67 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Ca(2+) signals are a core regulator of plant cell physiology and cellular responses to the environment. The channels, pumps, and carriers that underlie Ca(2+) homeostasis provide the mechanistic basis for generation of Ca(2+) signals by regulating movement of Ca(2+) ions between subcellular compartments and between the cell and its extracellular environment. The information encoded within the Ca(2+) transients is decoded and transmitted by a toolkit of Ca(2+)-binding proteins that regulate transcription via Ca(2+)-responsive promoter elements and that regulate protein phosphorylation. Ca(2+)-signaling networks have architectural structures comparable to scale-free networks and bow tie networks in computing, and these similarities help explain such properties of Ca(2+)-signaling networks as robustness, evolvability, and the ability to process multiple signals simultaneously.
[Show abstract][Hide abstract] ABSTRACT: Xu et al. were unable to measure circadian oscillations of cyclic adenosine diphosphate ribose (cADPR). Their experiments showing
very low concentrations of cADPR lack appropriate controls, which suggests that technical limitations might explain their
negative result. Xu et al. also report that chemically induced ADP ribosyl cyclase did not alter clock function, which exactly replicates our findings.
[Show abstract][Hide abstract] ABSTRACT: Increases in cytosolic free Ca(2+) ([Ca(2+)](cyt)) are common to many stress-activated signalling pathways, including the response to saline environments. We have investigated the nature of NaCl-induced [Ca(2+)](cyt) signals in whole Arabidopsis thaliana seedlings using aequorin. We found that NaCl-induced increases in [Ca(2+)](cyt) are heterogeneous and mainly restricted to the root. Both the concentration of NaCl and the composition of the solution bathing the root have profound effects on the magnitude and dynamics of NaCl-induced increases in [Ca(2+)](cyt). Alteration of external K(+) concentration caused changes in the temporal and spatial pattern of [Ca(2+)](cyt) increase, providing evidence for Na(+)-induced Ca(2+) influx across the plasma membrane. The effects of various pharmacological agents on NaCl-induced increases in [Ca(2+)](cyt) indicate that NaCl may induce influx of Ca(2+) through both plasma membrane and intracellular Ca(2+)-permeable channels. Analysis of spatiotemporal [Ca(2+)](cyt) dynamics using photon-counting imaging revealed additional levels of complexity in the [Ca(2+)](cyt) signal that may reflect the oscillatory nature of NaCl-induced changes in single cells.
[Show abstract][Hide abstract] ABSTRACT: We have reported that Arabidopsis might have genetically distinct circadian oscillators in multiple cell-types.1 Rhythms of CHLOROPHYLL A/B BINDING PROTEIN2 (CAB2) promoter activity are 2.5 h longer in phytochromeB mutants in constant red light and in cryptocrome1 cry2 double mutant (hy4-1 fha-1) in constant blue light than the wild-type.2 However, we found that cytosolic free Ca(2+) ([Ca(2+)](cyt)) oscillations were undetectable in these mutants in the same light conditions.1 Furthermore, mutants of CIRCADIAN CLOCK ASSOCIATED1 (CCA1) have short period rhythms of leaf movement but have arrhythmic [Ca(2+)](cyt) oscillations. More important, the timing of cab1-1 (toc1-1) mutant has short period rhythms of CAB2 promoter activity ( approximately 21 h) but, surprisingly, has a wild-type period for circadian [Ca(2+)](cyt) oscillations ( approximately 24 h). In contrast, toc1-2, a TOC1 loss-of-function mutant, has a short period of both CAB2 and [Ca(2+)](cyt) rhythms ( approximately 21 h). Here we discuss the difference between the phenotypes of toc1-1 and toc1-2 and how rhythms of CAB2 promoter activity and circadian [Ca(2+)](cyt) oscillations might be regulated differently.
[Show abstract][Hide abstract] ABSTRACT: Transcriptional feedback loops are a feature of circadian clocks in both animals and plants. We show that the plant circadian clock also incorporates the cytosolic signaling molecule cyclic adenosine diphosphate ribose (cADPR). cADPR modulates the circadian oscillator's transcriptional feedback loops and drives circadian oscillations of Ca2+ release. The effects of antagonists of cADPR signaling, manipulation of cADPR synthesis, and mathematical simulation of the interaction of cADPR with the circadian clock indicate that cADPR forms a feedback loop within the plant circadian clock.
[Show abstract][Hide abstract] ABSTRACT: Plants have circadian oscillations in the concentration of cytosolic free calcium ([Ca(2+)](cyt)). To dissect the circadian Ca(2+)-signaling network, we monitored circadian [Ca(2+)](cyt) oscillations under various light/dark conditions (including different spectra) in Arabidopsis thaliana wild type and photoreceptor and circadian clock mutants. Both red and blue light regulate circadian oscillations of [Ca(2+)](cyt). Red light signaling is mediated by PHYTOCHROME B (PHYB). Blue light signaling occurs through the redundant action of CRYPTOCHROME1 (CRY1) and CRY2. Blue light also increases the basal level of [Ca(2+)](cyt), and this response requires PHYB, CRY1, and CRY2. Light input into the oscillator controlling [Ca(2+)](cyt) rhythms is gated by EARLY FLOWERING3. Signals generated in the dark also regulate the circadian behavior of [Ca(2+)](cyt). Oscillations of [Ca(2+)](cyt) and CHLOROPHYLL A/B BINDING PROTEIN2 (CAB2) promoter activity are dependent on the rhythmic expression of LATE ELONGATED HYPOCOTYL and CIRCADIAN CLOCK-ASSOCIATED1, but [Ca(2+)](cyt) and CAB2 promoter activity are uncoupled in the timing of cab1 (toc1-1) mutant but not in toc1-2. We suggest that the circadian oscillations of [Ca(2+)](cyt) and CAB2 promoter activity are regulated by distinct oscillators with similar components that are used in a different manner and that these oscillators may be located in different cell types in Arabidopsis.
The Plant Cell 12/2007; 19(11):3474-90. DOI:10.1105/tpc.106.046011 · 9.34 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Circadian clocks are signalling networks that enhance an organism's relationship with the rhythmic environment. The plant circadian clock modulates a wide range of physiological and biochemical events, such as stomatal and organ movements, photosynthesis and induction of flowering. Environmental signals regulate the phase and period of the plant circadian clock, which results in an approximate synchronization of clock outputs with external events. One of the consequences of circadian control is that stimuli of the same strength applied at different times of the day can result in responses of different intensities. This is known as 'gating'. Gating of a signal may allow plants to better process and react to the wide range and intensities of environmental signals to which they are constantly subjected. Light signalling, stomatal movements and low-temperature responses are examples of signalling pathways that are gated by the circadian clock. In this review, we describe the many levels at which the circadian clock interacts with responses to the environment. We discuss how environmental rhythms of temperature and light intensity entrain the circadian clock, how photoperiodism may be regulated by the relationship between environmental rhythms and the phasing of clock outputs, and how gating modulates the sensitivity of the clock and other responses to environmental and physiological signals. Finally, we describe evidence that the circadian clock can increase plant fitness.
[Show abstract][Hide abstract] ABSTRACT: Stomata are the major route of gas exchange between the atmosphere and the leaf interior. The size of the stomatal pore is controlled by movements of the stomatal guard cells. The guard cells close the stomatal pore to conserve water during stress. Under more favourable conditions, the stomatal movements optimise CO 2 uptake whilst minimising water loss. The movements of stomata are controlled by an extensive network of signalling pathways responding to diverse stimuli. One of the regulators of stomata is the circadian clock. We discuss the physiological mechanisms by which the clock may regulate stomatal movements, and the benefits that circadian regulation of stomatal behaviour may confer to the plant.
[Show abstract][Hide abstract] ABSTRACT: We tested the hypothesis that the circadian clock modulates Ca2+-based signalling pathways, using low-temperature (LT)-induced Ca2+ signals. We investigated the relationship between diurnal and circadian modulation of LT-induced increases in cytosolic-free calcium ([Ca2+]cyt), and regulation of [Ca2+]cyt-dependent outputs of the LT-signalling network (RD29A transcript abundance and stomatal closure). We measured [Ca2+]cyt non-invasively using aequorin, and targeted aequorin to the guard cell using a guard cell-specific GAL4-green fluorescent protein enhancer trap line. LT caused transient increases in whole plant and guard cell [Ca2+]cyt. In guard cells, the LT-induced [Ca2+]cyt elevation preceded stomatal closure. In whole plants, the magnitude of LT-induced [Ca2+]cyt transients, measured from the entire plant or specifically the guard cell, varied with the time of day: LT-induced [Ca2+]cyt transients were significantly higher during the mid-photoperiod than at the beginning or end. Diurnal variation in LT-induced guard cell [Ca2+]cyt increases was not correlated to diurnal variation in LT-induced stomatal closure. There was circadian modulation of LT-induced whole plant [Ca2+]cyt increases, which were correlated to the circadian pattern of RD29A induction. In order to understand the significance of LT-induced [Ca2+]cyt increases, we used a computer simulation to demonstrate that, in guard cells, LT-induced [Ca2+]cyt increases measured from a population of cells are likely to represent the summation of cold-induced single-cell [Ca2+]cyt oscillations.
[Show abstract][Hide abstract] ABSTRACT: Plants, like all eukaryotes and most prokaryotes, have evolved sophisticated mechanisms for anticipating predictable environmental changes that arise due to the rotation of the Earth on its axis. These mechanisms are collectively termed the circadian clock. Many aspects of plant physiology, metabolism and development are under circadian control and a large proportion of the transcriptome exhibits circadian regulation. In the present review, we describe the advances in determining the molecular nature of the circadian oscillator and propose an architecture of several interlocking negative-feedback loops. The adaptive advantages of circadian control, with particular reference to the regulation of metabolism, are also considered. We review the evidence for the presence of multiple circadian oscillator types located in within individual cells and in different tissues.
[Show abstract][Hide abstract] ABSTRACT: Circadian clocks are believed to confer an advantage to plants, but the nature of that advantage has been unknown. We show that a substantial photosynthetic advantage is conferred by correct matching of the circadian clock period with that of the external light-dark cycle. In wild type and in long- and short-circadian period mutants of Arabidopsis thaliana, plants with a clock period matched to the environment contain more chlorophyll, fix more carbon, grow faster, and survive better than plants with circadian periods differing from their environment. This explains why plants gain advantage from circadian control.