RAPTOR Controls Developmental Growth Transitions by Altering the Hormonal and Metabolic Balance.

Vegetative growth requires the systemic coordination of numerous cellular processes, which are controlled by regulatory proteins that monitor extracellular and intracellular cues and translate them into growth decisions. In eukaryotes, one of the central factors regulating growth is the serine/threonine protein kinase Target of Rapamycin (TOR), which forms complexes with regulatory proteins. To understand the function of one such regulatory protein, Regulatory-Associated Protein of TOR 1B (RAPTOR1B), in plants, we analyzed the effect of raptor1b mutations on growth and physiology in Arabidopsis (Arabidopsis thaliana) by detailed phenotyping, metabolomic, lipidomic, and proteomic analyses. Mutation of RAPTOR1B resulted in a strong reduction of TOR kinase activity, leading to massive changes in central carbon and nitrogen metabolism, accumulation of excess starch, and induction of autophagy. These shifts led to a significant reduction of plant growth that occurred nonlinearly during developmental stage transitions. This phenotype was accompanied by changes in cell morphology and tissue anatomy. In contrast to previous studies in rice (Oryza sativa), we found that the Arabidopsis raptor1b mutation did not affect chloroplast development or photosynthetic electron transport efficiency; however, it resulted in decreased CO2 assimilation rate and increased stomatal conductance. The raptor1b mutants also had reduced abscisic acid levels. Surprisingly, abscisic acid feeding experiments resulted in partial complementation of the growth phenotypes, indicating the tight interaction between TOR function and hormone synthesis and signaling in plants.

Are there tides within trees?

Background: Tree stem diameters and electrical stem potentials exhibit rhythmic variations with periodicities of 24-25 h. Under free-running conditions of constant light or darkness these rhythms were suggested to be mediated by the lunisolar gravitational force. Scope: To further unravel the regulation of tree stem diameter dilatations, many of the published time courses of diameter variations were re-evaluated in conjunction with the contemporaneous time courses of the lunisolar tidal acceleration. This was accomplished by application of the Etide program, which estimates, with high temporal resolution, local gravitational changes as a consequence of the diurnal variations of the lunisolar gravitational force due to the orbits and relative positions of Earth, Moon and Sun. In all instances investigated, it was evident that a synchronism exists between the times of the turning points of both the lunisolar tide and stem diameter variations when the direction of extension changes. This finding of synchrony documents that the lunisolar tide is a regulator of the tree stem diameter dilatations. Conclusions: Under the described experimental conditions, rhythms in tree stem diameter dilations and electrical stem potentials are controlled by the lunisolar gravitational acceleration.

Control of plant leaf movements by the lunisolar tidal force.

Background: Investigations into the diurnal ascent and descent of leaves of beans and other species, as well as experimental interventions into these movements, such as exposures to light at different times during the movement cycle, led to the concept of an endogenous 'clock' as a regulator of these oscillations. The causal origin of leaf movement can be traced to processes that modulate cell volume in target tissues of the pulvinus and petiole. However, these elements of the leaf-movement process do not sufficiently account for the rhythms that are generated following germination in constant light or dark conditions, or when plants are transferred to similar free-running conditions. Scope: To further unravel the regulation of leaf-movement rhythms, many of the published time courses of leaf movements that provided evidence for the concept of the endogenous clock were analysed in conjunction with the contemporaneous time courses of the lunisolar tidal acceleration. This was accomplished by application of the Etide program, which estimates, with high temporal resolution, local gravitational changes as a consequence of the diurnal variations of the lunisolar gravitational force due to the orbits and relative positions of Earth, Moon and Sun. To substantiate the results obtained in earthbound laboratories additional experiments were performed in the International Space Station (ISS). Tidal recurrence within the ISS exhibited a periodicity of 45 min. In all instances investigated, it was evident that a synchronism exists between the times of the turning points of both the lunisolar tide and of the leaftide when the direction of leaf movement changes. This finding of synchrony documents that the lunisolar tide is a regulator of the leaftide, and that the rhythm of leaf movement is not of endogenous origin but is an expression of an exogenous lunisolar clock impressed upon the leaf-movement apparatus. Conclusions: A huge number of correlations between leaftide and Etide time courses were established for leaf movement rhythms in natural conditions of the greenhouse, in conditions of constant light or dark, and under the microgravity conditions of the International Space Station. Even the apparently spontaneous short-period, small-amplitude rhythms recorded from leaves under unusual growth conditions are consistent with the hypothesis of a lunisolar zeitgeber. Synchronism between leaftide and Etide is discussed in terms of classical and quantum mechanics.

A proposal to explain how the circatidal rhythm of the Arabidopsis thaliana root elongation rate could be mediated by the lunisolar gravitational force: a quantum physical approach.

Background and Aims: Roots of Arabidopsis thaliana exhibit a 24.8 h oscillation of elongation rate when grown under free-running conditions. This growth rhythm is synchronized with the time course of the local lunisolar tidal acceleration. The present study aims at a physiological/physical model to describe the interaction of weak gravitational fields with cellular water dynamics that mediate rhythmic root growth profiles. Methods: Fundamental physical laws are applied to model the water dynamics within single plant cells in an attempt to mimic the 24.8 h oscillations in root elongation growth. In particular, a quantum gravitational description of the time course in root elongation is presented, central to which is the formation of coherent assemblies of mass due to the lunisolar gravitational field. Mathematical equations that characterize lunisolar gravity-induced coherent assemblies of water molecules are derived and related to the mass of cellular water within roots of A. thaliana. Key Results: The derived physical model of gravitationally modulated water assemblies is capable of accounting for the experimentally observed arabidopsis root growth kinetics under free-running conditions. The close analogy between the derived time-dependent lunisolar effect upon coherent molecular states of water within single cells and the coherent assemblies of electrons that characterize the quantum Hall effect is emphasized. Conclusions: The dynamics of the lunisolar-induced variation in coherent water assemblies provide a possible mechanism to describe the observed 24.8 h oscillation of root growth rate of A. thaliana. Therefore, this mechanism could function as an independent timekeeper to control cell elongation.

Lunisolar tidal force and its relationship to chlorophyll fluorescence in Arabidopsis thaliana.

The yield of chlorophyll fluorescence Ft was measured in leaves of Arabidopsis thaliana over periods of several days under conditions of continuous illumination (LL) without the application of saturating light pulses. After linearization of the time series of the chlorophyll fluorescence yield (ΔFt), oscillations became apparent with periodicities in the circatidal range. Alignments of these linearized time series ΔFt with the lunisolar tidal acceleration revealed high degrees of synchrony and phase congruence. Similar congruence with the lunisolar tide was obtained with the linearized quantum yield of PSII (ΔФII), recorded after application of saturating light pulses. These findings strongly suggest that there is an exogenous timekeeper which is a stimulus for the oscillations detected in both the linearized yield of chlorophyll fluorescence (ΔFt) and the linearized quantum yield of PSII (ΔФII).

Lunar gravity affects leaf movement of Arabidopsis thaliana in the International Space Station.

MAIN CONCLUSION: Cyclic leaf ascent and descent occur in synchrony and phase congruence with the lunisolar tidal force under a broad range of conditions. Digitized records of the vertical leaf movements of Arabidopsis thaliana were collected under space flight conditions in the International Space Station (ISS). Oscillations of leaf movements with periods of 45 and 90 min were found under light-adapted conditions, whereas in darkness, the periods were 45, 90, and 135 min. To demonstrate the close relationship between these oscillations and cyclical variations of the lunisolar gravitational force, we estimated the oscillations of the in-orbit lunisolar tide as they apply to the ISS, with the aid of the Etide software application. In general, in-orbit lunisolar gravitational profiles exhibited a periodicity of 45 min. Alignment of these in-orbit oscillations with the oscillations of Arabidopsis leaf movement revealed high degrees of synchrony and a congruence of phase. These data corroborate previous results which suggested a correlative relationship and a possible causal link between leaf movement rhythms obtained on ground and the rhythmic variation of the lunisolar tidal force.

Sulphate fertilization ameliorates long-term aluminum toxicity symptoms in perennial ryegrass (Lolium perenne).

Effects of the oxanion sulphate on plant aluminum (Al(3+)) detoxification mechanisms are not well understood. Therefore, holistic physiological and biochemical modifications induced by progressively increased doses of sulphate fertilization in the presence of long-term Al(3+) stress were investigated in the aluminum sensitive perennial ryegrass (Lolium perenne L. cvJumbo). Plant growth inhibition induced by Al(3+) was decreased in response to increasing doses of sulphate supply. Aluminum concentrations measured in roots of perennial ryegrass by atomic absorption spectrometry declined significantly with increasing sulphate concentrations. In parallel, we determined a rise of sulphur in shoots and roots of perennial ryegrass. Inclusion of up to 360 µM of sulphate enhanced cysteine and glutathione biosynthesis in Al(3+) (1.07 µM) treated plants. This increase of thiol-containing compounds favored all modifications in the glutathione redox balance, declining lipid peroxidation, decreasing the activity of superoxide dismutase, and modifying the expression of proteins involved in the diminution of Al(3+) toxicity in roots. In particular, proteome analysis by 1D-SDS-PAGE and LC-MS/MS allowed to identify up (e.g. vacuolar proton ATPase, proteosome ß subunit, etc) and down (Glyoxilase I, Ascorbate peroxidase, etc.) regulated proteins induced by Al(3+) toxicity symptoms in roots. Although, sulphate supply up to 480 µM caused a reduction in Al(3+) toxicity symptoms, it was not as efficient as compared to 360 µM sulphate fertilization. These results suggest that sulphate fertilization ameliorates Al(3+) toxicity responses in an intracellular specific manner within Lolium perenne.

Expression of Sucrose Transporter cDNAs Specifically in Companion Cells Enhances Phloem Loading and Long-Distance Transport of Sucrose but Leads to an Inhibition of Growth and the Perception of a Phosphate Limitation.

Sucrose (Suc) is the predominant form of carbon transported through the phloem from source to sink organs and is also a prominent sugar for short-distance transport. In all streptophytes analyzed, Suc transporter genes (SUTs or SUCs) form small families, with different subgroups evolving distinct functions. To gain insight into their capacity for moving Suc in planta, representative members of each clade were first expressed specifically in companion cells of Arabidopsis (Arabidopsis thaliana) and tested for their ability to rescue the phloem-loading defect caused by the Suc transporter mutation, Atsuc2-4. Sequence similarity was a poor indicator of ability: Several genes with high homology to AtSUC2, some of which have phloem-loading functions in other eudicot species, did not rescue the Atsuc2-4 mutation, whereas a more distantly related gene, ZmSUT1 from the monocot Zea mays, did restore phloem loading. Transporter complementary DNAs were also expressed in the companion cells of wild-type Arabidopsis, with the aim of increasing productivity by enhancing Suc transport to growing sink organs and reducing Suc-mediated feedback inhibition on photosynthesis. Although enhanced Suc loading and long-distance transport was achieved, growth was diminished. This growth inhibition was accompanied by increased expression of phosphate (P) starvation-induced genes and was reversed by providing a higher supply of external P. These experiments suggest that efforts to increase productivity by enhancing sugar transport may disrupt the carbon-to-P homeostasis. A model for how the plant perceives and responds to changes in the carbon-to-P balance is presented.

NAC transcription factor speedy hyponastic growth regulates flooding-induced leaf movement in Arabidopsis.

In rosette plants, root flooding (waterlogging) triggers rapid upward (hyponastic) leaf movement representing an important architectural stress response that critically determines plant performance in natural habitats. The directional growth is based on localized longitudinal cell expansion at the lower (abaxial) side of the leaf petiole and involves the volatile phytohormone ethylene (ET). We report the existence of a transcriptional core unit underlying directional petiole growth in Arabidopsis thaliana, governed by the NAC transcription factor speedy hyponastic growth (SHYG). Overexpression of SHYG in transgenic Arabidopsis thaliana enhances waterlogging-triggered hyponastic leaf movement and cell expansion in abaxial cells of the basal petiole region, while both responses are largely diminished in shyg knockout mutants. Expression of several expansin and xyloglucan endotransglycosylase/hydrolase genes encoding cell wall-loosening proteins was enhanced in SHYG overexpressors but lowered in shyg. We identified ACC oxidase5 (ACO5), encoding a key enzyme of ET biosynthesis, as a direct transcriptional output gene of SHYG and found a significantly reduced leaf movement in response to root flooding in aco5 T-DNA insertion mutants. Expression of SHYG in shoot tissue is triggered by root flooding and treatment with ET, constituting an intrinsic ET-SHYG-ACO5 activator loop for rapid petiole cell expansion upon waterlogging.

Swarms, swarming and entanglements of fungal hyphae and of plant roots.

There has been recent interest in the possibility that plant roots can show oriented collective motion, or swarming behavior. We examine the evidence supportive of root swarming and we also present new observations on this topic. Seven criteria are proposed for the definition of a swarm, whose application can help identify putative swarming behavior in plants. Examples where these criteria are fulfilled, at many levels of organization, are presented in relation to plant roots and root systems, as well as to the root-like mycelial cords (rhizomorphs) of fungi. The ideas of both an "active" swarming, directed by a signal which imposes a common vector on swarm element aggregation, and a "passive" swarming, where aggregation results from external constraint, are introduced. Active swarming is a pattern of cooperative behavior peculiar to the sporophyte generation of vascular plants and is the antithesis of the competitive behavior shown by the gametophyte generation of such plants, where passive swarming may be found. Fungal mycelial cords could serve as a model example of swarming in a multi-cellular, non-animal system.

Salt-responsive ERF1 regulates reactive oxygen species-dependent signaling during the initial response to salt stress in rice.

Early detection of salt stress is vital for plant survival and growth. Still, the molecular processes controlling early salt stress perception and signaling are not fully understood. Here, we identified salt-responsive ERF1 (SERF1), a rice (Oryza sativa) transcription factor (TF) gene that shows a root-specific induction upon salt and hydrogen peroxide (H2O2) treatment. Loss of SERF1 impairs the salt-inducible expression of genes encoding members of a mitogen-activated protein kinase (MAPK) cascade and salt tolerance-mediating TFs. Furthermore, we show that SERF1-dependent genes are H2O2 responsive and demonstrate that SERF1 binds to the promoters of MAPK kinase kinase6 (MAP3K6), MAPK5, dehydration-responsive element bindinG2A (DREB2A), and zinc finger protein179 (ZFP179) in vitro and in vivo. SERF1 also directly induces its own gene expression. In addition, SERF1 is a phosphorylation target of MAPK5, resulting in enhanced transcriptional activity of SERF1 toward its direct target genes. In agreement, plants deficient for SERF1 are more sensitive to salt stress compared with the wild type, while constitutive overexpression of SERF1 improves salinity tolerance. We propose that SERF1 amplifies the reactive oxygen species-activated MAPK cascade signal during the initial phase of salt stress and translates the salt-induced signal into an appropriate expressional response resulting in salt tolerance.

Arabidopsis thaliana root elongation growth is sensitive to lunisolar tidal acceleration and may also be weakly correlated with geomagnetic variations.

BACKGROUND: Correlative evidence suggests a relationship between the lunisolar tidal acceleration and the elongation rate of arabidopsis roots grown under free-running conditions of constant low light. METHODS: Seedlings of Arabidopsis thaliana were grown in a controlled-climate chamber maintained at a constant temperature and subjected to continuous low-level illumination from fluorescent tubes, conditions that approximate to a 'free-running' state in which most of the abiotic factors that entrain root growth rates are excluded. Elongation of evenly spaced, vertical primary roots was recorded continuously over periods of up to 14 d using high temporal- and spatial-resolution video imaging and were analysed in conjunction with geophysical variables. KEY RESULTS AND CONCLUSIONS: The results confirm the lunisolar tidal/root elongation relationship. Also presented are relationships between the hourly elongation rates and the contemporaneous variations in geomagnetic activity, as evaluated from the disturbance storm time and ap indices. On the basis of time series of root elongation rates that extend over ≥4 d and recorded at different seasons of the year, a provisional conclusion is that root elongation responds to variation in the lunisolar force and also appears to adjust in accordance with variations in the geomagnetic field. Thus, both lunisolar tidal acceleration and the geomagnetic field should be considered as modulators of root growth rate, alongside other, stronger and more well-known abiotic environmental regulators, and perhaps unexplored factors such as air ions. Major changes in atmospheric pressure are not considered to be a factor contributing to oscillations of root elongation rate.

High-throughput phenotyping of root growth dynamics.

Plant organ phenotyping by noninvasive video imaging techniques provides a powerful tool to assess physiological traits, circadian and diurnal rhythms, and biomass production. In particular, growth of individual plant organs is known to exhibit a high plasticity and occurs as a result of the interaction between various endogenous and environmental processes. Thus, any investigation aiming to unravel mechanisms that determine plant or organ growth has to accurately control and document the environmental growth conditions. Here we describe challenges in establishing a recently developed plant root monitoring platform (PlaRoM) specially suited for noninvasive high-throughput plant growth analysis with highest emphasis on the detailed documentation of capture time, as well as light and temperature conditions. Furthermore, we discuss the experimental procedure for measuring root elongation kinetics and key points that must be considered in such measurements. PlaRoM consists of a robotized imaging platform enclosed in a custom designed phytochamber and a root extension profiling software application. This platform has been developed for multi-parallel recordings of root growth phenotypes of up to 50 individual seedlings over several days, with high spatial and temporal resolution. Two Petri dishes are mounted on a vertical sample stage in a custom designed phytochamber that provides exact temperature control. A computer-controlled positioning unit moves these Petri dishes in small increments and enables continuous screening of the surface under a binocular microscope. Detection of the root tip is achieved by applying thresholds on image pixel data and verifying the neighbourhood for each dark pixel. The growth parameters are visualized as position over time or growth rate over time graphs and averaged over consecutive days, light-dark periods and 24 h day periods. This setup enables the investigation of root extension profiles of different genotypes in various growth conditions (e.g., light protocol, temperature, growth media) and is especially suited for the detection of diurnal or circadian growth rhythms.

Arabidopsis thaliana root growth kinetics and lunisolar tidal acceleration.

• All living organisms on Earth are continually exposed to diurnal variations in the gravitational tidal force due to the Sun and Moon. • Elongation of primary roots of Arabidopsis thaliana seedlings maintained at a constant temperature was monitored for periods of up to 14 d using high temporal- and spatial-resolution video imaging. The time-course of the half-hourly elongation rates exhibited an oscillation which was maintained when the roots were placed in the free-running condition of continuous illumination. • Correlation between the root growth kinetics collected from seedlings initially raised under several light protocols but whose roots were subsequently in the free-running condition and the lunisolar tidal profiles enabled us to identify that the latter is the probable exogenous determinant of the rhythmic variation in root elongation rate. Similar observations and correlations using roots of Arabidopsis starch mutants suggest a central function of starch metabolism in the response to the lunisolar tide. The periodicity of the lunisolar tidal signal and the concomitant adjustments in root growth rate indicate that an exogenous timer exists for the modulation of root growth and development. • We propose that, in addition to the sensitivity to Earthly 1G gravity, which is inherent to all animals and plants, there is another type of responsiveness which is attuned to the natural diurnal variations of the lunisolar tidal force.

Lunisolar tidal force and the growth of plant roots, and some other of its effects on plant movements.

BACKGROUND: Correlative evidence has often suggested that the lunisolar tidal force, to which the Sun contributes 30 % and the Moon 60 % of the combined gravitational acceleration, regulates a number of features of plant growth upon Earth. The time scales of the effects studied have ranged from the lunar day, with a period of approx. 24.8 h, to longer, monthly or seasonal variations. SCOPE: We review evidence for a lunar involvement with plant growth. In particular, we describe experimental observations which indicate a putative lunar-based relationship with the rate of elongation of roots of Arabidopsis thaliana maintained in constant light. The evidence suggests that there may be continuous modulation of root elongation growth by the lunisolar tidal force. In order to provide further supportive evidence for a more general hypothesis of a lunisolar regulation of growth, we highlight similarly suggestive evidence from the time courses of (a) bean leaf movements obtained from kymographic observations; (b) dilatation cycles of tree stems obtained from dendrograms; and (c) the diurnal changes of wood-water relationships in a living tree obtained by reflectometry. CONCLUSIONS: At present, the evidence for a lunar or a lunisolar influence on root growth or, indeed, on any other plant system, is correlative, and therefore circumstantial. Although it is not possible to alter the lunisolar gravitational force experienced by living organisms on Earth, it is possible to predict how this putative lunisolar influence will vary at times in the near future. This may offer ways of testing predictions about possible Moon-plant relationships. As for a hypothesis about how the three-body system of Earth-Sun-Moon could interact with biological systems to produce a specific growth response, this remains a challenge for the future. Plant growth responses are mainly brought about by differential movement of water across protoplasmic membranes in conjunction with water movement in the super-symplasm. It may be in this realm of water movements, or even in the physical forms which water adopts within cells, that the lunisolar tidal force has an impact upon living growth systems.

NFX1-LIKE2 (NFXL2) suppresses abscisic acid accumulation and stomatal closure in Arabidopsis thaliana.

The NFX1-LIKE1 (NFXL1) and NFXL2 genes were identified as regulators of salt stress responses. The NFXL1 protein is a nuclear factor that positively affects adaptation to salt stress. The nfxl1-1 loss-of-function mutant displayed reduced survival rates under salt and high light stress. In contrast, the nfxl2-1 mutant, defective in the NFXL2 gene, and NFXL2-antisense plants exhibited enhanced survival under these conditions. We show here that the loss of NFXL2 function results in abscisic acid (ABA) overaccumulation, reduced stomatal conductance, and enhanced survival under drought stress. The nfxl2-1 mutant displayed reduced stomatal aperture under all conditions tested. Fusicoccin treatment, exposition to increasing light intensities, and supply of decreasing CO(2) concentrations demonstrated full opening capacity of nfxl2-1 stomata. Reduced stomatal opening presumably is a consequence of elevated ABA levels. Furthermore, seedling growth, root growth, and stomatal closure were hypersensitive to exogenous ABA. The enhanced ABA responses may contribute to the improved drought stress resistance of the mutant. Three NFXL2 splice variants were cloned and named NFXL2-78, NFXL2-97, and NFXL2-100 according to the molecular weight of the putative proteins. Translational fusions to the green fluorescent protein suggest nuclear localisation of the NFXL2 proteins. Stable expression of the NFXL2-78 splice variant in nfxl2-1 plants largely complemented the mutant phenotype. Our data show that NFXL2 controls ABA levels and suppresses ABA responses. NFXL2 may prevent unnecessary and costly stress adaptation under favourable conditions.

Tomato fruit photosynthesis is seemingly unimportant in primary metabolism and ripening but plays a considerable role in seed development.

Fruit of tomato (Solanum lycopersicum), like those from many species, have been characterized to undergo a shift from partially photosynthetic to truly heterotrophic metabolism. While there is plentiful evidence for functional photosynthesis in young tomato fruit, the rates of carbon assimilation rarely exceed those of carbon dioxide release, raising the question of its role in this tissue. Here, we describe the generation and characterization of lines exhibiting a fruit-specific reduction in the expression of glutamate 1-semialdehyde aminotransferase (GSA). Despite the fact that these plants contained less GSA protein and lowered chlorophyll levels and photosynthetic activity, they were characterized by few other differences. Indeed, they displayed almost no differences in fruit size, weight, or ripening capacity and furthermore displayed few alterations in other primary or intermediary metabolites. Although GSA antisense lines were characterized by significant alterations in the expression of genes associated with photosynthesis, as well as with cell wall and amino acid metabolism, these changes were not manifested at the phenotypic level. One striking feature of the antisense plants was their seed phenotype: the transformants displayed a reduced seed set and altered morphology and metabolism at early stages of fruit development, although these differences did not affect the final seed number or fecundity. Taken together, these results suggest that fruit photosynthesis is, at least under ambient conditions, not necessary for fruit energy metabolism or development but is essential for properly timed seed development and therefore may confer an advantage under conditions of stress.

Mutations in leaf starch metabolism modulate the diurnal root growth profiles of Arabidopsis thaliana.

Roots of Arabidopsis thaliana exhibit stable diurnal growth profiles that are controlled by the circadian clock. Here we describe the effects of mutations in leaf starch metabolism on the diurnal root growth characteristics of Arabidopsis thaliana. High temporal and spatial resolution video imaging was performed to quantify the growth kinetics of Arabidopsis wild-type as well as pgm, sex1, mex1, dpe1 and dpe2 starch metabolism mutants grown in three different photoperiods. As a result, root growth patterns of all genotypes displayed characteristic modifications in their diurnal kinetics that were also affected by the photoperiod. To further investigate the role of starch derived substrate deficiency on root growth, the effect of 0.05% extracellular sucrose was studied in 12 h-12 h light-dark cycles.

Circadian control of root elongation and C partitioning in Arabidopsis thaliana.

Plants grow in a light/dark cycle. We have investigated how growth is buffered against the resulting changes in the carbon supply. Growth of primary roots of Arabidopsis seedlings was monitored using time-resolved video imaging. The average daily rate of growth is increased in longer light periods or by addition of sugars. It responds slowly over days when the conditions are changed. The momentary rate of growth exhibits a robust diel oscillation with a minimum 8-9 h after dawn and a maximum towards the end of the night. Analyses with starch metabolism mutants show that starch turnover is required to maintain growth at night. A carbon shortfall leads to an inhibition of growth, which is not immediately reversed when carbon becomes available again. The diel oscillation persists in continuous light and is strongly modified in clock mutants. Central clock functions that depend on CCA1/LHY are required to set an appropriate rate of starch degradation and maintain a supply of carbon to support growth through to dawn, whereas ELF3 acts to decrease growth in the light period and promote growth in the night. Thus, while the overall growth rate depends on the carbon supply, the clock orchestrates diurnal carbon allocation and growth.