Chloroplast Movement Research Articles

Light stress-induced chloroplast movement and midday depression of photosynthesis in sorghum leaves

ABSTRACTPlants are exposed to high light intensity, high leaf temperatures and high air-to-leaf water vapor pressure deficit (ALVPD) during the day. These environmental stresses cause stomatal closure and photoinhibitory damage, leading to midday depression of photosynthesis. Chloroplast positioning is essential for the efficient operation of photosynthesis. However, chloroplast behavior before, during, and even after the midday depression of photosynthesis remains unknown. We investigated changes in the intracellular positioning of chloroplasts and photosynthetic traits under a diurnal pattern of light. Sorghum leaves were exposed to a 12-h regime of light mimicking the natural light environment, with constant leaf temperature and ALVPD. Net photosynthetic rate (Pn) showed a diurnal pattern, and midday depression in Pn was observed at 3.8 h of irradiation. Depression in Pn was attributed to stomatal limitation because the decrease in Pn was in accordance with the decrease in stomatal conductance. The maximum efficiency of photosystem II decreased with the increase in light intensity and remained low after 12 h of irradiation. Bundle sheath chloroplasts swelled after 8 h of irradiation, representing the accumulation of starch. Conversely, mesophyll chloroplasts exhibited avoidance response after 4 h of irradiation, and the avoidance position was maintained during the remainder of the daytime. These data suggest that chloroplasts are subject to light stress during and after the midday depression of photosynthesis. The intensity of natural light is excessive for most of the day and this light stress induces chloroplast avoidance response and depression of photosynthesis.

Slow induction of chlorophyll a fluorescence excited by blue and red light in Tradescantia leaves acclimated to high and low light.

Tradescantia is a good model for assaying induction events in higher plant leaves. Chlorophyll (Chl) fluorescence serves as a sensitive reporter of the functional state of photosynthetic apparatus in chloroplasts. The fluorescence time-course depends on the leaf growth conditions and actinic light quality.In this work, we investigated slow induction of Chl a fluorescence (SIF) excited by blue light (BL, λmax = 455nm) or red light (RL, λmax = 630nm) in dark-adapted leaves of Tradescantia fluminensis acclimated to high light (~ 1000µmol photons m-2s-1; HL) or low light (~ 100µmol photons m-2s-1; LL). Our special interest was focused on the contribution of the avoidance response to SIF kinetics. Bearing in mind that BL and RL have different impacts on photoreceptors that initiate chloroplast movements within the cell (accumulation/avoidance responses), we have compared the SIF patterns during the action of BL and RL. The time-courses of SIF and kinetics of non-photochemical quenching (NPQ) of Chl a fluorescence revealed a certain difference when leaves were illuminated by BL or RL. In both cases, the yield of fluorescence rose to the maximal level P and then, after the lag-phase P-S-M1, the fluorescence level decreased toward the steady state T (via the intermediate phases M1-M2 and M2-T). In LL-acclimated leaves, the duration of the P-S-M1 phase was almost two times longer that in HL-grown plants. In the case of BL, the fluorescence decay included the transient phase M1-M2. This phase was obscure during the RL illumination. Non-photochemical quenching of Chl a fluorescence has been quantified as [Formula: see text], where [Formula: see text] and [Formula: see text] stand for the fluorescence response to saturating pulses of light applied to dark-adapted and illuminated samples, respectively. The time-courses of such a formally determined NPQ value were markedly different during the action of RL and BL. In LL-grown leaves, BL induced higher NPQ as compared to the action of RL. In HL-grown plants, the difference between the NPQ responses to BL and RL illumination was insignificant. Comparing the peculiarities of Chl a fluorescence induced by BL and RL, we conclude that the avoidance response can provide a marked contribution to SIF and NPQ generation. The dependence of NPQ on the quality of actinic light suggests that chloroplast movements within the cell have a noticeable impact on the formally determined NPQ value. Analyzing kinetics of post-illumination decay of NPQ in the context of solar stress resistance, we have found that LL-acclimated Tradescantia leaves are more vulnerable to strong light than the HL-grown leaves.

Seasonal movement of chloroplasts in mesophyll cells of two Picea species.

Cold acclimation in evergreen conifers of temperate zone is associated with seasonal structural changes of mesophyll cells. Photoprotective reactions include the movement of chloroplasts from summer position when they are located along the cell walls to winter arrangement with their aggregation in one part of the cell. Special spatial arrangement of mesophyll in Picea species with chloroplasts located along the two opposite cell walls causes the very specific pattern of chloroplast movement. To reveal the intracellular apparatus involved in the seasonal organelle position changes, 3D reconstruction of mesophyll cell structure was applied. Two Picea species, P. obovata and P. pungens, from two geographic regions were studied in a 3-year course. The involvement of small transparent vacuoles in the development of cytoplasmic strands penetrating through the central vacuole and connecting two opposite lateral sides of the cell was shown. The nucleus and cytoplasmic organelles including chloroplasts move inwards the strand forming the cytoplasmic conglomerate enclosed by the vacuole at the cell center. Two Picea species have distinct differences in spatial organization of winter mesophyll cells and in structural events leading to its formation. Analysis of Picea species from two geographic regions over 3years of monitoring reveals dependence of seasonal organelle movement on the dynamics of temperature decline in autumn.

Relationship between relocation of phototropin to the chloroplast periphery and the initiation of chloroplast movement in Marchantia polymorpha.

The blue‐light photoreceptor kinase phototropin (phot) mediates chloroplast movement in response to light and temperature. Phot predominantly localizes at the plasma membrane, but also resides in the cytosol and the chloroplast periphery. Although the phot localized to the chloroplast periphery is thought to mediate chloroplast movement, the localization mechanism is unknown. In this study, we found that chloroplast movement does not occur in 0‐day‐old gemma cells of the liverwort Marchantia polymorpha but that the movement is induced in 1‐day‐old gemmaling cells. Along with this physiological change, the subcellular localization of phot also changed: In 0‐day‐old gemma cells, phot localized at the plasma membrane and the cytosol, but in 1‐day‐old gemmaling cells, the phot disappeared from the cytosol and appeared at the chloroplast periphery. When the relocalization was tracked using a photoconvertible fluorescent protein, the cytosolic phot relocated to the plasma membrane, and the plasma membrane‐resident phot relocated to the chloroplast periphery. The blue‐light‐dependent activation of phot kinase activity enhanced this relocalization. Mutated phot deficient in blue‐light reception or kinase activity had a severely reduced ability to localize at the chloroplast periphery. These findings suggest that photoactivated phot localizes at the chloroplast periphery to initiate chloroplast movement.

Chlorophyll fluorescence parameters to assess utilization of excitation energy in photosystem II independently of changes in leaf absorption

Measurement of Pulse-Amplitude-Modulated (PAM) chlorophyll a fluorescence is widely used method for obtaining information on the functional state of photosystem II (PSII). Recently, it has been shown that some of long-established fluorescence parameters must be interpreted with caution, when the light-induced chloroplast movements occur. In our work we have analyzed the effect of chloroplast movements on these parameters. We have derived new parameters that are independent of the change in PSII absorption occurring during measurement. To verify whether there is a need for new parameters or the difference between the parameters commonly used and the newly derived ones is insignificant, we conducted an experiment with Arabidopsis thaliana wild type plants and its phot1 phot2 mutant defective in chloroplast movement. Plants were exposed to light of different qualities (450, 470, 550 or 660 nm) and quantities (100, 400 or 1200 μmol m−2 s−1) for up to 40 min. Since the blue light-induced chloroplast avoidance reaction is a photoprotective mechanism, we expected that phot1 phot2 mutant will compensate the lack of this mechanism by increasing non-photochemical quenching. However, using the light at both 450 and 470 nm, the calculation of commonly used parameter, ΦNPQ (quantum yield of regulated light-induced thermal energy dissipation in PSII) based on Hendrickson et al. [L. Hendrickson, R.T. Furbank, W.S. Chow, Photosynth. Res. 82 (2004) 73–81] showed the opposite. On the other hand, the results obtained using our newly proposed formulae to determine quantum yield of PSII thermal energy dissipation were in line with our assumption. Thus, the experimental data showed that some formulae of fluorescence parameters are dependent on the change in PSII absorption and need to be interpreted carefully. On the contrary, the formulae introduced by us can remove the effect of changes in PSII absorption that occur during measurement, without additional measurements, and give the real estimate of light-induced non-photochemical quenching.

Phototropins Mediate Chloroplast Movement in Phalaenopsis aphrodite (Moth Orchid).

Chloroplast movement is important for plants to avoid photodamage and to perform efficient photosynthesis. Phototropins are blue light receptors in plants that function in chloroplast movement, phototropism, stomatal opening, and they also affect plant growth and development. In this study, full-length cDNAs of two PHOTOTROPIN genes, PaPHOT1 and PaPHOT2, were cloned from a moth orchid Phalaenopsis aphrodite, and their functions in chloroplast movement were investigated. Phylogenetic analysis showed that PaPHOT1 and PaPHOT2 orthologs were highly similar to PHOT1 and PHOT2 of the close relative Phalaenopsis equestris, respectively, and clustered with monocots PHOT1 and PHOT2 orthologs, respectively. Phalaenopsis aphrodite expressed a moderate level of PaPHOT1 under low blue light of 5 μmol�m-2�s-1 (BL5) and a high levels of PaPHOT1 at >BL100. However, PaPHOT2 was expressed at low levels at <BL50 but expressed at high levels at > BL100. Analysis of light-induced chloroplast movements using the SPAD method indicated that orchid accumulated chloroplasts at <BL10. The chloroplast avoidance response was detectable at >BL25 and significant chloroplast avoidance movement was observed at >BL100. Virus-induced gene silencing of PaPHOTs in orchids showed decreased gene expression of PaPHOTs and reduced both chloroplast accumulation and avoidance responses. Heterologous expression of PaPHOT1 in Arabidopsis phot1phot2 double mutant recovered chloroplast accumulation response at BL5, but neither PaPHOT1 nor PaPHOT2 was able to restore mutant chloroplast avoidance at BL100. Overall, this study showed that phototropins mediate chloroplast movement in Phalaenopsis orchid is blue light-dependent but their function is slightly different from Arabidopsis which might be due to gene evolution.

Advancing Terrestrial Ecosystem Science With a Novel Automated Measurement System for Sun‐Induced Chlorophyll Fluorescence for Integration With Eddy Covariance Flux Networks

AbstractSun‐induced chlorophyll fluorescence (SIF) provides critical information on the dynamics of gross primary productivity, a unique role not readily achievable using other methods. Long‐term continuous SIF observations have the potential to advance terrestrial ecosystem science. Realizing this potential, however, requires synergistic implementation of SIF measurements within eddy covariance (EC) flux networks. There is a need for SIF systems that can integrate seamlessly with EC instrumentation to maximize synergistic use of obtained data. Here we introduce the Fluorescence Auto‐Measurement Equipment (FAME) and protocol that fulfill such a purpose. FAME is designed specifically for plug‐and‐play integration with existing EC data acquisition systems. Its innovative hardware and software designs provide versatility, extensibility, autonomous operation, and ease of maintenance for acquiring SIF data of high quality and quantity. A major novel feature of FAME is its synchronized sampling of spectral irradiance and environmental variables, allowing for more precise interpretation of the SIF signal. FAME has been deployed since September 2016 at the Missouri Ozark AmeriFlux site, providing high‐quality measurements even when air temperatures approached 40 °C. Results reveal that canopy SIF saturated or even slightly decreased at high light, similar to leaf‐level photosynthesis. Clear diurnal hysteresis was observed: For the same light, morning SIF was higher than afternoon. Dynamic energy dissipation processes and stress‐induced movements of chloroplasts and leaves may explain the observed pattern. The technology and measurement protocol introduced here advances the coordinated observation of SIF and EC fluxes and represents a step change in observational ecosystem and carbon cycle science research.

Functional characterization of GhPHOT2 in chloroplast avoidance of Gossypium hirsutum

Chloroplast movement mediated by the plant-specific phototropin blue light photoreceptors is crucial for plants to cope with fluctuating light conditions. While chloroplasts accumulate at weak light-illuminated areas, chloroplast avoidance response mediated primarily by the phototropin2 (phot2) receptor is induced by strong light illumination. Although extensive studies have been performed on phot2-mediated chloroplast avoidance in the model plant Arabidopsis, little is known on the role of the corresponding PHOT2 orthologs in chloroplast movement in cotton. In this study, we found that chloroplast avoidance movement also occurs in the tetraploid G. hirsutum and two diploid species, G. arboreum and G. raimondii, albeit with distinct features. Further bioinformatics and genetic analysis identified the cotton PHOT2 ortholog, GhPHOT2-1, which retained a conserved role in plant chloroplast avoidance movement under strong blue light. Ghphot2-1was localized in the plasma membrane and formed aggregates after high blue light irradiation. Constitutive expression of GhPHOT2-1 restored chloroplast avoidance and accumulation response, as well as phototropism, and leaf flattening characteristics of the Arabidopsis phot2 or phot1 phot2 mutants. On the contrary, silencing of GhPHOT2-1 by virus-induced gene silencing (VIGS) disrupted high blue light-induced chloroplast avoidance movement and caused photo damage in cotton leaves. Taken together, these findings demonstrated that GhPHOT2-1 is a conserved PHOT2 ortholog in regulating chloroplast avoidance and the other aforementioned phot2-mediated responses, implicating its potential role for improving high light tolerance in cotton cultivars.

Chloroplast Accumulation Response Enhances Leaf Photosynthesis and Plant Biomass Production.

Under high light intensity, chloroplasts avoid absorbing excess light by moving to anticlinal cell walls (avoidance response), but under low light intensity, chloroplasts accumulate along periclinal cell walls (accumulation response). In most plant species, these responses are induced by blue light and are mediated by the blue light photoreceptor, phototropin, which also regulates phototropism, leaf flattening, and stomatal opening. These phototropin-mediated responses could enhance photosynthesis and biomass production. Here, using various Arabidopsis (Arabidopsis thaliana) mutants deficient in chloroplast movement, we demonstrated that the accumulation response enhances leaf photosynthesis and plant biomass production. Conspicuously, phototropin2 mutant plants specifically defective in the avoidance response but not in other phototropin-mediated responses displayed a constitutive accumulation response irrespective of light intensities, enhanced leaf photosynthesis, and increased plant biomass production. Therefore, our findings provide clear experimental evidence of the importance of the chloroplast accumulation response in leaf photosynthesis and biomass production.

Phototropins of the moss Physcomitrella patens function as blue-light receptors for phototropism in Arabidopsis

ABSTRACTFour phototropin genes (PHOTA1, PHOTA2, PHOTB1, PHOTB2) have been isolated in the moss Physcomitrella patens. These genes encode phototropins that mediate blue-light–induced chloroplast movement. However, the individual functions of these phototropins, including the function of mediating blue-light–induced phototropism, remain unclear. To elucidate the individual functions of P. patens phototropins, each of these phototropin genes was expressed in a phototropin-deficient mutant of Arabidopsis (phot1-5 phot2-1). In addition, fluorescence of GFP fused to these phototropins was examined to determine the subcellular localization of each phototropin. Our results demonstrate that all four P. patens phototropins mediate blue-light–induced phototropism and are associated with the plasma membrane in Arabidopsis.Abbreviations GFP: green fluorescent protein; Pp_phot: Physcomitrella patens phototropin

Dynamic coordination of plastid morphological change by cytoskeleton for chloroplast-nucleus communication during plant immune responses

ABSTRACTConsidering their sessile life, plants must efficiently coordinate their resources and energy for maintaining their presence in normal living conditions and for defending themselves against environmental threats. Collaboration between multiple subcellular compartments is a common strategy in several biological processes to modify cells’ architecture for their growth and development and to respond to acute changes in the environment. When plants defend themselves against microbial pathogens, chloroplasts generate tubular structures – so-called stromules- to facilitate chloroplast movement towards nuclei during innate immunity. Morphological changes and movements of stromules are directed by interactions with microtubule and actin cytoskeleton. Microtubules provide a direction for the stromule extension, while actin filaments restrict stromule retraction which provides a driving force for repositioning of chloroplast near nucleus during plant immune responses. These findings implicated a critical role for stromules in signal transduction from chloroplast to the nucleus in plant defense.

Leaf anatomy does not explain apparent short-term responses of mesophyll conductance to light and CO2 in tobacco.

Mesophyll conductance to CO2 (gm ), a key photosynthetic trait, is strongly constrained by leaf anatomy. Leaf anatomical parameters such as cell wall thickness and chloroplast area exposed to the mesophyll intercellular airspace have been demonstrated to determine gm in species with diverging phylogeny, leaf structure and ontogeny. However, the potential implication of leaf anatomy, especially chloroplast movement, on the short-term response of gm to rapid changes (i.e. seconds to minutes) under different environmental conditions (CO2 , light or temperature) has not been examined. The aim of this study was to determine whether the observed rapid variations of gm in response to variations of light and CO2 could be explained by changes in any leaf anatomical arrangements. When compared to high light and ambient CO2 , the values of gm estimated by chlorophyll fluorescence decreased under high CO2 and increased at low CO2 , while it decreased with decreasing light. Nevertheless, no changes in anatomical parameters, including chloroplast distribution, were found. Hence, the gm estimated by analytical models based on anatomical parameters was constant under varying light and CO2 . Considering this discrepancy between anatomy and chlorophyll fluorescence estimates, it is concluded that apparent fast gm variations should be due to artefacts in its estimation and/or to changes in the biochemical components acting on diffusional properties of the leaf (e.g. aquaporins and carbonic anhydrase).

Partitioning of absorbed light energy within photosystem II in barley can be affected by chloroplast movement

Plants have developed many ways to protect reaction centres of photosystems against overexcitation. One of the mechanisms involves reduction of the leaf absorption cross-section by light-induced chloroplast avoidance reaction. Decrease in the probability of photon absorption by the pigments bound within photosystem II (PSII) complexes leads to the increase in quantum yield of PSII photochemistry (ΦPSII). On the other hand, the decrease of PSII excitation probability causes reduction of chlorophyll a fluorescence intensity which is manifested as the apparent increase of determined quantum yield of regulated light-induced non-photochemical quenching (ΦNPQ). Absorption of different light intensity by phototropins led to the different chloroplast distribution within barley leaves, estimated by measurement of the leaf transmittance. Due to a weak blue light used for transmittance measurements, leaves exposed to actinic light with wavelengths longer than 520 nm undergo chloroplast accumulation reaction, in contrast with leaves exposed to light with shorter wavelengths, that showed a different extent of chloroplast avoidance reaction. Based on the ΦNPQ action spectra measured simultaneously with the transmittance, the influence of different chloroplast distribution on ΦNPQ was assessed. The analysis of results showed that decrease in the leaf absorption cross-section due to increasing part of chloroplasts reaching profile position significantly affected the partitioning of excitation energy within PSII and such rearrangement also distorted measured ΦNPQ and cannot be neglected in its interpretation. When the majority of chloroplasts reached profile position, the photoprotective effect appeared to be the most prominent for strong blue light that has the highest absorption in the upper leaf layers in comparison with green or red ones.

A chemical genetic approach to engineer phototropin kinases for substrate labeling

Protein kinases (PKs) control many aspects of plant physiology by regulating signaling networks through protein phosphorylation. Phototropins (phots) are plasma membrane-associated serine/threonine PKs that control a range of physiological processes that collectively serve to optimize photosynthetic efficiency in plants. These include phototropism, leaf positioning and flattening, chloroplast movement, and stomatal opening. Despite their identification over two decades ago, only a handful of substrates have been identified for these PKs. Progress in this area has been hampered by the lack of a convenient means to confirm the identity of potential substrate candidates. Here we demonstrate that the kinase domain of Arabidopsis phot1 and phot2 can be successfully engineered to accommodate non-natural ATP analogues by substituting the bulky gatekeeper residue threonine for glycine. This approach circumvents the need for radioactivity to track phot kinase activity and follow light-induced receptor autophosphorylation in vitro by incorporating thiophosphate from N6-benzyl-ATPγS. Consequently, thiophosphorylation of phot substrate candidates can be readily monitored when added or co-expressed with phots in vitro Furthermore, gatekeeper-modified phot1 retained its functionality and its ability to accommodate N6-benzyl-ATPγS as a phosphodonor when expressed in Arabidopsis We therefore anticipate that this chemical genetic approach will provide new opportunities for labeling and identifying substrates for phots and other related AGC kinases under in vitro and near-native in vivo conditions.

Corrigendum: Blue light-dependent changes in loosely bound calcium in Arabidopsis mesophyll cells: an X-ray microanalysis study.

Calcium is involved in the signal transduction pathway from phototropins, the blue light photoreceptor kinases which mediate chloroplast movements. The chloroplast accumulation response in low light is controlled by both phot1 and phot2, while only phot2 is involved in avoidance movement induced by strong light. Phototropins elevate cytosolic Ca(2+) after activation by blue light. In higher plants, both types of chloroplast responses depend on Ca(2+), and internal calcium stores seem to be crucial for these processes. Yet, the calcium signatures generated after the perception of blue light by phototropins are not well understood. To characterize the localization of calcium in Arabidopsis mesophyll cells, loosely bound (exchangeable) Ca(2+) was precipitated with potassium pyroantimonate and analyzed by transmission electron microscopy followed by energy-dispersive X-ray microanalysis. In dark-adapted wild-type Arabidopsis leaves, calcium precipitates were observed at the cell wall, where they formed spherical structures. After strong blue light irradiation, calcium at the apoplast prevailed, and bigger, multilayer precipitates were found. Spherical calcium precipitates were also detected at the tonoplast. After red light treatment as a control, the precipitates at the cell wall were smaller and less numerous. In the phot2 and phot1phot2 mutants, calcium patterns were different from those of wild-type plants. In both mutants, no elevation of calcium after blue light treatment was observed at the cell periphery (including the cell wall and a fragment of cytoplasm). This result confirms the involvement of phototropin2 in the regulation of Ca(2+) homeostasis in mesophyll cells.

Corrigendum: Fine tuning chloroplast movements through physical interactions between phototropins.

Corrigendum: Fine tuning chloroplast movements through physical interactions between phototropins.

Palisade cell shape affects the light-induced chloroplast movements and leaf photosynthesis

Leaf photosynthesis is regulated by multiple factors that help the plant to adapt to fluctuating light conditions. Leaves of sun-light-grown plants are thicker and contain more columnar palisade cells than those of shade-grown plants. Light-induced chloroplast movements are also essential for efficient leaf photosynthesis and facilitate efficient light utilization in leaf cells. Previous studies have demonstrated that leaves of most of the sun-grown plants exhibited no or very weak chloroplast movements and could accomplish efficient photosynthesis under strong light. To examine the relationship between palisade cell shape, chloroplast movement and distribution, and leaf photosynthesis, we used an Arabidopsis thaliana mutant, angustifolia (an), which has thick leaves that contain columnar palisade cells similar to those in the sun-grown plants. In the highly columnar cells of an mutant leaves, chloroplast movements were restricted. Nevertheless, under white light condition (at 120 µmol m−2 s−1), the an mutant plants showed higher chlorophyll content per unit leaf area and, thus, higher light absorption by the leaves than the wild type, which resulted in enhanced photosynthesis per unit leaf area. Our findings indicate that coordinated regulation of leaf cell shape and chloroplast movement according to the light conditions is pivotal for efficient leaf photosynthesis.

Stromule extension along microtubules coordinated with actin-mediated anchoring guides perinuclear chloroplast movement during innate immunity.

Dynamic tubular extensions from chloroplasts called stromules have recently been shown to connect with nuclei and function during innate immunity. We demonstrate that stromules extend along microtubules (MTs) and MT organization directly affects stromule dynamics since stabilization of MTs chemically or genetically increases stromule numbers and length. Although actin filaments (AFs) are not required for stromule extension, they provide anchor points for stromules. Interestingly, there is a strong correlation between the direction of stromules from chloroplasts and the direction of chloroplast movement. Stromule-directed chloroplast movement was observed in steady-state conditions without immune induction, suggesting it is a general function of stromules in epidermal cells. Our results show that MTs and AFs may facilitate perinuclear clustering of chloroplasts during an innate immune response. We propose a model in which stromules extend along MTs and connect to AF anchor points surrounding nuclei, facilitating stromule-directed movement of chloroplasts to nuclei during innate immunity.

Actin-mediated movement of chloroplasts.

Plants are sessile and require diverse strategies to adapt to fluctuations in the surrounding light conditions. Consequently, the photorelocation movement of chloroplasts is essential to prevent damages that are induced by intense light (avoidance response) and to ensure efficient photosynthetic activities under weak light conditions (accumulation response). The mechanisms that underlie chloroplast movements have been revealed through analysis of the behavior of individual chloroplasts and it has been found that these organelles can move in any direction without turning. This implies that any part of the chloroplast periphery can function as the leading or trailing edge during movement. This ability is mediated by a special structure, which consists of short actin filaments that are polymerized at the leading edge of moving chloroplasts and are specifically localized in the space between the chloroplast and the plasma membrane, and is called chloroplast-actin. In addition, several of the genes that encode proteins that are involved in chloroplast-actin polymerization or maintenance have been identified. In this Review, we discuss the mechanisms that regulate chloroplast movements through polymerization of the chloroplast-actin and propose a model for actin-driven chloroplast photorelocation movement.

Dynamic modelling of limitations on improving leaf CO2 assimilation under fluctuating irradiance.

A dynamic model of leaf CO2 assimilation was developed as an extension of the canonical steady-state model, by adding the effects of energy-dependent non-photochemical quenching (qE), chloroplast movement, photoinhibition, regulation of enzyme activity in the Calvin cycle, metabolite concentrations, and dynamic CO2 diffusion. The model was calibrated and tested successfully using published measurements of gas exchange and chlorophyll fluorescence on Arabidopsis thaliana ecotype Col-0 and several photosynthetic mutants and transformants affecting the regulation of Rubisco activity (rca-2 and rwt43), non-photochemical quenching (npq4-1 and npq1-2), and sucrose synthesis (spsa1). The potential improvements on CO2 assimilation under fluctuating irradiance that can be achieved by removing the kinetic limitations on the regulation of enzyme activities, electron transport, and stomatal conductance were calculated in silico for different scenarios. The model predicted that the rates of activation of enzymes in the Calvin cycle and stomatal opening were the most limiting (up to 17% improvement) and that effects varied with the frequency of fluctuations. On the other hand, relaxation of qE and chloroplast movement had a strong effect on average low-irradiance CO2 assimilation (up to 10% improvement). Strong synergies among processes were found, such that removing all kinetic limitations simultaneously resulted in improvements of up to 32%.