Plant Cell-Cell Transport via Plasmodesmata Is Regulated by Light and the Circadian Clock.

Plasmodesmata (PD) are essential for plant development, but little is known about their regulation. Several studies have linked PD transport to chloroplast-centered signaling networks, but the physiological significance of this connection remains unclear. Here, we show that PD transport is strongly regulated by light and the circadian clock. Light promotes PD transport during the day, but light is not sufficient to increase rates of PD transport at night, suggesting a circadian gating mechanism. Silencing expression of the core circadian clock gene, LHY/CCA1, allows light to strongly promote PD transport during subjective night, confirming that the canonical plant circadian clock controls the PD transport light response. We conclude that PD transport is dynamically regulated during the day/night cycle. Due to the many roles of PD in plant biology, this discovery has strong implications for plant development, physiology, and pathogenesis.

HSFA2 Functions in the Physiological Adaptation of Undifferentiated Plant Cells to Spaceflight.

Heat Shock Factor A2 (HsfA2) is part of the Heat Shock Factor (HSF) network, and plays an essential role beyond heat shock in environmental stress responses and cellular homeostatic control. Arabidopsis thaliana cell cultures derived from wild type (WT) ecotype Col-0 and a knockout line deficient in the gene encoding HSFA2 (HSFA2 KO) were grown aboard the International Space Station (ISS) to ascertain whether the HSF network functions in the adaptation to the novel environment of spaceflight. Microarray gene expression data were analyzed using a two-part comparative approach. First, genes differentially expressed between the two environments (spaceflight to ground) were identified within the same genotype, which represented physiological adaptation to spaceflight. Second, gene expression profiles were compared between the two genotypes (HSFA2 KO to WT) within the same environment, which defined genes uniquely required by each genotype on the ground and in spaceflight-adapted states. Results showed that the endoplasmic reticulum (ER) stress and unfolded protein response (UPR) define the HSFA2 KO cells' physiological state irrespective of the environment, and likely resulted from a deficiency in the chaperone-mediated protein folding machinery in the mutant. Results further suggested that additional to its universal stress response role, HsfA2 also has specific roles in the physiological adaptation to spaceflight through cell wall remodeling, signal perception and transduction, and starch biosynthesis. Disabling HsfA2 altered the physiological state of the cells, and impacted the mechanisms induced to adapt to spaceflight, and identified HsfA2-dependent genes that are important to the adaption of wild type cells to spaceflight. Collectively these data indicate a non-thermal role for the HSF network in spaceflight adaptation.

Mitogen-Activated Protein Kinase Phosphatases Affect UV-B-Induced Stomatal Closure via Controlling NO in Guard Cells.

Ultraviolet B (UV-B) radiation induces the activation of MITOGEN-ACTIVATED PROTEIN KINASE PHOSPHATASE1 (MKP1) and its targets MPK3 and MPK6, but whether they participate in UV-B guard cell signaling is not clear. Here, evidence shows that UV-B-induced stomatal closure in Arabidopsis (Arabidopsis thaliana) is antagonistically regulated by MKP1 and MPK6 via modulating hydrogen peroxide (H2O2)-induced nitric oxide (NO) production in guard cells. The mkp1 mutant was hypersensitive to UV-B-induced stomatal closure and NO production in guard cells but not to UV-B-induced H2O2 production, suggesting that MKP1 negatively regulates UV-B-induced stomatal closure via inhibiting NO generation in guard cells. Moreover, MPK3 and MPK6 were activated by UV-B in leaves of the wild type and hyperactivated in the mkp1 mutant, but the UV-B-induced activation of MPK3 and MPK6 was largely inhibited in mutants for ATRBOHD and ATRBOHF but not in mutants for NIA1 and NIA2 mpk6 mutants showed defects of UV-B-induced NO production and stomatal closure but were normal in UV-B-induced H2O2 production, while stomata of mpk3 mutants responded to UV-B just like those of the wild type. The defect of UV-B-induced stomatal closure in mpk6 mutants was rescued by exogenous NO but not by exogenous H2O2 Furthermore, double mutant mkp1/mpk6 and the single mutant mpk6 showed the same responses to UV-B in terms of either stomatal movement or H2O2 and NO production. These data indicate that MPK6, but not MPK3, positively regulates UV-B-induced stomatal closure via acting downstream of H2O2 and upstream of NO, while MKP1 functions negatively in UV-B guard cell signaling via down-regulation of MPK6.

Signaling Mechanisms Regulating Diverse Plant Cell Responses to UVB Radiation.

UVB radiation (290-320 nm) causes diverse effects in plant cells that vary with the fluence rate of exposure. High fluence rates of UVB radiation cause damage to DNA and formation of reactive oxygen species in mitochondria and chloroplasts, which lead to oxidation of membrane proteins and lipids and inhibition of cellular functions. In response to oxidative stress, mitochondrial transmembrane potential dissipates, resulting in cytochrome c release and activation of metacaspases. This leads to the apoptosis-like cell death. The signaling mechanism based on UVB DNA damage includes checkpoint activation, cell-cycle arrest, and finally programmed cell death with characteristic DNA fragmentation and morphological hallmarks typical of apoptotic cells. Recently, it was shown that among the components of this signaling mechanism the transcriptional factor SOG1 (suppressor of gamma response 1) plays a key role in regulation of programmed cell death in plants. In contrast to its damaging effects, UVB radiation at low fluence rates can act as a regulatory signal that is specifically perceived by plants to promote acclimation and survival in sunlight. The protective action of UVB is based on expression of various genes, including those encoding flavonoid synthesis enzymes that provide a UVB-absorbing sunscreen in epidermal tissues and DNA photorepair enzymes. These processes are mediated by the UVB photoreceptor UVR8, which has been recently characterized at the molecular level. Now progress is made in uncovering the UVR8-mediated signaling pathway mechanism in the context of UVB photon perception and revealing the biochemical components of the early stages of light signal transduction. In this review, attention is focused on the achievements in studying these UVB-induced signaling processes.

The Direct Involvement of Dark-Induced Tic55 Protein in Chlorophyll Catabolism and Its Indirect Role in the MYB108-NAC Signaling Pathway during Leaf Senescence in Arabidopsis thaliana.

The chloroplast relies on proteins encoded in the nucleus, synthesized in the cytosol and subsequently transported into chloroplast through the protein complexes Toc and Tic (Translocon at the outer/inner membrane of chloroplasts). A Tic complex member, Tic55, contains a redox-related motif essential for protein import into chloroplasts in peas. However, Tic55 is not crucial for protein import in Arabidopsis. Here, a tic55-II-knockout mutant of Arabidopsis thaliana was characterized for Tic55 localization, its relationship with other translocon proteins, and its association with plant leaf senescence when compared to the wild type. Individually darkened leaves (IDLs) obtained through dark-induced leaf senescence were used to demonstrate chlorophyll breakdown and its relationship with plant senescence in the tic55-II-knockout mutant. The IDLs of the tic55-II-knockout mutant contained higher chlorophyll concentrations than those of the wild type. Our microarray analysis of IDLs during leaf senescence identified seven senescence-associated genes (SAGs) that were downregulated in the tic55-II-knockout mutant: ASP3, APG7, DIN2, DIN11, SAG12, SAG13, and YLS9. Real-time quantitative PCR confirmed the reliability of microarray analysis by showing the same expression patterns with those of the microarray data. Thus, Tic55 functions in dark-induced aging in A. thaliana by indirectly regulating downstream SAGs expression. In addition, the expression of four NAC genes, including ANAC003, ANAC010, ANAC042, and ANAC075 of IDL treated tic55-II-knockout mutant appeared to be downregulated. Yeast one hybrid assay revealed that only ANAC003 promoter region can be bound by MYB108, suggesting that a MYB-NAC regulatory network is involved in dark-stressed senescence.

TCP2 positively regulates HY5/HYH and photomorphogenesis in Arabidopsis.

Light regulates plant growth and development via multiple photoreceptors including phytochromes and cryptochromes. Although the functions of photoreceptors have been studied extensively, questions remain regarding the involvement of cryptochromes in photomorphogenesis. In this study, we identified a protein, TEOSINTE-LIKE1, CYCLOIDEA, and PROLIFERATING CELL FACTOR 2 (TCP2), which interacts with the cryptochrome 1 (CRY1) protein in yeast and plant cells via the N-terminal domains of both proteins. Transgenic plants overexpressing TCP2 displayed a light-dependent short hypocotyl phenotype, especially in response to blue light. Moreover, light affected TCP2 expression in a wavelength-dependent manner and TCP2 positively regulates mRNA expression of HYH and HY5. These results support the hypothesis that TCP2 is a transcription activator which acts downstream of multiple photoreceptors, including CRY1.

Natural strategies for photosynthetic light harvesting.

Photosynthetic organisms are crucial for life on Earth as they provide food and oxygen and are at the basis of most energy resources. They have a large variety of light-harvesting strategies that allow them to live nearly everywhere where sunlight can penetrate. They have adapted their pigmentation to the spectral composition of light in their habitat, they acclimate to slowly varying light intensities and they rapidly respond to fast changes in light quality and quantity. This is particularly important for oxygen-producing organisms because an overdose of light in combination with oxygen can be lethal. Rapid progress is being made in understanding how different organisms maximize light harvesting and minimize deleterious effects. Here we summarize the latest findings and explain the main design principles used in nature. The available knowledge can be used for optimizing light harvesting in both natural and artificial photosynthesis to improve light-driven production processes.

Cell dedifferentiation, callus induction and somatic embryogenesis in Crataegus spp.

The present study describes the effects of light conditions, different kinds and concentrations of auxins [Naphthylacetic acid (NAA) and dichlorophenoxyacetic acid (2,4-D)] with cytokinin (Kin) in MS medium on callus induction and embryogenesis in Crataegus pseudoheterophylla, C. aronia and C.meyeri. At first leave explants sections were cultured on different combinations of plant growth regulators in dark and light for callus initiation and light conditions to evaluation the percentage and duration of survival, callus diameter, callus fresh weight and dry. Results of effects of plant growth regulators and light conditions on callus initiation revealed that highest percentage of callus initiation leaves in treatment (0.5 mg/l 2.4-D+0.5 mg/l KIN) for species C.pseudoheterophylla in dark conditions (100%). Dark conditions (100%) were more effective on callogenesis than light conditions (Photoperiodicity of 16-h and at light intensity of 40 µmol m-2 s-1). The callus induction of in vitro (64-100%) leaves was better than the ex vitro ones (0-100%). The combination of 2,4-D and Kin of in vitro leaves callogenesis has been indicated faster (one weeks) than the other combinations. The results also showed that the highest percentage (100%) and survival duration (6 months) was found in species C. pseudoheterophylla and C. meyeri in 0.1 mg/l 2,4.D + 0.5 mg/l KIN and 0.5 mg/l 2,4.D + 0.5 mg/l Kin. The minimum survival (0%) was absorbed in species C. aronia in 1 mg/l NAA. Maximum callus (10.63 and 10.00 mm respectively) was shown in 0.1 mg/l 2,4.D + 0.5 mg/l Kin and 0.5 mg/l 2,4.D + 0.5 mg/l Kin and was not significant differences after five week among species. The results showed that the highest fresh (1081.49 mg) and dry weight (506.88 and 506.98 mg respectively) was absorbed in species C. pseudoheterophylla in 0.1 mg/l 2,4.D + 0.5 mg/l Kin and 0.5 mg/l 2,4.D + 0.5 mg/l Kin. The embryogenesis was not occurred in any plant growth regulator combinations and species. The results of this study suggested that using 2,4-D with cytokinin (Kin) would be more beneficial for callogenesis.

Expression and testing in plants of ArcLight, a genetically-encoded voltage indicator used in neuroscience research.

BACKGROUND: It is increasingly appreciated that electrical controls acting at the cellular and supra-cellular levels influence development and initiate rapid responses to environmental cues. An emerging method for non-invasive optical imaging of electrical activity at cell membranes uses genetically-encoded voltage indicators (GEVIs). Developed by neuroscientists to chart neuronal circuits in animals, GEVIs comprise a fluorescent protein that is fused to a voltage-sensing domain. One well-known GEVI, ArcLight, undergoes strong shifts in fluorescence intensity in response to voltage changes in mammalian cells. ArcLight consists of super-ecliptic (SE) pHluorin (pH-sensitive fluorescent protein) with an A227D substitution, which confers voltage sensitivity in neurons, fused to the voltage-sensing domain of the voltage-sensing phosphatase of C iona i ntestinalis (Ci-VSD). In an ongoing effort to adapt tools of optical electrophysiology for plants, we describe here the expression and testing of ArcLight and various derivatives in different membranes of root cells in Arabidopsis thaliana. RESULTS: Transgenic constructs were designed to express ArcLight and various derivatives targeted to the plasma membrane and nuclear membranes of Arabidopsis root cells. In transgenic seedlings, changes in fluorescence intensity of these reporter proteins following extracellular ATP (eATP) application were monitored using a fluorescence microscope equipped with a high speed camera. Coordinate reductions in fluorescence intensity of ArcLight and Ci-VSD-containing derivatives were observed at both the plasma membrane and nuclear membranes following eATP treatments. However, similar responses were observed for derivatives lacking the Ci-VSD. The dispensability of the Ci-VSD suggests that in plants, where H(+) ions contribute substantially to electrical activities, the voltage-sensing ability of ArcLight is subordinate to the pH sensitivity of its SEpHluorin base. The transient reduction of ArcLight fluorescence triggered by eATP most likely reflects changes in pH and not membrane voltage. CONCLUSIONS: The pH sensitivity of ArcLight precludes its use as a direct sensor of membrane voltage in plants. Nevertheless, ArcLight and derivatives situated in the plasma membrane and nuclear membranes may offer robust, fluorescence intensity-based pH indicators for monitoring concurrent changes in pH at these discrete membrane systems. Such tools will assist analyses of pH as a signal and/or messenger at the cell surface and the nuclear periphery in living plants.

Inhibition by acrolein of light-induced stomatal opening through inhibition of inward-rectifying potassium channels in Arabidopsis thaliana.

Acrolein is a reactive α,ß-unsaturated aldehyde derived from lipid peroxides, which are produced in plants under a variety of stress. We investigated effects of acrolein on light-induced stomatal opening using Arabidopsis thaliana. Acrolein inhibited light-induced stomatal opening in a dose-dependent manner. Acrolein at 100 µM inhibited plasma membrane inward-rectifying potassium (Kin) channels in guard cells. Acrolein at 100 µM inhibited Kin channel KAT1 expressed in a heterologous system using Xenopus leaves oocytes. These results suggest that acrolein inhibits light-induced stomatal opening through inhibition of Kin channels in guard cells.

Gustav Senn (1875-1945): the pioneer of chloroplast movement research.

Gustav Senn analyzed for the first time light-induced movement and arrangement of chloroplasts. Using many plant species he performed physiological analyses of chloroplast migration in response to external stimuli, with emphasis on light. He determined light paths within a cell by measuring refractive indices and optical thickness of cellular compartments and confirmed that chloroplasts migrate towards the region where the light intensity is optimum. After 6 to 7 years' concentrated study, Senn published the famous monograph "Die Gestalts- und Lageveränderung der Pflanzen- Chromatophoren" (The Changes in Shape and Position of Plant Chloroplasts) in 1908. This book has stimulated many plant physiologists and photobiologists, because Senn not only thoroughly classified and defined various types of light-induced chloroplast migration but also already described possible interaction of different photoreceptor systems in Mougeotia more than 50 years before the discovery of phytochrome. This book also contains still useful experimental hints and overlooked findings on the interaction between light and other factors, such as temperature, water content, and nourishment. After publishing this book, Senn retreated from the study of chloroplasts and became a researcher of the Greek philosopher, Theophrastus. In this review, I introduce his biographical background and then summarize some of his key research accomplishment.

Blue-light-induced rapid chloroplast de-anchoring in Vallisneria epidermal cells.

In the outer periclinal cytoplasm of leaf epidermal cells of an aquatic angiosperm Vallisneria, blue light induces "chloroplast de-anchoring", a rapid decline in the resistance of chloroplasts against centrifugal force. Chloroplast de-anchoring is known induced within 1 min of irradiation with high-fluence-rate blue light specifically, preceding the commencement of chloroplasts migration toward the anticlinal cytoplasm. However, its regulatory mechanism has remained elusive, although pharmacological analysis suggested that a calcium release from intracellular calcium stores is necessary for the response. In search of the responsible photoreceptors, immunoblotting analysis using antibodies against phototropins demonstrated that cross-reactive polypeptides of 120-kDa exist in the plasma-membrane fraction prepared from the leaves. In vitro phosphorylation analysis revealed that 120-kDa polypeptides were phosphorylated by exposure to blue light in a fluence-dependent manner. The blue-light-induced phosphorylation activity was sensitive to a Ser/Thr kinase inhibitor, staurosporine, and unusually was retained at a high level for a long time in darkness. Furthermore, phototropin gene homologs (Vallisneria PHOTOTROPIN1 and PHOTOTROPIN2) expressed in leaves were isolated. We propose that calcium-regulated chloroplast de-anchoring, possibly mediated by phototropins, is an initial process of the blue-light-induced avoidance response of chloroplasts in Vallisneria.

Overexpression of rice OsREX1-S, encoding a putative component of the core general transcription and DNA repair factor IIH, renders plant cells tolerant to cadmium- and UV-induced damage by enhancing DNA excision repair.

Screening of 40,000 Arabidopsis FOX (Full-length cDNA Over-eXpressor gene hunting system) lines expressing rice full-length cDNAs brings us to identify four cadmium (Cd)-tolerant lines, one of which carried OsREX1-S as a transgene. OsREX1-S shows the highest levels of identity to Chlamydomonas reinhardtii REX1-S (referred to as CrREX1-S, in which REX denotes Required for Excision) and to yeast and human TFB5s (RNA polymerase II transcription factor B5), both of which are components of the general transcription and DNA repair factor, TFIIH. Transient expression of OsREX1-S consistently localized the protein to the nucleus of onion cells. The newly generated transgenic Arabidopsis plants expressing OsREX1-S reproducibly displayed enhanced Cd tolerance, confirming that the Cd-tolerance of the initial identified line was conferred solely by OsREX1-S expression. Furthermore, transgenic Arabidopsis plants expressing OsREX1-S exhibited ultraviolet-B (UVB) tolerance by reducing the amounts of cyclobutane pyrimidine dimers produced by UVB radiation. Moreover, those transgenic OsREX1-S Arabidopsis plants became resistant to bleomycin (an inducer of DNA strand break) and mitomycin C (DNA intercalating activity), compared to wild type. Our results indicate that OsREX1-S renders host plants tolerant to Cd, UVB radiation, bleomycin and mitomycin C through the enhanced DNA excision repair.

Proton cellular influx as a probable mechanism of variation potential influence on photosynthesis in pea.

Electrical signals (action potential and variation potential, VP) caused by environmental stimuli are known to induce various physiological responses in plants, including changes in photosynthesis; however, their functional mechanisms remain unclear. In this study, the influence of VP on photosynthesis in pea (Pisum sativum L.) was investigated and the proton participation in this process analysed. VP, induced by local heating, inactivated photosynthesis and activated respiration, with the initiation of the photosynthetic response connected with inactivation of the photosynthetic dark stage; however, direct VP influence on the light stage was also probable. VP generation was accompanied with pH increases in apoplasts (0.17-0.30 pH unit) and decreases in cytoplasm (0.18-0.60 pH unit), which probably reflected H(+) -ATPase inactivation and H(+) influx during this electrical event. Imitation of H(+) influx using the protonophore carbonyl cyanide m-chlorophenylhydrazone (CCCP) induced a photosynthetic response that was similar with a VP-induced response. Experiments on chloroplast suspensions showed that decreased external pH also induced an analogous response and that its magnitude depended on the magnitude of pH change. Thus, the present results showed that proton cellular influx was the probable mechanism of VP's influence on photosynthesis in pea. Potential means of action for this influence are discussed.

Plant nuclear photorelocation movement.

Organelle movement and positioning are essential for proper cellular function. A nucleus moves dynamically during cell division and differentiation and in response to environmental changes in animal, fungal, and plant cells. Nuclear movement is well-studied and the mechanisms have been mostly elucidated in animal and fungal cells, but not in plant cells. In prothallial cells of the fern Adiantum capillus-veneris and leaf cells of the flowering plant Arabidopsis thaliana, light induces nuclear movement and nuclei change their position according to wavelength, intensity, and direction of light. This nuclear photorelocation movement shows some common features with the photorelocation movement of chloroplasts, which is one of the best-characterized plant organelle movements. This review summarizes nuclear movement and positioning in plant cells, especially plant-specific nuclear photorelocation movement and discusses the relationship between nuclear photorelocation movement and chloroplast photorelocation movement.

Cell hydration as a biomarker for estimation of biological effects of nonionizing radiation on cells and organisms.

"Changes in cell hydration" have been hypothesized as an input signal for intracellular metabolic cascade responsible for biological effects of nonionizing radiation (NIR). To test this hypothesis a comparative study on the impacts of different temperature and NIR (infrasound frequency mechanical vibration (MV), static magnetic field (SMF), extremely low frequency electromagnetic field (ELF EMF), and microwave (MW)) pretreated water on the hydration of barley seeds in its dormant and germination periods was performed. In dormant state temperature sensitivity (Q 10) of seed hydration in distilled water (DW) was less than 2, and it was nonsensitive to NIR treated DW, whereas during the germination period (48-72 hours) seeds hydration exhibited temperature sensitivity Q 10 > 2 and higher sensitivity to NIR treated DW. Obtained data allow us to suggest that the metabolic driving of intracellular water dynamics accompanied by hydrogen bonding and breaking is more sensitive to NIR-induced water structure changes in seed bathing aqua medium than the simple thermodynamic processes such as osmotic gradient driven water absorption by seeds in dormant state. Therefore, cell hydration is suggested to be a universal and extrasensitive biomarker for detection of biological effects of NIR on cells and organisms.

Nitric oxide functions as a signal in ultraviolet-B-induced baicalin accumulation in Scutellaria baicalensis suspension cultures.

Stress induced by ultraviolet-B (UV-B) irradiation stimulates the accumulation of various secondary metabolites in plants. Nitric oxide (NO) serves as an important secondary messenger in UV-B stress-induced signal transduction pathways. NO can be synthesized in plants by either enzymatic catalysis or an inorganic nitrogen pathway. The effects of UV-B irradiation on the production of baicalin and the associated molecular pathways in plant cells are poorly understood. In this study, nitric oxide synthase (NOS) activity, NO release and the generation of baicalin were investigated in cell suspension cultures of Scutellaria baicalensis exposed to UV-B irradiation. UV-B irradiation significantly increased NOS activity, NO release and baicalin biosynthesis in S. baicalensis cells. Additionally, exogenous NO supplied by the NO donor, sodium nitroprusside (SNP), led to a similar increase in the baicalin content as the UV-B treatment. The NOS inhibitor, Nω-nitro-l-arginine (LNNA), and NO scavenger, 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO) partially inhibited UV-B-induced NO release and baicalin accumulation. These results suggest that NO is generated by NOS or NOS-like enzymes and plays an important role in baicalin biosynthesis as part of the defense response of S. baicalensis cells to UV-B irradiation.

Production of specialized metabolites in plant cell and organo-cultures: the role of gamma radiation in eliciting secondary metabolism.

PURPOSE: To provide an updated summary of recent advances in the application of gamma irradiation to elicit secondary metabolism and for induction of mutations in plant cell and organ cultures for the production of industrially important specialized metabolites (SMs). CONCLUSIONS: Research on the application of gamma radiation with plants has contributed a lot to microbial decontamination of seeds, and the promotion of physiological processes such as seed germination, seedling vigor, plant growth, and development. Various studies have demonstrated the influence of gamma rays on the morphology, physiology, and biochemistry of plants. Recent research efforts have also shown that low-dose gamma (5-100 Gy) irradiation can be utilized as an expedient solution to alleviate the deleterious effect of abiotic stresses and to obtain better yields of plants. Inducing mutagenesis using gamma irradiation has also evolved as a better option for inducing genetic variability in crops, vegetables, medicinal and ornamentals for their genetic improvement. Plant SMs are gaining increasing importance as pharmaceutical, therapeutic, cosmetic, and agricultural products. Plant cell, tissue, and organ cultures represent an attractive alternative to conventional methods of procuring useful SMs. Among the varied approaches the elicitor-induced in vitro culture techniques are considered an efficient tool for studying and improving the production of SMs. This review focuses on the utilization of low-dose gamma irradiation in the production of high-value SMs such as phenolics, terpenoids, and alkaloids. Furthermore, we present varied successful examples of gamma-ray-induced mutations in the production of SMs.

UV-Induced cell death in plants.

Plants are photosynthetic organisms that depend on sunlight for energy. Plants respond to light through different photoreceptors and show photomorphogenic development. Apart from Photosynthetically Active Radiation (PAR; 400-700 nm), plants are exposed to UV light, which is comprised of UV-C (below 280 nm), UV-B (280-320 nm) and UV-A (320-390 nm). The atmospheric ozone layer protects UV-C radiation from reaching earth while the UVR8 protein acts as a receptor for UV-B radiation. Low levels of UV-B exposure initiate signaling through UVR8 and induce secondary metabolite genes involved in protection against UV while higher dosages are very detrimental to plants. It has also been reported that genes involved in MAPK cascade help the plant in providing tolerance against UV radiation. The important targets of UV radiation in plant cells are DNA, lipids and proteins and also vital processes such as photosynthesis. Recent studies showed that, in response to UV radiation, mitochondria and chloroplasts produce a reactive oxygen species (ROS). Arabidopsis metacaspase-8 (AtMC8) is induced in response to oxidative stress caused by ROS, which acts downstream of the radical induced cell death (AtRCD1) gene making plants vulnerable to cell death. The studies on salicylic and jasmonic acid signaling mutants revealed that SA and JA regulate the ROS level and antagonize ROS mediated cell death. Recently, molecular studies have revealed genes involved in response to UV exposure, with respect to programmed cell death (PCD).

The efficiency of the CO2-concentrating mechanism during single-cell C4 photosynthesis.

The photosynthetic efficiency of the CO(2)-concentrating mechanism in two forms of single-cell C(4) photosynthesis in the family Chenopodiaceae was characterized. The Bienertioid-type single-cell C(4) uses peripheral and central cytoplasmic compartments (Bienertia sinuspersici), while the Borszczowioid single-cell C(4) uses distal and proximal compartments of the cell (Suaeda aralocaspica). C(4) photosynthesis within a single-cell raises questions about the efficiency of this type of CO(2) -concentrating mechanism compared with the Kranz-type. We used measurements of leaf CO(2) isotope exchange (Δ(13) C) to compare the efficiency of the single-cell and Kranz-type forms of C(4) photosynthesis under various temperature and light conditions. Comparisons were made between the single-cell C(4) and a sister Kranz form, S. eltonica[NAD malic enzyme (NAD ME) type], and with Flaveria bidentis[NADP malic enzyme (NADP-ME) type with Kranz Atriplicoid anatomy]. There were similar levels of Δ(13) C discrimination and CO(2) leakiness (Φ) in the single-cell species compared with the Kranz-type. Increasing leaf temperature (25 to 30 °C) and light intensity caused a decrease in Δ(13) C and Φ across all C(4) types. Notably, B. sinuspersici had higher Δ(13) C and Φ than S. aralocaspica under lower light. These results demonstrate that rates of photosynthesis and efficiency of the CO(2) -concentrating mechanisms in single-cell C(4) plants are similar to those in Kranz-type.