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Iron (Fe) is an essential micronutrient whose availability is limiting in many soils. During Fe deficiency, plants alter the expression of many genes to increase Fe uptake, distribution, and utilization. In a genetic screen for suppressors of Fe sensitivity in the E3 ligase mutant bts-3, we isolated an allele of the bHLH transcription factor (TF) ILR3, ilr3-4. We identified a striking leaf bleaching phenotype in ilr3 mutants that was suppressed by limiting light intensity, indicating that ILR3 is required for phototolerance during Fe deficiency. Among its paralogs that are thought to be partially redundant, only ILR3 was required for phototolerance as well as repression of genes under Fe deficiency. A mutation in the gene-encoding PYE, a known transcriptional repressor under Fe deficiency, also caused leaf bleaching. We identified singlet oxygen as the accumulating reactive oxygen species (ROS) in ilr3-4 and pye, suggesting photosensitivity is due to a PSII defect resulting in ROS production. During Fe deficiency, ilr3-4 and pye chloroplasts retain normal ultrastructure and, unlike wild type (WT), contain stacked grana similar to Fe-sufficient plants. Additionally, we found that the D1 subunit of PSII is destabilized in WT during Fe deficiency but not in ilr3-4 and pye, suggesting that PSII repair is accelerated during Fe deficiency in an ILR3- and PYE-dependent manner. Collectively, our results indicate that ILR3 and PYE confer photoprotection during Fe deficiency to prevent the accumulation of singlet oxygen, potentially by promoting reduction of grana stacking to limit excitation and facilitate repair of the photosynthetic machinery.

Singlet oxygen ( O ) is an excited state of molecular oxygen with an electron spin shift in the molecular orbitals, which is extremely unstable and highly reactive. In plants, O is primarily generated as a byproduct of photosynthesis in the photosystem II reaction center (PSII RC) and the light-harvesting antenna complex (LHC) in the grana core (GC). This occurs upon the absorption of light energy when the excited chlorophyll molecules in the PSII transfer the excess energy to molecular oxygen, thereby generating O . As a potent oxidant, O promotes oxidative damage. However, at sub-lethal levels, it initiates chloroplast-to-nucleus retrograde signaling to contribute to plant stress responses, including acclimation and cell death. The thylakoid membranes comprise two spatially separated O sensors: β-carotene localized in the PSII RC in the GC and the nuclear-encoded chloroplast protein EXECUTER1 (EX1) residing in the non-appressed grana margin (GM). Finding EX1 in the GM suggests the existence of an additional source of O in the GM and the presence of two distinct O -signaling pathways. In this review, we mainly discuss the genesis and impact of O in plant physiology.

Significance The fitness and robustness of plants crucially depend on the molecular repair of the vulnerable photosystem II (PS II) supercomplex, embedded in photosynthetic thylakoid membranes. To maintain photosynthetic performance, plants evolved an efficient multistep PS II repair cycle. The PS II repair cycle relies on a well-defined order of reactions and partial separation of individual repair steps. By combining biochemical, spectroscopic, and ultrastructural techniques, we discover that plants establish reaction order and separation by confinement of the enzymes that catalyze the individual steps to spatially separated thylakoid subcompartments—grana, grana margins, and stroma lamellae—formed by the stacked membranes. Structural flexibility of the thylakoid architecture allows controlled access of the damaged PS II by the repair machinery. A crucial component of protein homeostasis in cells is the repair of damaged proteins. The repair of oxygen-evolving photosystem II (PS II) supercomplexes in plant chloroplasts is a prime example of a very efficient repair process that evolved in response to the high vulnerability of PS II to photooxidative damage, exacerbated by high-light (HL) stress. Significant progress in recent years has unraveled individual components and steps that constitute the PS II repair machinery, which is embedded in the thylakoid membrane system inside chloroplasts. However, an open question is how a certain order of these repair steps is established and how unwanted back-reactions that jeopardize the repair efficiency are avoided. Here, we report that spatial separation of key enzymes involved in PS II repair is realized by subcompartmentalization of the thylakoid membrane, accomplished by the formation of stacked grana membranes. The spatial segregation of kinases, phosphatases, proteases, and ribosomes ensures a certain order of events with minimal mutual interference. The margins of the grana turn out to be the site of protein degradation, well separated from active PS II in grana core and de novo protein synthesis in unstacked stroma lamellae. Furthermore, HL induces a partial conversion of stacked grana core to grana margin, which leads to a controlled access of proteases to PS II. Our study suggests that the origin of grana in evolution ensures high repair efficiency, which is essential for PS II homeostasis.

In photosynthetic electron transport, large multiprotein complexes are connected by small diffusible electron carriers, the mobility of which is challenged by macromolecular crowding. For thylakoid membranes of higher plants, a long-standing question has been which of the two mobile electron carriers, plastoquinone or plastocyanin, mediates electron transport from stacked grana thylakoids where photosystem II (PSII) is localized to distant unstacked regions of the thylakoids that harbor PSI. Here, we confirm that plastocyanin is the long-range electron carrier by employing mutants with different grana diameters. Furthermore, our results explain why higher plants have a narrow range of grana diameters since a larger diffusion distance for plastocyanin would jeopardize the efficiency of electron transport. In the light of recent findings that the lumen of thylakoids, which forms the diffusion space of plastocyanin, undergoes dynamic swelling/shrinkage, this study demonstrates that plastocyanin diffusion is a crucial regulatory element of plant photosynthetic electron transport.

Thylakoids of land plants have a bipartite structure, consisting of cylindrical grana stacks, made of membranous discs piled one on top of the other, and stroma lamellae which are helically wound around the cylinders. Protein complexes predominantly located in the stroma lamellae and grana end membranes are either bulky [photosystem I (PSI) and the chloroplast ATP synthase (cpATPase)] or are involved in cyclic electron flow [the NAD(P)H dehydrogenase (NDH) and PGRL1–PGR5 heterodimers], whereas photosystem II (PSII) and its light-harvesting complex (LHCII) are found in the appressed membranes of the granum. Stacking of grana is thought to be due to adhesion between Lhcb proteins (LHCII or CP26) located in opposed thylakoid membranes. The grana margins contain oligomers of CURT1 proteins, which appear to control the size and number of grana discs in a dosage- and phosphorylation-dependent manner. Depending on light conditions, thylakoid membranes undergo dynamic structural changes that involve alterations in granum diameter and height, vertical unstacking of grana, and swelling of the thylakoid lumen. This plasticity is realized predominantly by reorganization of the supramolecular structure of protein complexes within grana stacks and by changes in multiprotein complex composition between appressed and non-appressed membrane domains. Reversible phosphorylation of LHC proteins (LHCPs) and PSII components appears to initiate most of the underlying regulatory mechanisms. An update on the roles of lipids, proteins, and protein complexes, as well as possible trafficking mechanisms, during thylakoid biogenesis and the de-etiolation process complements this review.

Photosynthetic light harvesting in plants is regulated by phosphorylation-driven state transitions: functional redistributions of the major trimeric light-harvesting complex II (LHCII) to balance the relative excitation of photosystem I and photosystem II. State transitions are driven by reversible LHCII phosphorylation by the STN7 kinase and PPH1/TAP38 phosphatase. LHCII trimers are composed of Lhcbi, Lhcb2, and Lhcb3 proteins in various trimeric configurations. Here, we show that despite their nearly identical amino acid composition, the functional roles of Lhcb1 and Lhcb2 are different but complementary. Arabidopsis thaliana plants lacking only Lhcb2 contain thylakoid protein complexes similar to wild-type plants, where Lhcb2 has been replaced by Lhcb1. However, these do not perform state transitions, so phosphorylation of Lhcb2 seems to be a critical step. In contrast, plants lacking Lhcb1 had a more profound antenna remodeling due to a decrease in the amount of LHCII trimers influencing thylakoid membrane structure and, more indirectly, state transitions. Although state transitions are also found in green algae, the detailed architecture of the extant seed plant light-harvesting antenna can now be dated back to a time after the divergence of the bryophyte and spermatophyte lineages, but before the split of the angiosperm and gymnosperm lineages more than 300 million years ago.

• In plants, the chilling response involves decreased photosynthetic capacity and increased starch accumulation in chloroplasts. However, the mechanisms that modulate these processes remain unclear. • We found that the SlWHY1 gene is significantly induced by chilling stress (4°C) in tomato. Three SlWHY1 overexpression (OE) lines grew better than the wild type (WT) under chilling stress; the OE plants retained intact photosynthetic grana lamellae and showed enhanced hydrolysis of starch. By contrast, RNAi lines that inhibited SlWHY1 were more affected than the corresponding WT cultivar. Their grana lamellae were damaged and starch content increased. • The psbA gene encodes the key photosystem II (PSII) protein D1. We show that SlWHY1 binds to the upstream region (A/GTTACCCT/A) of SlpsbA and enhances the de novo synthesis of D1 in chloroplasts. Additionally, SlWHY1 regulates the expression of the starchdegrading enzyme α-amylase (SlAMY3-L) and the starch synthesis-related enzyme isoamylase gene (SlISA2) in the nucleus, thus modulating the starch content in chloroplasts. • We demonstrate that SlWHY1 enhances the resistance of tomato to chilling stress by maintaining the function of PSII and degrading starch. Thus, overexpression of WHY1 may be an effective strategy for enhancing resistance to chilling stress of chilling-sensitive crops in agricultural production.

Background As damage to the ecological environment continues to increase amid unreasonable amounts of irrigation, soil salinization has become a major challenge to agricultural development. Melatonin (MT) is a pleiotropic signal molecule and indole hormone, which alleviates the damage of abiotic stress to plants. MT has been confirmed to eliminate reactive oxygen species (ROS) by improving the antioxidant system and reducing oxidative damage under adversity. However, the mechanism by which exogenous MT mediates salt tolerance by regulating the photosynthetic capacity and ion balance of cotton seedlings still remains unknown. In this study, the regulatory effects of MT on the photosynthetic system, osmotic modulators, chloroplast, and anatomical structure of cotton seedlings were determined under 0–500 μM MT treatments with salt stress induced by treatment with 150 mM NaCl. Results Salt stress reduces the chlorophyll content, net photosynthetic rate, stomatal conductance, intercellular CO 2 concentration, transpiration rate, PSII photochemical efficiency, PSII actual photochemical quantum yield, the apparent electron transfer efficiency, stomata opening, and biomass. In addition, it increases non-photochemical quenching. All of these responses were effectively alleviated by exogenous treatment with MT. Exogenous MT reduces oxidative damage and lipid peroxidation by reducing salt-induced ROS and protects the plasma membrane from oxidative toxicity. MT also reduces the osmotic pressure by reducing the salt-induced accumulation of Na + and increasing the contents of K + and proline. Exogenous MT can facilitate stomatal opening and protect the integrity of cotton chloroplast grana lamella structure and mitochondria under salt stress, protect the photosynthetic system of plants, and improve their biomass. An anatomical analysis of leaves and stems showed that MT can improve xylem and phloem and other properties and aides in the transportation of water, inorganic salts, and organic substances. Therefore, the application of MT attenuates salt-induced stress damage to plants. Treatment with exogenous MT positively increased the salt tolerance of cotton seedlings by improving their photosynthetic capacity, stomatal characteristics, ion balance, osmotic substance biosynthetic pathways, and chloroplast and anatomical structures (xylem vessels and phloem vessels). Conclusions Our study attributes help to protect the structural stability of photosynthetic organs and increase the amount of material accumulation, thereby reducing salt-induced secondary stress. The mechanisms of MT-induced plant tolerance to salt stress provide a theoretical basis for the use of MT to alleviate salt stress caused by unreasonable irrigation, fertilization, and climate change.

Grana Padano (GP) and Parmigiano Reggiano (PR) are the two most important Italian PDO cheeses. To improve the environmental sustainability of their production, a Life Cycle Assessment (LCA) was completed on 84 dairy farms located in the province of Mantova (Northern Italy). In particular, 33 farms delivered milk for GP production, whereas 51 farms to dairies for PR production. In GP farms, maize silage represented 33.7% of total farmland and alfalfa represented 28.1%. While in PR farms, alfalfa represented 63.6% of total farmland. Fat and Protein Corrected Milk (FPCM) and Dairy Efficiency (DE, calculated as kg of produced FPCM per kg of DM intake) were different in the two production system: FPCM was 30.2 ± 4.32 kg/d in GP farms and 25.0 ± 4.71 kg/d in PR farms; DE was 1.35 ± 0.26 in GP farms, and 1.15 ± 0.22 in PR farms. Mitigation strategies to improve both environmental and economic sustainability were suggested focussing on forage crop production, milk production, herd management and off-farm purchased feed. From the preliminary results, there is evidence that improvements are needed. Climate Change (kg CO 2 eq/kg FPCM) and Land Use (kg Carbon deficit/kg FPCM) were similar (1.38 ± 0.33 and 19.3 ± 7.08 for GP system; 1.46 ± 0.37 and 21.8 ± 11.4 for PR system). The most efficient farms in terms of milk production and DE generally showed the best environmental and economic sustainability, while the others show worse outcomes, mainly due to poor DE, livestock-management issues, feed purchase and ration composition. Highlights 84 farms producing milk for Grana Padano and Parmigiano Reggiano cheese were studied in the Province of Mantova. Life Cycle Assessment was used to quantify the environmental impacts of farms and statistical analysis was helpful to identify 6 clusters. Farm, animal and milk efficiencies were poor and mitigation strategies to improve the sustainability of milk production were suggested.

The machinery that conducts the light-driven reactions of oxygenic photosynthesis is hosted within specialized paired membranes called thylakoids. In higher plants, the thylakoids are segregated into two morphological and functional domains called grana and stroma lamellae. A large fraction of the luminal volume of the granal thylakoids is occupied by the oxygen-evolving complex of photosystem II. Electron microscopy data we obtained on dark- and light-adapted Arabidopsis thylakoids indicate that the granal thylakoid lumen significantly expands in the light. Models generated for the organization of the oxygen-evolving complex within the granal lumen predict that the light-induced expansion greatly alleviates restrictions imposed on protein diffusion in this compartment in the dark. Experiments monitoring the redox kinetics of the luminal electron carrier plastocyanin support this prediction. The impact of the increase in protein mobility within the granal luminal compartment in the light on photosynthetic electron transport rates and processes associated with the repair of photodamaged photosystem II complexes is discussed.