Rethinking Guard Cell Metabolism

Stomata control gaseous fluxes between the internal leaf air spaces and the external atmosphere and, therefore, play a pivotal role in regulating CO 2 uptake for photosynthesis as well as water loss through transpiration. Guard cells, which flank the stomata, undergo adjustments in volume, resulting in changes in pore aperture. Stomatal opening is mediated by the complex regulation of ion transport and solute biosynthesis. Ion transport is exceptionally well understood, whereas our knowledge of guard cell metabolism remains limited, despite several decades of research. In this review, we evaluate the current literature on metabolism in guard cells, particularly the roles of starch, sucrose, and malate. We explore the possible origins of sucrose, including guard cell photosynthesis, and discuss new evidence that points to multiple processes and plasticity in guard cell metabolism that enable these cells to function effectively to maintain optimal stomatal aperture. We also discuss the new tools, techniques, and approaches available for further exploring and potentially manipulating guard cell metabolism to improve plant water use and productivity

Acclimation to fluctuating light impacts the rapidity and diurnal rhythm of stomatal conductance.

Plant acclimation to growth light environment has been studied extensively, however, the majority of these studies have focused on light intensity and photo-acclimation, with few studies exploring the impact of dynamic growth light on stomatal acclimation and behavior. In order to assess the impact of growth light regime on stomatal acclimation, we grew plants in three different lighting regimes (with the same average daily intensity); fluctuating with a fixed pattern of light, fluctuating with a randomized pattern of light (sinusoidal), and non-fluctuating (square wave), to assess the effect of light regime dynamics on gas exchange. We demonstrated that gs acclimation is influenced by both intensity and light pattern, modifying the stomatal kinetics at different times of the day resulting in differences in the rapidity and magnitude of the gs response. We also describe and quantify response to an internal signal that uncouples variation in A and gs over the majority of the diurnal period, and represents 25% of the total diurnal gs. This gs response can be characterized by a Gaussian element and when incorporated into the widely used Ball-Berry Model greatly improved the prediction of gs in a dynamic environment. From these findings we conclude that acclimation of gs to growth light could be an important strategy for maintaining carbon fixation and overall plant water status, and should be considered when inferring responses in the field from laboratory based experiments

Diurnal Variation in Gas Exchange: The Balance between Carbon Fixation and Water Loss

Stomatal control of transpiration is critical for maintaining important processes, such as plant water status, leaf temperature, as well as permitting sufficient CO2 diffusion into the leaf to maintain photosynthetic rates (A). Stomatal conductance often closely correlates with A and is thought to control the balance between water loss and carbon gain. It has been suggested that a mesophyll-driven signal coordinates A and stomatal conductance responses to maintain this relationship; however, the signal has yet to be fully elucidated. Despite this correlation under stable environmental conditions, the responses of both parameters vary spatially and temporally and are dependent on species, environment, and plant water status. Most current models neglect these aspects of gas exchange, although it is clear that they play a vital role in the balance of carbon fixation and water loss. Future efforts should consider the dynamic nature of whole-plant gas exchange and how it represents much more than the sum of its individual leaf-level components, and they should take into consideration the long-term effect on gas exchange over time

Overexpression of the RieskeFeS protein increasese electron transport rates and biomass yield

In this study, we generated transgenic Arabidopsis (Arabidopsis thaliana) plants overexpressing the Rieske FeS protein (PetC), a component of the cytochrome b6f (cyt b6f) complex. Increasing the levels of this protein resulted in concomitant increases in the levels of cyt f (PetA) and cyt b6 (PetB), core proteins of the cyt b6f complex. Interestingly, an increase in the levels of proteins in both the photosystem I (PSI) and PSII complexes also was seen in the Rieske FeS overexpression plants. Although the mechanisms leading to these changes remain to be identified, the transgenic plants presented here provide novel tools to explore this. Importantly, overexpression of the Rieske FeS protein resulted in substantial and significant impacts on the quantum efficiency of PSI and PSII,electron transport, biomass, and seed yield in Arabidopsis plants. These results demonstrate the potential for manipulating electron transport processes to increase crop productivity

Multigene manipulation of photosynthetic carbon assimilation increases CO2 fixation and biomass yield in tobacco

Over the next 40 years it has been estimated that a 50% increase in the yield of grain crops such as wheat and rice will be required to meet the food and fuel demands of the increasing world population. Transgenic tobacco plants have been generated with altered combinations of sedoheptulose-1,7-bisphosphatase, fructose-1,6-bisphosphate aldolase, and the cyanobacterial putative-inorganic carbon transporter B, ictB, of which have all been identified as targets to improve photosynthesis based on empirical studies. It is shown here that increasing the levels of the three proteins individually significantly increases the rate of photosynthetic carbon assimilation, leaf area, and biomass yield. Furthermore, the daily integrated measurements of photosynthesis showed that mature plants fixed between 12-19% more CO2 than the equivalent wild-type plants. Further enhancement of photosynthesis and yield was observed when sedoheptulose-1,7-bisphosphatase, fructose-1,6-bisphosphate aldolase, and ictB were over-expressed together in the same plant. These results demonstrate the potential for the manipulation of photosynthesis, using multigene-stacking approaches, to increase crop yields

An integrated response of Trichodesmium erythraeum IMS101 growth and photo-physiology to Iron, CO₂, and light intensity

We have assessed how varying CO 2 (180, 380, and 720 μatm) and growth light intensity (40 and 400 μmol photons m -2 s -1 ) affected Trichodesmium erythraeum IMS101 growth and photophysiology over free iron (Fe') concentrations between 20 and 9,600 pM. We found significant iron dependencies of growth rate and the initial slope and maximal relative PSII electron transport rates (rP m ). Under iron-limiting concentrations, high-light increased growth rates and rPm; possibly indicating a lower allocation of resources to iron-containing photosynthetic proteins. Higher CO 2 increased growth rates across all iron concentrations, enabled growth to occur at lower Fe' concentrations, increased rPm and lowered the iron half saturation constants for growth (K m ). We attribute these CO 2 responses to the operation of the CCM and the ATP spent/saved for CO 2 uptake and transport at low and high CO 2 , respectively. It seems reasonable to conclude that T. erythraeum IMS101 can exhibit a high degree of phenotypic plasticity in response to CO 2 , light intensity and iron-limitation. These results are important given predictions of increased dissolved CO 2 and water column stratification (i.e., higher light exposures) over the coming decades

GIANT CHLOROPLAST 1 Is Essential for Correct Plastid Division in Arabidopsis

AbstractPlastids are vital plant organelles involved in many essential biological processes [1, 2]. Plastids are not created de novo but divide by binary fission mediated by nuclear-encoded proteins of both prokaryotic and eukaryotic origin [3–7]. Although several plastid division proteins have been identified in plants [8–17], limited information exists regarding possible division control mechanisms. Here, we describe the identification of GIANT CHLOROPLAST 1 (GC1), a new nuclear-encoded protein essential for correct plastid division in Arabidopsis. GC1 is plastid-localized and is anchored to the stromal surface of the chloroplast inner envelope by a C-terminal amphipathic helix. In Arabidopsis, GC1 deficiency results in mesophyll cells harbouring one to two giant chloroplasts, whilst GC1 overexpression has no effect on division. GC1 can form homodimers but does not show any interaction with the Arabidopsis plastid division proteins AtFtsZ1-1, AtFtsZ2-1, AtMinD1, or AtMinE1. Analysis reveals that GC1-deficient giant chloroplasts contain densely packed wild-type-like thylakoid membranes and that GC1-deficient leaves exhibit lower rates of CO2 assimilation compared to wild-type. Although GC1 shows similarity to a putative cyanobacterial SulA cell division inhibitor, our findings suggest that GC1 does not act as a plastid division inhibitor but, rather, as a positive factor at an early stage of the division process

Arabidopsis CP12 mutants have reduced levels of phosphoribulokinase and impaired function of the Calvin–Benson cycle

CP12 is a small, redox-sensitive protein, the most detailed understanding of which is the thioredoxin-mediated regulation of the Calvin–Benson cycle, where it facilitates the formation of a complex between glyceraldehyde-3-phosphate dehydrogenase (GAPDH) and phosphoribulokinase (PRK) in response to changes in light intensity. In most organisms, CP12 proteins are encoded by small multigene families, where the importance of each individual CP12 gene in vivo has not yet been reported. We used Arabidopsis thaliana T-DNA mutants and RNAi transgenic lines with reduced levels of CP12 transcript to determine the relative importance of each of the CP12 genes. We found that single cp12-1, cp12-2, and cp12-3 mutants do not develop a severe photosynthetic or growth phenotype. In contrast, reductions of both CP12-1 and CP12-2 transcripts lead to reductions in photosynthetic capacity and to slower growth and reduced seed yield. No clear phenotype for CP12-3 was evident. Additionally, the levels of PRK protein are reduced in the cp12-1, cp12-1/2, and multiple mutants. Our results suggest that there is functional redundancy between CP12-1 and CP12-2 in Arabidopsis where these proteins have a role in determining the level of PRK in mature leaves and hence photosynthetic capacity

Day length as a key factor moderating the response of coccolithophore growth to elevated pCO2

The fate of coccolithophores in the future oceans remains uncertain, in part due to key factors having not been standardized across experiments. A potentially moderating role for differences in day length (photoperiod) remains largely unexplored. We therefore cultured four different geographical isolates of the species Emiliania huxleyi, as well as two additional species, Gephyrocapsa oceanica (tropical) and Coccolithus braarudii (temperate), to test for interactive effects of pCO2 with the light : dark (L : D) cycle. We confirmed a general regulatory effect of photoperiod on the pCO2 response, whereby growth and particulate inorganic carbon and particulate organic carbon (PIC : POC) ratios were reduced with elevated pCO2 under 14 : 10 h L : D, but these reductions were dampened under continuous (24 h) light. The dynamics underpinning this pattern generally differed for the temperate vs. tropical isolates. Reductions in PIC : POC with elevated pCO2 for tropical taxa were largely through reduced calcification and enhanced photosynthesis under 14 : 10 h L : D, with differences dampened under continuous light. In contrast, reduced PIC : POC for temperate strains reflected increases of photosynthesis that outpaced increases in calcification rates under 14 : 10 h L : D, with both responses again dampened under continuous light. A multivariate analysis of 35 past studies of E. huxleyi further demonstrated that differences in photoperiod account for as much as 40% (strain B11/92) to 55% (strain NZEH) of the variance in reported pCO2-induced reductions to growth but not PIC : POC. Our study thus highlights a critical role for day length in moderating the effect of ocean acidification on coccolithophore growth and consequently how this response may play out across latitudes and seasons in future oceans