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

Thermography methods to assess stomatal behaviour in a dynamic environment.

Although thermography allows rapid, non-invasive measurements of large numbers of plants, it has not been used extensively due to the difficulty in deriving biologically relevant information such as leaf transpiration (E) and stomatal conductance (gsw) from thermograms. Methods normalizing leaf temperature using temperatures from reference materials (e.g. with and without evaporative flux) to generate stress indices are generally preferred due to their ease of use to assess plant water status. Here, a simplified method to solve dynamic energy balance equations is presented, which enables the calculation of 'wet' and 'dry' leaf temperatures in order to derive stress indices, whilst providing accurate estimates of E and gsw. Comparing stress indices and gas exchange parameters highlights the limitation of stress indices in a dynamic environment and how this problem can be overcome using artificial leaf references with known conductance. Additionally, applying the equations for each pixel of a thermogram to derive the rapidity of stomatal response over the leaf lamina in wheat revealed the spatial heterogeneity of stomatal behaviour. Rapidity of stomatal movements is an important determinant of water use efficiency, and our results showed 'patchy' responses that were linked to both the spatial and temporal response of gsw

State Legislative Update

As a response to a growing number of states enacting legislation regarding mediations 0 , the National Conference of Commissioners on Uniform State Laws (NCCUSL) and the Dispute Resolution section of the American Bar Association, drafted the Uniform Mediation Act (UMA). The goal of the drafters in creating the UMA was to promote uniformity in an area of law that varied greatly from state to state. After observing the approach of several states , the drafters chose to place a broad confidentiality privilege for all mediation participants at the heart of the Act. In addition, the UMA requires the mediator to make special disclosures regarding any potential conflicts of interest the mediator may have in the dispute, in order to help enhance both parties awareness of the mediator\u27s ability to act impartially

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

Calvin cycle and guard cell metabolism impact stomatal function.

Stomatal conductance (gs) determines CO2 uptake for photosynthesis (A) and water loss through transpiration, which is essential for evaporative cooling and maintenance of optimal leaf temperature as well as nutrient uptake. Stomata adjust their aperture to maintain an appropriate balance between CO2 uptake and water loss and are therefore critical to overall plant water status and productivity. Although there is considerable knowledge regarding guard cell (GC) osmoregulation (which drives differences in GC volume and therefore stomatal opening and closing), as well as the various signal transduction pathways that enable GCs to sense and respond to different environmental stimuli, little is known about the signals that coordinate mesophyll demands for CO2. Furthermore, chloroplasts are a key feature in GCs of many species, however, their role in stomatal function is unclear and a subject of debate. In this review we explore the current evidence regarding the role of these organelles in stomatal behaviour, including GC electron transport and Calvin-Benson-Bassham (CBB) cycle activity as well as their possible involvement correlating gs and A along with other potential mesophyll signals. We also examine the roles of other GC metabolic processes in stomatal function

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

Natural genetic variation in photosynthesis: an untapped resource to increase crop yield potential?

Raising crop yield potential is a major goal to ensure food security for the growing global population. Photosynthesis is the primary determinant of crop productivity and any gain in photosynthetic CO2 assimilation per unit of leaf area (A) has the potential to increase yield. Significant intraspecific variation in A is known to exist in various autotrophic organs that represent an unexploited target for crop improvement. However, the large number of factors that influence photosynthetic rates often makes it difficult to measure or estimate A under dynamic field conditions (i.e. fluctuating light intensities or temperatures). This complexity often results in photosynthetic capacity, rather than realized photosynthetic rates being used to assess natural variation in photosynthesis. Here we review the work on natural variation in A, the different factors determining A and their interaction in yield formation. A series of drawbacks and perspectives are presented for the most common analyses generally used to estimate A. The different yield components and their determination based on different photosynthetic organs are discussed with a major focus on potential exploitation of various traits for crop improvement. To conclude, an example of different possibilities to increase yield in wheat through enhancing A is illustrated

From green to gold: agricultural revolution for food security

Fifty years ago, Norman Borlaug was awarded the Nobel Peace Prize in recognition of his efforts to increase cereal production. The changes, which became known as the 'Green Revolution', included the introduction of dwarfing genes that enabled dramatic increases in yield through the use of fertilizer and irrigation, coupled with disease resistance. Since the human population has continued to rise dramatically, maintaining a constant per capita cereal production has required a continual increase in global cereal production. Now a Golden Revolution in agriculture is needed to sustainably feed the future population and meet their rising expectations.We acknowledge financial support from the Australian Government through the Australian Research Council Centre of Excellence for Translational Photosynthesis (CE140100015) and The Australian National Universit