The use of chlorophyll fluorescence to monitor photosynthetic performance in algae and plants is now widespread. This review examines how fluorescence parameters can be used to evaluate changes in photosystem II (PSII) photochemistry, linear electron flux, and CO(2) assimilation in vivo, and outlines the theoretical bases for the use of specific fluorescence parameters. Although fluorescence parameters can be measured easily, many potential problems may arise when they are applied to predict changes in photosynthetic performance. In particular, consideration is given to problems associated with accurate estimation of the PSII operating efficiency measured by fluorescence and its relationship with the rates of linear electron flux and CO(2) assimilation. The roles of photochemical and nonphotochemical quenching in the determination of changes in PSII operating efficiency are examined. Finally, applications of fluorescence imaging to studies of photosynthetic heterogeneity and the rapid screening of large numbers of plants for perturbations in photosynthesis and associated metabolism are considered.

Chlorophyll fluorescence: a probe of photosynthesis in vivo

Summary Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, pre- diction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric

/pdf/mechanisms-of-plant-survival-and-mortality-during-drought-5rza45sn4g.pdf

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

In the last decade, our understanding of the processes underlying plant response to drought, at the molecular and whole-plant levels, has rapidly progressed. Here, we review that progress. We draw attention to the perception and signalling processes (chemical and hydraulic) of water deficits. Knowledge of these processes is essential for a holistic understanding of plant resistance to stress, which is needed to improve crop management and breeding techniques. Hundreds of genes that are induced under drought have been identified. A range of tools, from gene expression patterns to the use of transgenic plants, is being used to study the specific function of these genes and their role in plant acclimation or adaptation to water deficit. However, because plant responses to stress are complex, the functions of many of the genes are still unknown. Many of the traits that explain plant adaptation to drought - such as phenology, root size and depth, hydraulic conductivity and the storage of reserves - are those associated with plant development and structure, and are constitutive rather than stress induced. But a large part of plant resistance to drought is the ability to get rid of excess radiation, a concomitant stress under natural conditions. The nature of the mechanisms responsible for leaf photoprotection, especially those related to thermal dissipation, and oxidative stress are being actively researched. The new tools that operate at molecular, plant and ecosystem levels are revolutionising our understanding of plant response to drought, and our ability to monitor it. Techniques such as genome-wide tools, proteomics, stable isotopes and thermal or fluorescence imaging may allow the genotype-phenotype gap to be bridged, which is essential for faster progress in stress biology research.

Understanding plant responses to drought — from genes to the whole plant

http://nhjy.hzau.edu.cn/kech/ssyy/pdf/drought/4.PDF

Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell

Physiological and environmental regulation of stomatal conductance, photosynthesis and transpiration: a model that includes a laminar boundary layer

. Stomatal responses to humidity were studied in several species using normal air and a helium: oxygen mixture (79:21 v/v, with CO2 and water vapour added), which we termed ‘helox’. Since water vapour diffuses 2.33 times faster in helox than in air, it was possible to vary the water-vapour concentration difference between the leaf and the air at the leaf surface independently of the transpiration rate and vice versa. The CO2 concentration at the evaporating surfaces (ci), leaf temperature and photon flux density were kept constant throughout the experiments. The results of these experiments were consistent with a mechanism for Stomatal responses to humidity that is based on the rate of water loss from the leaf. Stomata apparently did not directly sense and respond to either the water vapour concentration at the leaf surface or the difference in water vapour concentration between the leaf interior and the leaf surface. In addition, stomatal responses that caused reductions in transpiration rate at low humidities were accompanied by decreases in photosynthesis at constant ci, suggesting heterogeneous (patchy) stomatal closure.

Stomatal responses to humidity in air and helox

A mathematical model of stomatal conductance is presented. It is based on whole-plant and epidermal hydromechanics, and on two hypotheses: (1) the osmotic gradient across guard cell membranes is proportional to the concentration of ATP in the guard cells; and (2) the osmotic gradient that can be sustained per unit of ATP is proportional to the turgor pressure of adjacent epidermal cells. In the present study, guard cell [ATP] is calculated using a previously published model that is based on a widely used biochemical model of C 3 mesophyll photosynthesis. The conductance model for Vicia faba L. is parameterized and tested As with most other stomatal models, the present model correctly predicts the stomatal responses to variations in transpiration rate, irradiance and intercellular CO 2 . Unlike most other models, however, this model can predict the transient stomatal opening often observed before conductance declines in response to decreases in humidity, soil water potential, or xylem conductance. The model also explicitly accommodates the mechanical advantage of the epidermis and correctly predicts that stomata are relatively insensitive to the ambient partial pressure of oxygen, as a result of the assumed dependence on ATP concentration.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.2003.01094.x

A hydromechanical and biochemical model of stomatal conductance

Most studies on stomatal responses to CO 2 assume that guard cells respond only to intercellular CO 2 concentration and are insensitive to the CO 2 concentrations in the pore and outside the leaf. If stomata are sensitive to the CO 2 concentration at the surface of the leaf or in the stomatal pore, the stomatal response to intercellular CO 2 concentration will be incorrect for a `normally9 operating leaf (where ambient CO 2 concentration is a constant). In this study asymmetric CO 2 concentrations for the two surfaces of amphistomatous leaves were used to vary intercellular and leaf surface CO 2 concentrations independently in Xanthium strumarium L. and Helianthus annuus L. The response of stomata to intercellular CO 2 concentration when the concentration at the leaf surface was held constant was found to be the same as the response when the surface concentration was varied. In addition, stomata did not respond to changes in leaf surface CO 2 concentration when the intercellular concentration for that surface was held constant. It is concluded that stomata respond to intercellular CO 2 concentration and are insensitive to the CO 2 concentration at the surface of the leaf and in the stomatal pore.

Do Stomata Respond to CO2 Concentrations Other than Intercellular

Stomatal conductance (gs) typically declines in response to increasing intercellular CO2 concentration (ci). However, the mechanisms underlying this response are not fully understood. Recent work suggests that stomatal responses to ci and red light (RL) are linked to photosynthetic electron transport. We investigated the role of photosynthetic electron transport in the stomatal response to ci in intact leaves of cocklebur (Xanthium strumarium) plants by examining the responses of gs and net CO2 assimilation rate to ci in light and darkness, in the presence and absence of the photosystem II inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), and at 2% and 21% ambient oxygen. Our results indicate that (1) gs and assimilation rate decline concurrently and with similar spatial patterns in response to DCMU; (2) the response of gs to ci changes slope in concert with the transition from Rubisco- to electron transport-limited photosynthesis at various irradiances and oxygen concentrations; (3) the response of gs to ci is similar in darkness and in DCMU-treated leaves, whereas the response in light in non-DCMU-treated leaves is much larger and has a different shape; (4) the response of gs to ci is insensitive to oxygen in DCMU-treated leaves or in darkness; and (5) stomata respond normally to RL when ci is held constant, indicating the RL response does not require a reduction in ci by mesophyll photosynthesis. Together, these results suggest that part of the stomatal response to ci involves the balance between photosynthetic electron transport and carbon reduction either in the mesophyll or in guard cell chloroplasts.

Evidence for involvement of photosynthetic processes in the stomatal response to CO2.

It has been suggested that some biological processes are equivalent to computation, but quantitative evidence for that view is weak. Plants must solve the problem of adjusting stomatal apertures to allow sufficient CO2 uptake for photosynthesis while preventing excessive water loss. Under some conditions, stomatal apertures become synchronized into patches that exhibit richly complicated dynamics, similar to behaviors found in cellular automata that perform computational tasks. Using sequences of chlorophyll fluorescence images from leaves of Xanthium strumarium L. (cocklebur), we quantified spatial and temporal correlations in stomatal dynamics. Our values are statistically indistinguishable from those of the same correlations found in the dynamics of automata that compute. These results are consistent with the proposition that a plant solves its optimal gas exchange problem through an emergent, distributed computation performed by its leaves.

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Keith A. Mott

Papers

Stomatal responses to humidity in air and helox

A hydromechanical and biochemical model of stomatal conductance

Do Stomata Respond to CO2 Concentrations Other than Intercellular

Evidence for involvement of photosynthetic processes in the stomatal response to CO2.

Evidence for complex, collective dynamics and emergent, distributed computation in plants