typically written from a biophysicist’s or a molecular plant physiologist’s point of view (Horton and Bowyer, Chlorophyll fluorescence analysis has become one of 1990; Krause and Weis, 1991; Govindjee, 1995). The aim the most powerful and widely used techniques avail- of this review is to provide a simple, practical guide to able to plant physiologists and ecophysiologists. This chlorophyll fluorescence for those beginners who are review aims to provide an introduction for the novice interested in applying the technique in both field and into the methodology and applications of chlorophyll laboratory situations. Whilst the principles behind the fluorescence. After a brief introduction into the theor- measurements will be discussed briefly, the emphasis will etical background of the technique, the methodology be on the applications and limitations of this technique and some of the technical pitfalls that can be encoun- in plant ecophysiology. tered are explained. A selection of examples is then used to illustrate the types of information that fluorescence can provide. The basis of chlorophyll fluorescence measurements

/pdf/chlorophyll-fluorescence-a-practical-guide-22dx8txaml.pdf

Chlorophyll fluorescence—a practical guide

BIOPHYSICAL BASIS O F FLUORESCENCE EMISSION FROM CHLOROPLASTS . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . 314 Fluorescence as a Reaction Competing in the Deactivation of Excited Chlorophyll . . . . . . . . . . ... . . .. . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314 Lifetimes of Fluorescence . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . 317 Origin of Fluorescence Emission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 1 Fluorescence of PS 11 and PS I at Ambient and Low Temperatures . . . . . . . . . . . . . . . . . . . 323 FLUORESCENCE INDUCTION AND PS II HETEROGENEITy 325 Fluorescence Transient from Fo to FM . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325 The FI Level and Inactive PS11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... .... . .. . . . .. . . . . . . . . . 326 Fluorescence Induction in High Ught .. . . . . . . . . 327 Rise in the Presence of DCMU and a/{3 Heterogeneity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327 FLUORESCENCE QUENCHING 329 Resolution of Quenching Components . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ........ . .. . . . . . . . . . . . . . . 330 Mechanism of Energy·Dependent Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 1 Quenching Related t o State Transition . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. . ......... . . .. .. . . . . . . . 334 Photoinhibitory Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 Further Quenching Mechanisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338 Physiological Aspects of Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 338 CONCLUSIONS AND PERSPECTiVES 341

Chlorophyll Fluorescence and Photosynthesis: The Basics

Numerical assessments of global air quality and potential changes in atmospheric chemical constituents require estimates of the surface fluxes of a variety of trace gas species. We have developed a global model to estimate emissions of volatile organic compounds from natural sources (NVOC). Methane is not considered here and has been reviewed in detail elsewhere. The model has a highly resolved spatial grid (0.5° × 0.5° latitude/longitude) and generates hourly average emission estimates. Chemical species are grouped into four categories: isoprene, monoterpenes, other reactive VOC (ORVOC), and other VOC (OVOC). NVOC emissions from oceans are estimated as a function of geophysical variables from a general circulation model and ocean color satellite data. Emissions from plant foliage are estimated from ecosystem specific biomass and emission factors and algorithms describing light and temperature dependence of NVOC emissions. Foliar density estimates are based on climatic variables and satellite data. Temporal variations in the model are driven by monthly estimates of biomass and temperature and hourly light estimates. The annual global VOC flux is estimated to be 1150 Tg C, composed of 44% isoprene, 11% monoterpenes, 22.5% other reactive VOC, and 22.5% other VOC. Large uncertainties exist for each of these estimates and particularly for compounds other than isoprene and monoterpenes. Tropical woodlands (rain forest, seasonal, drought-deciduous, and savanna) contribute about half of all global natural VOC emissions. Croplands, shrublands and other woodlands contribute 10–20% apiece. Isoprene emissions calculated for temperate regions are as much as a factor of 5 higher than previous estimates.

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A global model of natural volatile organic compound emissions

Salt tolerance and salinity effects on plants: a review.

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

PHOTO PROTECTION 604 Prevention oj Excessive Light Absorption... 604 Removal of Excess Excitation Energy Directly within the Light-Capturing System ......... ...... . . ..... ..... . .... . ..... ...... .... . .. . .. . . ..... . . . ... ... . 604 Removal oj Active Oxygen Formed in the Photochemical Apparatus ........ . . .. . . . . . . 605 INACTIV A TIONiTURNOVER OF PS II 606 THE XANTHOPHYLL CYCLE AND THERMAL ENERGY DISSIPATION: A PHOTOPROTECTIVE RESPONSE 608 Characteristics oj the Xanthophyll Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . 608 Association Among the De-epoxidized State oj the Xanthophyll Cycle, Thermal Energy Dissipation. and Photoprotection .. .. . . . .. . . ...... .. .. ... ... 609 Operation of the Xanthophyll Cycle in the Field . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . .... . . . .. . . . . 611 CONCLUSIONS 618

Photoprotection and Other Responses of Plants to High Light Stress

The role of xanthophyll cycle carotenoids in the protection of photosynthesis

In the present study we explored the possibility of assessing the allocation of photons absorbed by photosystem II (PSII) antennae to thermal energy dissipation and photosynthetic electron transport in leaves of several plant species under field conditions. Changes in chlorophyll fluorescence parameters were determined in situ over the course of an entire day in the field in sun-exposed leaves of two species with different maximal rates of photosynthesis, Helianthus annuus (sunflower) and Vinca major. Leaves of Vinca minor (periwinkle) growing in a deeply shaded location were also monitored. We propose using diurnal changes in the efficiency of open PSII centers (F′v/F′m) in these sun and shade leaves to (a) assess diurnal changes in the allocation of absorbed light to photochemistry and thermal energy dissipation and, furthermore, (b) make an estimate of changes in the rate of thermal energy dissipation, an analogous expression to the rate of photochemistry. The fraction of light absorbed in PSII antennae that is dissipated thermally (D) is proposed to be estimated from D = 1-F′v/F′m, in analogy to the widely used estimation of the fraction of light absorbed in PSII antennae (P) that is utilized in PSII photochemistry from P = F′v/F′m× qP (where qP is the coefficient for photochemical quenching; Genty, B., Briantais, J.-M. & Baker, N. R. 1989. Biochim. Biophys. Acta 990: 87-92). The rate of thermal dissipation is consequently given by D × PFD (photon flux density), again in analogy to the rate of photochemistry P × PFD, both assuming a matching behavior of photosystems I and II. Characterization of energy dissipation from the efficiency of open PSII centers allows an assessment from a single set of measurements at any time of day; this is particularly useful under field conditions where the fully relaxed reference values of variable or maximal fluorescence needed for the computation of nonphotochemical quenching may not be available. The usefulness of the assessment described above is compared with other currently used parameters to quantify nonphotochemical and photochemical chlorophyll fluorescence quenching.

Using chlorophyll fluorescence to assess the fraction of absorbed light allocated to thermal dissipation of excess excitation

This review places photoprotection into the context of ecology and species diversity. The focus is on photoprotection via the safe removal - as thermal energy - of excess solar energy absorbed by the light collecting system, which counteracts the formation of reactive oxygen species. An update on the surprisingly complex, multiple variations of thermal energy dissipation is presented, placing these different forms into ecological and genetic contexts. Zeaxanthin-facilitated, flexible thermal dissipation associated with the PsbS protein and controlled by the trans-thylakoid pH gradient apparently occurs ubiquitously in plants, and can become sustained (and thus less flexible) at low temperatures. Long-lived, slow-growing plants with low intrinsic capacities for photosynthesis have greater capacities for this flexible dissipation than short-lived, fast-growing species. Furthermore, potent, but inflexible (zeaxanthin-facilitated) thermal dissipation, prominent in evergreen species under prolonged environmental stress, is characterized with respect to the involvement of photosystem II core rearrangement and/or degradation as well as the absence of control by trans-thylakoid pH and, possibly, PsbS. A role of PsbS-related proteins in photoprotection is discussed.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2006.01835.x

Photoprotection in an ecological context: the remarkable complexity of thermal energy dissipation.

The harnessing of solar energy by photosynthesis depends on a safety valve that effectively eliminates hazardous excess energy and prevents oxidative damage to the plant cells. Many of the compounds that protect plant cells also protect human cells. Improving plant resistance to stress may thus have the beneficial side effect of also improving the nutritional quality of plants in the human diet. The pathways that synthesize these compounds are becoming amenable to genetic manipulation, which may yield benefits as widespread as improved plant stress tolerance and improved human physical and mental health.

https://science.sciencemag.org/content/sci/298/5601/2149.full.pdf

William W. Adams

Papers

Photoprotection and Other Responses of Plants to High Light Stress

The role of xanthophyll cycle carotenoids in the protection of photosynthesis

Using chlorophyll fluorescence to assess the fraction of absorbed light allocated to thermal dissipation of excess excitation

Photoprotection in an ecological context: the remarkable complexity of thermal energy dissipation.

Antioxidants in photosynthesis and human nutrition.