Quantifying and monitoring functional Photosystem II and the stoichiometry of the two photosystems in leaf segments: Approaches and approximations

Given its unique function in light-induced water oxidation and its susceptibility to photoinactivation during photosynthesis, photosystem II (PS II) is often the focus of studies of photosynthetic structure and function, particularly in environmental stress conditions. Here we review four approaches for quantifying or monitoring PS II functionality or the stoichiometry of the two photosystems in leaf segments, scrutinizing the approximations in each approach. (1) Chlorophyll fluorescence parameters are convenient to derive, but the information-rich signal suffers from the localized nature of its detection in leaf tissue. (2) The gross O2 yield per single-turnover flash in CO2-enriched air is a more direct measurement of the functional content, assuming that each functional PS II evolves one O2 molecule after four flashes. However, the gross O2 yield per single-turnover flash (multiplied by four) could overestimate the content of functional PS II if mitochondrial respiration is lower in flash illumination than in darkness. (3) The cumulative delivery of electrons from PS II to P700? (oxidized primary donor in PS I) after a flash is added to steady background far-red light is a whole-tissue measurement, such that a single linear correlation with functional PS II applies to leaves of all plant species investigated so far. However, the magnitude obtained in a simple analysis (with the signal normalized to the maximum photo-oxidizable P700 signal), which should equal the ratio of PS II to PS I centers, was too small to match the independently-obtained photosystem stoichiometry. Further, an under-estimation of functional PS II content could occur if some electrons were intercepted before reaching PS I. (4) The electrochromic signal from leaf segments appears to reliably quantify the photosystem stoichiometry, either by progressively photoinactivating PS II or suppressing PS I via photo-oxidation of a known fraction of the P700 with steady far-red light. Together, these approaches have the potential for quantitatively probing PS II in vivo in leaf segments, with prospects for application of the latter two approaches in the field

Measuring the dynamic photosynthome

Background: Photosynthesis underpins plant productivity and yet is notoriously sensitive to small changes inenvironmental conditions, meaning that quantitation in nature across different time scales is not straightforward. The ‘dynamic’ changes in photosynthesis (i.e. the kinetics of the various reactions of photosynthesis in response to environmental shifts) are now known to be important in driving crop yield. Scope: It is known that photosynthesis does not respond in a timely manner, and even a small temporal “mismatch” between a change in the environment and the appropriate response of photosynthesis toward optimality can result in a fall in productivity. Yet the most commonly measured parameters are still made at steady state or a temporary steady state (including those for crop breeding purposes), meaning that new photosynthetic traits remain undiscovered. Conclusions: There is a great need to understand photosynthesis dynamics from a mechanistic and biological viewpoint especially when applied to the field of ‘phenomics’ which typically uses large genetically diverse populations of plants. Despite huge advances in measurement technology in recent years, it is still unclear whether we possess the capability of capturing and describing the physiologically relevant dynamic features of field photosynthesis in sufficient detail. Such traits are highly complex, hence we dub this the ‘photosynthome’. This review sets out the state of play and describes some approaches that could be made to address this challenge with reference to the relevant biological processes involved

Advantages and disadvantages on photosynthesis measurement techniques: A review

Through photosynthesis, green plants and cyanobacteria are able to transfer sunlight energy to molecular reaction centers for conversion into chemical energy with nearly 100% efficiency. Speed is the key as the transfer of the solar energy takes place almost instantaneously such that little energy is wasted as heat. How photosynthesis achieves this near instantaneous energy transfer is a longstanding mystery that may have finally been solved. Measurements of this process are useful in order to understand how it might be controlled and how the phytomonitoring of plant development to increase productivity can be carried out. Techniques in this sense have evolved and nowadays several have been used for this purpose. Thus, the aim of this paper is to present a review of the various methods and principles that have been used in measuring photosynthesis presenting the advantages and disadvantages of various existing measurement methodologies in order to recommend the most appropriate method according to the needs of specific investigations

Application of modulated chlorophyll fluorescence and modulated chlorophyll fluorescence imaging to study the environmental stress effect

Chlorophyll (Chl) a fluorescence is a widely used tool to monitor the photosynthetic process in plants subjected to environmental stresses. this review reports the theoretical bases of Chl fluorescence, and the significance of the most important Chl fluorescence parameters. it also reports how these parameters can be utilised to estimate changes in photosystem (Ps) ii photochemistry, linear electron flux and dissipation mechanisms. the relation between actual Psii photochemistry and Co2 assimilation is discussed, as is the role of photochemical and non-photochemical quenching in inducing changes in Psii activity. the application of Chl fluorescence imaging to study heterogeneity on leaf lamina is also considered. this review summarises only some of the results obtained by this methodology to study the effects of different environmental stresses, namely water availability, nutrients, pollutants, temperature and salinity

In vivo label-free mapping of the effect of a photosystem II inhibiting herbicide in plants using chlorophyll fluorescence lifetime

Background In order to better understand and improve the mode of action of agrochemicals, it is useful to be able to visualize their uptake and distribution in vivo, non-invasively and, ideally, in the field. Here we explore the potential of plant autofluorescence (specifically chlorophyll fluorescence) to provide a readout of herbicide action across the scales utilising multiphoton-excited fluorescence lifetime imaging, wide-field single-photon excited fluorescence lifetime imaging and single point fluorescence lifetime measurements via a fibre-optic probe. Results Our studies indicate that changes in chlorophyll fluorescence lifetime can be utilised as an indirect readout of a photosystem II inhibiting herbicide activity in living plant leaves at three different scales: cellular (~μm), single point (~1 mm2) and macroscopic (~8 × 6 mm2 of a leaf). Multiphoton excited fluorescence lifetime imaging of Triticum aestivum leaves indicated that there is an increase in the spatially averaged chlorophyll fluorescence lifetime of leaves treated with Flagon EC—a photosystem II inhibiting herbicide. The untreated leaf exhibited an average lifetime of 560 ± 30 ps while the leaf imaged 2 h post treatment exhibited an increased lifetime of 2000 ± 440 ps in different fields of view. The results from in vivo wide-field single-photon excited fluorescence lifetime imaging excited at 440 nm indicated an increase in chlorophyll fluorescence lifetime from 521 ps in an untreated leaf to 1000 ps, just 3 min after treating the same leaf with Flagon EC, and to 2150 ps after 27 min. In vivo single point fluorescence lifetime measurements demonstrated a similar increase in chlorophyll fluorescence lifetime. Untreated leaf presented a fluorescence lifetime of 435 ps in the 440 nm excited chlorophyll channel, CH4 (620–710 nm). In the first 5 min after treatment, mean fluorescence lifetime is observed to have increased to 1 ns and then to 1.3 ns after 60 min. For all these in vivo plant autofluorescence lifetime measurements, the plants were not dark-adapted. Conclusions We demonstrate that the local impact of a photosystem II herbicide on living plant leaves can be conveniently mapped in space and time via changes in autofluorescence lifetime, which we attribute to changes in chlorophyll fluorescence. Using portable fibre-optic probe instrumentation originally designed for label-free biomedical applications, this capability could be deployed outside the laboratory for monitoring the distribution of herbicides in growing plants

Frequently asked questions about chlorophyll fluorescence, the sequel

Using chlorophyll (Chl) a fluorescence many aspects of the photosynthetic apparatus can be studied, both in vitro and, noninvasively, in vivo. Complementary techniques can help to interpret changes in the Chl a fluorescence kinetics. Kalaji et al. (Photosynth Res 122:121–158, 2014a) addressed several questions about instruments, methods and applications based on Chl a fluorescence. Here, additional Chl a fluorescence-related topics are discussed again in a question and answer format. Examples are the effect of connectivity on photochemical quenching, the correction of FV/FM values for PSI fluorescence, the energy partitioning concept, the interpretation of the complementary area, probing the donor side of PSII, the assignment of bands of 77 K fluorescence emission spectra to fluorescence emitters, the relationship between prompt and delayed fluorescence, potential problems when sampling tree canopies, the use of fluorescence parameters in QTL studies, the use of Chl a fluorescence in biosensor applications and the application of neural network approaches for the analysis of fluorescence measurements. The answers draw on knowledge from different Chl a fluorescence analysis domains, yielding in several cases new insights.La lista completa de autores que integran el documento puede consultarse en el archivo.Este documento tiene una corrección (ver documento relacionado).Instituto de Fisiología Vegeta

Frequently asked questions about chlorophyll fluorescence, the sequel

[EN] Using chlorophyll (Chl) a fluorescence many aspects of the photosynthetic apparatus can be studied, both in vitro and, noninvasively, in vivo. Complementary techniques can help to interpret changes in the Chl a fluorescence kinetics. Kalaji et al. (Photosynth Res 122: 121-158, 2014a) addressed several questions about instruments, methods and applications based on Chl a fluorescence. Here, additionalChl a fluorescence-related topics are discussed again in a question and answer format. Examples are the effect of connectivity on photochemical quenching, the correction of F-V/F-M values for PSI fluorescence, the energy partitioning concept, the interpretation of the complementary area, probing the donor side of PSII, the assignment of bands of 77 K fluorescence emission spectra to fluorescence emitters, the relationship between prompt and delayed fluorescence, potential problems when sampling tree canopies, the use of fluorescence parameters in QTL studies, the use of Chl a fluorescence in biosensor applications and the application of neural network approaches for the analysis of fluorescence measurements. The answers draw on knowledge fromdifferent Chl a fluorescence analysis domains, yielding in several cases new insights.Kalaji, H.; Schansker, G.; Brestic, M.; Bussotti, F.; Calatayud, A.; Ferroni, L.; Goltsev, V.... (2017). Frequently asked questions about chlorophyll fluorescence, the sequel. 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Multi-colour fluorescence imaging of photosynthetic activity and plant stress

Imaging the four fluorescence bands of leaves, the red (F$_{690}$) and far-red (F$_{740}$) chlorophyll (Chl) fluorescence as well as the blue (F$_{440}$) and green (F$_{520}$) fluorescence of leaves and the corresponding fluorescence ratios is a fast and excellent nondestructive technique to detect the photosynthetic activity and capacity of leaves, of gradients over the leaf area as well as the effect of various strain and stress parameters on plants. This review primarily deals with the first and pioneering multi-colour fluorescence imaging results obtained since the mid-1990s in a cooperation with French colleagues in Strasbourg and in my laboratory in Karlsruhe. Together we introduced not only the joint imaging of the red and far-red Chl fluorescence but also of the blue and green fluorescence of leaves. The two instrumental setups composed for this purpose were (1) the Karlsruhe–Strasbourg UV-Laser Fluorescence Imaging System (Laser-FIS) and (2) the Karlsruhe Flash-Light Fluorescence Imaging System (FL-FIS). Essential results obtained with these instruments are summarized as well as the basic principles and characteristics of multi-colour fluorescence imaging. The great advantage of fluorescence imaging is that the fluorescence yield in the four fluorescence bands is sensed of several thousand up to 200,000 pixels per leaf area in one image. The multi-colour FIS technique allows to sense many physiological parameters and stress effects in plants at an early stage before a damage of leaves is visually detectable. Various examples of plant stress detection by the multi-colour FIS technique are given. Via imaging the Chl fluorescence ratio F690/F740 it is even possible to determine the Chl content of leaves. The FIS technique also allows to follow the successive uptake of diuron and loss of photosynthetic function and to screen the ripening of apples during storage. Particularly meaningful and of high statistical relevance are the fluorescence ratio images red/far-red (F$_{690}$/ F$_{740}$), blue/red (F$_{440}$/F$_{690}$), and blue/green (F$_{440}$/F$_{520}$) as well as images of the fluorescence decrease ratio R$_{Fd}$, which is an indicator of the net CO$_{2}$ assimilation rates of leaves

Multimodal optical spectroscopy for application in the life sciences

Many optical modalities are being investigated, applied, and further developed for non-invasive analysis and sensing in the life sciences. Often, the combination of two or more modalities is required for in depth analysis because of the complexity of the study objects and questions in this field. The work presents multimodal sensing concepts for applications ranging from non-invasive quantification of biomolecules in the living organism to supporting medical diagnosis showing the combined capabilities of Raman spectroscopy, optical coherence tomography, and optoacoustic