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      Arbuscular Mycorrhizal Symbiosis Enhances Photosynthesis in the Medicinal Herb Salvia fruticosa by Improving Photosystem II Photochemistry

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          Abstract

          We investigated the influence of Salvia fruticosa colonization by the arbuscular mycorrhizal fungi (AMF) Rhizophagus irregularis on photosynthetic function by using chlorophyll fluorescence imaging analysis to evaluate the light energy use in photosystem II (PSII) of inoculated and non-inoculated plants. We observed that inoculated plants used significantly higher absorbed energy in photochemistry (Φ PSII ) than non-inoculated and exhibited significant lower excess excitation energy (EXC). However, the increased Φ PSII in inoculated plants did not result in a reduced non-regulated energy loss in PSII (Φ NO ), suggesting the same singlet oxygen ( 1O 2) formation between inoculated and non-inoculated plants. The increased Φ PSII in inoculated plants was due to an increased efficiency of open PSII centers to utilize the absorbed light (F v’/F m’) due to a decreased non-photochemical quenching (NPQ) since there was no difference in the fraction of open reaction centers (q p ). The decreased NPQ in inoculated plants resulted in an increased electron-transport rate (ETR) compared to non-inoculated. Yet, inoculated plants exhibited a higher efficiency of the water-splitting complex on the donor side of PSII as revealed by the increased F v/F o ratio. A spatial heterogeneity between the leaf tip and the leaf base for the parameters Φ PSII and Φ NPQ was observed in both inoculated and non-inoculated plants, reflecting different developmental zones. Overall, our findings suggest that the increased ETR of inoculated S. fruticosa contributes to increased photosynthetic performance, providing growth advantages to inoculated plants by increasing their aboveground biomass, mainly by increasing leaf biomass.

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          Chlorophyll Fluorescence and Photosynthesis: The Basics

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            Chlorophyll fluorescence analysis: a guide to good practice and understanding some new applications.

            Chlorophyll fluorescence is a non-invasive measurement of photosystem II (PSII) activity and is a commonly used technique in plant physiology. The sensitivity of PSII activity to abiotic and biotic factors has made this a key technique not only for understanding the photosynthetic mechanisms but also as a broader indicator of how plants respond to environmental change. This, along with low cost and ease of collecting data, has resulted in the appearance of a large array of instrument types for measurement and calculated parameters which can be bewildering for the new user. Moreover, its accessibility can lead to misuse and misinterpretation when the underlying photosynthetic processes are not fully appreciated. This review is timely because it sits at a point of renewed interest in chlorophyll fluorescence where fast measurements of photosynthetic performance are now required for crop improvement purposes. Here we help the researcher make choices in terms of protocols using the equipment and expertise available, especially for field measurements. We start with a basic overview of the principles of fluorescence analysis and provide advice on best practice for taking pulse amplitude-modulated measurements. We also discuss a number of emerging techniques for contemporary crop and ecology research, where we see continual development and application of analytical techniques to meet the new challenges that have arisen in recent years. We end the review by briefly discussing the emerging area of monitoring fluorescence, chlorophyll fluorescence imaging, field phenotyping, and remote sensing of crops for yield and biomass enhancement.
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              New fluorescence parameters for the determination of q(a) redox state and excitation energy fluxes.

              A number of useful photosynthetic parameters are commonly derived from saturation pulse-induced fluorescence analysis. We show, that q(P), an estimate of the fraction of open centers, is based on a pure 'puddle' antenna model, where each Photosystem (PS) II center possesses its own independent antenna system. This parameter is incompatible with more realistic models of the photosynthetic unit, where reaction centers are connected by shared antenna, that is, the so-called 'lake' or 'connected units' models. We thus introduce a new parameter, q(L), based on a Stern-Volmer approach using a lake model, which estimates the fraction of open PS II centers. We suggest that q(L) should be a useful parameter for terrestrial plants consistent with a high connectivity of PS II units, whereas some marine species with distinct antenna architecture, may require the use of more complex parameters based on intermediate models of the photosynthetic unit. Another useful parameter calculated from fluorescence analysis is Phi(II), the yield of PS II. In contrast to q(L), we show that the Phi(II) parameter can be derived from either a pure 'lake' or pure 'puddle' model, and is thus likely to be a robust parameter. The energy absorbed by PS II is divided between the fraction used in photochemistry, Phi(II), and that lost non-photochemically. We introduce two additional parameters that can be used to estimate the flux of excitation energy into competing non-photochemical pathways, the yield induced by downregulatory processes, Phi(NPQ), and the yield for other energy losses, Phi(NO).
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                Author and article information

                Journal
                Plants (Basel)
                Plants (Basel)
                plants
                Plants
                MDPI
                2223-7747
                30 July 2020
                August 2020
                : 9
                : 8
                : 962
                Affiliations
                [1 ]Department of Biology, Faculty of Science, Istanbul University, 34134 Istanbul, Turkey; gulrizb@ 123456istanbul.edu.tr (G.B.); hilarella87@ 123456gmail.com (H.E.)
                [2 ]Department of Botany, Aristotle University of Thessaloniki, GR-54124 Thessaloniki, Greece
                [3 ]Institute of Plant Breeding and Genetic Resources, Hellenic Agricultural Organization-Demeter, Thermi, 57001 Thessaloniki, Greece; ilektras@ 123456bio.auth.gr
                [4 ]Biology Division, Institute of Graduate Studies in Science, Istanbul University, 34134 Istanbul, Turkey
                Author notes
                [* ]Correspondence: moustak@ 123456bio.auth.gr (M.M.); eelefth@ 123456bio.auth.gr (E.P.E.)
                Author information
                https://orcid.org/0000-0003-0480-9387
                https://orcid.org/0000-0002-8755-0421
                Article
                plants-09-00962
                10.3390/plants9080962
                7463761
                32751534
                832cc586-3992-4408-bf63-66c6b2d5c895
                © 2020 by the authors.

                Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( http://creativecommons.org/licenses/by/4.0/).

                History
                : 28 May 2020
                : 28 July 2020
                Categories
                Article

                sage,inoculation,electron transport rate,rhizophagus irregularis,photoprotective mechanism,redox state,photosynthetic heterogeneity,chlorophyll fluorescence imaging,non-photochemical quenching,medicinal plants

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