Abstract
At the request of the editors, this chapter is based on the book section written by Babin (2008), however, much of the manuscript has been revised and updated to address different readerships. While Babin (2008) is aimed at a more general audience interested in understanding the basis of the measurement and the current instruments available, this chapter is aimed at those who will use the fluorescence tool and are interested in understanding more of its underlying theory, as well as the assumptions associated with it. In short, while the first chapter was aimed more at a beginning user, this one is aimed more at an intermediate user. Nevertheless, some sections have seen little changes.
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Notes
- 1.
Photosystem II is the location of the first photochemical reaction in the photosynthetic apparatus and location of water splitting and oxygen evolution (see Section 2.3.1).
- 2.
Variation in the fluxes going to these alternative paths is the proposed mechanisms for the rapid fluorescence transients observed upon nutrient supply of nutrient stressed algae (see Chapter 11 – Shelly et al.).
- 3.
β-carotene is thought to be involved in the de-excitation of active forms of oxygen that occur under high light and produce damage to the cell.
- 4.
In prokaryotic phytoplankton, zeaxanthin is often present in larger concentration than chl a. However, it is located in the cell wall and, though it absorbs light, it does not appear to have a direct link to photosynthetic processes.
- 5.
The rapid switch between xanthophylls associated with the so-called xanthophylls cycle is not considered a break down or de novo synthesis and is included within the process of regulation.
- 6.
The presence of photodamage, which can arguably be considered a form of photoacclimation/photoprotection, should be conserved in this state. Removal of photodamage indeed requires break down and de novo synthesis of proteins.
- 7.
In low-light acclimated/adapted cells, this fraction can increase to near 50% since the fraction going to non-photosynthetic pigments is reduced.
- 8.
The model for the completely connected case (or lake model) is also expressed very simply as \({f}_{F}^{\prime }=\frac{{k}_{F}}{{k}_{F}+\mathbb{R}_{O}{k}_{P}+{k}_{D}+{k}_{qe}}\). It thus leads to equivalent expressions for the fluorescence yield when the reaction centres are all open (\(\mathbb{R}_{O}\) = 1) or all closed (\(\mathbb{R}_{O}\) = 0)
- 9.
Rohacek uses a multiplicative factor to represent the increased heat dissipation under high light as \(d{k}_{D}\)where d is termed the “dissipation factor”. In our representation (which we used to portray two different types of dissipation) we use k D+k qe to describe the same processes. It follows that d=1+ k qe/k D.
- 10.
The k qe is equivalent to what Oxborough and Baker (2000) call the “non-radiative decay through Stern-Volmer quenching”, instead of using a “variable rate constant”, they express it as a rate constant k SV multiplied by a concentration of quencher [SV]. The equivalence is k qe=k SV[SV].
- 11.
The completely connected model is probably not appropriate in the case of damaged reaction centres as these are generally separated from the complex of connected PSII in the grana (when it exist) and transported to the stroma for resynthesis and reconstruction (Aro et al. 2005).
- 12.
We assume that these photosystems are still capable of increased heat dissipation due to energy-dependent quenching as it seem to be regulated by events that are relatively far from the reaction centre, furthermore, there is no obvious reason why they couldn’t “feel” the transthylakoid pH gradient (see section 2.9.3 for more details).
- 13.
Exception do arise, however, such as when the plastoquinone pool is reduced in the dark by chlororespiration, the maximal level of fluorescence is then observed in low light (see Kromkamp and Forster 2003 and references therein).
- 14.
In this section we will describe the state of a reaction centre with respect to the reduction state of the two quinones QA and QB. The following successive states are possible from the dark-regulated (fully oxidized) state: no charge separation \({Q}_{A}{Q}_{B}\); one charge separation \({Q}_{A}^{-}{Q}_{B}\); \({Q}_{A}{Q}_{B}^{-}\); two charge separations: \({Q}_{A}^{-}{Q}_{B}^{-}\), \({Q}_{A}{Q}_{B}{H}_{2}\), three charge separations \({Q}_{A}^{-}{Q}_{B}{H}_{2}\). The inflexion points are, here, associated with the transfer of the electron from the state where most reaction centres are in the \({Q}_{A}\) reduced the state to the state where the \({\rm {Q\_B}}\) is reduced after one and two charge separations (the J and I inflexion respectively).
- 15.
Light measured at the detector but not originating from the process that is being measured. In fluorometers, this can originate from, for example, scattered light in the measuring volume that is not perfectly filtered by the emission filter.
- 16.
When phytoplankton samples are extracted in an organic solvent, these assumptions are generally met such that the method of chlorophyll determination in vitro is much more accurate once the influence of other fluorescing substances can be excluded (mostly chlorophyll b and pheopigments, Welschmeyer 1994).
- 17.
We note that similar sources of variability are present for fluorometers designed to estimate the biomass of cyanobacteria (whether through phycobilosomes excitation or emission) in particular the presence of NPQ (Karapetyan 2007).
- 18.
Kd(PAR) is the vertical diffuse attenuation coefficient for the downwelling planar irradiance. It is a good proxy of K(PAR) and much better described in the literature.
- 19.
Here instead of defining these five measurements with respect to a protocol, we define them with respect to their theoretical meaning.
- 20.
The fluorescence parameters in the recovery phase following illumination by an actinic light are usually denoted with double primes (e.g. \({F}_{m}^{\prime \prime }\)) this will not be discussed herein.
- 21.
In the hypothetical case of non-quenching damaged reaction centers (kI = 0), F v /F m becomes a linear function of the fraction of damaged reaction centre.
- 22.
The fully connected case also reduces to relatively simple expressions, while intermediate levels of connectivity, probably more true of the reality, are more complex.
- 23.
It takes at least two photons per reaction centre to measure the fluorescence of a closed reaction center; the first photon closes the reaction centre and the second measures or “feels” its closed state. At sufficiently low irradiance the second photon arrives after the reaction centre has reopened.
- 24.
The advantage of the high intensity of the flash is that fluorescence can be measured in low chlorophyll concentration as the fluorescence is proportional to the excitation light (see Eq. 2) while the short duration makes the flashlet subsaturating.
- 25.
This corresponds to the “puddle” model.
- 26.
This corresponds to the connected model and when fully connected often referred to the “lake” model.
- 27.
The Fasttracka fluorometer shows a systematic artefactual transient in the measured fluorescence signal over the series of ∼100 flashes used to close reaction centres. This artefact can be observed using a fluorophore solution that shows no variable fluorescence (e.g. extracted chl a, rhodamine B).
- 28.
Parkhill et al. (2001) make the following distinctions regarding the growth conditions and nutrient status of phytoplankton. When nutrients are available in concentrations that do not limit the growth rate and other environmental factors are constant, the physiological condition is said nutrient replete and phytoplankton assume balanced growth. That is, over a daily cycle the growth rate will be the same if measured by the concentrations of different cellular components (e.g. DNA, chlorophyll, carbon). When a nutrient is limiting growth, the physiological condition of phytoplankton is said to be nutrient stressed. This refers to two nutritional states: nutrient limitation and nutrient starvation . In the former, a nutrient is in short supply, but the fluxes are steady and sufficient to allow the phytoplankton to assume a balanced, albeit reduced, growth rate. Under nutrient starvation, the availability of the nutrient decreases with time relative to the demand so that phytoplankton cannot acclimate physiologically and their growth is unbalanced. Beyond nutrient stress, unbalanced growth is also generally assumed when environmental conditions change.
- 29.
This protocol is in practice somewhat difficult to apply because some parameters cannot be estimated easily simply with fluorometry. The main advantage of the more recent version is that it does not require an estimate of the turnover time for electron transport through the photosynthetic chain and carbon fixation.
- 30.
One recent alternative protocol was proposed by Smyth et al. (2004) using profiles of fluorescence properties.
- 31.
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Acknowledgements
We are very thankful to John Cullen for sharing with us his insights on chlorophyll fluorescence over the years. This work was funded by a CNES grant to M. Babin and a CNES fellowship to Y. Huot.
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Huot, Y., Babin, M. (2010). Overview of Fluorescence Protocols: Theory, Basic Concepts, and Practice. In: Suggett, D., Prášil, O., Borowitzka, M. (eds) Chlorophyll a Fluorescence in Aquatic Sciences: Methods and Applications. Developments in Applied Phycology, vol 4. Springer, Dordrecht. https://doi.org/10.1007/978-90-481-9268-7_3
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