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The production of reactive oxygen species (ROS) is an unavoidable consequence of oxygenic photosynthesis and represents a major cause of oxidative stress in phototrophs, having detrimental effects on the photosynthetic apparatus, limiting cell growth, and productivity. Several methods have been developed for the quantification of cellular ROS, however, most are invasive, requiring the destruction of the sample. Here, we present a new methodology that allows the concurrent quantification of ROS and photosynthetic activity, using the fluorochrome dichlorofluorescein (DCF) and in vivo chlorophyll a fluorescence, respectively. Both types of fluorescence were measured using an imaging Pulse Amplitude Modulation (PAM) fluorometer, modified by adding a UVA-excitation light source (385 nm) and a green bandpass emission filter (530 nm) to enable the sequential capture of red chlorophyll fluorescence and green DCF fluorescence in the same sample. The method was established on Phaeodactylum tricornutum Bohlin, an important marine model diatom species, by determining protocol conditions that permitted the detection of ROS without impacting photosynthetic activity. The utility of the method was validated by quantifying the effects of two herbicides (DCMU and methyl viologen) on the photosynthetic activity and ROS production in P. tricornutum and of light acclimation state in Navicula cf. recens Lange-Bertalot, a common benthic diatom. The developed method is rapid and non-destructive, allowing for the high-throughput screening of multiple samples over time.

Photosynthesis vs. light curves (LCs) have played a central role in photosynthesis research for decades. They are the commonest form of describing how photosynthesis responds to changes in light, being frequently used for characterizing photoacclimation. However, LCs are often interpreted exclusively regarding the response to light intensity, the effects of time of exposure not being explicitly considered. This study proposes the use of 'hysteresis light curves' (HLC), an experimental protocol focused on the cumulative effects of light exposure to obtain information on the time dependence of photosynthetic light responses. HLC are generated by exposing samples to a symmetrical sequence of increasing and decreasing light levels. The comparison of the light-increasing and the light-decreasing phases allows the quantification of the hysteresis caused by high-light exposure, the magnitude and direction of which inform on the activation, and subsequent relaxation of high-light-induced photosynthetic processes. HLCs of the chlorophyll fluorescence indices rETR (relative electron transport rate of photosystem II) and Y(NPQ) (index of non-photochemical quenching) were measured on cyanobacteria, algae, and plants, with the aim of identifying main patterns of hysteresis and their diversity. A non-parametric index is proposed to quantify the magnitude and direction of hysteresis in HLCs of rETR and Y(NPQ). The results of this study show that HLCs can provide additional relevant information on the time dependence of the light response of photosynthetic samples, not obtainable from conventional LCs, useful for phenotyping photosynthetic traits, including photoacclimation state and kinetics of light activation and relaxation of electron flow and energy dissipation processes.

Sacoglossa are known for stealing photosynthetically active chloroplasts from their macroalgal food and incorporating them into their cytosol. The nutritional support these alien organelles (kleptoplasts) provide to the slugs is still debatable. Comparing slugs starved in continuous darkness (non-photosynthetic condition) and light (photosynthetic condition) is often used to understand the contribution of the kleptoplasts to the slugs' metabolism. Here, we examined the slugs' side of starvation in darkness to better understand the effects of darkness on the slugs. We compared the gene expression profile and digestive activity of Elysia viridis, starved for one week under ambient light and continuous darkness. Starvation in darkness led to the up-regulation of genes related to glucose deficiency, while genes involved in the development, cellular organization, and reproduction were down-regulated. This specific gene expression may counteract reduced nutrient availability under non-photosynthetic conditions. Under photosynthetic conditions, kleptoplasts may have a higher nutritional value and may be able to support some metabolic processes. It appears that the slugs can only access kleptoplast photosynthates through autophagy during starvation. Nevertheless, autophagy and length reduction in darkness are highly elevated compared to light conditions, suggesting that more slug tissue is needed to satisfy the nutritional demands under non-photosynthetic conditions. Since we did not detect a gene expression related to the export of photosynthates to the slugs, our results support the hypothesis that slugs use kleptoplasts as larders accessible via autophagy. As long as the kleptoplasts are functional, they provide an energetic support, helping the slugs to reduce starvation-induced stress.

Recent advances in imaging studies and treatment approaches have greatly improved our knowledge about Giant Cell Arteritis (GCA). Previously thought of as a predominantly cranial disease, we now know that GCA is a systemic disease that may involve other medium and large vessel territories. Several imaging studies have shown that between 30 and 70% of patients with GCA present with large-vessel vasculitis. Moreover, a significant proportion of patients present large-vessel disease in the absence of cranial involvement. Extra-cranial disease also poses management challenges as these patients may have a more refractory-relapsing disease course and need additional therapies. Aortic dilation and aneurysms are well-described late complications of GCA involving the large artery territories. In this chapter, we discuss the clinical picture of extra-cranial involvement in GCA, focusing on improved diagnostic protocols and suitable treatment strategies.

Berghia stephanieae (Nudibranchia, Cladobranchia) is a photosymbiotic sea slug that feeds exclusively on sea anemones from the genus Exaiptasia. It then specifically incorporates dinoflagellates belonging to the Symbiodiniaceae obtained from their prey. Here, we present the complete mitochondrial genome sequence of B. stephanieae combining Oxford Nanopore long read and Illumina short-read sequencing data. The mitochondrial genome has a total length of 14,786 bp, it contains the 13 protein-encoding genes, 23 tRNAs, and two rRNAs and is similar to other nudibranchs except for the presence of a duplicated tRNA-Ser 1.

Functional kleptoplasty is a photosymbiotic relationship, in which photosynthetically active chloroplasts serve as an intracellular symbiont for a heterotrophic host. Among Metazoa, functional kleptoplasty is only found in marine sea slugs belonging to the Sacoglossa and recently described in Rhabdocoela worms. Although functional kleptoplasty has been intensively studied in Sacoglossa, the fundamentals of the specific recognition of the chloroplasts and their subsequent incorporation are unknown. The key to ensure the initiation of any symbiosis is the ability to specifically recognize the symbiont and to differentiate a symbiont from a pathogen. For instance, in photosymbiotic cnidarians, several studies have shown that the host innate immune system, in particular scavenger receptors (SRs) and thrombospondin‐type‐1 repeat (TSR) protein superfamily, is playing a major role in the process of recognizing and differentiating symbionts from pathogens. In the present study, SRs and TSRs of three Sacoglossa sea slugs, Elysia cornigera, Elysia timida, and Elysia chlorotica, were identified by translating available transcriptomes into potential proteins and searching for receptor specific protein and/or transmembrane domains. Both receptors classes are highly diverse in the slugs, and many new domain arrangements for each receptor class were found. The analyses of the gene expression of these three species provided a set of species‐specific candidate genes, that is, SR‐Bs, SR‐Es, C‐type lectins, and TSRs, that are potentially relevant for the recognition of kleptoplasts. The results set the base for future experimental studies to understand if and how these candidate receptors are indeed involved in chloroplast recognition., We investigated the abundance and gene expression of scavenger receptors and thrombospondin‐type‐1 repeat protein superfamily, known to be involved in symbiont recognition, in three plastid‐bearing sea slugs. We could identify candidate receptors, similar to those found in cnidarians for Symbiodinium recognition, that might be relevant in plastid recognition.

Botryosphaeriaceae fungi are phytopathogens and human opportunists. The influence of temperature on the phytotoxicity and cytotoxicity of culture filtrates of five Botryosphaeriaceae species was investigated. All culture filtrates of fungi grown at 25 °C were phytotoxic: symptoms were evaluated based on visual inspection of necrosis areas and on the maximum quantum yield of photosystem II, Fv/Fm. Diplodia corticola and Neofusicoccum kwambonambiense were the most phytotoxic, followed by Neofusicoccum parvum CAA704 and Botryosphaeria dothidea. Phytotoxicity dramatically decreased when strains were grown at 37 °C, except for B. dothidea. All strains, except N. parvum CAA366 and Neofusicoccum eucalyptorum, grown either at 25 °C or 37 °C, were toxic to mammalian cells; at 25 °C and at 37°C, D. corticola and B. dothidea were the most cytotoxic, respectively. Although the toxicity of B. dothidea to both cell lines and of N. kwambonambiense to Vero cells increased with temperature, the opposite was found for the other species tested. Our results suggest that temperature modulates the expression of toxic compounds that, in a scenario of a global increase of temperature, may contribute to new plant infections but also human infections, especially in the case of B. dothidea.

BACKGROUND: When the human population increases, so does the need to explore a wider range of feedstocks and biomasses, such as cyanobacteria. However, a deeper understanding of the growth patterns and pigment production is required to support the selection of the most beneficial species and conditions for industrial production. The growth and pigment production (i.e., chlorophyll a and C-phycocyanin) of three cyanobacterium species were evaluated following a three-fold aim. The first goal was to compare among a species commonly selected for exploitation (Arthrospira platensis) and two alternative species (Anabaena cylindrica and Nostoc muscorum). The second goal was analyzing pigment production in the long-term. The last goal involved comparing different methods (spectrophotometry and fluorimetry) to understand whether there is an appropriate proxy of biomass increase and pigment production that can be used for monitoring purposes. RESULTS: All species showed high longevity and proved capable of growing for more than 100 days without any additional supplementation. However, the maximum quantum yield of PS II (Fv /Fm) revealed that their photosynthetic efficiency varied over time with a clear decrease after 2 months. Pigment analysis showed a heterogeneous pattern during the growth periods of all three species that could only be captured by the parameter Fv /Fm, but the pattern was only present for A. cylindrica and N. muscorum in some stages of the culture period. CONCLUSION: N. muscorum was found to be the best chlorophyll a and C-phycocyanin producer, with the production peaking for all species at defined time periods within the growth profile. © 2022 The Authors. Journal of Chemical Technology and Biotechnology published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry (SCI). published

Microphytobenthos (MPB) are a key primary producer of intertidal mudflats. MPB face strong variability in incident irradiance during low tides. Despite photoprotection and photoacclimation, such variations can translate into photoinhibition of MPB cells. This study explores the effect of photoinhibition on MPB primary production (PP) over a large and productive temperate mudflat (Brouage mudflat, NW France). We used a regional and high-resolution tri-dimensional hydrodynamic model coupled to a MPB model with or without photoinhibition. Photoinhibition leads to a 20% (-0.79 t C) decrease of the simulated MPB PP over the entire mudflat. As the upper shore is exposed to light more frequently and longer than the lower shore, the decrease of MPB PP is higher on the upper shore (-29%) than on the lower shore (-5%). With the highest photosynthetically active radiation cumulated over the mudflat, the decrease of MPB PP due to photoinhibition is the highest during spring and spring tides (-22% and -23%, respectively). The model suggests MPB photoinhibition is sensitive to the photoacclimation status of MPB cells through the light saturation parameter. This first modeling attempt to account for MPB photoinhibition is highly constrained by our current theoretical knowledge and limitations on the MPB growth physiology, but it suggests that this process can have a substantial impact on the MPB PP. As such, assessing the MPB photosynthetic response to the highly variable environmental conditions that prevail in large and productive intertidal mudflats is a real challenge for quantifying MPB PP from a synoptic to inter-annual time scale. Plain Language Summary Benthic micro-algae or microphytobenthos (MPB) inhabiting the surficial sediment sustain the high biological production of intertidal mudflats. MPB achieve photosynthesis by aggregating into a dense biofilm at the mud surface during daytime low tides. As MPB can be exposed to short-term variations and high light levels, they change their short-term physiology and position within the sediment to protect themselves. However, such strategies can be outbalanced by a too long stressful light exposure. In this study, we explore with a numerical model the impact of photoinhibition on MPB primary production (PP) over a large and very productive mudflat (NW France). The model suggests that photoinhibition can strongly impact MPB PP. With photoinhibition, the yearly PP decreases by 20 % over the whole mudflat. The model suggests MPB PP is sensitive to the photoacclimation status of MPB cells, i.e. their light use efficiency at a given light level. This first modeling attempt to account for MPB photoinhibition is highly constrained by our current theoretical knowledge and limitations on the MPB growth physiology, but it suggests that this process can have a substantial impact on the MPB PP.

Most studies on sediment-inhabiting microphytobenthos are based on the biomass present on the surface layers of intertidal flats. However, large amounts of microalgal biomass are known to exist below the surface. This study tested the role of subsurface microalgal biomass as a potential source of photosynthetically active cells for the biofilm on the surface. The resilience of buried cells was evaluated by exposing samples from various depths to surface conditions and investigating the recovery of photosynthetic activity. Additionally, vertical migration by subsurface epipelic diatoms was followed at sub-millimeter scales to evaluate its role for transporting cells to the vicinity of the sediment surface. Finally the relative importance of subsurface microalgal biomass was assessed by estimating the proportion of subsurface:surface biomass for different types of sediments from the Ria de Aveiro. Vertical profiles of chlorophyll a, 10 cm-deep, were measured on samples from three intertidal sites, representative of the range of sediment characteristics found in this estuary. The ratio of total biomass to surface biomass (‘subsurface biomass fraction’) based on total biomass (0–10 cm depth interval; Csub, total) and on viable biomass (between the surface and the maximum depth with significant photosynthetic recovery; Csub, viable). The experiments showed that buried cells were able to recover photosynthetic activity within 1.5–3 h of light exposure, with the rate of recovery being dependent on depth and type of sediment. Furthermore, subsurface vertical migration was found to enable motile cells to reach the surface from layers deeper than 1 mm within a low tide period. Overall, the results showed that surface biomass (0–0.5 cm) only accounted for one fifth to one third of the total biomass present between the surface and 10 cm, and that the amount of subsurface viable biomass reached 2–3 times the biomass present at the surface. Applying the estimates of Csub, total and Csub, viable to the whole intertidal area of the Ria de Aveiro, spatially-weighted averages for subsurface biomass fractions were found to reach 3.8 and 2.1 respectively.

In the last years the environmental presence of pharmaceuticals has gained increasing attention. Research data show that these compounds can cause toxicological effects in different species of fish, mollusks and macroinvertebrates. However, the literature is scarce in terms of ecotoxicity data especially focusing on plants as test organisms. Ecotoxicological plant-based tests following the standard OEDC guideline 221 (OECD, 2006) are strongly restricted due to the recommended end-points: growth and yield of plants. It is necessary to develop and validate alternative macrophyte-based tests (non-standard endpoints), more sensible and providing additional information about the chemical contamination effects in plants. To attain this purpose, species from the Lemna genus were selected. Thus, the aim of this study was to analyze the toxic effects of pharmaceuticals in non-standard endpoints on two macrophyte species, Lemna minor and Lemna gibba. To this purpose an acute assay (96 h) was performed with L. minor and L. gibba exposed to chlorpromazine (CPZ), paracetamol (APAP), and diclofenac (DCF), in the following concentration ranges: 0 to 20 μg/L, 0 to 125 μg/L, and 0 to 100 μg/L, respectively. The analyzed endpoints were: levels of chlorophyll a and b, total chlorophyll, carotenoids, anthocyanins; chlorophyll fluorescence; and catalase activity. In general, higher concentrations of the tested pharmaceuticals caused significant effects on both Lemna species in terms of the different endpoints analyzed. In conclusion, acute exposures to CPZ, APAP, and DCF differently affected the defensive system of the tested species; among chlorophylls, chlorophyll b content was more affected, but pharmaceutical exposure was not able to cause alterations on chlorophyll fluorescence.

Background Autoimmune inner ear disease (AIED) represents less than 1% of all cases of sensorineural hearing loss (SNHL) but its frequency may be underestimated due to lack of specific clinical and laboratory criteria. AIED can be associated with a systemic autoimmune disease (SAID) in 15%-30% of the cases. The objective of the present study was to characterize the clinical and prognostic factors of a cohort of patients with AIED. Materials and methods The authors conducted a retrospective descriptive analysis of a cohort of patients with AIED referred from the otorhinolaryngology department to a systemic immune-mediated diseases unit between March 2013 and November 2020. A consecutive sample of 39 patients with suspected AIED was referred. SNHL was defined as a fall of the hearing threshold of at least 30 decibels in three consecutive frequencies. Eight patients were excluded for not meeting the audiometric criteria or having confounding factors. The remaining 31 patients were included with a total of 50 affected ears. To classify the intensity of hearing loss, an arithmetic mean of pure tone was calculated. Normal hearing or mild hearing loss at the last pure tone audiometry of the follow-up were an indicator of good prognosis and were considered the outcome of interest. Results Thirty-two percent of the patients had an associated SAID. There were no differences regarding demographic and clinical characteristics when comparing patients with AIED alone and patients with AIED and a SAID, except for the positivity of antinuclear antibodies (ANA). ANA positivity was superior in patients with the association of AIED and a SAID when compared with patients with AIED alone (90% vs 50%; p=0.037). The SAID was diagnosed after the AIED in 70% of the patients, in which diagnosis of the SAID occurred a median of 4,2 (IQR 0.8-5.1) years after the diagnosis of the AIED. Normal audiometric evaluation or a mild hearing loss was achieved in 31% of the ears at the last audiometric evaluation. A normal audiometry or a mild hearing loss at the time of diagnosis was independently associated with a better outcome (31%, 14%, CI 1.71-273.69; p=0.018). Bilateral hearing loss was independently associated with a worse outcome (54%, 79%, CI 0.01-0.84; p=0.035). The use of systemic corticosteroids (p=0.941), transtympanic corticosteroids (p=0.700) and non-steroid immunomodulator drugs (p=0.986) did not affect prognosis. The presence of a SIAD did not affect the prognosis (p=0.986). Conclusions In this cohort, SAID was present in one-third of the patients with AIED. A good prognosis was achieved in one-third of the patients. A normal audiometry or mild disease at presentation was associated with a good outcome, whilst bilateral involvement was associated with a bad one. Association of a SAID did not seem to influence the hearing-related prognosis. Positivity of ANA antibodies may justify performing a complementary investigation to determine the presence of a SAID.

Thanks are due to FCT/MCTES for the financial support to CESAM (UIDP/50017/2020 + UIDB/50017/2020), through national funds, to JS, and to the NERC Blue-coast project to DMP (NE/N016009/1).

This study aimed to assess the toxicity of glyphosate (GLY; 0, 10, 20 and 30?mg?kg-1) in Solanum lycopersicum L., particularly focusing on the photosynthetic metabolism. By combining ecophysiological, ultrastructural, biochemical and molecular tools, the results revealed that the exposure of tomato plants to GLY led to alterations in leaf water balance regulation [increasing stomatal conductance (gs) and decreasing water use efficiency (WUEi) at higher concentrations] and induced slight alterations in the structural integrity of cells, mainly in chloroplasts, accompanied by a loss of cell viability. Moreover, the transcriptional and biochemical control of several photosynthetic-related parameters was reduced upon GLY exposure. However, in vivo chlorophyll fluorometry and IRGA gas-exchange studies revealed that the photosynthetic yield of S. lycopersicum was not repressed by GLY. Overall, GLY impacts cellular and subcellular homeostasis (by affecting chloroplast structure, reducing photosynthetic pigments and inhibiting photosynthetic-related genes transcription), and leaf structure, but is not reducing the carbon flow on a leaf area basis. Altogether, these results suggest a trade-off effect in which GLY-induced toxicity is compensated by a higher photosynthetic activity related to GLY-induced dysfunction in gs and an increase in mesophyll thickness/density, allowing the viable leaf cells to maintain their photosynthetic capacity., FCT -Fundação para a Ciência e a Tecnologia(SFRH/BD/115643/2016), info:eu-repo/semantics/publishedVersion

Oxygenic photoautotrophs are, paradoxically, subject to photoinhibition of their photosynthetic apparatus, in particular one of its major components, the Photosystem II (PSII). Photoinhibition is generalized across species, light conditions and habitats, imposing substantial metabolic costs that lower photosynthetic productivity and constrain the niches of photoautotrophy. As a process driven by light reaching PSII, light attenuation in optically thick samples influences both the actual extent, and the detection, of photoinhibition. Chlorophyll fluorescence is widely used to measure photoinhibition, but fluorescence-based parameters are affected by light attenuation of both downwelling incident radiation traversing the sample to reach PSII, and emitted fluorescence upwelling through the sample. We used modelling, experimental manipulation of within-sample light attenuation, and meta-analysis of published data, to show substantial, differential effects of light attenuation and depth-integration of emitted fluorescence upon measurements of photoinhibition. Numerical simulations and experimental manipulation of light attenuation indicated that PSII photoinactivation tracked using chlorophyll fluorescence can appear to be over three times lower than the inherent cellular susceptibility to photoinactivation, in optically-dense samples such as leaves or biofilms. The meta-analysis of published data showed that this general trend was unknowingly present in the literature, revealing an overall difference of more than five times between optically thick leaves and optically thin cell suspensions. Although fluorescence-based parameters may provide ecophysiologically relevant information for characterizing the sample as a whole, light attenuation and depth integration can vary between samples independently of their intrinsic physiology. They should be used with caution when aiming to quantify in absolute terms inherent photoinhibition-related parameters in optically thick samples.

Pine pitch canker (PPC), caused by Fusarium circinatum Nirenberg and O’Donnell, represents an important threat to conifer forests worldwide, being associated with significant economic losses. Although essential to develop disease mitigation strategies, little research focused on host susceptibility/resistance mechanisms has been conducted. We aimed to explore the response of a highly susceptible (Pinus radiata D. Don) and a relatively resistant (Pinus pinea L.) species to F. circinatum infection at different stages of infection. Morpho-physiological, hormonal and oxidative stress-related changes were assessed for each pine species and sampling point. Most of the changes found occurred in symptomatic P. radiata, for which an increased susceptibility to photoinhibition was detected together with decreased superoxide dismutase activity. Abscisic acid catabolism was activated by F. circinatum inoculation in both pine species, leading to the accumulation of the inactive dihydrophaseic acid in P. radiata and of the less-active phaseic acid in P. pinea. Hormone confocal analysis revealed that this strategy may be of particular importance at 6 d.p.i. in P. pinea, which together with photosynthesis maintenance to fuel defense mechanism, could in part explain the species resistance to PPC. These results are of great interest for the development of hormone-based breeding strategies or for the use of hormone application as inducers of resistance to F. circinatum infection.

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Neofusicoccum parvum is a fungal pathogen associated with a wide range of plant hosts. Despite being widely studied, the molecular mechanism of infection of N. parvum is still far from being understood. Analysis of N. parvum genome lead to the identification of six putative genes encoding necrosis and ethylene-inducing proteins (NLPs). The sequence of NLPs genes (NprvNep 1-6) were analyzed and four of the six NLP genes were successfully cloned, expressed in E. coli and purified by affinity chromatography. Pure recombinant proteins were characterized according to their phytotoxic and cytotoxic effects to tomato leaves and to mammalian Vero cells, respectively. These assays revealed that all NprvNeps tested are cytotoxic to Vero cells and also induce cell death in tomato leaves. NprvNep2 was the most toxic to Vero cells, followed by NprvNep1 and 3. NprvNep4 induced weaker, but, nevertheless, still significant toxic effects to Vero cells. A similar trend of toxicity was observed in tomato leaves: the most toxic was NprvNep 2 and the least toxic NprvNep 4. This study describes for the first time an overview of the NLP gene family of N. parvum and provides additional insights into its pathogenicity mechanism.

Microbially induced calcification is an ancient, community-driven mineralisation process that produces different types of microbialites. Symbiolites are photosynthesis-induced microbialites, formed by calcifying co-cultures of dinoflagellates from the family Symbiodiniaceae and bacteria. Symbiolites encase the calcifying community as endolithic cells, pointing at an autoendolithic niche of symbiotic dinoflagellates, and provide a rare opportunity to study the role of bacteria in bacterial-algal calcification, as symbiodiniacean cultures display either distinct symbiolite-producing (SP) or non-symbiolite-producing (NP) phenotypes. Using Illumina sequencing, we found that the bacterial communities of SP and NP cultures differed significantly in the relative abundance of 23 genera, 14 families, and 2 phyla. SP cultures were rich in biofilm digesters from the phylum Planctomycetes and their predicted metagenomes were enriched in orthologs related to biofilm formation. In contrast, NP cultures were dominated by biofilm digesters from the Bacteroidetes, and were inferred as enriched in proteases and nucleases. Functional assays confirmed the potential of co-cultures and bacterial isolates to produce biofilms and point at acidic polysaccharides as key stimulators for mineral precipitation. Hence, bacteria appear to influence symbiolite formation primarily through their biofilm-producing and modifying activity and we anticipate that symbiolite formation, as a low-complexity in vitro model, will significantly advance our understanding of photosynthesis-induced microbial calcification processes.

All the diverse lineages of phytoplankters use conserved Photosystem II complexes to mediate photochemical extraction of electrons from water, with concomitant release of oxygen. Photosystem II suffers light-dependent photoinactivation through multiple mechanisms, which is in turn countered by protein subunit removal and replacement. This cycle imposes both direct costs and also delayed opportunity costs. The effective quantum yield for photoinactivation of Photosystem II depends upon taxa, cell size, prior acclimation state, the instantaneous spectral regime and light level. Low temperature or constraints upon metabolism can drive up excitation pressure and photoinactivation, while slowing counteracting repair. Environmental shifts in nutrients or CO2 alter resource allocations and electron flows, with sometimes counter-intuitive effects upon susceptibility to photoinactivation across taxa. These interacting factors generate a wide range of susceptibilities to photoinactivation across phytoplankton cells. Some phytoplankters engage in luxury or pre-emptive recycling of Photosystem II proteins before photoinactivation occurs, and some accumulate large pools of Photosystem II proteins outside active complexes, which may relate to the diverse thylakoid architectures across taxa. These interacting factors generate complex cost-benefit tradeoffs to maintain Photosystem II, and thus the range of conditions where oxygenic photosynthesis is feasible for a given taxa.

The Symbiodiniaceae are a family of marine dinoflagellates known mostly for their endosymbiotic interactions with invertebrates and protists, but facultatively and exclusively free-living life histories in this family are also evident. A recent systematic revision of the Symbiodiniaceae replaced the clade-based nomenclature of seven divergent lineages of "Symbiodinium" sensu lato with one based on formally described genera. The revised taxonomy was not extended to the whole group because type species to describe a new genus for each of the remaining clades and subclades were lacking. In an effort to characterize benthic habitats of symbiodiniaceans in sediments at Heron Island (Great Barrier Reef, Australia), we isolated >100 monoclonal Symbiodiniaceae cultures. Four of these belonged to Symbiodiniaceae 'subclade' Fr3, and three to Clade H, based on nucleotide sequence similarity (ITS2, LSU, cp23S, and mtCOB), representing the first cultures of these taxa. Based on these isolates, we propose two new genera: Freudenthalidium gen. nov. and Halluxium gen. nov., circumscribing Clades Fr3 and H, respectively. Three new species are described: Freudenthalidium heronense, F. endolithicum, and Halluxium pauxillum. Kofoidian tabulations of motile cells confirm previous observations that amphiesmal vesicle arrangements are generally conserved across the family. These descriptions are an important step toward completing the systematic revision of the Symbiodiniaceae. That this contribution was enabled by isolates from an endopsammic habitat highlights the potential of discovering new symbiodiniacean species in the environment, the study of which will lead to a deeper understanding of free-living versus symbiotic life histories in this ecologically important family of dinoflagellates.