The ∼407-myr-old Rhynie chert of Scotland contains exquisite body fossils of land plants, animals and microorganisms, which provide our earliest reasonably complete snapshot of a Phanerozoic terrestrial ecosystem. These fossils have been instrumental to our understanding of the ‘greening of the land’, a major transition in the history of the Earth–life system. Among the primary producers preserved in the chert are cyanobacteria, of which only a fraction have been formally described. Here, we report the occurrence of the colony-forming cyanobacterium Eoentophysalis in the Rhynie chert. To our knowledge, this represents the first bona fide record of Entophysalidaceae from any post-Cambrian fossil assemblage or any non-marine fossil assemblage of any age. The Rhynie Eoentophysalis appears remarkably similar in appearance both to modern marine and freshwater Entophysalis ssp. and to Eoentophysalis belcherensis, a shallow-marine fossil from the ∼2 Ga Belcher Group of Canada that is perhaps the oldest convincing cyanobacterium on record. Darkened cell envelopes in the Rhynie Eoentophysalis correspond well with both E. belcherensis and modern Entophysalis, whose cell envelopes often contain the photoprotective brown pigment scytonemin. The occurrence of Eoentophysalis in the Rhynie chert supports previous claims that the fossilisable traits of entophysalid cyanobacteria are evolutionarily static through geological time. These organisms may be such effective generalists that major changes in their environment – in this case, the transition to a fully non-marine habitat – have not imposed significant selection pressure on these traits.

Hair lichens are distinctive for their capillary growth and typically arboreal occurrence, especially in temperate and boreal forests. They consist of two morphogroups based on cortical pigments: a brown-black group with fungal melanin and a pale yellow-green group with usnic acid. Here we test the hypothesis that these morphogroups are ecologically distinct and thus appropriately regarded as functional groups. We examine their respective horizontal occurrence in the lower canopy of 60-year-old conifer forests on a 250 m tall volcanic cone in south-central British Columbia. Trees on open south-facing slopes and near the summit were found to support mainly melanic hair lichens (Bryoria and Nodobryoria), whereas more densely spaced trees on north-facing slopes and at the base had higher cover values of usnic lichens (especially Alectoria sarmentosa and Ramalina thrausta). The cover of melanic hair lichens was strongly correlated with canopy openness but not for their usnic counterparts. We suggest that investment in light-absorbing melanic pigments is an extreme form of specialization for high light, favouring persistence in dry, sun-exposed canopies of otherwise cool forests. By contrast, the cortex of pendent usnic hair lichens appears to facilitate optimum light transmission to underlying photobionts in shaded sites, though at the cost of sensitivity to light in open habitats, especially in rather dry regions.

Photosynthetic organisms have evolved a great variety of mechanisms to optimize their use of sunlight. Some of the clearest examples of adaptations can be seen by comparing photosynthesis in different species and in different individuals of the same species that grow under high and low light levels. While the adaptations of sun and shade higher plants have been relatively well studied, much less information is available on the photobionts of lichenized Ascomycetes. An important adaptation that can protect photosynthetic organisms from the potentially harmful effects of excess light is non-photochemical quenching (NPQ); NPQ can dissipate unused light energy as heat. Here we used chlorophyll fluorescence to compare the induction and relaxation of NPQ and the induction of electron transport (rETR) in collections of the same lichen species from exposed and from more shaded locations. All species have trebouxioid photobionts and normally grow in more exposed microhabitats but can also be readily collected from more shaded locations. Shade forms display generally higher NPQ, presumably to protect lichens from occasional rapid increases in light that occur during sunflecks. Furthermore, the NPQ of shade forms relaxes quickly when light levels are reduced, presumably to ensure efficient photosynthesis after a sunfleck has passed. The maximal relative electron transport rate is lower in shade than sun collections, probably reflecting a downregulation of photosynthetic capacity to reduce energy costs. We also compared collections of pale and melanized thalli from three species of shade lichens with Symbiochloris as their photobiont. Interestingly, NPQ in melanized thalli from slightly more exposed microhabitats induced and relaxed in a way that resembled shade rather than sun forms of the trebouxioid lichens. This might suggest that in some locations melanization induced during a temporary period of high light may be excessive and could potentially reduce photosynthesis later in the growing season. Taken together, the results suggest that lichen photobionts can flexibly adjust the amount and type of NPQ, and their levels of rETR in response to light availability.

Non-photochemical quenching (NPQ) plays an important role in protecting photosynthetic organisms from photoinhibition by dissipating excess light energy as heat. However, excess NPQ can greatly reduce the quantum yield of photosynthesis at lower light levels. Recently, there has been considerable interest in understanding how plants balance NPQ to ensure optimal productivity in environments in which light levels are rapidly changing. In the present study, chlorophyll fluorescence was used to study the induction and relaxation of non-photochemical quenching (NPQ) in the dark and the induction of photosynthesis in ten species of lichens, five sampled from exposed and five sampled from shaded habitats. Here we show that the main difference between sun and shade lichens is the rate at which NPQ relaxes in the dark, rather than the speed that photosynthesis starts upon illumination. During the first two minutes in the dark, NPQ values in the five sun species declined only by an average of 2%, while by contrast, in shade species the average decline was 40%. For lichens growing in microhabitats where light levels are rapidly changing, rapid relaxation of NPQ may enable their photobionts to use the available light most efficiently.

This research studies whether photoprotection mechanisms are able to counterbalance the short-term effect of two herbicides, acifluorfen methyl (AFM) and paraquat (PQ), that generate photo-oxidative stress in different subcellular locations. Duckweed plants grown under three light intensities (high-, medium-, and low light), and consequently expressing three levels of photoprotection, were exposed to both herbicides under the same light regime. Oxidative damage induced by AFM originated mainly from the cytosolic accumulation of protoporphyrin IX, leading to a process of plasma membrane disruption, a progressive and slow degradation of ascorbate and photosynthetic pigments, and glutathione accumulation. As most photoprotective mechanisms (antioxidants and xanthophylls-cycle-related energy dissipation) operate mainly within the chloroplast, these systems were unable to protect plants from AFM damage irrespective of the level of light acclimation. Paraquat effects developed more rapidly and to a greater extent than AFM in treated plants. Irrespective of the light intensity, the same sequence of degradation was observed: ascorbate followed by glutathione, α-tocopherol, pigments, and membrane disruption. In PQ-treated plants the generation of oxidative stress occurred mainly in the chloroplast, and cellular damage developed more slowly in highly photoprotected plants (high light); in fact, electrolyte leakage can be used as a marker for PQ tolerance. The effects of both herbicides indicate that the xanthophyll cycle is an early protective mechanism and confirms the central role of ascorbate in early photoprotection response. High levels of lipophilic and hydrophilic antioxidant contents did not lead to attenuated phytotoxicity of acifluorfen methyl and thus are not the basis to explain differential susceptibilities among duckweed plants.

Light adaptive strategies were studied in a comparative analysis of the congeneric seagrass species Zostera muelleri and Zostera marina at two case study areas in New Zealand and Germany. Surveys in intertidal seagrass meadows were conducted from pre-dawn until sunset on days when either low or high tide coincided with noon. The results show marked fluctuations of photophysiology (optimum and effective quantum yield, non-photochemical quenching, cycling of xanthophyll cycle (XC) pigments) over daily and tidal cycles. At both locations, we observed a near complete conversion (de-epoxidation) of violaxanthin to zeaxanthin at times with high irradiance and a rapid and complete re-epoxidation under subsequent lower light conditions. At the New Zealand site we also observed significantly larger XC-pigment pool sizes in seagrass leaves sampled in a week when low tide coincided with noon (larger daily fluence and higher maximum irradiance), compared with leaves sampled in a week when high tide was at noon. This dynamic adjustment of xanthophyll pool size has not been previously reported for intertidal seagrasses. It adds to our understanding of an important adaptive feature in a highly dynamic light environment and to the general ecology and adaptability of seagrasses.

Photosynthesis research employs several biophysical methods, including the detection of fluorescence. Even though fluorescence is a key method to detect photosynthetic efficiency, it has not been applied/adapted to single-cell confocal microscopy measurements to examine photosynthetic microorganisms. Experiments with photosynthetic cells may require automation to perform a large number of measurements with different parameters, especially concerning light conditions. However, commercial microscopes support custom protocols (through Time Controller offered by Olympus or Experiment Designer offered by Zeiss) that are often unable to provide special set-ups and connection to external devices (e.g., for irradiation). Our new system combining an Arduino microcontroller with the Cell⊕Finder software was developed for controlling Olympus FV1000 and FV1200 confocal microscopes and the attached hardware modules. Our software/hardware solution offers (1) a text file-based macro language to control the imaging functions of the microscope; (2) programmable control of several external hardware devices (light sources, thermal controllers, actuators) during imaging via the Arduino microcontroller; (3) the Cell⊕Finder software with ergonomic user environment, a fast selection method for the biologically important cells and precise positioning feature that reduces unwanted bleaching of the cells by the scanning laser. Cell⊕Finder can be downloaded from http://www.alga.cz/cellfinder. The system was applied to study changes in fluorescence intensity in Synechocystis sp. PCC6803 cells under long-term illumination. Thus, we were able to describe the kinetics of phycobilisome decoupling. Microscopy data showed that phycobilisome decoupling appears slowly after long-term (>1 h) exposure to high light.

The symbiotic association between the acoel flatworm Symsagittifera roscoffensis and the prasinophyte microalgae Tetraselmis convolutae was studied regarding its photophysiology and photobehaviour. The photoacclimation status and the photophysiological responses to high light of the algal endosymbiont were studied non-destructively on individual S. roscoffensis using pulse amplitude modulated fluorometry. Specimens collected in an intertidal sandy shore were characterized regarding the maximum quantum yield of photosystem II (PSII), Fv/Fm, and the light response of photosynthetic activity, by constructing rapid light-response curves of the relative electron transport rate of PSII, rETR. The studied population could be considered as high light-acclimated when compared with other intertidal photosynthetic organisms (e.g. macroalgae), with the light-saturation parameter Ek averaging 250 μmol m−2 s−1. Light stress experiments showed S. roscoffensis to be able to withstand the exposure to high light without displaying signs of photoinhibition, suggesting the operation of efficient physiological photoprotective processes. The photobehaviour of S. roscoffensis was studied by characterizing the distribution of the flatworms under a light gradient, using a custom-made photoaccumulation chamber. The results showed a photoaccumulation pattern evidencing a clear avoidance of extreme low or high light levels, and with maximum photoaccumulation values being found for a range of irradiances (150–400 μmol m−2 s−1) that generally coincided with the optima for photosynthetic activity. This matching between the optimum light levels for photosynthetic activity and photoaccumulation suggested that S. roscoffensis may use vertical migration as a form of behavioural photoprotection. This behavioural response may be used to rapidly and flexibly control light exposure, avoiding photodamage to the endosymbiont photosynthetic apparatus by direct exposure to sunlight.

This study explored reflectance indices sampled with a ‘leaf reflectometer’ as measures of pigment content for leaves of contrasting light history, developmental stage and functional type (herbaceous annual versus sclerophyllous evergreen). We employed three reflectance indices: a modified normalized difference vegetation index (NDVI), an index of chlorophyll content; the red/green reflectance ratio (RRED[ratio ]RGREEN), an index of anthocyanin content; and the change in photochemical reflectance index upon dark–light conversions (ΔPRI), an index of xanthophyll cycle pigment activity. In Helianthus annuus (sunflower), xanthophyll cycle pigment amounts were linearly related to growth light environment; leaves in full sun contained approximately twice the amount of xanthophyll cycle pigments as leaves in deep shade, and at midday a larger proportion of these pigments were in the photoprotective, de-epoxidized forms relative to shade leaves. Reflectance indices also revealed contrasting patterns of pigment development in leaves of contrasting structural types (annual versus evergreen). In H. annuus sun leaves, there was a remarkably rapid increase in amounts of both chlorophyll and xanthophyll cycle pigments along a leaf developmental sequence. This pattern contrasted with that of Quercus agrifolia (coast live oak, a sclerophyllous evergreen), which exhibited a gradual development of both chlorophyll and xanthophyll cycle pigments along with a pronounced peak of anthocyanin pigment content in newly expanding leaves. These temporal patterns of pigment development in Q. agrifolia leaves suggest that anthocyanins and xanthophyll cycle pigments serve complementary photoprotective roles during early leaf development. The results illustrate the use of reflectance indices for distinguishing divergent patterns of pigment activity in leaves of contrasting light history and functional type.

Two brown algae, Pelvetia canaliculata and Laminaria saccharina, from the higher and lower mediolittoral belts respectively, have been tested for their capacity to overcome high-light stress in water and in air (in both fully hydrated and desiccated states). When exposed to supersaturating light irradiance in water, the two species developed non-photochemical quenching of fluorescence (NPQ) which was correlated with an increase in the de-epoxidation ratio (DR) of the xanthophyll cycle carotenoids (violaxanthin, antheraxanthin and zeaxanthin) and was followed by a slower decrease in oxygen evolution. NPQ reached values of up to 9 in P. canaliculata but only 4·5 in L. saccharina, at DRs of 0·65 and 0·5, respectively. In air, the xanthophyll cycle was also operative but the efficiency of de-epoxidation decreased linearly with the degree of hydration of the thallus. Photoprotection capacities in air also appeared higher in P. canaliculata than in L. saccharina, probably due to the higher molar content of the xanthophyll cycle pool size relative to chlorophyll a (Chl a) in the former (nearly double of that L. saccharina at 19 carotenoid molecules per 100 Chl a), which may be associated with a higher DR at the same level of desiccation. The concomitant higher accumulation of zeaxanthin in P. canaliculata might divert a higher percentage of the incident energy from the reaction centres, as demonstrated by the levels of NPQ, with steady-state fluorescence reduced to below the initial F0 level. Such differences, together with the unequal resistance to desiccation of the operation of the xanthophyll cycle, should be considered as possible factors responsible for the distribution for these two species on the shore.

Responses of gas exchange and photosynthesis to changes in CO2 concentration and PPFD were examined in well watered plants of Delosperma tradescantioides Bgr. to establish the relative importance of these environmental changes on the photosynthetic machinery in this CAM-cycling species which grows naturally in both exposed and partly shaded environments. Plants were grown at two PPFDs (220 [LL] and 550 [HL] μmol m−2 s−1). HL plants had larger leaves with higher specific weight, water content and diurnal malic acid fluctuation. Photosynthetic PPFD responses were typically those of sun and shade species for HL and LL plants, both under 21% O2 and non-photorespiratory (2% O2) conditions. The CO2 compensation point in the absence of non-photorespirational CO2 evolution in the light (Γ∗) was c. 30 μmol mol−1. Irradiation reduced mitochondrial respiration by >50%. Comparison of the PPFD responses of linear electron flow rates derived from gas exchange measurements and from fluorescence analysis ([Jscr ]F) indicated effective photosynthetic control. [Jscr ]F was always larger than electron flow rates calculated from gas exchange, indicating that processes other than carboxylation and oxygenation were consistently important in energy consumption under all sampled environmental conditions. Regardless of PPFD during growth, electron flow to carboxylation and [Jscr ]F were linearly correlated, demonstrating that the photosynthetic apparatus was well adapted to PPFD during growth. In HL plants, non-photochemical quenching increased, and photochemical quenching and the quantum yield of linear electron transport through PS II decreased more slowly with increasing PPFD than in LL plants. In plants of both treatments non-photochemical energy dissipation seemed to be exhausted when the proportion of photons not utilizable by photochemistry exceeded 0·7. Results illustrate a pronounced ability of D. tradescantioides to acclimate to a 100% change in the prevailing PPFD and lend support to the hypothesis that CAM cycling might act as a photoprotective process.