Some lichenized ascomycetes synthesize melanic pigments in their upper cortices when exposed to ultraviolet light and high solar radiation. Our previous work showed that melanized chloro- and cyanolichens from both high light and more shaded habitats were less photoinhibited than pale ones during controlled exposure to high light. However, protection from high light might not necessarily be the consequence of just sun-screening by melanins in upper cortices. An inherent problem with earlier experiments was that the photobionts of melanized thalli might have received more light than those beneath pale cortices. The photobionts may therefore have possessed other light-induced tolerance mechanisms that gave protection from photoinhibition. Here, we aimed to test directly the inherent tolerance of lichen photobionts to photoinhibition. The method involved removing the lower cortices and medullas of three lichen species, Cetraria islandica, Crocodia aurata and Lobaria pulmonaria, and exposing the photobionts to light from below. Results confirmed that most of the improvement in tolerance to photoinhibition in melanized lichens derives from fungal melanization in the upper cortex. However, in C. islandica, the most heavily melanized species, algae from melanized thalli possessed a significantly higher tolerance to photoinhibition than those from pale thalli, suggesting that photobionts can also adapt themselves to high light.

Terminal fluorescence (FT) was monitored following root or foliar uptake of metribuzin in soybean, velvetleaf, and smooth pigweed seedlings grown in solution culture. These species are known to be tolerant, moderately susceptible, and susceptible, respectively, to metribuzin under field conditions. The indirect monitoring of herbicide metabolism by fluorescence is complicated by several factors (particularly photoinhibition) and is difficult to interpret when plants are kept in the light. Rapid declines in FT at metribuzin concentrations causing high photosystem II reaction center inhibition were interpreted, in conjunction with fresh weight and injury data, as resulting from photoinhibition in velvetleaf and smooth pigweed, and from a combination of photoinhibition and herbicide metabolism in soybeans. At lower concentrations, FT declines were apparently not affected by photoinhibition but were probably representative of herbicide metabolism. These results suggested a significant rate of metribuzin metabolism in soybeans, a minor degree of metabolism in smooth pigweed beginning after 12 h, and no detectable metabolism in velvetleaf. FT monitored over a 24-h dark period following foliar absorption showed declines in soybeans indicating a modest degree of metribuzin metabolism (at intermediate herbicide doses) but no FT declines attributable to herbicide metabolism in velvetleaf or smooth pigweed. Results indicate that the inhibition of a high percentage of photosystem II reaction centers is required before photoinhibition and photooxidative leaf damage can result from photosystem II inhibitor herbicides such as metribuzin.

Light conditions provide important information about the best time and place for seedling establishment. Photoinhibition of seed germination (PISG), defined as the partial or complete suppression of germination under white light, has been interpreted as a physiological adaptation to avoid germination at or near the soil surface. This review is the first report of an all-inclusive, fully quantitative analysis of PISG in seed plants. Pertinent data available from the published literature for 301 taxa from 59 families and 27 orders were assessed. The association of PISG with several plant and seed traits allowed us to consider the adaptive significance of PISG in relation to plant life histories and the natural environments. As no gymnosperm has been found to be truly photoinhibited, it seems that PISG is apomorphic to flowering plants (especially monocots). Seeds of most taxa with PISG have a dark colour and intermediate mass, mostly in the range 1 to 27 mg. PISG is absent from humid tropical regions and from cold climates, but it is strongly associated with open, disturbed and dry habitats. An intriguing implication of PISG is the formation of a soil-surface seed bank. Taken together, these results clearly indicate that PISG is a physiological adaptation to avoid germination on the soil surface, where conditions are not suitable for seedling establishment. PISG is probably much more frequent in seed plants than previously thought. Thus, laboratory experiments should be conducted under well-characterized light and dark conditions.

Mediterranean characteristics are attributed to the genus Crocus, which is inadequately studied in terms of seed germination. An afterripening requirement is very common in environments with warm and dry periods, and photoinhibition has been detected in many angiosperms inhabiting dry and open areas. The effects of afterripening and light on seed germination were investigated for the first time in 23 native Greek Crocus taxa, collected from various localities with either a Mediterranean or a temperate climate. Germination experiments were conducted in continuous darkness and in light at the optimal temperature for each taxon, with both freshly collected and afterripened seeds; warm stratification (20°C, darkness) was also examined in 22 taxa. A number of selected taxa were additionally investigated with respect to afterripening outdoors, afterripening and warm stratification at higher temperatures (35 and 25°C, respectively), stratification at 20/10°C, dry storage at low temperatures, response to gibberellic acid and phenology of embryo growth. It was postulated that an afterripening requirement is a characteristic of the genus Crocus, and we found that it can be fulfilled in nature during the Mediterranean dry summer. Also, for the vast majority of the taxa, warm stratification and stratification at 20/10°C can both meet the afterripening requirement. Embryos of the taxa studied are underdeveloped and have to grow prior to germination. Intrageneric differences of seed germination were observed only towards light, with photoinhibition being predominant in taxa from drier environments.

Seeds of watermelon (Citrullus lanatus cv. Sugar Baby) germinated optimally in the dark, throughout the range 20–40°C. Germination was inhibited by continuous (c) irradiation with farred (FR), blue or white incandescent light; short pulses were ineffective. Intermittent FR could fully substitute for cFR; the effect of alternating red and FR pulses during 30-min dark cycles depended upon the timing of the illuminations within each cycle. It is concluded that germination in watermelon is controlled by the low-energy reaction of phytochrome. However, continuous and intermittent red light resulted in partial reductions in germinability. Opening the seed coat at the radicle end enhanced germination in the dark and reduced photosensitivity towards FR. The outer, lignified part of the testa exerted a mechanically restrictive force upon the expanding radicle; this force was estimated to be equivalent to 0.3 MPa. The kinetics, at 25°C in the dark, of the time course of germination and the escape from the inhibitory actions of cFR and osmoticum (0.5m mannitol) were all sigmoid curves, which, upon transformation to normal distributions, had different means, but statistically similar variances. The cFR-irreversible activation of germination by phytochrome and the mannitol-irreversible onset of radicle elongation preceded radicle protrusion (mean at 32 h, 25°C) by 8 and 6 h, respectively. From the results and data on imbition, it is concluded that activation of germination in watermelon takes place during the second, stationary phase of imbibition (20–30 h after sowing at 25°C).

Ecophysiological aspects of seed germination were investigated in four Mediterranean geophytes of the genus Muscari (Liliaceae): M. comosum (tassel hyacinth), M. neglectum (common grape hyacinth), M. commutatum and M. weissii. Experiments were performed at constant temperatures in the dark and under temperature and light conditions simulating those prevailing in nature during November–January, i.e. well into the rainy season of the Mediterranean climate. In all species, no primary dormancy was revealed, and germination occurred in a rather narrow range of cool temperatures (optimum at 10 or 15°C) and at a remarkably slow rate; both germination characteristics seem to be associated with autumn/winter seed germination and seedling establishment. Such a postulated strategy is ecologically advantageous within an unpredictable rainfall regime, known to prevail during the start of the rainy period of the Mediterranean climate. This strategy may also explain the spread of germination of M. comosum seeds over two consecutive years, observed by Theophrastus. Far-red light, simulating light conditions under a dense canopy, resulted in only a slight delay of germination compared to dark controls. Diurnal white light, qualitatively simulating natural daylight, caused a significant decrease of the germination rate in all four species studied. Moreover, white light was found to suppress considerably final seed germination (photoinhibition) in M. weissii and M. neglectum; in the latter species, prolonged imbibition under white light also led to the induction of secondary dormancy.

An individual-based model is presented which describes the spatial and temporal evolution of phytoplankton growth in terms of a Lagrangian ensemble of cells affected by various physical and biological forcing factors. The motion of cells develops according to a turbulent velocity field which simulates the Antarctic mixed layer during the summer. The cell growth is a function of the irradiance regime, nutrient availability and the vertical position of the individual with respect to the other cells (in order to evaluate the self-shading effect). The values of photosynthetic parameters used to simulate the photophysiological response of the organisms are derived from measurements collected in the Ross Sea. In contrast to previous individual-based descriptions, all the physical and biological processes involved are explicitly reproduced in their dynamical features. Coupling different mixing levels with photoacclimation strategies leads to a wide range of photophysiological responses which underline the role of individual physiological histories in determining the growth of the population as a whole. Simulated photosynthetic parameters, chlorophyll a concentrations and integrated primary production correspond closely to in situ data and confirm that photoacclimation to low irradiance and strong mixing regimes may be considered as crucial factors in the photosynthetic performance of Antarctic phytoplankton.

Two shade-adapted (Lobaria pulmonaria and Cetraria islandica) and two sun-adapted lichen populations (Xanthoria parietina and Cetraria nivalis) were exposed to three irradiance regimes (1: photosynthetic radiation—PAR, 2: PAR+UV-A, 3: PAR+UV-A+UV-B) and two hydration regimes (1: no hydration, 2: daily hydration) in a growth cabinet for three weeks. Shade-adapted thalli had transparent upper cortices without coloured pigments, whereas sun-adapted thalli had coloured UV-B absorbing cortical pigments masking the photobiont. Manipulation of pigment concentration was the third factor used in the factorial design (1: pigments intact, 2: pigments non-destructively extracted from air-dry living thalli by acetone). Inhibition of the photobiont due to PAR alone was severe in the two shade-adapted populations, but no applied UV wavelength bands caused additional aggravation of photoinhibition. Shade-adapted thalli of the ubiquitous C. islandica were more PAR-susceptible than of the rare old forest lichen L. pulmonaria, suggesting that screening by the mycobiont rather than photobiont characteristics, account for their different success in sun-exposed localities. Hydration of shade-adapted species during exposure reduced their photoinhibition substantially, probably because of moisture-activated repair mechanisms. On the contrary, the sun-adapted X. parietina was most phototolerant in the desiccated state, whereas hydration caused increased photoinhibition. When removing the orange cortical pigment parietin, the photoinhibition in moist thalli was aggravated, confirming a PAR-protective function of parietin. No effects of irradiance treatment, pigment extraction (usnic acid), or hydration level were observed in C. nivalis.

A mechanistic model was developed, to simulate the main facets of photoinhibition in phytoplankton. Photoinhibition is modelled as a time dependent decrease in the initial slope of a photosynthesis versus irradiance curve, related to D1 (photosystem II reaction centre protein) damage and non-photochemical quenching. The photoinhibition model was incorporated into an existing ammonium-nitrate nutrition interaction model capable of simulating photoacclimation and aspects of nitrogen uptake and utilization. Hence the current model can simulate the effects of irradiance on photosynthesis from sub-saturating to inhibitory photon flux densities, during growth on different nitrogen sources and under nutrient stress. Model output conforms well to experimental data, allowing the extent of photoinhibition to be predicted under a range of nutrient and light regimes. The ability of the model to recreate the afternoon depression of photosynthesis and the enhancement of photosynthesis during fluctuating light suggests that these two processes are related to photoinhibition. The model may be used to predict changes in biomass and/or carbon fixation under a wide range of oceanographic situations, and it may also help to explain the progression to dominance of certain algal species, and bloom formation under defined irradiance and nutrient conditions.

The effects of enhanced UV-B radiation were investigated in the carnivorous plant Pinguicula vulgaris in a field experiment performed in Abisko, North Sweden (68° 21′ N, 18° 49′ E, 380 m above sea level). Potted plants were exposed to either ambient or ambient plus supplemental UV-B radiation, simulating a 15% ozone depletion. No effect was observed on either the epicuticular (external) or cellular (internal) UV absorbing capacity of the leaves. However, the anthocyanin content was more than doubled by supplemental UV-B radiation. In laboratory experiments, the anthocyanin rich, UV-B treated leaves were less susceptible to a low temperature/high light photoinhibitory treatment, as judged by in vivo chlorophyll fluorescence measurements. Yet, this potential benefit did not considerably affect the growth of the plant in the field (leaf area and dry mass, reproductive dry mass, flowering frequency, senescence rates, dry mass of winter buds). However, there was a marginally significant increase in root dry mass and in the root to shoot ratio, which may underlie the significant increase in the nitrogen content of the leaves. We suggest that P. vulgaris is resistant against UV-B radiation damage and that the possible negative effects of additional UV-B radiation on the growth of these plants may have been effectively counterbalanced by the lower risk of photoinhibition, due to the concomitant increase in anthocyanins.

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.

Photoinhibition of photosynthesis in the red alga Chondrus crispus from different shore levels was investigated under laboratory and field conditions. The underwater light climate at Roscoff (French Atlantic coast), characterized by transmittance spectra, was shown to be intermediate between Jerlov coastal type 1 and oceanic type III. Chondrus crispus thalli were collected at four depths (3·5–8·5 m below high tide level) and exposed to daily courses of artificial and natural light. Filters were used to discriminate between the effects of photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) under the photoinhibitory treatments. In the presence of natural UVR, photoinhibition at midday was increased by up to 60%. Recovery from photoinhibition was measured at different times throughout the light stress. Recovery kinetics differed between morning and early afternoon in all treatments, and were dependent on the light dose. Whereas biphasic kinetics were observed in all treatments in the morning, the recovery in early afternoon was significantly slower. The influence of UVR on recovery was apparent after exposure to higher light doses. In all investigations, sensitivity to photoinhibition corresponded to zonation of the algae, with deeper-growing algae showing a greater depression of fluorescence yield, and slower recovery (indicating acclimation to the growth irradiance). An increase in the phycoerythrin content and increasing photosynthetic efficiency with increasing growth depth were observed.

Decreases in the concentration of ozone in the stratosphere are predicted to result in more biologically damaging ultraviolet-B radiation (UVBR) reaching the Earth's surface in the near future. The consequences for plant life of this small change in the solar spectrum are currently uncertain; this is particularly true for aquatic macrophytic algae. The aim of this review is to summarize the current knowledge of UVBR effects on macrophytes, and to highlight areas for future research. An overview of published underwater UVBR measurements shows that penetration of this waveband is low in the coastal waters where most macroalgae are found. Knowledge of UVBR targets in macroalgae is poor, and biological weighting functions have yet to be established. Studies of higher plants and phytoplankton are used to illustrate the many possible targets and effects of UVBR. The role of photosynthetically active radiation in causing dynamic and chronic photoinhibition in macroalgae and the interactions of PAR with UVR are examined in detail. These interactions are central to interpretation of field measurements of photoinhibition under natural solar radiation conditions. Potential strategies and mechanisms for repair of UVBR damage are discussed. Before the long-term effects of UVBR on macroalgal production and community structure can be determined, a greater knowledge of the specific targets of UVBR in macrophytes and the mechanisms of protection and repair must be determined. The ability of macrophytes to acclimate to higher UVBR and the interaction of UVBR with other stresses, such as high PAR, temperature and desiccation, must be assessed. Studies are needed which encompass effects on all stages of the macrophyte life cycle, from recruitment to maturity.