Exposure to solar radiation increases damage to both host tissues and algal symbionts of corals during thermal stress

Elevated seawater temperatures have long been accepted as the principal stressor causing the loss of symbiotic algae in corals and other invertebrates with algal symbionts (i.e., bleaching). A secondary factor associated with coral bleaching is solar irradiance, both its visible (PAR: 400–700 nm) and ultraviolet (UVR: 290–400 nm) portions of the spectrum. Here we examined the synergistic role of solar radiation on thermally induced stress and subsequent bleaching in a common Caribbean coral, Montastraea faveolata. Active fluorescent measurements show that steady-state quantum yields of photosystem II (PSII) fluorescence in the zooxanthellae are markedly depressed when exposed to high solar radiation and elevated temperatures, and the concentration of D1 protein is significantly lower in high light when compared to low light treatments under the same thermal stress. Both photosynthetic pigments and mycosporine-like amino acids (MAAs) are also depressed after experimental exposure to high solar radiation and thermal stress. Host DNA damage is exacerbated under high light conditions and is correlated with the expression of the cell cycle gene p 53, a cellular gatekeeper that modulates the fate of damaged cells between DNA repair processes and apoptotic pathways. These markers of cellular stress in the host and zooxanthellae have in common their response to the enhanced production of reactive oxygen species during exposure to high irradiances of solar radiation and elevated temperatures. Taking these results and previously published data into consideration, we conclude that thermal stress during exposure to high irradiances of solar radiation, or irradiances higher than the current photoacclimatization state, causes damage to both photochemistry and carbon fixation at the same time in zooxanthellae, while DNA damage, apoptosis, or necrosis are occurring in the host tissues of symbiotic cnidarians.Abbreviations _PSII Functional absorption cross-section for PSII - F_o, F_m Minimum and maximum yields of chlorophyll a fluorescence measured after dark acclimation (relative units) - F_v Variable fluorescence after dark acclimation (=F_m–F_o), dimensionless - F_v/F_m Maximum quantum yield of photochemistry in PSII measured after dark acclimation, dimensionless - F, F_m Steady-state and maximum yields of chlorophyll a fluorescence measured under ambient light (relative units) - F/F_m Quantum yield of photochemistry in PSII measured at steady state under ambient light Communicated by R.C. Carpenter     

Photosynthesis of two Arctic macroalgae under different ambient radiation levels and their sensitivity to enhanced UV radiation

The change in optimal quantum efficiency (F _v/F _m) of the Arctic species Laminaria saccharina and Palmaria palmata was investigated in a long-term experiment in situ under different radiation levels during the summer of 1997 in the Kongsfjord (Ny-Ålesund, Spitsbergen, Norway, 78°55.5′N, 11°56.0′E). Whole plants were incubated in an open box system made of UV-transparent Perspex and exposed to solar radiation (λ>295?nm), solar radiation excluding UVB (λ?>?320?nm) and solar radiation excluding UVA?+ UVB (λ?>?400?nm). Increasing radiation levels were simulated by transplantation of the pre-adapted algae from their growth depth at 2?m to a water depth of 1?m. Sensitivity to artificially increased UV radiation was determined by exposure of algae from the three treatments to 6?h of strong UV radiation. P. palmata was relatively insensitive to increasing UV radiation and recovered very fast and almost completely in 2?h. Even plants pre-cultured in ambient radiation levels excluding UVA?+?UVB or UVB only showed no photoinhibition after exposure to extra UV radiation in the laboratory. L. saccharina was, in comparison to P. palmata, more sensitive and showed photoinhibition under solar radiation and solar minus UVB radiation after transplantation from 2 to 1?m water depth. However, after 3?weeks at 1?m depth, F _v/F _m of L. saccharina was equal in all treatments and restored to the original values at the start of the experiment. Sensitivity to extra UV radiation in the laboratory increased in time, although recovery was also fast and occurred within 20?h.     

Inhibition of photosynthesis by solar radiation in Dunaliella salina: relative efficiencies of UV-B, UV-A and PAR

Inhibition of photosynthesis after exposure to solar radiation was investigated in the marine green alga Dunaliella salina by monitoring photosynthetic optimal quantum yield F_v/F_m and efficiency of oxygen production. Samples were exposed to solar radiation in Ancient Korinth, Greece (37°58′ N, 23°0′ E) in August 1994. Within 30 min, F_v/F_m and efficiency of oxygen production decreased with similar kinetics with increasing exposure time. The inhibition, however, diminished when ultraviolet radiation was progressively excluded by means of colour filter glasses. Samples exposed for 3 h showed complete or partial recovery of photosynthesis, with almost the same rate under all irradition conditions. The fit of the experimental data with an analytical model describing inhibition of photosynthesis as a function of a linear combination of the photon fluence in the UV-B, UV-A and PAR allows one to estimate the relative mean effectiveness for inhibition by the three spectral ranges [about 2 × 10^?4, 4 × 10^?6 and 2 × 10^?7 (μmol photons m^?2)^?1 for UV-B, UV-A and PAR, respectively].     

Growth enhancement of soybean (Glycine max) upon exclusion of UV-B and UV-B/A components of solar radiation: characterization of photosynthetic parameters in leaves

Exclusion of UV (280–380 nm) radiation from the solar spectrum can be an important tool to assess the impact of ambient UV radiation on plant growth and performance of crop plants. The effect of exclusion of UV-B and UV-A from solar radiation on the growth and photosynthetic components in soybean (Glycine max) leaves were investigated. Exclusion of solar UV-B and UV-B/A radiation, enhanced the fresh weight, dry weight, leaf area as well as induced a dramatic increase in plant height, which reflected a net increase in biomass. Dry weight increase per unit leaf area was quite significant upon both UV-B and UV-B/A exclusion from the solar spectrum. However, no changes in chlorophyll a and b contents were observed by exclusion of solar UV radiation but the content of carotenoids was significantly (34–46%) lowered. Analysis of chlorophyll (Chl) fluorescence transient parameters of leaf segments suggested no change in the F _v/F _m value due to UV-B or UV-B/A exclusion. Only a small reduction in photo-oxidized signal I (P700⁺)/unit Chl was noted. Interestingly the total soluble protein content per unit leaf area increased by 18% in UV-B/A and 40% in UV-B excluded samples, suggesting a unique upregulation of biosynthesis and accumulation of biomass. Solar UV radiation thus seems to primarily affect the photomorphogenic regulatory system that leads to an enhanced growth of leaves and an enhanced rate of net photosynthesis in soybean, a crop plant of economic importance. The presence of ultra-violet components in sunlight seems to arrest carbon sequestration in plants. An erratum to this article can be found at     

Photoinhibition by visible and ultraviolet radiation in the red macroalga Porphyra umbilicalis grown in the laboratory

Photosynthetic oxygen production and PAM fluorescence measurements were used to follow photoinhibition in the red macroalga Porphyra umbilicalis. Exposure to simulated solar radiation caused inhibition of the effective photosynthetic quantum yield from which the thalli partially recovered in the shade in subsequent hours. There were no significant differences between samples exposed to unfiltered radiation and those exposed to radiation from which increasing portions of UV radiation had been removed indicating that the thalli are well adapted to current levels of solar PAR and UV radiation. This notion was supported by the finding of high concentrations of UV screening pigments which were even enhanced by exposure to increased UV radiation. However, when exposed to (only) UV radiation about 50% higher than that encountered by the organisms in their natural habitat, the photosynthetic yield decreased slowly and did not show any recovery even when the degree of inhibition did not exceed 10%.     

Differential responses of Nostoc sphaeroides and Arthrospira platensis to solar ultraviolet radiation exposure

During October to December 2003 we carried out experiments to assess the impact of high solar radiation levels (as normally occurring in a tropical region of Southern China) on the cyanobacteria Nostoc sphaeroides and Arthrospira (Spirulina) platensis. Two types of experiments were done: a) Short-term (i.e., 20 min) oxygen production of samples exposed to two radiation treatments (i.e., PAR+UVR—280–700 nm, and PAR only—400–700 nm, PAB and P treatments, respectively), and b) Long-term (i.e., 12 days) evaluation of photosynthetic quantum yield (Y) of samples exposed to three radiation treatments (i.e., PAB; PA (PAR+UV-A, 320–700 nm) and P treatments, respectively). N. sphaeroides was resistant to UVR, with no significant differences (P>0.05) in oxygen production within 20 min of exposure, but with a slight inhibition of Y within hours. A fast recovery of Y was observed after one day even in samples exposed to full solar radiation. A. platensis, on the other hand, was very sensitive to solar radiation (mainly to UV-B), as determined by oxygen production and Y measurements. A. platensis had a circadian rhythm of photosynthetic inhibition, and during the first six days of exposure to solar radiation, it varied between 80 and 100% at local noon, but cells recovered significantly during afternoon hours. There was a significant decrease in photosynthetic inhibition after the first week of exposure with values less than 50% at local noon in samples receiving full solar radiation. Samples exposed to PA and P treatments recovered much faster (within 2–3 days), and there were no significant differences in Y between the three radiation treatments when irradiance was low (late afternoon to early morning). Long-term acclimation seems to be important in A. platensis to cope with high UVR levels however, it is not attained through the synthesis of UV-absorbing compounds but it seems to be mostly related to adaptive morphological changes.     

Cellular responses to elevated light levels in Fucus spiralis embryos during the first days after fertilization

Cellular responses of 1‐, 2‐ and 4‐d‐old Fucus spiralis embryos subjected to a single dose of elevated photosynthetically active photon flux density (PPFD), with or without ultraviolet (UV) radiation, were investigated by measuring the effects on the effective quantum yield of photosystem II (ΔF / F_m′) and intracellular production of active oxygen species (AOS). Production of AOS was determined by the in vivo conversion of 5‐(and‐6)‐chloromethyl‐2′,7′‐dichlorodihydrofluorescein diacetate (CM‐DCFH₂‐DA) to the fluorescent compound dichlorofluorescein (DCF) using confocal laser scan microscopy (CLSM) and image analysis. The role of xanthophyll cycle pigments in photoprotection was also assessed. A rapid decline in ΔF / F_m′ was observed under all elevated light conditions. A correlation was found between non‐photochemical quenching and the de‐epoxidation ratio zeaxanthin/(zeaxanthin + violoxanthin). Active oxygen formation increased with PPFD and was higher in older embryos and when UVB was present. Two photoinhibition responses were recognized: (i) a rapid decline of the PSII yield due to the violoxanthin–zeaxanthin cycle (photoprotection), and (ii) a slower second‐phase decline, correlated with active oxygen production. Electron transport rate (ETR) increased with embryo age, and was correlated with AOS production. As a result of enhanced AOS production, there was a slow recovery of the PSII yield, in particular with increased effective UV dose. In general, embryos were able to recover from the imposed light conditions, but UVB had a more damaging effect. Overall, our data suggest that under natural conditions, embryos of F. spiralis are susceptible to elevated light levels, and that UVB radiation is an important stress factor.     

Ultraviolet-B inhibition of motility in green and dark bleachedEuglena gracilis

The motility of green and dark bleachedEuglena gracilis was studied under artificial and solar UV-B radiation. The percentage of motile cells in the population was drastically impaired after exposure to unfiltered sunlight for only a few hours. Dark bleached cells were even more affected than green organisms. The effect is caused mainly by the solar UV-B component, since filtering the sunlight by either a layer of ozone or a UV-B-absorbing filter substantially increased the survival rate. Addition ofp-quinone, a scavenger of free radicals produced in a type I photodynamic reaction, did not relieve the UV-B effects, but was cytotoxic at higher concentrations. Likewise, 1,4-diazobicyclo[2,2,2]octane and imidazole, which quench singlet oxygen (¹O₂) generated in a type II photodynamic reaction, did not prolong the survival in UV-B irradiation. D₂O, which, in contrast, prolongs the lifetime of¹O₂, is tolerated by the cells but does not aggravate the UV-B inhibition. Thus, photodynamic processes of both type I and II can be ruled out as possible mechanisms of UV-B inhibition of motility inEuglena gracilis.     

Effects of solar radiation and solar radiation deprived of UV-B and total UV on photosynthetic oxygen production and pulse amplitude modulated fluorescence in the brown alga Padina pavonia

Abstract: The effects of solar radiation on photosynthetic oxygen production and pulse amplitude modulated (PAM) fluorescence were measured in the marine brown macroalga Padina pavonia harvested from different depths from the Greek coast near Korinth. In fluence rate-response curves the light compensation point for photosynthetic oxygen production increased and the saturation level decreased with increasing exposure time to solar radiation. Cutting off the UV-B wavelength range (280–315 nm) from solar radiation reduced the inhibition of photosynthesis, and the organisms were less affected when all of the UV radiation was filtered out. Algae collected from 7 m depth were much more prone to photoinhibition than those harvested from rock pools exposed to unfiltered solar radiation. During continuous exposure to solar radiation, rock pool algae showed photoinhibition after longer periods of time than specimens from 7 m or from dark adapted habitats. When subjected to unfiltered solar radiation the ratio of the variable fluorescence to the maximal fluorescence     (F_v = F_m− F_o) rapidly declined with increasing exposure time. However, again algae from 7 m depth were more prone to photoinhibition than rock pool algae. The differences between the two ecological strains were less obvious when UV-B or total UV was removed from solar radiation. Only in the latter case a complete recovery was observed after 2 h while, when exposed to unifiltered sunlight, only the rock pool algae recovered completely within that time.     

Photoinhibition and recovery in tropical plant species: response to disturbance

Disturbance or rainforest is often followed by mass mortality of understorey seedlings. Transitions of shade grown plants to full sunlight can cause reductions in the efficiency with which light is used in photosynthesis, called photoinhibition. In order to assess the influence of photoinhibition on mortality and growth after rainforest disturbance this study examined photoinhibition in both simulated and real forest disturbances in northern Papua New Guinea. In an experiment simulating rainforest disturbance, exposure of shade-grown plants to full sunlight resulted in abrupt decreases in the chlorophyll fluorescence parameter F _v/F _m that is characteristic of photoinhibition. However, in the well-watered plants used in these experiments there were no fatalities during 3 weeks after exposure to full sunlight. Thus, it is unlikely that photoinhibition, alone, is responsible for seedling fatalities after rainforest disturbances, but more likely that fatalities are due to photoinhibition in conjunction with other environmental stress. There were differences between the response of species to the simulated disturbance that concurred with their preferred habitats. For example, species form the genus Barringtonia, which is commonly found in shaded understorey environments, underwent greater reductions in F _v/F _m and were slower to recover than species that usually inhabit high solar radiation environments. The extent of photoinhibition and the rate of recovery appeared to be dependent on avoidance of direct solar radiation by altering leaf angles and on increasing maximum photosynthetic rates. A field survey of photoinhibition in man-made rainforest gaps corroborated the findings of the simulated disturbance experiment showing that plant species commonly found in shaded environments showed a greater degree of photoinhibition in forest gaps at midday than those species which are classified as species that benefit from gaps or specialist gap inhabitors.     

IN VIVO MEASUREMENT OF ACTIVE OXYGEN PRODUCTION IN THE BROWN ALGA FUCUS EVANESCENS USING 2′,7′-DICHLOROHYDROFLUORESCEIN DIACETATE1

Intracellular production of active oxygen in the brown alga Fucus evanescens C. Ag. was studied by measuring the capacity for in vivo conversion of 2′,7′-dichlorohydrofluorescein diacetate (DCFH-DA) to the fluorescent dye 2′,7′-dichlorofluorescein (DCF), both in emersed and immersed seaweeds. Algae were incubated in seawater containing DCFH-DA under a range of conditions, and it was also possible to load algae with DCFH-DA and then follow subsequent DCF production in emersed tissue. DCF formation was linear for at least 2 h in both darkness and light, with the rate of formation increasing with the light level. DCF formation was temperature dependent. It also increased when algae were treated with H₂O₂ or methyl viologen (paraquat), which disrupts photosystem 1 electron transport and increases O^?₂ production. Exogenous catalase reduced in vivo DCF production, presumably by lowering cellular concentrations of H₂O₂. Hydrogen peroxide was released into the seawater by illuminated algae resulting in external dye conversion to DCF. However, this does not interfere with in vivo measurement of DCF by loaded, washed algae because DCF leakage appeared to be negligible. Internal DCF did not affect photosynthetic oxygen production relative to untreated controls. Overall, our data suggest that DCFH-DA is a potentially very useful probe for studying active oxygen metabolism in seaweeds subjected to environmental stresses.     

Influence of ultraviolet-B (UV-B) radiation on photosynthetic and growth characteristics in field-grown cassava (Manihot esculentum Crantz)

The effects of ultraviolet-B (UV-B between 290 and 320 nm) on photosynthesis and growth characteristics were investigated in field grown cassava (Manihot esculentum Crantz). Plants were grown at ambient and ambient plus a 5.5kJ m^?2 d^?1 supplementation of UV-B radiation for 95 d. The supplemental UV-B fluence used in this experiment simulated a 15% depletion in stratospheric ozone at the equator (0°N). Carbon dioxide exchange, oxygen evolution, and the ratio of variable to maximum fluorescence (F_v/F_m) were determined for fully expanded leaves after 64–76 d of UV-B exposure. AH plants were harvested after 95 d of UV-B exposure, assayed for chlorophyll and UV-B absorbing compounds, and separated into leaves, petioles, stems and roots. Exposure to UV-B radiation had no effect on in situ rates of photosynthesis or dark respiration. No difference in the concentration of UV-B absorbing compounds was observed between treatments. A 2-d daytime diurnal comparison of F_v to F_m ratios indicated a significant decline in F_v/F_m ratios and a subsequent increase in photoinhibition under enhanced UV-B radiation if temperature or PPF exceeded 35°C or 1800μmol m^?2 s^?1, respectively. However, UV-B effects on fluorescence kinetics appeared to be temporal since maximal photosynthetic rates as determined by oxygen evolution at saturated CO₂ and PPF remained unchanged. Although total biomass was unaltered with UV-B exposure, alterations in the growth characteristics of cassava grown with supplemental UV-B radiation are consistent with auxin destruction and reduced apical dominance. Changes in growth included an alteration of biomass partitioning with a significant increase in shoot/root ratio noted for plants receiving supplemental UV-B radiation. The increase in shoot/root ratio was due primarily to a significant decrease in root weight (–32%) with UV-B exposure. Because root production determines the harvest-able portion of cassava, UV-B radiation may still influence the yield of an important tropical agronomic species, even though photosynthesis and total dry biomass may not be directly affected.     

The effects of solar radiation and nutrient solution temperature on the uptake of oxygen by submerged roots of mature tomato plants

Summary A statistical analysis of data on oxygen uptake by submerged roots of mature tomato plants, from a nutrient solution containing sufficient oxygen, has demonstrated that variations in oxygen uptake are positively related to variations in solar radiaton, particularly at radiation levels below 2.0 MegaJoules.h⁻¹.m⁻². Above this level, however, anegative relationship is indicated, suggesting that intense solar radiation reduces the ability of plant roots to obtain sufficient oxygen for optimum growth processes. It also appears that there is atime lag of about one to two hours between the effects of solar radiation on oxygen uptake. Greenhouse air temperature is positively correlated with radiation, with little or no time lag, and variations in greenhouse air temperature effect variations in solution temperature with a time lag of 2 to 3 hours. Greenhouse air temperature affects respiration in the submerged roots indirectly, with a 1 to 2 hour lag, by effecting the nutrient solution temperature, and possibly by affecting the growth processes in thesubaerial roots, which in turn may result, in some way, in increased oxygen uptake by the submerged roots, partly bypassing the restraining effects of lagging nutrient solution temperature. Variations in nutrient solution temperature do not appear to be causally related in effecting variations in oxygen uptake in a major way, probably because a considerable time lag of around 3 hours seems to exist between variations in radiation and solution temperature, in this experiment. As such, radiation and greenhouse air temperature appear to be rather important and dominant variables, effecting changes in oxygen uptake. In addition, the results suggest thatdaytime, variable heating of the nutrient solution, in step with the variations of solar radiation, as well as shading of the tomato crop under conditions of intense solar radiation would likely be beneficial for plant growth and crop production.     

Photoinhibition in common atlantic macroalgae measured on site in Gran Canaria

The photosynthetic quantum yield was analysed in four common atlantic macroalgae, the Rhodophytes Gelidium arbuscula and Halopithys incurvus and the Phaeophytes Halopteris scoparia and Lobophora variegata in Gran Canaria, Canary Islands at their growth site. The fluorescence parameters were measured using a portable pulse amplitude modulated (PAM) fluorometer (PAM 2000) instrument and a diving PAM under water without removing the thalli from their growth sites. Solar radiation was monitored continuously above and under water during the whole experimental period using two three-channel dosimeters (European light dosimeter network; ELDONET) (Real Time Computer, M?hrendorf, Germany). These instruments measure solar radiation in three wavelength ranges, ultraviolet (UV)-A, UV-B and photosynthetic active radiation (PAR). In all four algae the effective photosynthetic quantum yield decreased significantly from the optimal values measured after dark adaptation due to exposure to 15 min solar radiation, but at least partially recovered subsequently in the shade within several hours. Increasing the exposure period to 30 min intensified the photoinhibition. In some algae no recovery was observed after this treatment and in others no significant recovery could be detected. Exposure to unfiltered solar radiation caused a significantly higher photoinhibition than PAR-only radiation or PAR plus UV-A. A substantial inhibition was found in all algae at their growth sites in the water column when the sun was at high angles, as measured with the diving PAM. Received in revised form: 15 May 2000 Electronic Publication     

Ocean acidification stimulates particulate organic carbon accumulation in two Antarctic diatom species under moderate and high natural solar radiation

Impacts of rising atmospheric CO₂ concentrations and increased daily irradiances from enhanced surface water stratification on phytoplankton physiology in the coastal Southern Ocean remain still unclear. Therefore, in the two Antarctic diatoms Fragilariopsis curta and Odontella weissflogii, the effects of moderate and high natural solar radiation combined with either ambient or future pCO₂ on cellular particulate organic carbon (POC) contents and photophysiology were investigated. Results showed that increasing CO₂ concentrations had greater impacts on diatom physiology than exposure to increasing solar radiation. Irrespective of the applied solar radiation regime, cellular POC quotas increased with future pCO₂ in both diatoms. Lowered maximum quantum yields of photochemistry in PSII (F_v/F_m) indicated a higher photosensitivity under these conditions, being counteracted by increased cellular concentrations of functional photosynthetic reaction centers. Overall, our results suggest that both bloom‐forming Antarctic coastal diatoms might increase carbon contents under future pCO₂ conditions despite reduced physiological fitness. This indicates a higher potential for primary productivity by the two diatom species with important implications for the CO₂ sequestration potential of diatom communities in the future coastal Southern Ocean.     

Resurrection kinetics of photosynthesis in desiccation‐tolerant terrestrial green algae (Chlorophyta) on tree bark

The rough bark of orchard trees (Malus) around Darmstadt is predominantly covered in red to purple‐brown layers (biofilms) of epiphytic terrestrial alga of Trentepohlia umbrina. The smooth bark of forest trees (Fagus sylvatica L. and Acer sp.) in the same area is covered by bright green biofilms composed of the green algae Desmococcus, Apatococcus and Trebouxia, with a few cells of Coccomyxa and ‘Chlorella’ trebouxioides between them. These algae are desiccation tolerant. After samples of bark with the biofilms were kept in dry air in darkness for various periods of time, potential quantum yield of PSII, F_v/F_m, recovered during rehydration upon rewetting. The kinetics and degree of recovery depended on the length of time that the algae were kept in dry air in the desiccated state. Recovery was better for green biofilm samples, i.e. quite good even after 80 days of desiccation (F_v/F_m = ca. 50% of initial value), than the red samples, where recovery was only adequate up to ca. 30–40 days of desiccation (F_v/F_m = ca. 20–55% of initial value). It is concluded that the different bark types constitute different ecophysiological niches that can be occupied by the algae and that can be distinguished by their capacity to recover from desiccation after different times in the dry state.     

The influence of micro-algae on the oxygen dynamics in a brackish ditch

Summary Seasonal changes in the activity of phytoplankton and benthic algae in relation to diurnal oxygen pulses were investigated in a 120 cm deep, brackish hypertrophic ditch. A vertical chloride gradient was built up by saline seepage and drain-water effluent. The stable chloride gradient could lead to oxygen stratification near the sediment, and to oxygen gradients towards the water surface. The oxygen gradients were rather unstable, depending on the chloride gradient and the wind velocity.Light was limiting photosynthesis both in summer and in winter. Surface oxygen maxima increased with solar radiation during summer.In summer the diatomCyclotella caused surface oxygen maxima at light saturation in the late afternoon. Simultaneously, the dominant flagellatesPeridinium andChlamydomonas produced oxygen in dim light, probably choosing their favourite light energy level by vertical migration. Oxygen fluctuations ranged from 0 to 34 mg O₂.l^–1 in a 100 cm vertical profile above a 20 cm anoxic layer. The amplitude of the diurnal oxygen maxima varied from 10 to 34 mg O₂.l^–1.In winter the water became very clear. The oxygen gradient was inverted during the day showing a characteristic oxygen maximum above the bottom, produced by benthicAchnanthes colonies.Communication no. 193.     

Signatures of diversifying selection at EST‐SSR loci and association with climate in natural Eucalyptus populations

Understanding the environmental parameters that drive adaptation among populations is important in predicting how species may respond to global climatic changes and how gene pools might be managed to conserve adaptive genetic diversity. Here, we used Bayesian F_ST outlier tests and allele–climate association analyses to reveal two Eucalyptus EST‐SSR loci as strong candidates for diversifying selection in natural populations of a southwestern Australian forest tree, Eucalyptus gomphocephala (Myrtaceae). The Eucalyptus homolog of a CONSTANS‐like gene was an F_ST outlier, and allelic variation showed significant latitudinal clinal associations with annual and winter solar radiation, potential evaporation, summer precipitation and aridity. A second F_ST outlier locus, homologous to quinone oxidoreductase, was significantly associated with measures of temperature range, high summer temperature and summer solar radiation, with important implications for predicting the effect of temperature on natural populations in the context of climate change. We complemented these data with investigations into neutral population genetic structure and diversity throughout the species range. This study provides an investigation into selection signatures at gene‐homologous EST‐SSRs in natural Eucalyptus populations, and contributes to our understanding of the relationship between climate and adaptive genetic variation, informing the conservation of both putatively neutral and adaptive components of genetic diversity.     

Contribution of the UV component of natural sunlight to photoinhibition of photosynthesis in six species of subtidal brown and red seaweeds

Field‐collected specimens of three species of Laminaria and three species of subtidal red algae (Delesseria sanguinea, Plocamium cartilagineum and Phyllophora pseudoceranoides) were exposed to natural summer sunlight on Helgoland (southern North Sea) for up to 4 h at 15 °C. Dark‐adapted variable fluorescence (F_v : F_m) was measured immediately after these treatments, and following 6, 24 and 48 h of recovery in moderate irradiances of white light. The response of plants to the full spectrum of natural sunlight was compared with that to PAR alone, UV‐A + visible, UV‐A + UV‐B, or UV‐A alone. The F_v : F_m values of all species were reduced to minimal values after 4 h in all of these treatments, but those of the more resistant species (Laminaria spp. and P. pseudoceranoides) were higher after shorter exposures to UV radiation alone than to PAR with or without UV. The recovery of F_v : F_m in all species was also more rapid in the two treatments that contained UV radiation alone than in those that included PAR. These results suggest that it is the high irradiances of PAR in natural sunlight which are responsible for the photoinhibition of photosynthesis of subtidal seaweeds and that the current ambient irradiances of UV radiation (either UV‐B or UV‐A) in northern temperate latitudes would not contribute significantly to this photoinhibition.     

Evidence that UV-B tolerance of the photosynthetic apparatus in microalgae is related to the D1-turnover mediated repair cycle in vivo

The present study was conceived to elucidate the potential importance of the D1 turnover-mediated repair mechanism in UV-B tolerance of the photosynthetic apparatus in microalgae. To this end, the lab-identified UV-B sensitive and tolerant species of Chlorophyte and Chromophyte algae was used to examine photosynthetic response to UV-B exposure in the presence vs. the absence of streptomycin, an inhibitor of chloroplast protein synthesis. Measurements of photosynthetic O₂ evolution capacity and chlorophyll fluorescence parameters (F_v/F_m, Φ_PSII) illustrated species-specific UV-B sensitivity of the photosynthetic apparatus. Addition of the inhibitor streptomycin caused significant enhancements of UV-B-caused depression of photosynthesis in UV-B tolerant species, while little effect was observed in the sensitive species. In the tolerant species, recovery from UV-B induced 20 percnt; decline in F_v/F_m reached completion within 2 hours, much faster than that in the sensitive species. Immunoblotting revealed that exposure to UV-B radiation caused substantial degradation of the D1 protein in the sensitive Heterococcus brevicellularis, which was little enhanced by addition of the inhibitor. The same UV-B exposure lead to less D1 degradation in the tolerant Scenedesmus sp., which was significantly enhanced by addition of the inhibitor. This study shows that UV-B tolerance of the photosynthetic apparatus in microalgae was associated with a strong capacity for recovery from the UV-B-induced damage and this capacity related to the D1 turnover-mediated repair cycle, and largely determined UV-B tolerance of the photosynthetic apparatus in these organisms.