EFFECTS OF INCREASING UV‐B RADIATION DUE TO OZONE DEPLETION ON PHOTOSYNTHESIS OF ANTARCTIC CYANOBACTERIA

Ground level ultraviolet‐B (UV‐B; 290–320 nm) fluxes in Antarctica have been increasing due to stratospheric ozone depletion. Although mat‐forming cyanobacteria are major component of freshwater algal biomass in Antarctica, little is known about their response to increasing ultraviolet radiation (UVR). The present study evaluated the sensitivity to UVR of two strains of mat‐forming cyanobacteria with different cell size, Phormidium murrayi (6.0 x 3.2 μm) and Schizothrix calcicola (2.2 x 2.3 μm). Cyanobacterial photosynthesis was measured under different UV spectral quality and quantity achieved by polychromatic filters with different cutoff wavelengths and neutral density screens. The productivity and irradiance data were used to generate biological weighting functions (BWF) for the assessment of UV inhibition on photosynthesis. The kinetics of UV inhibition, as determined by PAM fluorometry, differed between the two species so that inhibition of P. murrayi and S. calcicola were modeled based on UV‐irradiance and cumulative exposure, respectively. After a one hour exposure, BWF's did not differ between the two isolates of cyanobacteria despite their differences in cell size. To evaluate the negative impact of increased UV‐B exposure due to ozone depletion on cyanobacteria, the BWF's were applied to two solar spectra obtained from McMurdo Station, one on a day when the ozone hole was prominent (O₃ = 170 Dobson units; DU = 10‐3 cm O₃), and the other on a day with high ozone concentration (O₃ = 328 DU). The decrease in ozone level would reduce productivity by 3–8%. Seasonal variation of UVR has a bigger impact on cyanobacterial productivity than ozone depletion.     

Inhibition by ultraviolet and photosynthetically available radiation lowers model estimates of depth‐integrated picophytoplankton photosynthesis: global predictions for Prochlorococcus and Synechococcus

Phytoplankton photosynthesis is often inhibited by ultraviolet (UV) and intense photosynthetically available radiation (PAR), but the effects on ocean productivity have received little consideration aside from polar areas subject to periodic enhanced UV‐B due to depletion of stratospheric ozone. A more comprehensive assessment is important for understanding the contribution of phytoplankton production to the global carbon budget, present and future. Here, we consider responses in the temperate and tropical mid‐ocean regions typically dominated by picophytoplankton including the prokaryotic lineages, Prochlorococcus and Synechococcus. Spectral models of photosynthetic response for each lineage were constructed using model strains cultured at different growth irradiances and temperatures. In the model, inhibition becomes more severe once exposure exceeds a threshold (E_max) related to repair capacity. Model parameters are presented for Prochlorococcus adding to those previously presented for Synechococcus. The models were applied to estimate midday, water column photosynthesis based on an atmospheric model of spectral radiation, satellite‐derived spectral water transparency and temperature. Based on a global survey of inhibitory exposure severity, a full‐latitude section of the mid‐Pacific and near‐equatorial region of the east Pacific were identified as representative regions for prediction of responses over the entire water column. Comparing predictions integrated over the water column including versus excluding inhibition, production was 7–28% lower due to inhibition depending on strain and site conditions. Inhibition was consistently greater for Prochlorococcus compared to two strains of Synechococcus. Considering only the surface mixed layer, production was inhibited 7–73%. On average, including inhibition lowered estimates of midday productivity around 20% for the modeled region of the Pacific with UV accounting for two‐thirds of the reduction. In contrast, most other productivity models either ignore inhibition or only include PAR inhibition. Incorporation of E_max model responses into an existing spectral model of depth‐integrated, daily production will enable efficient global predictions of picophytoplankton productivity including inhibition.     

Bacterial cyclic β‐(1,2)‐glucans sequester iron to protect against iron‐induced toxicity

Cellular iron homeostasis is critical for survival and growth. Bacteria employ a variety of strategies to sequester iron from the environment and to store intracellular iron surplus that can be utilized in iron‐restricted conditions while also limiting the potential for the production of iron‐induced reactive oxygen species (ROS). Here, we report that membrane‐derived oligosaccharide (mdo) glucan, an intrinsic component of Gram‐negative bacteria, sequesters the ferrous form of iron. Iron‐binding, uptake, and localization experiments indicated that both secreted and periplasmic β‐(1,2) ‐ glucans bind iron specifically and promote growth under iron‐restricted conditions. Xanthomonas campestris and Escherichia coli mutants blocked in the production of β‐(1,2) ‐ glucan accumulate low amounts of intracellular iron under iron‐restricted conditions, whereas they exhibit elevated ROS production and sensitivity under iron‐replete conditions. Our results reveal a critical role of glucan in intracellular iron homeostasis conserved in Gram‐negative bacteria.     

Size‐spectrum based differential response of phytoplankton to nutrient and iron‐organic matter combinations in microcosm experiments in a Chilean Patagonian Fjord

The Patagonian fjords have been recognized as a major region of relatively high primary productivity systems during spring–summer bloom periods, where iron‐organic matter forms may be essential complexes involved in key growth processes connected to the carbon and nitrogen cycles. We used two dissolved organic matter (DOM) types, marine polysaccharide and siderophore, as a model to understand how they affect the bioavailability of Fe to phytoplankton and bacteria and to assess their ecological role in fjord systems. A 10‐day microcosm study was performed in the Comau Fjord during summer conditions (March 2012). Pico‐, nano‐, and microphytoplankton abundance, total chlorophyll‐a and bacteria abundance, and bacterial secondary production estimates were analyzed in five treatments: (i) control (no additions), (ii) only nutrients (NUT: PO₄, NO₃, Si), (iii) nutrients + Fe(II), (iv) polysaccharide (natural diatoms extracted: 1–3 beta Glucan), and (v) Hexandentate Desferroxiamine B (DFB, siderophore). Our results showed that while DFB reduced Fe bioavailability for almost all phytoplankton assemblages in the fjord, polysaccharide did not have effects on the iron bioavailability. At Nutrients + Fe and Polysaccharide treatments, chlorophyll‐a concentration abruptly increased from 0.9 to 20 mg m^?3 during the first 4–6 days of the experimental period. Remarkably, at the Nutrients + Fe treatment, the development of the bloom was accompanied by markedly high abundances of Synechococcus, picoeukaryotes, and autotrophic nanoflagellates within the first 4 days of the experiment. Our study indicated that small plankton (phytoplankton <20 μm and bacteria) were the first to respond to dissolved Nutrients + Fe compared to large sized micro‐phytoplankton cells (>20 μm). This could be at least partially attributed to biological utilization of Fe (2 to 3 nM) by <20 μm phytoplankton and bacteria through the interaction with organic ligands released by bacteria that eventually could increase solubility of the Fe dissolved fraction thus having a positive effect on the small‐sized phytoplankton community.     

Evidence for iron‐regulated cyanobacterial predominance in oligotrophic lakes

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Increased risk of cyanobacterial blooms in northern high‐latitude lakes through climate warming and phosphorus enrichment

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SIDEROPHORE‐INDEPENDENT IRON UPTAKE BY IRON‐LIMITED CELLS OF THE CYANOBACTERIUM ANABAENA FLOS‐AQUAE1

Iron acquisition by iron‐limited cyanobacteria is typically considered to be mediated mainly by siderophores, iron‐chelating molecules released by iron‐limited cyanobacteria into the environment. In this set of experiments, iron uptake by iron‐limited cells of the cyanobacterium Anabaena flos‐aquae (L.) Bory was investigated in cells resuspended in siderophore‐free medium. Removal of siderophores decreased iron‐uptake rates by ～60% compared to siderophore‐replete conditions; however, substantial rates of iron uptake remained. In the absence of siderophores, Fe(III) uptake was much more rapid from a weaker synthetic chelator [N‐(2‐hydroxyethyl)ethylenediamine‐N,N′,N′‐triacetic acid (HEDTA); log K_cond = 28.64 for Fe(III)HEDTA(OH)^?] than from a very strong chelator [N,N′‐bis(2‐hydroxybenzyl)‐ethylenediamine‐N,N′‐diacetic acid (HBED); log K_cond = 31.40 for Fe(III)HBED^?], and increasing chelator:Fe(III) ratios decreased the Fe(III)‐uptake rate; these results were evident in both short‐term (4 h; absence of siderophores) and long‐term (116 h; presence of siderophores) experiments. However, free (nonchelated) Fe(III) provided the most rapid iron uptake in siderophore‐free conditions. The results of the short‐term experiments are consistent with an Fe(III)‐binding/uptake mechanism associated with the cyanobacterial outer membrane that operates independently of extracellular siderophores. Iron uptake was inhibited by temperature‐shock treatments of the cells and by metabolically compromising the cells with diphenyleneiodonium; this finding indicates that the process is dependent on active metabolism to operate and is not simply a passive Fe(III)‐binding mechanism. Overall, these results point to an important, siderophore‐independent iron‐acquisition mechanism by iron‐limited cyanobacterial cells.     

Nitric oxide is involved in integration of UV‐B absorbing compounds among parts of clonal plants under a heterogeneous UV‐B environment

In nature, ultraviolet‐B (UV‐B) radiation is highly heterogeneous, both spatially and temporally. Plants exposed to UV‐B radiation produce UV‐B absorbing compounds that function as a protective filter. For clonal plants under heterogeneous UV‐B radiation conditions, integration among ramets can allow irradiated ramets to benefit un‐irradiated ramets by causing them to increase their UV‐B absorbing compounds content. In this study, we evaluated integration between pairs of clonal ramets of Glechoma longituba under heterogeneous or homogeneous UV‐B conditions. We determined the levels of UV‐B absorbing compounds, nitric oxide (NO) and hydrogen peroxide (H₂O₂) and measured the activity of phenylalanine ammonia‐lyase (PAL) in connected ramet pairs under homogeneous or heterogeneous UV‐B conditions. Under heterogeneous UV‐B conditions, the UV‐B absorbing compounds content increased in leaves of irradiated and un‐irradiated ramets, but not in the connecting stolons. The NO content increased in irradiated and un‐irradiated leaves and stolons, but the H₂O₂ content did not. Application of NO synthesis inhibitors and an NO blocker to irradiated ramets blocked the increase in UV‐B absorbing compounds and PAL activity in un‐irradiated ramets. These results suggested that NO is involved in the integration process for UV‐B absorbing compounds among ramets. Our findings suggested that a UV‐B‐induced increase in NO transmits a signal to un‐irradiated ramets via the stolon, leading to an increase in PAL activity and UV‐B absorbing compounds content. The internal translocation of signal enables members of clonal networks to function as a whole unit and to mount an efficient defensive response to localized UV‐B radiation.     

Differences in growth and physiological traits of two poplars originating from different altitudes as affected by UV‐B radiation and nutrient availability

Cuttings of Populus kangdingensis and Populus cathayana originating from altitudes of 3500 and 1500 m in southwestern China, respectively, were grown for one growing season in the field under ambient or ambient plus supplemental ultraviolet‐B (UV‐B) radiation with two levels of nutrients. In both species, enhanced UV‐B radiation significantly increased UV‐B absorbing compounds and guaiacol peroxidase (GPX) activity, while no significant effects were observed in photosynthetic pigments and proline content. On the other hand, cuttings grown with high‐nutrient availability had larger leaf area, higher total biomass and GPX activity as well as higher water use efficiency (WUE) (as measured by stable carbon isotope composition, δ¹³C) when compared with low‐nutrient conditions, while UV‐B absorbing compounds and ascorbic acid (AsA) content significantly decreased. Differences in responses to enhanced UV‐B radiation and nutrient availability were observed between the two species. Nutrient‐induced increases in chlorophyll a, chlorophyll b and total chlorophyll as well as in carotenoids were greater in P. kangdingensis than in P. cathayana. In P. cathayana, enhanced UV‐B radiation significantly decreased leaf area and total biomass, while it significantly increased WUE and ascorbate peroxidase (APX). In contrast, such changes were not observed in P. kangdingensis. In addition, the effects of enhanced UV‐B radiation on leaf area, total biomass and UV‐B absorbing compounds were closely related to the nutrient status. Our results indicated that P. kangdingensis, which originates from the altitude of 3500 m and is apparently adapted to low‐nutrient and high‐UV‐B habitats, exhibits better tolerance to enhanced UV‐B radiation and greater growth under low‐nutrient availability than does P. cathayana originating from the altitude of 1500 m.     

Responses to UV-B radiation in Trifolium repens L. – physiological links to plant productivity and water availability

This study used comparisons across nine populations of Trifolium repens (white clover) in conjunction with drought to examine physiological responses to ultraviolet‐B radiation (UV‐B). Plants were exposed for 12 weeks to supplementation with 13.3 kJ m^?2 d^?1 UV‐B, accompanied by 4 weeks of drought under controlled environmental conditions. UV‐B increased the levels of UV‐B‐absorbing compounds and of flavonol glycosides and this effect was synergistically enhanced by water stress. These changes were more pronounced for the ortho‐dihydroxylated quercetin, rather than the monohydroxylated kaempferol glycosides. UV‐B increased leaf water potential (ψ_L) by 16% under drought and proline levels by 23% under well‐watered conditions. The intraspecific comparisons showed that higher UV‐B‐induced levels of UV‐B‐absorbing compounds, of quercetin glycosides and of ψ_L were linked to lower plant productivity and to higher UV‐B tolerance under well‐watered conditions. These findings suggest that: (1) slow‐growing T. repens ecotypes adapted to other stresses have higher capacity for physiological acclimation to UV‐B; and (2) that these attributes also contribute to decreased UV‐B sensitivity under drought.     

Multi‐wavelength Pulse Amplitude Modulated fluorometry (Phyto‐PAM) reveals differential effects of ultraviolet radiation on the photosynthetic physiology of phytoplankton pigment groups

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Cyanobacterial dynamics in shallow Lake Apopka (Florida,U.S.A.) before and after the shift from a macrophyte‐dominated to a phytoplankton‐dominated state

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Responses of photosynthetic properties and chloroplast ultrastructure of Bryum argenteum from a desert biological soil crust to elevated ultraviolet‐B radiation

Our understanding of plant responses to enhanced ultraviolet‐B (UV‐B) radiation has improved over recent decades. However, research on cryptogams is scarce and it remains controversial whether UV‐B radiation causes changes in physiology related to photosynthesis. To investigate the effects of supplementary UV‐B radiation on photosynthesis and chloroplast ultrastructure in Bryum argenteum Hedw., specimens were cultured for 10 days under four UV‐B treatments (2.75, 3.08, 3.25 and 3.41 W m^–2), simulating depletion of 0% (control), 6%, 9% and 12% of stratospheric ozone at the latitude of Shapotou, a temperate desert area of northwest China. Analyses showed malondialdehyde content significantly increased, whereas chlorophyll (Chl) fluorescence parameters and Chl contents decreased with increased UV‐B intensity. These results corresponded with changes in thylakoid protein complexes and chloroplast ultrastructure. Overall, enhanced UV‐B radiation leads to significant decreases in photosynthetic function and serious destruction of the chloroplast ultrastructure of B. argenteum. The degree of negative influences increased with the intensity of UV‐B radiation. These results may not only provide a potential mechanism for supplemental UV‐B effects on photosynthesis of moss crust, but also establish a theoretical basis for further studies of adaptation and response mechanisms of desert ecosystems under future ozone depletion.     

Solar UV‐B radiation and ethylene play a key role in modulating effective defenses against Anticarsia gemmatalis larvae in field‐grown soybean

Solar UV‐B radiation has been reported to enhance plant defenses against herbivore insects in many species. However, the mechanism and traits involved in the UV‐B mediated increment of plant resistance are unknown in crops species, such as soybean. Here, we studied defense‐related responses in undamaged and Anticarsia gemmatalis larvae‐damaged leaves of two soybean cultivars grown under attenuated or full solar UV‐B radiation. We determined changes in jasmonates, ethylene (ET), salicylic acid, trypsin protease inhibitor activity, flavonoids, and mRNA expression of genes related with defenses. ET emission induced by Anticarsia gemmatalis damage was synergistically increased in plants grown under solar UV‐B radiation and was positively correlated with malonyl genistin concentration, trypsin proteinase inhibitor activity and expression of IFS2, and the pathogenesis protein PR2, while was negatively correlated with leaf consumption. The precursor of ET, aminocyclopropane‐carboxylic acid, applied exogenously to soybean was sufficient to strongly induce leaf isoflavonoids. Our results showed that in field‐grown soybean isoflavonoids were regulated by both herbivory and solar UV‐B inducible ET, whereas flavonols were regulated by solar UV‐B radiation only and not by herbivory or ET. Our study suggests that, although ET can modulate UV‐B‐mediated priming of inducible plant defenses, some plant defenses, such as isoflavonoids, are regulated by ET alone.     

UV‐radiation and elevated temperatures induce formation of reactive oxygen species in gametophytes of cold‐temperate/Arctic kelps (Laminariales,Phaeophyceae)

Enhanced UV‐radiation (UVR) through stratospheric ozone depletion and global warming are crucial stressors to marine macroalgae. Damages may arise through formation of reactive oxygen species (ROS) in gametophytes of ecologically important kelps, brown algae of the order Laminariales, Such stress‐induced damages may have a negative impact on their fitness and further impact their following life stages. In our study, gametophytes of three kelp species Alaria esculenta (L.) Grev., Laminaria digitata (Huds.) Lamour., Saccharina latissima (L.) Lane, Mayes, Druehl, Saunders from the Arctic, and of L. hyperborea (Gunnerus) Foslie from the North Sea were exposed to photosynthetically active radiation, UV‐A, and UV‐B radiation and four temperatures (2–18°C). ROS are formed predominantly in the peripheral cytoplasm and in chloroplasts especially after exposure to UVR. Superoxide (O₂*^‐) is additionally formed in small, globular cytoplasmic structures, possibly mitochondria. In the surrounding medium O₂*^‐‐concentration increased markedly at elevated temperatures and under UV stress in some cases. Ultrastructural damage was negligible pointing to a high stress tolerance of this developmental stage. Our data indicate that stress tolerant gametophytes of three Arctic kelp species should sustain their crucial function as seed bank for kelp populations even under prospective rising environmental perturbations.     

Sexual differences in defensive and protective mechanisms of Populus cathayana exposed to high UV‐B radiation and low soil nutrient status

Previous studies have indicated that Populus cathayana Rehder females are more sensitive and less tolerant to stressful environments than males, but it is still unknown whether there are sexual differences in defensive and protective traits under high UV‐B (HUVB) radiation and low soil nutrient status. In this study, P. cathayana was employed as a model species to investigate sex‐related physiological and biochemical responses to UV‐B radiation under different soil nutrient conditions. Cuttings were exposed to two UV‐B radiation regimes (ambient UV‐B radiation and decreased UV‐B radiation) under two soil nutrient status (topsoil and deep soil) for 100 days over one growing season. Both HUVB radiation and low soil nutrient status induced greater decreases in plant growth, dry mass accumulation, gas exchange and leaf nitrogen use efficiency in females than in males, and greater increases in lipid peroxide and antioxidant enzyme activities, and secondary defense capacities in males than in females. Moreover, sexually different responses happened also in organelle ultrastructure. Our results showed that: (1) females suffered greater negative effects and exhibited lower defense capacities than did males under HUVB radiation, low soil nutrient status and their combination; (2) low soil nutrient status reduced plant's sensitivity to HUVB radiation by increasing allocation to defense and decreasing allocation to growth compared with high soil nutrient status. These results provide evidence that sexually different tradeoffs happen between growth and defense in P. cathayana under HUVB radiation and low soil nutrient status.     

Characterization of two Synechococcus sp. PCC7002‐related cyanobacterial strains in relation to 16S rDNA,crtR gene,lipids and pigments

Two Synechococcus strains from the Culture Collection of the Institute for Marine Sciences of Andalusia (Cádiz, Spain), namely Syn01 and Syn02, were found to be closely related to the model strain Synechococcus sp. PCC7002 according to 16S rDNA (99% identity). Pigment and lipid profiles and crtR genes of these strains were ascertained and compared. The sequences of the crtR genes of these strains were constituted by 888 bp, and showed 99% identity between Syn01 and Syn02, and 94% identity of Syn01 and Syn02 to Synechococcus sp. PCC7002. There was coincidence in photosynthetic pigments between the three strains apart from the pigment synechoxanthin, which could be only observed in Synechococcus sp. PCC7002. Species of sulfoquinovosyl‐diacyl‐glycerol (SQDG), phosphatidyl‐glycerol (PG), mono‐ and di‐galactosyl‐diacyl‐glycerol (MGDG and DGDG) were detected by high performance liquid chromatography‐mass spectrometry analysis of lipid extracts. The most abundant species within each lipid class were those containing C18:3 together with C16:0 fatty acyl substituents in the glycerol backbone of the same molecule. From these results it is concluded that these cyanobacterial strains belong to group 2 of the lipid classification of cyanobacteria.     

Effects of iron on the growth and minimal fluorescence yield of three marine Synechococcus strains (Cyanophyceae)

Two coastal Synechococcus stains PCC 7002 and CC9311 and one oceanic strain WH8102 were cultured with 4–1000 nM Fe in Aquil medium. Compared with those under iron‐replete conditions, their growth rates were significantly decreased by 59% for WH8102 at 15 nM Fe, by 37% for CC9311 at 15 nM Fe and by 57% for PCC 7002 at 4 nM Fe. Among these three strains, PCC 7002 was the most tolerant to iron limitation while WH8102 was the most sensitive to iron limitation. For each strain under the same iron concentration, the growth rates calculated from the minimal fluorescence yield and cell concentration showed no significant difference. The linear correlation was established between the minimal fluorescence yield and cell concentration although the minimal fluorescence yield per cell varied depending on the strains and iron levels. Under iron‐replete conditions, the minimal fluorescence yield per cell was 100‐fold higher for the phycoerythrin‐lacking strain PCC 7002 than two phycoerythrin‐containing strains WH8102 and CC9311. Under iron‐deplete conditions, it was increased respectively by 128% and 7% for WH8102 and CC9311 but was decreased by 30% for PCC 7002. Furthermore, the minimal fluorescence yield per cell for PCC 7002 and CC9311 showed little difference throughout the light and dark diel cycle. However, it was significantly higher for WH8102 in the daytime than in the dark.