Inclusion of ionizing radiation in a mathematical model for photosynthesis.

Ionizing radiation of astrophysical origin might have played an important role in biological evolution during the long course of Earth's evolution. Several phenomena might have induced intense fluctuations in background ionizing radiation, such as highly energetic stellar explosions. There might also be anthropogenic causes for environmental radiation fluctuations, resulting from nuclear industry activities. The inclusion of these effects in a mathematical model for photosynthesis provides a useful tool to account for the damages of the above-mentioned phenomena in vegetal life. Mathematical models for photosynthesis typically only consider ultraviolet radiation and photosynthetically active radiation, as they have been a ubiquitous physical factor in the settlement of vegetal life. In this work a mathematical model for aquatic photosynthesis is modified, from first principles, to include the action of particulate ionizing radiation on the photosynthetic process. After assuming an ansatz allowing to separate damage/repair kinetics of ultraviolet and ionizing radiations, a treatable mathematical expression of the model is obtained. This generalized model is presented as a function of radiometric and photometric magnitudes, making it prone to calibration and useful to apply to aquatic ecosystems under radiational stress due to gamma-ray bursts, cosmic ray bursts, solar storms, or other sources of ionizing radiations.

Direct Effects of Temperature on Growth of Different Tropical Phytoplankton Species.

Temperature increase may influence competition among phytoplankton species, potentially intensifying cyanobacteria blooms that can be favored by direct and indirect effects of temperature. In this study, we aimed to clarify how cyanobacteria can be favored by the direct effects of increased temperature compared to diatoms and chlorophytes. Strains of the most representative species of a eutrophic coastal lagoon (Microcystis aeruginosa, Planktothrix agardhii, Desmodesmus communis, and Cyclotella meneghiniana) were used to test the hypothesis that cyanobacteria would be favored by the direct effect of temperature increase. First, we evaluated the effect of temperature increase on growth in monocultures (batch and chemostats) at 25 and 30 °C and after in mixed cultures (chemostats). In batch monocultures, the cyanobacteria showed higher growth rates in 30 °C than in 25 °C. However, in continuous culture experiments (chemostats), growth rates of M. aeruginosa and P. agardhii were not affected by temperature, but the strains showed higher biovolume in steady-state with the temperature increase. In continuous mixed cultures, M. aeruginosa was always dominant and C. meneghiniana was excluded, regardless of temperature tested. D. communis was able to coexist with lower biomass. This study shows that rising temperatures can be detrimental to diatoms, even for a tropical strain. Although some studies indicate that the dominance of cyanobacteria in warmer climates may be due to the indirect effect of warming that will promote physical conditions in the environment more favorable to cyanobacteria, the outcomes of mixed cultures demonstrate that the direct effect of temperature can also favor the dominance of cyanobacteria.

Partitioning of the diffuse attenuation coefficient for photosynthetically available irradiance in a deep dendritic tropical lake.

We studied the effects of particulate and dissolved optically active components on the attenuation of photosynthetic active radiation (PAR) in a tropical lake. The temporal and spatial distribution of tripton, Chl-a and aCDOM(440) and their relative contribution to the diffuse PAR attenuation coefficient (Kd) was investigated at 21 sites (dry and wet seasons and two intermediate periods) and at monthly interval at 1 pelagic site. Higher values of ​​ Kd were observed during the mixing period, characterized by a higher concentration of tripton and Chl-a compared to the stratified rainy season. In the spatial sampling PAR attenuation was dominated by tripton absorption/scattering (average relative contribution of 79%), followed by Chl-a (average 11.6%). In the monthly sampling tripton and Chl-a accounted for most of the Kd with relative contributions of 47.8% and 35.6%, respectively. Multiple linear regression analysis showed that Chl-a and tripton in combination explained 97% of the monthly variation in Kd (p<0.001), but Chl-a had more influence (higher regression coefficient). Thus, although most of light attenuation was due to tripton, seasonal variations in phytoplankton abundance were responsible for most of the temporal fluctuations in Kd.

Partitioning of the diffuse attenuation coefficient for photosynthetically available irradiance in a deep dendritic tropical lake

ABSTRACT We studied the effects of particulate and dissolved optically active components on the attenuation of photosynthetic active radiation (PAR) in a tropical lake. The temporal and spatial distribution of tripton, Chl-a and aCDOM(440) and their relative contribution to the diffuse PAR attenuation coefficient (Kd) was investigated at 21 sites (dry and wet seasons and two intermediate periods) and at monthly interval at 1 pelagic site. Higher values of ​​ Kd were observed during the mixing period, characterized by a higher concentration of tripton and Chl-a compared to the stratified rainy season. In the spatial sampling PAR attenuation was dominated by tripton absorption/scattering (average relative contribution of 79%), followed by Chl-a (average 11.6%). In the monthly sampling tripton and Chl-a accounted for most of the Kd with relative contributions of 47.8% and 35.6%, respectively. Multiple linear regression analysis showed that Chl-a and tripton in combination explained 97% of the monthly variation in Kd (p<0.001), but Chl-a had more influence (higher regression coefficient). Thus, although most of light attenuation was due to tripton, seasonal variations in phytoplankton abundance were responsible for most of the temporal fluctuations in Kd.

Assessment of the Effects of Light Availability on Growth and Competition Between Strains of Planktothrix agardhii and Microcystis aeruginosa.

In this study, we tested the hypothesis that Planktothrix agardhii strains isolated from a tropical water body were better competitors for light than Microcystis aeruginosa strains. These cyanobacteria are common in eutrophic systems, where light is one of the main drivers of phytoplankton, and Planktothrix is considered more shade-adapted and Microcystis more high-light tolerant. First, the effect of light intensities on growth was studied in batch cultures. Next, the minimum requirement of light (I*) and the effect of light limitation on the outcome of competition was investigated in chemostats. All strains showed similar growth at 10 µmol photons m(-2) s(-1), demonstrating the ability of the two species to grow in low light. The optimum light intensity was lower for P. agardhii, but at the highest light intensity, Microcystis strains reached higher biovolume, confirming that P. agardhii has higher sensitivity to high light. Nonetheless, P. agardhii grew in light intensities considered high (500 µmol photons m(-2) s(-1)) for this species. M. aeruginosa showed a higher carrying capacity in light-limited condition, but I* was similar between all the strains. Under light competition, Microcystis strains displaced P. agardhii and dominated. In two cases, there was competitive exclusion and in the other two P. agardhii managed to remain in the system with a low biovolume (≈15%). Our findings not only show that strains of P. agardhii can grow under higher light intensities than generally assumed but also that strains of M. aeruginosa are better competitors for light than supposed. These results help to understand the co-occurrence of these species in tropical environments and the dominance of M. aeruginosa even in low-light conditions.

UV radiation simultaneously affects phototrophy and phagotrophy in nanoflagellate-dominated phytoplankton from an Andean shallow lake.

Mixotrophic nanoflagellates, that combine photosynthesis and phagotrophy, are important members of planktonic food webs in many aquatic environments depending on the balance among the different carbon and energy sources. We carried out field sampling and laboratory experiments with natural nanoflagellate assemblages from an Andean North-Patagonian lake exposing them or not to UVR, and measuring photosynthetic parameters and bacterivory. The effect of different light treatments on the photosynthetic efficiency was studied by the non-invasive, pulse amplitude-modulated (PAM) fluorescence technique, and bacterivory was assessed with fluorescently labeled bacteria (FLB). Mixotrophic nanoflagellates were clearly dominant (up to 90% of total phytoplankton and 88% of total nanoflagellate abundance), and in the experiments labeled bacteria were observed in more than 75% of mixotrophic cells. These results support the idea that these phytoflagellates were never entirely photosynthetic. The high light : phosphorus ratio and the high C : N : P ratio suggest a strong nutrient limitation towards P. Our results show that both functions, photosynthesis and bacteria ingestion, were simultaneously reduced by the same level of UVR. We estimated that UVR exposure of mixotrophic nanoflagellates reduced photosystem II activity between 23% and 31% while ingestion rates were reduced between 23% and 28%. Therefore, our results suggest that the different cell functions could be concurrently impacted by UVR, implying that patterns and rates of C transfer would be substantially altered in the microbial food web.

Quality of UVR exposure for different biological systems along a latitudinal gradient.

The exposure of organisms to ultraviolet radiation (UVR) is characterized by the climatology (annual cycle) and the variance (anomalies) of biologically-weighted irradiances at eight geographical locations in austral South America, from 1995-2002. The net effect of UVR on biological systems is a result of the balance of damage and repair which depends on intensity and duration of irradiance and is modulated by its variability. The emphasis in this study is on day-to-day variability, a time scale of importance to adaptive strategies that counteract UVR damage. The irradiances were weighted with DNA- and phytoplankton photosynthesis-action spectra. Low latitude sites show high average UVR. For all sites, the frequency of days with above average irradiances is higher than below average irradiances. Persistence in anomalies is generally low (

Combined effects of solar ultraviolet radiation and nutrients addition on growth, biomass and taxonomic composition of coastal marine phytoplankton communities of Patagonia.

Experiments (6-8 days) were conducted during late summer, late fall and late winter, 2003 with waters collected off Bahía Nueva, Chubut, Argentina (42.7 degrees S, 65 degrees W) to determine the combined effects of solar ultraviolet radiation (UVR, 280-400 nm) and nutrient addition on phytoplankton communities. Samples were put in UVR-transparent containers and incubated under two radiation treatments: (a) Samples exposed to full solar radiation (PAB treatment, 280-400 nm) and (b) Samples exposed only to PAR (PAR treatment, 400-700 nm). At the beginning of the experiments, nutrients (i.e., NaPO(4)H(2) and NaNO(3)) were added to one set of samples from each radiation treatment (N cultures) whereas in the other set, nutrients remained at the concentration of the seawater. Chlorophyll a, biomass, UV-absorbing compounds and taxonomic composition were recorded throughout the experiments. N cultures always had significantly higher growth rates (P<0.05) than that in non-enriched cultures. At the beginning of experiments, phytoplankton communities were generally dominated by monads and flagellates but by the end, diatoms comprised the bulk of biomass, with only one to four taxa dominating, suggesting a selection towards more tolerant/less sensitive species. Over the experimental time frame, the observed taxonomic changes were mostly due to nutrient availability, and to a lesser extent to solar UVR exposure. Overall, the results indicate that environmental conditions (i.e., light history, nutrient concentration) together with the physiological status of the cells play a very important role at the time to assess the combined effect of nutrient addition and solar radiation on coastal phytoplankton assemblages from Patagonia.

Interactive effects of ultraviolet radiation and nutrient addition on growth and photosynthesis performance of four species of marine phytoplankton.

Experiments (6-8 days) were carried out during the austral summer of 2005 in Chubut, Argentina (43 degrees S, 65 degrees W) to determine the interactive effects of solar UVR (280-400 nm) and nutrient addition on growth and chlorophyll fluorescence of four species of marine phytoplankton--the diatoms Thalassiosira fluviatilis Hustedt and Chaetoceros gracilis Schütt, and the dinoflagellates Heterocapsa triquetra (Ehrenberg) Stein and Prorocentrum micans (Ehrenberg). Samples were incubated under three radiation treatments (two sets of each radiation treatment): (a) samples exposed to full solar radiation (PAR+UVR, PAB treatment, 280-700 nm); (b) samples exposed to PAR and UV-A (PA treatment, 320-700 nm) and (c) samples exposed only to PAR (P treatment, 400-700 nm). At the beginning of the experiments, nutrients (i.e., NaPO(4)H(2) and NaNO(3)) were added to one set of samples from each radiation treatment ("N" cultures) whereas in the other, the nutrients concentration was that of the culture medium. At all times, the lowest growth rates (mu) were determined in the PAB treatments, where enriched cultures had significantly higher mu (P<0.05) than non-enriched cultures. Daily cycles of photochemical quantum yield (Y) displayed a pattern of relatively high values early in the morning with a sharp decrease at noon; recovery was observed late in the afternoon. In general, higher Y values were determined in enriched cultures than in non-enriched cultures. As the experiments progressed, acclimation (estimated as the difference between Y at noon and that at time zero) was observed in all species although in variable degree. All species displayed some degree of UVR-induced decrease in the photochemical quantum yield, although it was variable among treatments and species. However, this effect decreased with time, and this pattern was more evident in the dinoflagellates, as the concentration of UV-absorbing compounds increased. Thus, under conditions of nutrient enrichment as may occur by river input or by re-suspension by mixing, dinoflagellates outcompete with diatoms because they may have a higher fitness under UVR stress.

Photoacclimation to long-term ultraviolet radiation exposure of natural sub-Antarctic phytoplankton communities: Fixed surface incubations versus mixed mesocosms.

Solar UVB radiation (280-320 nm) is known to have detrimental effects on marine phytoplankton. Associated with the seasonal ozone hole in Antarctica, stratospheric ozone depletion occasionally influences the sub-Antarctic (Beagle Channel, Argentina) region, enhancing levels of UVB. The primary objective of this work was to study the effects of several (i.e. 6-10) days of exposure to UVB on the taxonomic composition and photosynthetic inhibition of local phytoplankton communities. For different light treatments, fixed-depth incubations placed in an outdoors water tank were compared with incubations in 1900 L mesocosms, where vertical mixing was present. Phytoplankton growth was inhibited by UV radiation (UVR) in fixed-depth experiments but not in the mixed mesocosms. Under fixed and mixed conditions alike, photosynthesis was significantly inhibited by UVB at the beginning of the experiment but no longer after several days of exposure, suggesting that cells had acclimated to radiation conditions. There was a change in species composition in response to UVR exposure in both experiments, which likely explained acclimation. In the community exposed to fixed conditions this change was from a phytoflagellate-dominated assemblage to a community with high relative abundance of diatoms after 6 days of exposure. UVA was responsible for most of the observed growth inhibition; however, the reduction in photosynthesis was produced by UVB. The reasons behind this variability in responses to UVR are associated with species-specific sensitivity and acclimation, and the previous light history of cells. In the community exposed in mesocosms, an assemblage codominated by phytoflagellates and diatoms was observed at the beginning of the experiments. After 10 days of exposure, green algae (Eutreptiella sp.) had increased, and phytoflagellates were the dominant group. The synthesis of mycosporine-like amino acids (MAAs), antioxidant enzymes and photosynthetic antenna pigments, in relation to repair and protection processes, may explain the reduced inhibition of both growth and photosynthesis that was observed in the phytoplankton community after several days of exposure. For environments such as the Beagle Channel seasonally exposed to the ozone hole, the results obtained from the fixed-depth experiments show that species can cope with UVR by means of MAA synthesis, while mixing would primarily promote a change in species composition and defense strategies.

Ultraviolet B-photoprotection efficiency of mesocosm-enclosed natural phytoplankton communities from different latitudes: Rimouski (Canada) and Ubatuba (Brazil).

Photoprotection against UV-B radiation (UVBR; 280-320 nm) was examined in natural phytoplankton communities from two coastal environments at different latitudes: temperate Rimouski (Canada) and tropical Ubatuba (Brazil). Mesocosm experiments were performed at these sites to examine the response of phytoplankton to increases in UVBR that corresponded to local depletions of 30% and 60% in atmospheric ozone levels (low and high UVBR treatments, respectively). A fluorescence method using a pulse amplitude modulation fluorometer (Xe-PAM, Walz, Germany) with selective UV filters was used to estimate photoprotection, and these results were compared with an index of mycosporine-like amino acid (MAA) concentrations determined using spectrophotometry of methanol extracts. The present study provided the first evidence, to our knowledge, of the suitability of this in vivo fluorescence method for the estimation of UV photoprotection efficiency in natural phytoplankton. No significant differences were found for most of the variables analyzed between the light treatments used at both sites, but differences were found between sites throughout the duration of the experiments. Vertical mixing, used to maintain cells in suspension, likely alleviated serious UVBR-induced damage during both experiments by reducing the length of time of exposure to the highest UVBR irradiances at the surface. In Rimouski, this was the main factor minimizing the effects of treatment, because optical properties of the coastal seawater rapidly attenuated UVBR throughout the water column of the ca 2 m deep mesocosms. In this location, synthesis of MAAs and photoprotective pigments likely contributed to the observed phototolerance of phytoplankton and, hence, to their growth; however, in a comparison of the UVBR treatments, these variables showed no differences. In Ubatuba, where nutrient concentrations were significantly lower than those in Rimouski, light attenuation was less than that in Rimouski and UVBR reached the bottom of the mesocosms. UVBR penetration and the forced vertical mixing of the cells, without the possibility of vertical migration below this photostress zone, resulted in photo-inhibition, because confinement in the mesocosms forced cells to remain constantly exposed to high levels of irradiance during the daytime. Hence, additional effects of UVBR were masked in this experiment, because cells were damaged too much and phytoplankton populations were rapidly declining. There was also an overall preservation of MAAs, in contrast with chlorophyll (Chl) degradation, in spite of the fact that this UV screening was not sufficient to counteract photo-inhibition, which suggests an important role for these molecules, either in the overall photoprotection strategy or in other physiological processes. Altogether, local water characteristics, namely attenuation, mixing, and nutrients concentration, can strongly modulate the photoprotection strategies used by natural phytoplankton populations in coastal environments.

The whole is more than the sum of its parts: Modeling community-level effects of UVR in marine ecosystems.

The effect of UVB radiation (UVBR, 290-320 nm) on the dynamics of the lower levels of the marine plankton community was modeled. The model was built using differential equations and shows a good fit to experimental data collected in mesocosms (defined as large enclosures of 1500 L filled with natural marine waters). Some unexpected results appear to be possible by indirect effects in prey (bacteria, phytoplankton and heterotrophic flagellates). In particular, apparent competition appears between small phytoplankton and bacteria. This effect is caused by a shared predator (ciliates). Another remarkable effect is an increase in bacteria and flagellates populations due to enhanced UVBR. This effect is similar to that observed under mesocosm experimental conditions and is related to the decrease of predation due to the direct damage to predators (ciliates) by UVBR. The effect of UVBR changing interaction coefficients may be dramatic on the community structure, producing big changes in equilibrium populations, as demonstrated by sensitivity analysis of the model. In order to generalize these results to field conditions it will be necessary to increase model complexity and include extra organic mater sources, mixing and sinking effects and predation by large zooplankton. This work shows that UVBR may produce community global responses that are consequence of both direct and indirect effects among populations.

Impact of solar ultraviolet radiation on marine phytoplankton of Patagonia, Argentina.

Patagonia area is located in close proximity to the Antarctic ozone "hole" and thus receives enhanced ultraviolet B (UV-B) radiation (280-315 nm) in addition to the normal levels of ultraviolet A (UV-A; 315-400 nm) and photosynthetically available radiation (PAR; 400-700 nm). In marine ecosystems of Patagonia, normal ultraviolet radiation (UVR) levels affect phytoplankton assemblages during the three phases of the annual succession: (1) prebloom season (late summer-fall), (2) bloom season (winter-early spring) and (3) postbloom season (late spring-summer). Small-size cells characterize the pre- and postbloom communities, which have a relatively high photosynthetic inhibition because of high UVR levels during those seasons. During the bloom, characterized by microplankton diatoms, photosynthetic inhibition is low because of the low UVR levels reaching the earth's surface during winter; this community, however, is more sensitive to UV-B when inhibition is normalized by irradiance (i.e. biological weighting functions). In situ studies have shown that UVR significantly affects not only photosynthesis but also the DNA molecule, but these negative effects are rapidly reduced in the water column because of the differential attenuation of solar radiation. UVR also affects photosynthesis versus irradiance (P vs E) parameters of some natural phytoplankton assemblages (i.e. during the pre- but not during the postbloom season). However, there is a significant temporal variability of P vs E parameters, which are influenced by the nutrient status of cells and taxonomic composition; taxonomic composition is in turn associated with the stratification conditions (e.g. wind speed and duration). In Patagonia, wind speed is one of the most important variables that conditions the development of the winter bloom by regulating the depth of the upper mixed layer (UML) and hence the mean irradiance received by cells. Studies on the interactive effects of UVR and mixing show that responses of phytoplankton vary according to the taxonomic composition and cell structure of assemblages; therefore cells use UVR if >90% of the euphotic zone is being mixed. In fact, cell size plays a very important role when estimating the impact of UVR on phytoplankton, with large cells being more sensitive when determining photosynthesis inhibition, whereas small cells are more sensitive to DNA damage. Finally, in long-term experiments, it was determined that UVR can shape the diatom community structure in some assemblages of coastal waters, but it is virtually unknown how these changes affect the trophodynamics of marine systems. Future studies should consider the combined effects of UVR on both phytoplankton and grazers to establish potential changes in biodiversity of the area.