Effects of solar UV-B radiation on aquatic ecosystems.

Solar UV degrades dissolved organic carbon photolytically so that they can readily be taken up by bacterioplankton. On the other hand solar UV radiation inhibits bacterioplankton activity. Bacterioplankton productivity is far greater than previously thought and is comparable to phytoplankton primary productivity. According to the "microbial loop hypothesis," bacterioplankton is seen in the center of a food web, having a similar function to phytoplankton and protists. The penetration of UV and PAR into the water column can be measured. Marine waters show large temporal and regional differences in their concentrations of dissolved and particulate absorbing substances. A network of dosimeters (ELDONET) has been installed in Europe ranging from Abisko in Northern Sweden to Gran Canaria. Cyanobacteria are capable of fixing atmospheric nitrogen which is then made available to higher plants. The agricultural potential of cyanobacteria has been recognized as a biological fertilizer for wet soils such as in rice paddies. UV-B is known to impair processes such as growth, survival, pigmentation, motility, as well as the enzymes of nitrogen metabolism and CO2 fixation. The marine phytoplankton represents the single most important ecosystem on our planet and produces about the same biomass as all terrestrial ecosystems taken together. It is the base of the aquatic food chain and any changes in the size and composition of phytoplankton communities will directly affect food production for humans from marine sources. Another important role of marine phytoplankton is to serve as a sink for atmospheric carbon dioxide. Recent investigations have shown a large sensitivity of most phytoplankton organisms toward solar short-wavelength ultraviolet radiation (UV-B); even at ambient levels of UV-B radiation many organisms seem to be under UV stress. Because of their requirement for solar energy, the phytoplankton dwell in the top layers of the water column. In this near-surface position phytoplankton will be exposed to solar ultraviolet radiation. This radiation has been shown to affect growth, photosynthesis, nitrogen incorporation and enzyme activity. Other targets of solar UV irradiation are proteins and pigments involved in photosynthesis. Whether or not screening pigments can be induced in phytoplankton to effectively shield the organisms from excessive UV irradiation needs to be determined. Macroalgae show a distinct pattern of vertical distribution in their habitat. They have developed mechanisms to regulate their photosynthetic activity to adapt to the changing light regime and protect themselves from excessive radiation. A broad survey was carried out to understand photosynthesis in aquatic ecosystems and the different adaptation strategies to solar radiation of ecologically important species of green, red and brown algae from the North Sea, Baltic Sea, Mediterranean, Atlantic, polar and tropical oceans. Photoinhibition was quantified by oxygen exchange and by PAM (pulse amplitude modulated) fluorescence measurements based on transient changes of chlorophyll fluorescence.     

Annual solar UV exposure and biological effective dose rates on the Martian surface.

The ultraviolet (UV) environment of Mars has been investigated to gain an understanding of the variation of exposure throughout a Martian year, and link this flux to biological effects and possible survival of organisms at the Martian surface. To gain an idea of how the solar UV radiation varies between different regions, including planned landing sites of two future Mars surface missions, we modelled the total solar UV surface flux throughout one Martian year for two different dust scenarios. To understand the degree of solar UV stress on micro-organisms and/or molecules essential for life on the surface of Mars, we also calculated the biologically effective dose (BED) for T7 and Uracil in relevant wavelength regions at the Martian surface as a function of season and latitude, and discuss the biological survival rates in the presence of Martian solar UV radiation. High T7/Uracil BED ratios indicate that even at high latitudes where the UV flux is significantly reduced, the radiation environment is still hostile for life due to the persisting UV-C component of the flux.     

Effects of UV-B radiation on photosynthesis activity of Wolffia arrhiza as probed by chlorophyll fluorescence transients

The higher plant Wolffia arrhiza is regarded to be well suited concerning the provision of photosynthetic products in the cycle of matter of a Controlled Ecological Life Support System (CELSS) to be established in the context of extraterrestrial, human-based colonization and long-term space flight. Since UV radiation is one major extraterrestrial environmental stress for growth of any plant, effects of UV-B radiation on W. arrhiza were assessed in the present study. We found that UV-B radiation significantly inhibited photosynthetic CO₂ assimilation activity, and the contents of chlorophyll a, chlorophyll b (Chl a, Chl b) and carotenoids considerably decreased when plants were exposed to UV-B radiation for 12 h. High UV-B radiation also declined the quantum yield of primary photochemistry (φ_po), the quantum yield for electron transport (φ_Eo) and the efficiency per trapped excitation (Ψ_o) in W. arrhiza simultaneously, while the amount of active PSII reaction centers per excited cross section (RC/CS) and the total number of active reaction centers per absorption (RC/ABS) had comparative changes. These results indicate that the effects of UV-B radiation on photosynthesis of W. arrhiza is due to an inhibition of the electron transport and via inactivation of reaction centers, but the inhibition may take place at more than one site in the photosynthetic apparatus.     

Stability of nucleic acid under the effect of UV radiation.

Nucleic acids (combined with protein molecules) are essential constituents of the living systems playing an important role in the early evolution of life as well. A specific feature of these molecules has been found and directly confirmed recently: under the influence of short-wavelength UV radiation bipyrimidine photoproducts (cyclobutane dimers and 6-4 bipyrimidines) are induced and the reversion of them can be provoked by the same photons. However, reversion is preferred by the shorter wavelengths. With increasing ratio of the longer wavelength components of the radiation (using artificial UV sources and solar light on the Earth's surface) the impact of the reversible photoproducts in the harmful biological effect decreases and other photoproducts are dominant. Assuming the photoinduced reactions (dimerisation and reversion) are statistical events, during the irradiation the chance for a number of nucleoprotein molecules to survive the radiation damage can be reality. The theoretical and experimental basis of these assumptions will be discussed in the case of bacteriophage T7 nucleoprotein.     

Stratospheric ozone loss, ulraviolet effects and action spectroscopy

The major effect of stratospheric ozone loss will be an increase in the amount of ultraviolet radiation reaching the ground. This increase will be entirely contained within the UV-B (290–320nm). How this will impact life on Earth will be determined by the UV-B photobiology of exposed organisms, including humans. One of the analytical methods useful in estimating these effects is Action Spectroscopy (biological effect as a function of wavelength). Carefully constructed action spectra will allow us to partially predict the increase in bio-effect due to additional UV exposure. What effect this has on the organism and the system in which the organism resides is of paramount importance. Suitable action spectra already exist for human skin cancer, human cell mutation and killing, and for one immune response. Comprehensive and widely applicable action spectra for terrestrial and aquatic plant responses are being generated but are not yet suitable for extensive analysis. There is little data available for animals, other than those experiments completed in the laboratory as model systems for human studies. Some polychromatic action spectra have proven useful in determining the possible impact of ozone loss on biological systems. The pitfalls and limits of this approach will be addressed.     

Biological space experiments for the simulation of Martian conditions: UV radiation and Martian soil analogues.

The survivability of resistant terrestrial microbes, bacterial spores of Bacillus subtilis, was investigated in the BIOPAN facility of the European Space Agency onboard of Russian Earth-orbiting FOTON satellites (BIOPAN I -III missions). The spores were exposed to different subsets of the extreme environmental parameters in space (vacuum, extraterrestrial solar UV, shielding by protecting materials like artificial meteorites). The results of the three space experiments confirmed the deleterious effects of extraterrestrial solar UV radiation which, in contrast to the UV radiation reaching the surface of the Earth, also contains the very energy-rich, short wavelength UVB and UVC radiation. Thin layers of clay, rock or meteorite material were shown to be only successful in UV-shielding, if they are in direct contact with the spores. On Mars the UV radiation climate is similar to that of the early Earth before the development of a protective ozone layer in the atmosphere by the appearance of the first aerobic photosynthetic bacteria. The interference of Martian soil components and the intense and nearly unfiltered Martian solar UV radiation with spores of B. subtilis will be tested with a new BIOPAN experiment, MARSTOX. Different types of Mars soil analogues will be used to determine on one hand their potential toxicity alone or in combination with solar UV (phototoxicity) and on the other hand their UV protection capability. Two sets of samples will be placed under different cut-off filters used to simulate the UV radiation climate of Mars and Earth. After exposure in space the survival of and mutation induction in the spores will be analyzed at the DLR, together with parallel samples from the corresponding ground control experiment performed in the laboratory. This experiment will provide new insights into the principal limits of life and its adaptation to environmental extremes on Earth or other planets which and will also have implications for the potential for the evolution and distribution of life.     

R3D-B2 – Measurement of ionizing and solar radiation in open space in the BIOPAN 5 facility outside the FOTON M2 satellite

Solar and space radiation have been monitored using the R3D-B2 radiation risks radiometer-dosimeter on board a recent space flight on the Russian satellite Foton M2 within the ESA Biopan 5 facility mounted on the outside of the satellite exposed to space conditions. The solar radiation has been assayed in four wavelength bands (UV-C, 170–280 nm, UV-B, 280–315 nm), UV-A (315–400 nm) and PAR (photosynthetic active radiation, 400–700 nm). The data show an increasing tumbling rotation of the satellite during the mission. The photodiodes do not show a cosine response to the incident light which has been corrected. After calibration of the signals using the extraterrestrial spectrum, doses have been calculated for each orbit, for each day and for the total mission as basic data for the biological material which has been exposed in parallel in the Biopan facility. Cosmic ionizing radiation has been monitored and separated in 256 deposited energy spectra, which were further used for determination of the absorbed dose rate and flux. Basic data tables were prepared to be used by other Biopan 5 experiments. The paper summarizes the results for the Earth radiation environment at the altitude (262–304 km) of the Foton M2 spacecraft. Comparisons with the predictions of NASA Earth radiation environment experimental models AE-8 and AP-8, and the PSB97 model are also presented, which calculate the fluxes of ionizing radiation from a simulation. AP-8 is a model for trapped radiation.     

Biological UV dosimeters in simulated space conditions.

Polycrystalline uracil thin layers participate in the phage and uracil response (PUR) experiment, assigned to the biological dosimetry of the extraterrestrial solar radiation on the International Space Station (ISS). In ground based experiments (experiment verification tests), the following space parameters were simulated and studied: temperature, vacuum and short wavelength UV (UV-C, down to 200 nm) radiation. The closed uracil samples proved to be vacuum-tight for 7 days. In the tested temperature range (from -20 to +40 degrees C) the uracil samples are stable. The kinetic of dimer formation (dimerization) and reversion (monomerization) of uracil dimers due to short wavelength UV radiation was detected, the monomerization efficiency of the polychromatic deuterium lamp is higher than that of the germicidal lamp. A mathematical model describing the kinetic of monomerization-dimerization was constructed. Under the influence of UV radiation the dimerization-monomerization reactions occur simultaneously, thus the additivity law of the effect of the various wavelengths is not applicable.     

Monitoring of environmental UV radiation by biological dosimeters.

As a consequence of the stratospheric ozone layer depletion biological systems can be damaged due to increased UV-B radiation. The aim of biological dosimetry is to establish a quantitative basis for the risk assessment of the biosphere. DNA is the most important target molecule of biological systems having special sensitivity against short wavelength components of the environmental radiation. Biological dosimeters are usually simple organisms, or components of them, modeling the cellular DNA. Phage T7 and polycrystalline uracil biological dosimeters have been developed and used in our laboratory for monitoring the environmental radiation in different radiation conditions (from the polar to equatorial regions). Comparisons with Robertson-Berger (RB) meter data, as well as with model calculation data weighted by the corresponding spectral sensitivities of the dosimeters are presented. Suggestion is given how to determine the trend of the increase in the biological risk due to ozone depletion.     

Biological dosimetry to determine the UV radiation climate inside the MIR station and its role in vitamin D biosynthesis

The vitamin D synthesis in the human skin, is absolutely dependent on UVB radiation. Natural UVB from sunlight is normally absent in the closed environment of a space station like MIR. Therefore it was necessary to investigate the UV radiation climate inside the station resulting from different lamps as well as from occasional solar irradiation behind a UV-transparent quartz window. Biofilms, biologically weighting and integrating UV dosimeters successfully applied on Earth (e.g. in Antarctica) and in space (D-2, Biopan I) were used to determine the biological effectiveness of the UV radiation climate at different locations in the space station. Biofilms were also used to determine the personal UV dose of an individual cosmonaut. These UV data were correlated with the concentration of vitamin D in the cosmonaut's blood and the dietary vitamin D intake. The results showed that the UV radiation climate inside the Mir station is not sufficient for an adequate supply of vitamin D, which should therefore be secured either by vitamin D supplementat and/or by the regular exposure to special UV lamps like those in sun-beds. The use of natural solar UV radiation through the quartz window for ‘sunbathing’ is dangerous and should be avoided even for short exposure periods.     

Solar UV irradiance variation during cycles 22 and 23

The Solar Ultraviolet Spectral Irradiance Monitor (SUSIM) aboard the Upper Atmosphere Research Satellite (UARS) has been measuring solar UV irradiances since October 1991, a period which includes the decline of solar cycle 22 followed by the rise of cycle 23. Daily solar measurements include scans over the wavelength range 115–410 nm at 1.1 nm resolution. As expected, the measured time series of UV irradiances exhibit strong periodicities in solar cycle and solar rotation. For all wavelengths, the UV irradiance time series are similar to that of the Mg II core-to-wing ratio. During solar cycle 22, the irradiance of the strong Ly- line varied by more than a factor of two. The peak-to-peak irradiance variation declined with increasing wavelength, reaching 10% just below the Al edge at 208 nm. Between the Al edge and 250 nm the variation was 6–7%. Above 250 nm, the variation declines further until none is observed above 290 nm. Preliminary results for the first portion of cycle 23 indicate that the far UV below the Al edge is rising at about the same rate as the Mg II index while the irradiances in the Ly- emission line and for wavelengths longer than the Al edge are rising more slowly — even after accounting for the lower level of activity of cycle 23.     

Survival of microorganisms in space protected by meteorite material: results of the experiment 'EXOBIOLOGIE' of the PERSEUS mission.

During the early evolution of life on Earth, before the formation of a protective ozone layer in the atmosphere, high intensities of solar UV radiation of short wavelengths could reach the surface of the Earth. Today the full spectrum of solar UV radiation is only experienced in space, where other important space parameters influence survival and genetic stability additionally, like vacuum, cosmic radiation, temperature extremes, microgravity. To reach a better understanding of the processes leading to the origin, evolution and distribution of life we have performed space experiments with microorganisms. The ability of resistant life forms like bacterial spores to survive high doses of extraterrestrial solar UV alone or in combination with other space parameters, e.g. vacuum, was investigated. Extraterrestrial solar UV was found to have a thousand times higher biological effectiveness than UV radiation filtered by stratospheric ozone concentrations found today on Earth. The protective effects of anorganic substances like artificial or real meteorites were determined on the MIR station. In the experiment EXOBIOLOGIE of the French PERSEUS mission (1999) it was found that very thin layers of anorganic material did not protect spores against the deleterious effects of energy-rich UV radiation in space to the expected amount, but that layers of UV radiation inactivated spores serve as a UV-shield by themselves, so that a hypothetical interplanetary transfer of life by the transport of microorganisms inside rocks through the solar system cannot be excluded, but requires the shielding of a substantial mass of anorganic substances.     

Reconstruction of the solar UV irradiance back to 1974

The variability of the solar UV irradiance has strong effects on the terrestrial atmosphere. In order to study the solar influence for times when no UV observations are available, it is necessary to reconstruct the variation of the UV irradiance with time on the basis of proxies. We present reconstructions of the solar UV irradiance based on the analysis of space-based and ground-based magnetograms of the solar disk going back to 1974. With COde for Solar Irradiance (COSI) we calculate solar intensity spectra for the quiet Sun and different active regions and combine them according to their fractional area on the solar disk, whereby their time-dependent contributions over the solar cycle lead to a variability in radiation. COSI calculates the continuum and line formation under conditions which are out of local thermodynamic equilibrium (non-LTE). The applied temperature and density structures include the chromosphere and transition region, which is particularly important for the UV. The reconstructions are compared with observations.     

Microscopic approach to analyze solar-sail space-environment effects

Near-sun space-environment effects on metallic thin films solar sails as well as hollow-body sails with inflation fill gas are considered. Analysis of the interaction of the solar radiation with the solar-sail materials is presented. This analysis evaluates worst-case solar radiation effects during solar-radiation-pressure acceleration. The dependence of the thickness of solar sail on temperature and on wavelength of the electromagnetic spectrum of solar radiation is investigated. Physical processes of the interactions of photons, electrons, protons and α-particles with sail material atoms and nuclei, and inflation fill gas molecules are analyzed. Calculations utilized conservative assumptions with the highest values for the available cross sections for interactions of solar photons, electrons and protons with atoms, nuclei and hydrogen molecules. It is shown that for high-energy photons, electrons and protons the beryllium sail is mostly transparent. Sail material will be partially ionized by solar UV and low-energy solar electrons. For a hollow-body photon sail effects including hydrogen diffusion through the solar-sail walls, and electrostatic pressure is considered. Electrostatic pressure caused by the electrically charged sail’s electric field may require mitigation since sail material tensile strength decreases with elevated temperature. It also can substitute inflation-gas pressure loss due to gas diffusion and perforation by micrometeoroids impact to keep the sail inflated.     

The variability of the sun's ultraviolet radiation

The intensity of continua and emission lines which form the solar UV spectrum below 2100 Å is variable. Continua and emission lines originating from different layers in the solar atmosphere show a different degree of variability. Coronal emission lines at short wavelengths are much more variable than continua at longer wavelengths which originate in lower layers of the solar atmosphere. Typical time-scales of solar UV variability are minutes (flare induced), days (birth of active regions), 27 days (solar rotation), 11 years (solar cycle) and perhaps centuries, caused by long-term changes of the solar activity. UV intensity variations have been determined by either absolute irradiance measurements or by contrast measurements of plages vs. the quiet sun. Plages are the main contributor to the solar UV variability. Typical values for the solar UV variability over a solar cycle are: <1% at wavelengths longer than 2100 Å, 8% at 2080 Å (continuum), 20% at 1900 Å (continuum), 70% at H Lyα, 200% in certain emission lines 1200 < λ < 1800 Å and more than a factor of 4 in coronal lines λ < 1000 Å. Plage models predict the variable component of the solar UV radiation within ±50%. Absolute fluxes are known within ±30%. Several efforts are underway to monitor the solar UV irradiance with a precision better than a few percent over a solar activity cycle.     

短波长紫外辐射变化与臭氧含量涨落之间的关系

位于低纬地区的印度同时测量了短波紫外辐射与臭氧含量,表明臭氧含量下降3%,短波紫外辐射通量(290nm)上升35%.但这个数值大于以往理论预期的数值.本文给出一个简单公式,这个公式描述了臭氧含量的变化与相应的紫外辐射变化之间的关系,用此公式计算得出的结果与印度测得的结果相符,很好地解释了低纬地区的印度测量的臭氧涨落与其相应的短波长紫外辐射变化之间的关系.      

Inhibition of denitrification by ultraviolet radiation.

It has been shown that UV-A (lambda=320-400 nm) and UV-B (lambda=280-320 nm) inhibit photosynthesis, nitrogen fixation and nitrification. The purpose of this study was to determine the effects, if any, on denitrification in a microbial community inhabiting the intertidal. The community studied is the microbial mat consisting primarily of Lyngbya that inhabits the Pacific marine intertidal, Baja California, Mexico. Rates of denitrification were determined using the acetylene blockage technique. Pseudomonas fluorescens (ATCC #17400) was used as a control organism, and treated similarly to the mat samples. Samples were incubated either beneath a PAR transparent, UV opaque screen (OP3), or a mylar screen to block UV-B, or a UV transparent screen (UVT) for 2 to 3 hours. Sets of samples were also treated with nitrapyrin to inhibit nitrification, or DCMU to inhibit photosynthesis and treated similarly. Denitrification rates were greater in the UV protected samples than in the UV exposed samples the mat samples as well as for the Ps fluorescens cultures. Killed controls exhibited no activity. In the DCMU and nitrapyrin treated samples denitrification rates were the same as in the untreated samples. These data indicate that denitrification is directly inhibited by UV radiation.     

Solar UV radiation variations and their stratospheric and climatic effects

NIMBUS-7 SBUV measurements of the short-term solar UV variations caused by solar rotation and active-region evolution have determined the amplitude and wavelength dependence for the active-region component of solar UV variations. Intermediate-term variations lasting several months are associated with rounds of major new active regions. The UV flux stays near the peak value during the current solar cycle variation for more than two years and peaks about two years later than the sunspot number. NIMBUS-7 measurements have observed the concurrent stratospheric ozone variations caused by solar UV variations. There is now no doubt that solar UV variations are an important cause of short- and long-term stratospheric variations, but the strength of the coupling to the troposphere and to climate has not yet been proven.     

Quantification of biologically effective environmental UV irradiance.

To determine the impact of environmental UV radiation on human health and ecosystems demands monitoring systems that weight the spectral irradiance according to the biological responses under consideration. In general, there are three different approaches to quantify a biologically effective solar irradiance. (i) weighted spectroradiometry where the biologically weighted radiometric quantities are derived from spectral data by multiplication with an action spectrum of a relevant photobiological reaction, e.g. erythema, DNA damage, skin cancer, reduced productivity of terrestrial plants and aquatic foodweb, (ii) wavelength integrating chemical-based or physical dosimetric systems with spectral sensitivities similar to a biological response curve, and (iii) biological dosimeters that directly weight the incident UV components of sunlight in relation to the effectiveness of the different wavelengths and to interactions between them. Most biological dosimeters, such as bacteria, bacteriophages, or biomolecules, are based on the UV sensitivity of DNA. If precisely characterized, biological dosimeters are applicable as field and personal dosimeters.     

Model simulations of the impact of the 27-day solar rotation period on stratospheric ozone and temperature

Results are presented from two-year simulations of the effects of short-term solar ultraviolet (UV) variability using the Met. Office coupled chemistry-climate model. The model extends from the ground to 0.1 mbar and contains a complete range of chemical reactions allowing representation of all the main ozone formation and destruction processes in the stratosphere. The simulations were achieved by incorporating a 27-day oscillation in the pre-calculated model photolysis rates. Amplitudes for this signal were determined using solar spectral UV observations from the SOLar STellar Irradiance Comparison Experiment (SOLSTICE) instrument. Two experiments were carried out, one in which the UV variability was included in both the photolysis and radiation schemes and one in which only the photolysis scheme was modified.

The model reproduced several main features of observed correlations between short-term solar UV variability and both ozone and temperature in the tropical upper stratosphere, including the downward propagation of the phase lag and sensitivities of ozone and temperature to solar UV which are similar in magnitude to those observed. In the lower stratosphere, the ozone response to solar UV variability has not been well characterised from observations. Both model runs show a reversal of the propagation of phase lag below 10mb. The model response was found to be different between the two runs indicating that radiatively induced dynamical effects may play a significant role in the ozone response to solar UV variability.