Surface solar ultraviolet radiation for paleoatmospheric levels of oxygen and ozone

Many investigators have concluded that the level of solar ultraviolet radiation (200–300 nm) reaching the surface was a key parameter in the origin and evolution of life on Earth. The level of solar ultraviolet radiation between 200 and 300 nm is controlled primarily by molecular absorption by ozone, whose presence is trongly coupled to the level of molecular oxygen. In this paper, we present a series of calculations of the solar ultraviolet radiation reaching the surface for oxygen levels ranging from 10^–4 present atmospheric level to the present level. The solar spectrum between 200 and 300 mn has been divided into 34 spectral intervals. For each spectral interval, we have calculated the solar ultraviolet radiation reaching the Earth's surface by considering the attenuation of the incoming beam due to ozone and oxygen absorption. A one-dimensional photochemical model of the atmosphere was used for these calculations.     

The early atmosphere: a new picture

Over the last several years, many of the fundamental ideas concerning the composition and chemical evolution of the Earth's early atmosphere have changed. While many aspects of this subject are clouded--either uncertain or unknown, a new picture is emerging. We are just beginning to understand how astronomical, geochemical, and atmospheric processes each contributed to the development of the gaseous envelope around the third planet from the sun some 4.6 billion years ago and how that envelope chemically evolved over the history of our planet. Simple compounds in that gaseous envelope, energized by atmospheric lightning and/or solar ultraviolet radiation, formed molecules of increasing complexity that eventually evolved into the first living systems on our planet. This process is called "chemical evolution" and immediately preceded biological evolution; once life developed and evolved, it began to alter the chemical composition of the atmosphere that provided the very essence of its creation. Photosynthetic organisms which have the ability to biochemically transform carbon dioxide and water to carbohydrates, which they use for food, produce large amounts of molecular oxygen (O2) as a by-product of the reaction. Atmospheric oxygen photochemically formed ozone, which absorbs ultraviolet radiation from the sun and shields the Earth's surface from this biologically lethal radiation. Once atmospheric ozone levels increased sufficiently, life could leave the safety of the oceans and go ashore for the first time. Throughout the history of our planet, there has been strong interaction between life and the atmosphere. Understanding our cosmic roots is particularly relevant as we embark on a search for life outside the Earth. At this very moment, several radio telescopes around the world are searching for extraterrestrial intelligence (SETI).     

Who, what, where and when-influences on cutaneous vitamin D synthesis

The synthesis of vitamin D in skin is a two-stage process that begins with the production of previtamin D after irradiation of 7-dehydrocholesterol by ultraviolet (UV) radiation. A number of personal and environmental factors control the probability of a suitable UV photon reaching a molecule of 7-dehydrocholesterol in the skin. These are astronomical factors that govern the solar zenith angle (SZA), and the local state of the atmosphere, determining the available solar UV radiation; skin pigmentation and age, determining competing absorbers of UV radiation and available 7-dehydrocholesterol; individual behaviour in the local surroundings, determining exposure of unprotected skin to available UV radiation. The only one of these influences that can be determined unequivocally for any situation is the SZA. The other influences must be considered either as individual case studies, or be represented by "typical" and "idealised" situations for the weather, skin and behaviour. At large SZAs there is insufficient solar UV radiation to initiate significant vitamin D synthesis. At smaller SZAs assessment of solar exposure necessary for vitamin D synthesis can only be indicative and application of any such assessment necessarily requires awareness of both self- and the local environment.     

Hornet peak flight activity is correlated with solar UV radiation: a multi-annual survey

This study deals with the effect which solar irradiation of short wavelength, particularly ultraviolet (UV), exerts on the activities of hornets. The findings are based on multi-annual observations carried out during the years 1985, 1989 and 1998 on hornet nests in the field. At the peak of UV radiation, which occurs at noon, hornet activity is greater by 1-2 orders of magnitude than that during the morning or evening hours. The main visible hornet activity appears to be the removal of soil particles from the nest so as to enlarge its volume, enable the building of additional combs and also increase the size of existing combs. Hornet flight during peak insolation hours is characterized by its briefness (5-20 seconds only) and brevity (to distances of 5-10 meters only) as compared to flights at other hours of the day. These prolonged, multi-annual observations lead to the conclusion that hornets are capable of converting the energy of UV radiation into a form amenable to metabolic usage. In this respect the hornet cuticle behaves as a thermophotovoltaic device, i.e., a semiconductor diode that converts photons radiating from the sunlight into electrical energy.     

Antarctic terrestrial ecosystem and role of pigments in enhanced UV-B radiations

The terrestrial ecosystem of Antarctica are among the most extreme on earth, challenging the communities and making their existence difficult by rapidly increasing annual summer influx of solar ultraviolet radiations (UV-R), extremely cold conditions and lesser availability of nutrients. Spring time ozone depletion is due to release of chlorofluorocarbons in the earth atmosphere and is a serious cause of concern among environmentalists. Antarctic continent is mostly dominated by cryptogamic plants with limited distribution in different parts of the icy continent however; their distribution is mostly confined to Sub-Antarctic region. By the virtue of light requirement, cryptogams are exposed to extreme seasonal fluctuation in photosynthetically active radiation (PAR), and ultraviolet (UV) radiation which are closely associated with photosynthetic pigments in photoautotrophic organisms. Antarctic cryptogams cope up the stress imposed by UV radiation by the development of efficient systems for repairing damage by synthesis of screening compounds such as UV-B absorbing pigments and anthocyanin compounds. A major part of the UV absorbing compounds are appeared to be constitutive in lichens which are usnic acid, perlatolic acid and fumarphotocetraric acid which is particularly induced by UV-B. Secondary metabolites such as phenolics, atranorin, parietin and melanin also enhance the plant defense, by different molecular targets in specific solar irradiance and potential for increased antioxidative protection to UV induced vulnerability.     

Effect of Vacuum Ultraviolet Irradiation on Abiogenous Synthesis of Adenine Nucleotides in the Presence of Lunar Ground

Vacuum ultraviolet (VUV) radiation, a powerful energy source in free Space, have been established to promote synthesis of natural nucleotides. It is shown that lunar ground protects from destruction (up to 1.5–2.0% ) the nucleotides formed after VUV-irradiation of dry films (adenosine and inorganic phosphate). Identification and quantitative determination of the products of synthesis and destruction is performed by the method of high performance liquid chromatography. The following products of synthesis are found: 5'-AMP > 2'3'-cAMP > 2'-AMP > 3'5'-cAMP > 3'-AMP. The obtained results are discussed from the viewpoint of hypothesis about the Space (extraterrestrial) origin of biologically important compounds that were initial for evolution on the Earth.     

Ultraviolet Radiation-Absorbing “Humic Pigments” of Cyanobacteria in Microbial Mats: Their Presumptive Photoprotective Function and Relevance to Early Precambrian Microbial Ecology and Evolution

Cyanobacteria that form the primary components of microbial mats in freshwater bogs and intertidal marine environments in the Bahamas produce water-soluble brown pigments whose spectral properties imply that they are a type of humic acid. These “humic pigments” are produced by vital processes of living cyanobacteria, not by decomposition of dead ones, as shown by decreases in the concentrations of humic pigments, ultraviolet (UV) radiation-absorbing photoprotective mycosporine-like amino acids (MAAs), and chlorophyll from upper to lower layers of the mats, and by the occurrence of humic pigments in cyanobacterial cultures. Unlike MAAs, which absorb UV radiation only within limited ranges of wavelengths, humic pigments absorb radiation spanning the entire UV spectrum, and absorbance increases with decreasing wavelength. These observations suggest that the biosynthesis of humic pigments originated as a photoprotective adaptation in the early Precambrian, enabling cyanobacteria to colonize shallow-water and terrestrial environments even though the atmosphere was virtually devoid of O₂ and O₃ and therefore transparent to all solar radiation in the UV region of the spectrum. Moreover, the evolution of this photoprotective mechanism may have been linked to the evolution of photosynthesis.     

RecA Expression in Response to Solar UVR in the Marine Bacterium Vibrio natriegens

Solar ultraviolet radiation may produce daily stress on marine and estuarine communities as cells are damaged and repair that damage. Reduction in the earth's stratospheric ozone layer has increased awareness of the potential effects that ultraviolet radiation may have in the environment, including how marine bacteria respond to changes in solar radiation. We examined the use of the bacterial RecA protein as an indicator of the potential of bacteria to repair DNA damage caused by solar UV irradiation using the marine bacterium Vibrio natriegens as a model. RecA is universally present in bacteria and is a regulator protein for the so-called Dark Repair Systems, which include excision repair, postreplication recombinational repair, and mutagenic or SOS repair. Solar UVB and UVA both reduced V. natriegens viability in seawater microcosms. After exposure to unfiltered solar radiation or radiation in which UVB was blocked, survival dropped below 1%, whereas visible light from which UVA and UVB had been filtered had no effect on survival. Using a RecA-specific antibody for detection, RecA protein was induced by solar radiation in a diel pattern in marine microcosms conducted in the Gulf of Mexico. Peak induction was observed at dusk each day. Although RecA expression was correlated with the formation of UVB-induced cyclobutyl pyrimidine dimers, longer wavelength UVA radiation also induced recA gene expression. Our results demonstrate that RecA-regulated, light-independent repair is an important component in the ability of marine bacteria to survive exposure to solar ultraviolet radiation and that RecA expression is a useful monitor of bacterial repair after exposure to solar UVR.     

Impact of solar ultraviolet-B radiation (290-320 nm) upon marine microalgae

For years scientists and laymen alike have casually noted the impact of solar ultraviolet radiation upon the non-human component of the biosphere. It was not until recently, when human activities were thought to threaten the protective stratospheric ozone shield, that researchers undertook intensive studies into the biological stress caused by the previously neglected short-wavelength edge of the global solar spectrum. Stratospheric ozone functions effectively as an ultraviolet screen by filtering out solar radiation in the 220–320 nm waveband as it penetrates through the atmosphere, thus allowing only small amounts of the longer wavelengths of radiation in this waveband to leak through to the surface of the earth. Although this ultraviolet radiation (UV-B radiation, 290–320 nm) comprises only a small fraction (less than 1%) of the total solar spectrum, it can have a major impact on biological systems due to its actinic nature. Many organic molecules, most notably DNA and proteins, absorb UV-B radiation which can initiate photochemical reactions. It is life's ability, or lack thereof, to cope with enhanced levels of solar UV-B radiation that has generated the concern over the potential depletion of stratospheric ozone. The defense mechanisms that serve to protect both plants and animals from current levels of UV-B radiation are quite varied. Whether these mechanisms will suffice for marine microalgae under conditions of enhanced levels of UV-B radiation is the subject of this review.     

Penetration and effects of solar UV-B on phytoplankton and macroalgae

The effects of short wavelength solar radiation on aquatic ecosystems were studied in several marine and freshwater systems. The spectral distribution and the penetration of solar radiation into different water types (coastal and oceanic waters of the Baltic Sea, North Sea, Atlantic and Mediterranean) were investigated. Penetration of short wavelength solar radiation strongly depends on the content of dissolved and particulate substances as well as the concentration of phytoplankton. The primary producers often show a typical vertical distribution within the euphotic zone and are reached as well as affected by the penetrating UV-B radiation. The effect of this radiation was both determined in phytoplankton and macroalgae. Measuring pulse amplitude modulated (PAM) fluorescence indicated that major biomass producers were severely inhibited by surface radiation and even impaired at their natural growth site. Likewise, photosynthetic oxygen production was affected by extended exposure to solar radiation.     

Motility and orientation of a dinoflagellate, Gymnodinium, impaired by solar and ultraviolet radiation

Abstract The effects of solar and artificial ultraviolet radiation on the motility and orientation of the dinoflagellate Y-100 were studied. The cells show a weak photokinesis but a pronounced phototaxis which is consistently positive between 1 and 100 klx (= 4 mW m⁻² to 400 mW m⁻²); the precision of orientation increases with the fluence rate. Unfiltered solar radiation as well as artificial ultraviolet radiation reduce the percentage of motile cells increasingly with exposure time but the velocity of the still motile cells is less affected. Unirradiated control cells show a negative gravitaxis. After short exposure to solar or artificial ultraviolet radiation the precision of gravitaxis decreases and after prolonged exposure the cells start to actively move downward in the water column (positive gravitaxis). Phototaxis is also strongly impaired by ultraviolet radiation.     

Wavelength-dependent induction of UV-absorbing mycosporine-like amino acids in the red alga Chondrus crispus under natural solar radiation

Polychromatic response spectra for the induction of UV absorbing mycosporine-like amino acids (MAAs) were calculated after exposing small thalli of the red alga Chondrus crispus under various cut-off filters to natural solar radiation on the North Sea island Helgoland, Germany. The laboratory-grown specimens typically contain only traces of palythine and synthesise five different MAAs rapidly and in high concentrations after being transplanted into shallow water. The resulting qualitative and quantitative patterns of MAA induction differed markedly with respect to spectral distribution. Furthermore, the wavebands effective for MAA induction vary within the MAA. UV-B radiation had a negative effect on the accumulation of the major MAAs shinorine (λ_max=334 nm) and palythine (λ_max=320 nm), while short wavelength UV-A exhibits the highest quantum efficiency on their synthesis. In contrast, the synthesis of asterina-330 (λ_max=330 nm), palythinol (λ_max=332 nm) and palythene (λ_max=360 nm) was mainly induced by UV-B radiation. Whether the synthesis of shinorine and palythine is induced by a photoreceptor with an absorption maximum in the short wavelength UV-A and whether a second photoreceptor absorbing UV-B radiation is responsible for the induction of asterina-330, palythinol and palythene remains to be studied.Our results show that C. crispus has a high capacity to adapt flexibly the qualitative and quantitative MAA concentration to the prevailing spectral distribution of irradiance. On one hand, this is regarded as an important aspect with respect to the acclimation of algae to increasing UV-B irradiance in the context of ongoing depletion of stratospheric ozone. On the other hand, the experiment demonstrates that UV-A irradiance is more important for the induction of the major MAAs shinorine and palythine than UV-B.     

Both near ultraviolet radiation and the oxidizing agent hydrogen peroxide induce a 32-kDa stress protein in normal human skin fibroblasts

We have analyzed the pattern of protein synthesis in solar near ultraviolet (334 nm, 365 nm) and near visible (405 nm) irradiated normal human skin fibroblasts. Two hours after irradiation we find that one major stress protein of approximately 32 kDa is induced in irradiated cells. This protein is not induced by ultraviolet radiation at wavelengths shorter than 334 nm and is not inducible by heat shock treatment of these cells. Although sodium arsenite, diamide, and menadione all induced a 32-kDa protein, they also induced the major heat shock proteins. In contrast, the oxidizing agent, hydrogen peroxide, induced the low molecular weight stress protein without causing induction of the major heat shock proteins. A comparison of the 32-kDa proteins induced by sodium arsenite, H2O2, and solar near ultraviolet radiation using chemical peptide mapping shows that they are closely related. These results imply that the pathways for induction of the heat shock response and the 32-kDa protein are not identical and suggest that, at least in the case of radiation and treatment with H2O2, the 32-kDa protein might be induced in response to cellular oxidative stress. This conclusion is supported by the observation that depletion of endogenous cellular glutathione prior to solar near ultraviolet irradiation lowers the fluence threshold for induction of the 32-kDa stress protein.     

Activation of mammalian gene expression by the UV component of sunlight – from models to reality

Ultraviolet radiation activates the expression of a wide variety of genes, by pathways which differ between the short non-solar ultraviolet C (UVC) wavelengths, which are strongly absorbed by nucleic acids, and the long solar ultraviolet A (UVA, 320–380 nm) wavelengths, which generate active oxygen intermediates. Intermediate solar ultraviolet (UV) wavelengths in the UVB (290–320 nm) range also contain an oxidative component, but more closely resemble UVC in their gene activating properties. Short wavelength UV, in common with other extracellular stimuli including growth factors, activates signal transduction events that involve both stress- and mitogen-activated protein kinase cascades. The extrapolation of the complex modulation of gene expression that ensues to the consequences of natural UV exposure requires careful attention to the details of doses and wavelength employed in the model experiments. Nevertheless, there is evidence that UVB irrradiation of skin can activate the expression of proteins including immunomodulating cytokines, ornithine decarboxylase and, to a limited extent, nuclear oncogene products, as well as lead to stabilisation of p53. Non-cytotoxic doses of UVA radiation also lead to the strong activation of several genes which would be expected to have functional relevance in vivo.     

UV effects on photosynthesis and phycobiliprotein composition in the flagellate Cyanophora paradoxa

Abstract The effects of solar and artificial ultraviolet radiation on photosynthetic oxygen production and phycobiliprotein composition were investigated in the freshwater flagellate, Cyanophora paradoxa . The phycobiliproteins of the cell acting as photosynthetic accessory pigments were found to be readily affected by even short exposure to ultraviolet radiation, while the membrane-bound chlorophyll protein complexes were hardly impaired as demonstrated by SDS polyacrylamide gel electrophoresis, isoelectric focussing and fast protein liquid chromatography (FPLC). The phycobilisomes are dissembled from the outside inwards and go from higher molecular weight components to hexamers (αβ)₆, trimers (αβ)₃ and finally monomers (αβ). Fluorescence spectra indicated that the energy transfer from accessory pigments to the photosystems was impaired by ultraviolet radiation. Photosynthetic oxygen production was affected on a much faster timescale than changes in the absorption spectra or at the protein level.     

UV effects on photosynthesis and phycobiliprotein composition in the flagellate Cyanophora paradoxa

Abstract The effects of solar and artificial ultraviolet radiation on photosynthetic oxygen production and phycobiliprotein composition were investigated in the freshwater flagellate, Cyanophora paradoxa . The phycobiliproteins of the cell acting as photosynthetic accessory pigments were found to be readily affected by even short exposure to ultraviolet radiation, while the membrane-bound chlorophyll protein complexes were hardly impaired as demonstrated by SDS polyacrylamide gel electrophoresis, isoelectric focussing and fast protein liquid chromatography (FPLC). The phycobilisomes are dissembled from the outside inwards and go from higher molecular weight components to hexamers ( αβ )₆, trimers ( αβ )₃ and finally monomers (αβ). Fluorescence spectra indicated that the energy transfer from accessory pigments to the photosystems was impaired by ultraviolet radiation. Photosynthetic oxygen production was affected on a much faster timescale than changes in the absorption spectra or at the protein level.     

Optimizing solar UV-radiation exposures for vitamin D3: comparing global and diffuse spectral UV radiation

Currently, there is a major gap in the knowledge that is needed to optimize the beneficial effects related to ultraviolet (UV) radiation at wavelengths that induce vitamin D(3) synthesis (UV(D3)) compared to reducing the biologically damaging overexposure to UV radiation. The aim of this study was to investigate the use of diffuse (radiation that is scattered from all directions) UV radiation to optimize exposures to UV(D3) radiation and maximize the reduction of exposure to UVA radiation. Data on global and diffuse solar UV-radiation spectra were collected at 10-min intervals in the Southern Hemisphere in the late spring and summer from 1 November 2006 to 28 February 2007. For a solar zenith angle (SZA) of approximately 5 degrees , the observed maximum UV(D3) irradiances were 0.80 W/m(2) and 0.46 W/m(2) for global and diffuse UV radiation, respectively. The observed maximum UVA irradiances were 79.0 W/m(2) and 36.2 W/m(2) for global and diffuse UV radiation, respectively. For diffuse UV radiation, the maximum ratio of vitamin D(3) to UVA radiation was 1.75% at a SZA of approximately 10 degrees , whereas the maximum ratio for global UV was 1.27% at 10 degrees . For SZAs of 25 degrees and less, more UV radiation is in the wavelength region contributing to vitamin D(3) synthesis (UV(D3)) than in the UVA region for diffuse UV radiation than for global UV radiation.     

A plant gene for photolyase: an enzyme catalyzing the repair of UV-light-induced DNA damage

Photolyases are thought to be critical components of the defense of plants against damage to DNA by solar ultraviolet light, but nothing is known about their molecular or enzymatic nature. The molecular cloning of a photolyase from mustard ( Sinapis alba ) described here is intended to increase the knowledge about this important repair mechanism in plant species at a molecular level. The gene encodes a polypeptide of 501 amino acids with a predicted molecular mass of 57 kDa. There is a strong sequence similarity to bacterial and yeast photolyases, with a close relationship to enzymes with a deazaflavin chromophor. The plant photolyase is shown to be functional in Escherichia coli which also indicates conservation of photolyases during evolution. It is demonstrated that photolyase expression in plants is light induced, thus providing good evidence for the adaptation of plants to their environment in order to diminish the harmful effects of sunlight.     

Factors that influence the cutaneous synthesis and dietary sources of vitamin D

The major sources of vitamin D for most humans are casual exposure of the skin to solar ultraviolet B (UVB; 290-315 nm) radiation and from dietary intake. The cutaneous synthesis of vitamin D is a function of skin pigmentation and of the solar zenith angle which depends on latitude, season, and time of day. In order to mimic the natural environment of skin to sunlight exposure, we therefore measured serum 25-hydroxyvitamin D levels in volunteers with different skin types following repeated UV irradiation. Because melanin pigment in human skin competes for and absorbs the UVB photons responsible for the photolysis of 7-dehydrocholesterol to previtamin D3, we also studied the effect of skin pigmentation on previtamin D3 production in a human skin model by exposing type II and type V skin samples to noon sunlight in June when the solar zenith angle is most acute. Vitamin D is rare in food. Among the vitamin D-rich food, oily fish are considered to be one of the best sources. Therefore, we analyzed the vitamin D content in several commonly consumed oily and non-oily fish. The data showed that farmed salmon had a mean content of vitamin D that was approximately 25% of the mean content found in wild caught salmon from Alaska, and that vitamin D2 was found in farmed salmon, but not in wild caught salmon. The results provide useful global guidelines for obtaining sufficient vitamin D3 by cutaneous synthesis and from dietary intake to prevent vitamin D deficiency and its health consequences, ensuing illness, especially, bone fractures in the elderly.     

Occurrence and induction of a ultraviolet‐absorbing substance in the cyanobacterium Fischerella muscicola TISTR8215

The filamentous cyanobacterium Fischerella muscicola TISTR8215 was tested for the presence of ultraviolet (UV)‐absorbing mycosporine‐like amino acids (MAAs) and their induction by UV radiation. Reverse‐phase high performance liquid chromatographic coupled with photodiode‐array detection studies revealed the presence of a MAA having an absorption maximum at 332 nm and a retention time of around 16.1 min. Based on absorption maximum, the compound was designated as M‐332. This is the first report for the occurrence of a MAA and its inducibility as influenced by UV radiation in Fischerella strains studied so far. Photosynthetically active radiation (PAR) had no significant impact on MAA induction. PAR + UV‐A radiation significantly induced the synthesis of M‐332; however, PAR + UV‐A + UV‐B radiation conferred highest impact on MAA synthesis. The cultures exposed to alternate light and dark conditions showed the induction of M‐332 synthesis mostly during the light period in contrast to the decreased levels of M‐322 during the dark period suggesting a circadian induction of its synthesis. Overall results indicate that F. muscicola may protect itself from deleterious short wavelength UV radiation by synthesizing the photoprotective compounds particularly during summer time in its natural brightly‐lit habitats.