Abstract Previous reports on the effects of enhanced UV-B radiation on bryophytes have been equivocal. This study shows that mosses not only respond to enhanced UV-B, but they are affected by changes in ambient radiation. The studies were conducted with two model species common in northern environments; red-stemmed feather moss (Pleurozium schreberi) and juniper haircap moss (Polytrichum juniperinum). Both species showed high concentrations of methanol-extractable UV-absorbing compounds (UACs) with high spring-time and early-summer UV, whereas in P. juniperinum, the concentration was affected by early-summer drought. The UACs of P. juniperinum increased again towards autumn suggesting a role in winter hardening. The (spring-time) cell wall-bound UV screen was important to both species. The fundamental adaptation of P. juniperinum to open and exposed environments was reflected in relatively higher concentrations of total UACs compared to P. schreberi. The enhanced UV-B experiments in situ were conducted over two years in Oulu and six years at the FUVIRC site in Sodankylä. Some of the effects of UV-B were seen within the first years of the experiments, or even within hours, while others were observed after several years. Five or six years of enhanced UV-B treatment increased the methanol-extractable UACs of P. schreberi and decreased the green shoot growth of P. juniperinum. The immediate light environment was proposed to have an impact on the varying UAC concentrations. Some mitigating effects of UV-A were observed as well. Off-site measured, reconstructed and modelled UV radiation data was used for comparisons of light environment in situ, or when performing a reconstructive research with historical samples. The environmental sample banks can provide a useful tool to study past environmental conditions, and even reconstruct past radiation levels. It was shown in this study that UACs in P. schreberi and P. juniperinum have fundamental roles as UV-B screens in the cell walls, but there is also a variable response with the soluble fraction that reacts and adapts to the changes in UV radiation. The responses to increasing UV-B radiation vary in magnitude and in time. As P. schreberi and P. juniperinum possess circumboreal and cosmopolitan distributions, the effects of UV-B on these species and consequently on ecosystems has a broad application.

When plants are exposed to multiple environmental stress factors, one form of stress can affect the response to another stress. This study used seedlings of a new cultivar of wheat(Triticum aestivum L. cv. 1862), grown under factorial combinations of two levels of ultraviolet-B (UV-B)radiation, two salinity regimes and two levels of iron treatment in chelator-buffered nutrient solutions in a growth chamber. A number of morphological and physiological measurements were made. The accumulation of chlorophyll, UVabsorbing compounds and proline in shoots, as well as phytosiderophores (PSs) in root exudates were measured. Feed value measurements included crude protein, water-soluble carbohydrates, acid detergent fibre and Fe in shoots and roots. After 21 days of stress exposure, results showed that Fe deficiency and NaCl stress generally decreased plant growth and function as well as nutritive value, but increased plant biochemical protection traits such as proline accumulation (16.3 fold under salinity stress) and release of PSs (2.4 fold under Fe deficiency). Interestingly, UV-B radiation affected belowground parameters, inducing a 47% reduction in PS release, together with decreasing root DM by 9% and Fe concentration in roots by 7%. When Fe deficiency and NaCl stress were combined, the results showed a decrease in PS release by 3.5 fold compared to unstressed plants. UV-B radiation synergistically increased UV-absorbing compound levels in combination with Fe deficiency, compared to plants grown under optimal Fe levels. This stress combination also resulted in a cumulative effect by decreasing Fe concentration in shoots and roots. However, salt stress did not interact with UV-B radiation for any of the traits measured. In addition, some three-way interactions were noted, with the Fe x NaCl x UV-B stress combination slightly decreasing PS release and resulting in a cumulative effect by decreasing Fe concentration in roots. In conclusion, this study found that aboveground stress factors such as UV-B can affect important aspects of belowground plant function, and that Fe deficiency can interact with UV-B and salinity stress in modifying plant responses to either stress alone.

Ultraviolet light is damaging but fish have evolved protective mechanisms, which allows them to live in shallow water reefs, high in UV radiation. This thesis details my investigation into the physiological ecology of solar ultraviolet (UV) absorbing compounds, known as mycosporine-like amino acids found in the external epithelial mucus, and examines the supporting role potentially played by a UV-induced DNA repair mechanism in coral reef fish of the Indo-Pacific. Using reverse phase chromatography and UV spectrophotometry, I examined whether the distribution of MAA compounds across different areas of the body is correlated with differential UV exposure. Comparisons were made between the MAA content and the absorbance spectra of mucus from the dorsal, ventral, caudal and head body surface areas in five species of Scaridae (Chlorurus sordidus, Scarus schlegeli, S. niger, S. psittacus and S. globiceps) from Ningaloo Reef, Coral Bay, Western Australia. All fish analysed had at least five MAAs present, and results showed that fish had increased UV absorbance in mucus over the dorsal area, which receives the brunt of UV radiation. Little UV protection was found in mucus from the ventral area, which receives the lower level of UV radiation mostly via reflection of the sand and reef surfaces. Furthermore, UV absorbance per mg dry mucus versus standard fish length showed that there is a positive relationship in C. sordidus with increasing size. I examined whether there is a difference in the quantity of UV screening compounds found in the mucus of fish along a longitudinal geographical gradient from inshore reefs (Lizard Island, Great Barrier Reef) to the outer edge reefs to oceanic reefs (Osprey Reef). MAA absorbance increased with longitudinal distance from the mainland landmass of Australia to more oligotrophic outer reefs, where UV attenuation is reduced and the ocean is more transparent to UV wavelength. I determined that fish living on inshore, more turbid reefs where UV attenuation in shallow waters is high have lower levels of MAA protection than fish from clear oceanic reefs. Furthermore, there seems to be a direct relationship between light attenuation and exposure with the quantity of protective sunscreening found in the mucus of reef fish. It is know that UV irradiation decreases with water depth and that mucus from fish with deep habitats absorbs less UV than that of fish from shallow habitats. It is unknown however, whether this UV protection is variable within the same individuals and if so, how fast changes 11 occur. To test this, I relocated 9 ambon damselfish from a deep reef (18 m) to a shallow reef (1.5 m) to expose fish to increased levels of UV and relocated another 7 fish from a shallow to a deep reef to expose fish to decreased levels of UV. One week after relocation, all fish were returned to their original reef site to determine whether MAA levels would return to their initial levels. Fish relocated to a shallower depth were recovered and had a 60% (SD+/-2%) increase in mucus UV absorbance. Conversely, the fish relocated to a deeper depth were recovered and had a 41% (SD+/-1%) decrease mucus UV absorbance. No difference was found between UV absorbance of relocated and original fish at both depth. Six days after fish were returned to their original reef, mucus UV absorbance levels had returned to 67% +/- 4% of the original level. These results show that mucus UV absorbance is variable in individual ambon damselfish and that the sunscreen protection typical for a certain depth is reached in relocated fish within just a few days of relocation. The rate of MAA loss is higher than the accumulation of MAAs suggesting that diet is not the sole determining factor involved in the sequestration of MAAs to mucus. The cleaner fish Labroides dimidiatus performs a mutualistic service by removing ectoparasites such as gnathiid isopods as well other dead infected tissue from its clients. Cleaner fish however are also known to feed on client mucus. The benefits of eating mucus until recently were unclear. In this study, we analysed the mucus of several cleaner fish clients to determine whether mucus feeding has a nutritional advantage over gnathiids and whether cleaner fish obtain their own MAA protection through this dietary mucus ingestion. Results show that host fish that are infected with gnathiids of poor nutritional value, in contrast to those that harbour gnathiids with higher nutritional value, continuously exude mucus that has both high nutritional value and high MAA content. These findings support the conclusion that in a competitive market for cleaners some host fish are forced to offer more than parasites to cleaners. Ultraviolet light that is not filtered by UV absorbing compounds such as MAA may still lead to DNA damage such as the formation of cyclobutane pyrimidine dimers (CPDs) or 6-4 photoproducts (6-4 PPs). However, coral reef fish have alternative mechanisms to overcome UV induced damage via the photolyase DNA repair mechanisms. We experimentally demonstrated for the first time that a coral reef fish species, the moon wrasse Thalassoma lunare has the ability to repair DNA damage via photoreactivation. Fish both with and without MAA protection were irradiated with UVB wavelength to induce DNA lesions. Half of the experimental fish were then exposed to photoreactivating wavelength to induce DNA repair 12 while the other fish were blocked from the repair mechanisms. Fish which had undergone DNA repair had the lowest number of lesions regardless of mucus MAA protection. When fish were blocked from photoreactivation wavelengths MAA sunscreens clearly served a photoprotective role. The amount of damage was greatest in fish which both lacked MAAs and which were also blocked from photoreactivating wavelengths. Thus for the overall UV protection of fish both the MAA sunscreens as well as the DNA repair system play a significant role in counteracting UV damage. Ultraviolet protection by MAA sunscreens is ubiquitous in marine fish. To date the same 5 MAA compounds (palythine (λmax 320 nm), asterina (λmax 330 nm), palythinol (λmax 332 nm), usujirene (λmax 357 nm) and palythene (λmax 360nm) have been identified in the mucus of several different species of reef fish from Australia. Here we report the first evidence of the presence of additional UV absorbing compounds found in the mucus of fish from Indonesia. Using UV spectroscopy the mucus of four species of fish was compared between both geographical regions. The presence of an additional peak between 294-296 nm wavelengths suggests the presence of gadusol and/or deoxygadusol, which are photoprotective compounds, thought to be the precursors of MAAs. Thus, UV protecting compounds in the mucus of fish may not be as conserved between different regions as previously assumed. Our knowledge concerning the effect of UV radiation has advanced considerably in the past decade and my research findings contribute to the better understanding of protective mechanisms of marine fish. The correlations I have found between UV attenuation/exposure, depth, and longitude of sampled individuals lead me to believe that mucus UV absorbing MAA compounds are a highly efficient adaptive defence.