European Center for Arctic Environmental Research Ny-Ålesund, Svalbard, Norway ARCFAC V: Project Proposal
All applicants must use this standard template for their project proposals. The project proposal must be maximum 8 pages, including the list of references. Standard page format is A4, with 3,17 cm margins, 12 point font. The proposal will be evaluated by a User Selection Panel involving both internal and external independent experts, in a peer review process emphasising scientific quality. Please refer to the specific eligibility and assessment criteria posted on the bsite. The Project Proposal shall be converted to PDF and sent along with CVs to: 0 Project Title: Thermal tolerance in congeneric Calanus: Implications for biogeographic distribution and ecosystem function in response to global warming.
1 Project Summary (max. ½ page): Differences in thermal tolerance between species and populations, and the ability of individuals to alter their tolerance limits, have been suggested as playing an important role in setting distribution limits and determining how organisms might be affected by climate change. As a contribution to a larger project being developed to investigate thermal tolerance in congeneric Calanus throughout their geographic ranges, we propose to undertake thermal stress experiments at Ny Alesund with the three species of Calanus (C. finmarchicus, C. glacialis, C. hyperboreus) that co-occur in Kongsfjorden to compare their thermal tolerance limits. We propose to use heat shock protein (hsp) expression as a bioindicator of thermal stress in Calanus by measuring the amount of hsp produced in response to exposure to different temperatures in the laboratory. We will also quantify hsp expression in Calanus resident in the fjord for comparison with individuals collected from other Arctic and sub-Arctic seas on IPY cruises in 2007 and 2008. We will expose Calanus collected from Kongsfjorden to a range of temperatures in the laboratory to determine the temperatures at which heat shock protein expression is turned on and off (thermal set- points) in each species. We hypothesise that differences in hsp expression between species and populations will provide an indication of whether Calanus are able to alter their temperature tolerance ranges, and therefore predict how they might respond to climate change, and how their distributions might change in the future. 2 Principal Objective and sub-goals (max ¼ page): The overall aims of this project are to determine the thermal tolerance limits of congeneric Calanus in the Arctic, and investigate differences in thermal tolerance between the species by quantifying hsp70 expression and investigating its adaptability to thermal stress. This will provide a key step in progressing our understanding of the physiological capability of Calanus to survive temperature stress, and improve our ability to predict the consequences of global warming on their future distributions. The project provides an opportunity to link physiological mechanisms with ecosystem function in an ecologically important genus in the marine zooplankton. Objective 1: To establish hsp70 expression as a biomarker for determining the physiological capability of Calanus to survive global warming by comparing thermal tolerance limits between co-occurring species and characterising inter-individual levels of variation in hsp expression. We will test the hypothesis that thermal tolerance limits and adaptability will be lower in the cold-water Calanus species. Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET 3 Scientific Introduction (max 2 pages): Please give a brief scientific introduction to the field, state of the art, reference to previous work, relevant research, role in larger research activities etc. Global Warming and Calanus Two important considerations in the assessment of how climate change will impact organisms are (1) how close they currently are to their thermal limits and, (2) the potential to respond to increasing habitat temperatures by adjusting their thermal sensitivity (Stillman, 2003). Understanding the biological bases that underlie the responses of organisms to increasing habitat temperature is the only way we can increase the predictive power of how climate change will impact on species and communities (Stillman, 2003). In common with other ectotherms, marine copepods will be sensitive to climate change as they are adapted to a characteristic temperature window in their natural environment in order to maintain phenological relationships, physiological processes and geographic ranges. Substantial changes in the biogeography of calanoid copepod assemblages in the eastern North Atlantic Ocean and European shelf seas between 1960 and 1999 have been correlated with changes in Northern Hemisphere temperature (NHT) anomalies, the North Atlantic Oscillation (NAO) index, and sea surface temperature (SST) (Beaugrand et al., 2002). Consistent long-term changes in species associations, a poleward increase by 10o latitude in pseudo-oceanic temperate species, and decreasing diversity of subarctic and arctic species in the north reflect a movement of marine ecosystems towards a warmer dynamical equilibrium (Beaugrand et al., 2002). In the western North Atlantic (the sub-polar gyre) and Labrador Sea, there has been a shift towards a colder dynamical equilibrium, with Arctic species diversity increasing. Members of the genus Calanus are among the largest calanoid copepods, and are the dominant herbivores, in subarctic and Arctic Seas. They play a key role in pelagic food webs, influencing energy flow and functioning of productive marine ecosystems (Tande, 1991; Longhurst, 1998). They can contribute >90% of the dry weight biomass in some areas, and their importance in the diet of the juvenile stages of various economically important fish (such as cod, haddock, herring, mackerel, capelin, polar cod) is well known. In polar regions, C. hyperboreus is also critically important in the diet of little Auks. Therefore, changes in the abundance and distribution of Calanus will affect ecosystem structure and function, as has already been illustrated in the North Sea, where the progressive decline of C. finmarchicus since the mid-1980s has modified the plankton community and affected cod recruitment (Beaugrand et al., 2003), with implications on ecosystem complexity and resilience (Jennings et al., 2002). Here we focus on three Calanus species, which have distinctive temperature preferences and whose biogeographic ranges correlate with environmental gradients. Calanus finmarchicus is a sub-Arctic species that occurs throughout the North Atlantic north of the Gulf Stream and wherever North Atlantic water penetrates further north such as in the Barents Sea, Labrador-Baffin region and the Arctic Ocean (Grainger, 1963; Jaschnov, 1970). C. hyperboreus occurs in oceanic Arctic waters, including the Arctic Ocean, the eastern Canadian Arctic archipelago and Labrador-Baffin region (Grainger, 1963; Conover, 1988). C. glacialis is also an Arctic species, but in contrast to C. hyperboreus, is generally found in shallower shelf regions in the Arctic Ocean, Greenland Sea, Barents Sea, the eastern Canadian Arctic archipelago, and the Labrador-Baffin region (Grainger, 1963; Conover, 1988). Head et al. (2003) suggest that differences in the distribution of Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
the two Arctic species in the Labrador Sea may relate to their temperature tolerances. Of the two, C. hyperboreus appears to be more temperature tolerant and is common in waters >3oC, whereas C. glacialis seems to be restricted to cooler waters <2oC. Because the environmental temperature ranges experienced by the three species are significantly different, we hypothesize that their thermal tolerance windows and adaptive capabilities will differ significantly, with consequent implications on subsequent biogeography. To test this hypothesis, we plan to investigate differences in thermal tolerance between Calanus species using heat shock proteins (hsp) as indicators of thermal stress, and biomarkers of adaptability to climate change. Heat Shock Proteins (hsp) and Thermal Tolerance in Calanus. Developments in biochemical and molecular techniques have improved our ability to investigate the mechanisms underlying the effects of environmental conditions on animal physiology. In particular, the use of inducible molecular chaperones as integrative biochemical indicators of organism stress is becoming more widely used (e.g. Dahlhoff, 2004; Helmuth et al., 2005). Organisms respond to extreme environmental conditions such as thermal stress with rapid changes in gene expression and up-regulation of stress-inducible molecular chaperones (hsp), which stabilise thermally denaturing proteins and reduce tissue levels of unfolded proteins by repairing them (Dahlhoff, 2004). Adjustments in gene and protein expression afford individuals an element of phenotypic plasticity, improving their ability to recover from environmental stress and to cope with subsequent stress under fluctuating environmental conditions (Buckley et al., 2001). However, the heat shock response (HSR) is also energetically costly to organisms, with over-expression of hsp’s having deleterious consequences on organism fitness by diverting energy away from vital processes such as growth and reproduction. Initiation of the HSR, and increased synthesis of hsp’s, generally results in a concomitant reduction in the synthesis of other proteins (Parsell and Lindquist, 1993). Thus, plasticity in the HSR may represent a trade-off between the costs of maintaining thermotolerance at elevated versus resting levels (Hamdoun et al., 2003). Patterns of hsp synthesis appear to be related to thermal tolerances at the organism level (Somero, 2002). The temperature at which hsps are activated, and the threshold of induction, can vary within the lifetime of an individual, and is subject to thermal acclimation and acclimatisation (Buckley et al. 2001), with natural variations in hsp induction correlating with differences in average environmental temperature and the degree of environmental heterogeneity across large and small scale temperature gradients (Dahlhoff, 2004). In marine fish and invertebrates, hsp threshold induction temperatures have been positively correlated with environmental temperatures, and vary according to season, thermal history, environmental gradient, and laboratory acclimation, as well as with genetic differences between populations and species (e.g. Roberts et al. 1997; Tomanek and Somero, 1999, 2002; Buckley et al. 2001; Hamdoun et al., 2003; Sorte and Hofmann, 2004). For example, in a study of congeneric Tegula snails, induction of hsps was lowest in the temperate low-intertidal to sub-tidal species, and highest in the subtropical intertidal species (Tomanek and Somero, 1999). The plasticity of the HSR response allows organisms to adapt to natural thermal stress, increasing their tolerance to otherwise lethal conditions. This response is particularly important in sessile species or species exposed to highly variable environmental
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conditions. The fact that genetically fixed differences in thermal set-points appear to exist in different species means they probably play an important role in setting their different thermal tolerance limits (Tomanek and Somero, 1999). Thus, differences in thermal tolerance limits between populations and species, and their capacities to adjust these limits, have been suggested as playing important roles in setting biogeographic distributions and determining how organisms might be affected by climate change (Somero, 2005). Hsp70 has been shown to be the primary family of hsp that is responsive to thermal stress (Tomanek and Somero, 1999). Characterising changes in hsp70 gene and protein expression in response to thermal stress will provide us with a method to assess physiological stress in subarctic and Arctic Calanus species and to determine the capacity of different Calanus populations to survive and acclimate to increasing environmental temperatures. Induction of hsp70 mRNA has been demonstrated in laboratory temperature stressed C. finmarchicus (Voznesensky et al. 2004), where a four-fold increase in mRNA expression was observed for individuals exposed to environmentally realistic temperatures. We are focussing our efforts on C. finmarchicus, C. glacialis and C. hyperboreus because they are key species in northern North Atlantic and Arctic plankton communities. 4 Work Content and implementation plan (max 2 pages): Please provide a detailed description of the planned research activities in Ny-Ålesund: Planned field activities, laboratory work, instrument operation etc. Use the Gantt chart for the implementation plan. List the type and nature of collaboration with existing research activities and research groups at the site. The proposed work will require access to facilities at Ny Alesund for a period of two weeks during the summer months (June to August) when a range of Calanus life stages are present in the fjord. The dates given in the following table are indicative only. 15/07/2007 29/07/2007
Laboratory thermal tolerance limits experiments
Field Activities: Collection of Calanus from Kongsfjorden will require access to a small boat for 1 to 2 hours each day, either daily or every two days, for a period of two weeks. Ideally, a small winch is required on the boat for deployment of a CTD and zooplankton net. SAMS can supply a plankton ring-net and self contained CTD if these are not available on site. Laboratory Experiments: Continuous access to wet lab facilities and experimental temperature rooms, and use of experimental tanks (aquaria) and microscopes will be required for the duration of the visit for the temperature stress experiments. Liquid nitrogen (if available) and access to a - 80oC freezer is also necessary. Experimental Protocol: Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
Laboratory temperature stress experiments at Ny Alesund will compare differences in: (1) lethal thermal tolerance and; (2) thermal tolerance limits between field-acclimatised individuals of the three Calanus species co-occurring in Kongsfjorden. (The results from Kongsfjorden will also be compared with lethal and thermal tolerance limits of C. finmarchicus collected from Loch Etive in Scotland, at its southern range limit; experiments will be undertaken at SAMS). Calanus will be collected from Kongsfjorden using vertical hauls of a 200 µm plankton net with a non-filtering cod-end and maintained in the laboratory at ambient temperature. Water column temperature profiles will be taken using a CTD when samples are collected. Temperature stress experiments will involve exposing individuals (ca. 20 per 4L chambers) to temperatures ranging from 2oC above ambient to ca. 15oC (depending on mortality rates at higher temperatures). Lethal experiments (1) will assess mortality in 24 h exposures. Thermal tolerance tests (2) will employ a range of exposure and post-exposure durations to assess synthesis and accumulation of hsp70 in each species, and differences in hsp expression between life-stages of each species. Following the thermal tolerance experiments, copepods will either be snap-frozen in liquid nitrogen or preserved in RNAlater for quantification of hsp70 protein and mRNA expression using Western Blotting and real-time PCR on return to SAMS. Quantification of hsp70 expression in individuals at the different exposure temperatures will allow us to identify thermal set-points (Ton, Tpeak, Toff) for the different Calanus species. 5 Integration, Collaboration and Networking (max. ½ page): Please provide information about existing and new collaboration activities and complementary value to existing research at the site, if known. Relevance with respect to the International Polar Year (IPY). Role and importance with respect to its link to larger research projects, etc. The proposed research will contribute valuable complementary information on Calanus thermotolerance to international and IPY programmes that are studying the physical factors controlling the distribution of key zooplankton species (and their implications for higher trophic levels). In particular, this proposal has direct collaborative links with (1) the Bedford Institute of Oceanography’s long-term monitoring programme in the Labrador Sea studying the relationship between species distributions and hydrography (collaborative partner Erica Head), and (2) Ecosystem Studies of Subarctic and Arctic Regions (ESSAR), in particular the Norwegian IPY contribution NESSAR, which aims to address how climate variability and change affects the marine ecosystems of subarctic and Arctic seas and their sustainability (collaborative partner Ken Drinkwater). SAMS is also a partner in the MariClim project studying the marine ecosystem consequences of climate induced changes in water masses off West Spitsbergen (, along with the Polish Institute for Oceanology (IO PAS; collaborative partner Slawek Kwasniewski). As part of the larger project investigating geographic differences in Calanus thermal tolerance, Calanus are being collected on various IPY cruises, including cruises within the ESSAR, ArcOD and SPACE programmes. Data on Calanus abundance and hydrographic parameters collected by the project partners mentioned above and on the IPY cruises will provide complimentary information to the thermotolerance data, Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
allowing us to test hypotheses relating to factors governing species distributions and their response to climate change. Data on long-term temperature conditions in Kongsfjorden will be available from temperature loggers on a permanent mooring in the fjord, which is maintained by SAMS and the Norwegian Polar Institute within the SAMS/NERC Northern Seas Programme and the MariClim project.
6 Publication & Outreach plans (max. ½ page) Please provide a list of planned publications in peer review journals as well as other publication & outreach activities. Specify its relevance/role with respect to the International Polar Year (IPY). The main objective of the proposal is to investigate the underlying mechanisms by which physiological adaptation to temperature governs distribution patterns in marine copepods, and how they might be affected by climate change. The proposal has strong multidisciplinary appeal and the results should be of interest to a wide range of beneficiaries including marine and evolutionary biologists, physiologists, modellers involved in predicting the likely impacts of climate change on marine ecosystems, and environmental managers and policy makers involved with protecting marine biodiversity and fisheries resources. In order to disseminate the results to as wide a range of likely beneficiaries as possible, we will submit manuscripts to peer-reviewed journals with a broad scientific readership, and short articles will be written for magazines such as the New Scientist. The work will be presented at scientific conferences, such as the proposed 2008 Arctic Frontiers conference, and used to promote awareness of the consequences of global warming during science fairs, school visits and laboratory open days. 8 Consortium and Participants Please give a brief presentation of the project team, use the Gantt chart to justify the activities and the research days applied for in Ny- Ålesund, specify whether the team members are new to Ny-Ålesund or not. It is anticipated that three SAMS staff can carry out the proposed work over a period of two weeks during the summer months (June to August). Kate Willis will lead the team, with field and experimental help from Kenny Black and Shona Magill. The dates given in the following table are indicative only. Project Team Members Nb of days New or Last visit: (dd.mm.yy) (dd.mm.yy) (Yes/Year) 9 References Beaugrand G et al. 2002. Science 296:1692-1694. Beaugrand G et al. 2003. Nature 426:661-664. Buckley BA et al. 2001. J Expt Biol 204:3571-3579. Conover RJ 1988. Hydrobiologia 167/168:127-142 Dahlhoff EP 2004. Annu Rev Physiol 66:183-207. Grainger EH 1963. In: Dunbar MJ (ed) Marine distributions. Toronto, University of Toronto, Canada. Hamdoun AM et al. 2003. Biol Bull 205:160-169. Head EJH et al. 2003. Prog Oceanogr 59:1-30. Helmuth B et al. 2005. Ann Rev Physiol 67:177-201. Hughes L 2000. TREE 15(2):56-61. Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
Jaschnov WA 1970. Int Rev Gesamten Hydrobiol 55:197-212. Jennings S et al. 2002. MEPS 226:77-86. Longhurst A 1998. Ecological Geography of the Sea. Academic Press, San Diego, CA, USA. Parsell DA, Lindquist S 1993. Annu Rev Genet 27:437-496. Roberts DA et al. 1997. Biol Bull 192:309-320. Somero GN 2002. Integ Comp Biol 42:780-789. Somero GN 2005. Frontiers in Zoology 2:1 Sorte JB, Hofmann GE 2004. Mar Ecol Prog Ser 274:263-268. Stillman JH 2003. Science 301:65 Tande KS 1991. Pol Res 10:389-407. Tomanek L 2006. J Expt Biol 208:3133-3143. Tomanek L, Somero GN 1999. J Expt Biol 202:2925-2936. Tomanek L, Somero GN 2002. J Expt Biol 205:677-685. Voznesensky M et al. 2004. J Expt Mar Biol Ecol 311:37-46. 10 Attachments Short CV (max 4 page) of Team leader and participants CVs for Kate Willis, Kenny Black and Shona Magill are attached. Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET Kate Willis Date of Birth: 7 12 1968 Zooplankton Ecologist QUALIFICATIONS (most recent first)
Year Qualification Awarding CAREER HISTORY (most recent first) Date Event Organisation
Trainee Medical Laboratory Scientific Officer
PUBLICATIONS Ashton, G.V., Stevens, M., Hart, M., Green, D.H., Burrows, M.T., Cook, E.J., Willis, K.J., Introduction pathways of Caprella mutica populations in the northern hemisphere: a phylogenetic analysis. Molecular Ecology. Submitted. Ashton, G.V., Willis, K.J., Burrows, M.T., Cook, E.J. Environmental tolerance of Caprella mutica: implications for its distribution as a marine non-native species. Marine Environmental Research. Submitted. Willis, K.J., Cottier, F., Kwasniewski, S., Wold, A., Falk-Petersen, S. (2006). The influence of advection on zooplankton community composition in an Arctic fjord (Kongsfjorden, Svalbard). Journal of Marine Systems.61: 39-54. Ashton, G.A., Willis, K.J., Cook, E.J., Burrows, M.T. Distribution of the introduced amphipod, Caprella mutica Schurin, 1935, on the west coast of Scotland and a review of its global distribution. Hydrobiologia. In Press. Cook, E.J., Willis, K.J., Lozano Fernandez, M. Survivorship, growth and reproduction of the non-native Caprella mutica Schurin (Crustacea: Amphipoda). Hydrobiologia. In Press. Willis, K.J., Gillibrand, P.A., Cromey, C.J., Black, K.D. (2005). Sea lice treatments on salmon farms have no adverse effect on zooplankton communities: A case study. Marine Pollution Bulletin. 50: 806-816. Ling, N., Willis, K.J. (2005). Impacts of mosquitofish, Gambusia affinis, on black mudfish, Neochanna diversus. New Zealand Journal of Marine and Freshwater Research. 39: 1215-1223. Willis, K.J., Cook, E.J., Lozano-Fernandez, M., Takeuchi, I. (2004). First record of the caprellid amphipod, Caprella mutica, in the U.K. Journal of the Marine Biological Association of the United Kingdom. 84: 1027-1028. Willis, K.J., Van den Brink, P.J., Green, J.D. (2004). Seasonal variation in plankton community responses of mesocosms dosed with pentachlorophenol. Ecotoxicology 13: 707-720. Willis, K.J., Ling, N. (2004). The toxicity of the aquaculture pesticide cypermethrin to planktonic marine copepods. Aquaculture Research. 35:263-270. Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET Willis, K.J., Ling, N. (2003). The toxicity of emamectin benzoate, an aquaculture pesticide, to planktonic marine copepods. Aquaculture 221: 289-297. Ling, N., Gleeson, D.M., Willis, K.J., Binzegger, S.U. (2001). Creating and destroying species: the `new' biodiversity and evolutionarily significant units among New Zealand's galaxiid fishes. Journal of Fish Biology 59:209-222. Willis, K.J. & Ling, N. (2000). Sensitivies of mosquitofish and black mudfish to a piscicide: Could rotenone be used to control mosquitofish in NZ wetlands? NZ Journal of Zoology 27(2): 85-91. Willis, K.J. (1999). Acute and chronic bioassays with NZ freshwater copepods using PCP. Environmental Toxicology & Chemistry 18: 2580-2586. Willis, K.J., Ling, N., Chapman, M.A. (1995). The effects of temperature and chemical formulation on the toxicity of pentachlorophenol to Simocephalus vetulus. NZ Journal of Marine and Freshwater Research 29: 289-294. ______________________________________________________________________________________ Kenneth D Black Date of Birth: 14 9 1963 Head of Ecology Department QUALIFICATIONS (most recent first) Year Qualification Awarding CAREER HISTORY (most recent first) Date Event Organisation
Higher Scientific Officer, Leader of Coastal Impacts Research Group.
PUBLICATIONS 1. Black, K. D. and Gunstone, F. D. (1990) The Synthesis and Spectroscopic Properties of Some Polyol
Esters and Ethers. Chemistry and Physics of Lipids,56 169-173.
2. Black, K. D., Ezzi, I. A., Kiemer, M. C. B. and Wallace, A. J. (1994) Preliminary Evaluation of the
Effects of Long-Term Periodic Sublethal Exposure to Hydrogen-Sulfide On the Health of Atlantic Salmon (Salmo- Salar L). Journal of Applied Ichthyology-Zeitschrift Fur Angewandte Ichthyologie,10 362-367.
3. Kiemer, M. C. B., Black, K. D., Lussot, D., Bullock, A. M. and Ezzi, I. (1995) The effects of chronic and
acute exposure to hydrogen sulfide on Atlantic salmon (Salmo salar L). Aquaculture,135 311- 327. Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
4. Black, K. D. and Gunstone, F. D. (1996) The Friedel-Crafts Adducts of Methyl Oleate With Benzene and
Toluene. Chemistry and Physics of Lipids,79 87-94.
5. Black, K. D. and Gunstone, F. D. (1996) Friedel-Crafts Alkylation of Benzene and Toluene With
Olefinic C-6 Hydrocarbons and Esters. Chemistry and Physics of Lipids,79 79-86.
6. Black, K. D., Kiemer, M. C. B. and Ezzi, I. A. (1996) Benthic impact, hydrogen sulphide and fish health:
field and laboratory studies., in Aquaculture and sea lochs (ed K. D. Black), Scottish Association for Marine Science, Oban, pp 16-26.
7. Black, K. D., Kiemer, M. C. B. and Ezzi, I. A. (1996) The relationships between hydrodynamics, the
concentration of hydrogen sulfide produced by polluted sediments and fish health at several marine cage farms in Scotland and Ireland. Journal of Applied Ichthyology,12 15-20.
8. Black, K. D., Fleming, S., Nickell, T. D. and Pereira, P. M. F. (1997) The effects of ivermectin, used to
control sea lice on caged farmed salmonids, on infaunal polychaetes. ICES Journal of Marine Science,54 276-279.
9. Kiemer, M. C. B. and Black, K. D. (1997) The effects of hydrogen peroxide on the gill tissues of Atlantic
salmon, Salmo salar L. Aquaculture,153 181-189.
10. Provost, P. G., Black, K. D., Davies, I. M. and Read, P. A. (1997) Antibiotics in fish farm sediments, in
Environmental Pollution: Assessment and Treatment (ed P. Read and J. Kinross), Napier University Press, Edinburgh, pp 7-20.
11. Heasman, M. S. and Black, K. D. (1998) The potential of Arctic charr Salvelinus alpinus (L.) for
mariculture. Aquaculture Research.,29 67-76.
12. Henderson, R. J., D.A.M. Forrest, Black, K. D. and Park, M. T. (1998) Environmental distribution of
lipids in sealoch sediments underlying marine fish cages. Aquaculture,158 69-83.
13. Carrie, R., Mitchell, L. & Black, K. (1998) Seasonal fatty acid fluctuations on the Hebridean Shelf
Edge. Organic Geochemistry,29 1583-1593.
14. Cromey, C.J., Black, K.D., Edwards, A. and Jack I.A. (1998) Modelling the deposition and biological
effects of organic carbon from marine sewage discharges. Estuarine Coastal and Shelf Science, 47, 295-308.
15. Thetmeyer, H., Waller, U., Black, K. D., Inselmann, S. and Rosenthal, H. (1999) Growth of European
sea bass (Dicentrarchus labrax L.) under hypoxic and oscillating oxygen conditions. Aquaculture, 174 355-367.
16. Mac Dougall, N. & Black, K. D. (1999) Determining sediment properties around a marine cage farm
using acoustic ground discrimination: RoxAnn. Aquaculture Research,30 1-8.
17. McKenzie, J. D., Black, K. D., Kelly, M. S., Newton, L. C., Handley, L. L., Scrimgeour, C. M., Raven,
J. A. and Henderson, R. J. (2000) Comparisons of fatty acid and stable isotope ratios in symbiotic and non-symbiotic brittlestars from Oban Bay, Scotland. Journal of the Marine Biological Association of the United Kingdom,80 311-320.
18. Cook, E. J., Bell, M. V., Black, K. D. and Kelly, M. S. (2000) Fatty acid composition of gonagal
material and diets of the sea urchin, Psammechinus miliaris: trophic and nutritional implications. Journal of Experimental Marine Biology and Ecology,255 261-274.
19. Pantazis, P. A., Kelly, M. S., Connolly, J. G. and Black, K. D. (2000) Effect of artificial diets on
growth, lipid utilization, and gonad biochemistry in the adult sea urchin Psammechinus miliaris. Journal of Shellfish Research,19 995-1001.
20. Black, K. D. and Mac Dougall, N., (2002). Hydrography of four Mediterranean marine cage sites.
Journal of Applied Ichthyology, 18, 1-5
21. Cromey, C. J., Nickell, T. D. & Black, K. D. (2002). DEPOMOD - modelling the deposition and
biological effects of waste solids from marine cage farms. Aquaculture214, 211-239.
22. Cromey, C. J., Nickell, T. D., Black, K. D., Provost, P. G. & Griffiths, C. R. (2002). Validation of a fish
farm waste resuspension model by use of a particulate tracer discharged from a point source in a coastal environment. Estuaries25, 916-929.
23. Nickell, L. A., K. D. Black, D. J. Hughes, J. Overnell, T. Brand, T. D. Nickell, E. Breuer and S. M.
Harvey (2003). "Bioturbation, sediment fluxes and benthic community structure around a salmon cage farm in Loch Creran, Scotland." Journal of Experimental Marine Biology and Ecology285: 221-233.
24. Pereira, P. M. F., Black, K. D., Mclusky, D. S. & Nickell, T. D. (2004). Recovery of sediments after
cessation of marine fish farm production. Aquaculture. 235, 315–330 Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET
25. Willis, K.J., Gillibrand, P.A., Cromey, C.J. and Black, K.D. (2005) Sea Lice Treatments at a Salmon
Farm have No Adverse Effects on Zooplankton Communities, Marine Pollution Bulletin, 50 806-816.
26. Lepper, P.A., Turner, V.L.G., Goodson, A.D. and Black, K.D.(2004)Source levels and spectra emitted
by three commercial aquaculture anti-predation devices. Proceedings of the Seventh European Conference on Underwater Acoustics, ECUA 2004Delft, The Netherlands, 5-8 July, 2004
27. Obajimi, O., Black, K. D., Macdonald, D. J., Boyle, R. M., Glen, I. and Ross, B. M. (2005) Differential
effects of eicosapentaenoic and docosahexaenoic acids upon oxidant-stimulated release and uptake of arachidonic acid in human lymphoma U937 cells. Pharmacoogy Research 52, 183-191.
28. Whitmarsh, D. J., Cook, E. J. and Black, K. D. (2006) Searching for sustainability in aquaculture: an
investigation into the economic prospects for an integrated salmon-mussel production system. Marine Policy 30, 293-298.
29. Hughes, A. D., Catarino, A. I., Kelly, M. S., Barnes, D. K. A. and Black, K. D. (2005) Gonad fatty
acids and trophic interactions of the echinoid Psammechinus miliaris. Marine Ecology Progress Series 305, 101-111.
30. Hughes, A. D., Kelly, M. S., Barnes, D. K. A., Catarino, A. I. and Black, K. D. (2006) The dual
functions of sea urchin gonads are reflected in the temporal variations of their biochemistry. Marine Biology 148, 789-798.
31. Cook, E.J. K.D.Black, M.D.J. Sayer, C.J. Cromey, D. Angel, E. Spanier, A. Tsemel, T. Katz, N. Eden,
I. Karakassis, M. Tsapakis, E. Apostolaki and A. Malej (2006) The influence of caged mariculture on the early development of sub-littoral fouling communities: a pan-European study. ICES Journal of Marine Science 63: 637-649
32. Dean, R. J., Shimmield, T. M. and Black, K. D. (2007). Copper, zinc and cadmium in marine cage fish
farm sediments: an extensive survey. Environmental Pollution145, 84-95.
33. Obajimi, O., Black, K. D., Glen, I. and Ross, B. M. (in press) Antioxidant modulation of oxidant-
stimulated uptake and release of arachidonic acid in eicosapentaenoic acid supplemented human lymphoma U937 cells. Prostaglandins, Leukotrienes and Essential Fatty Acids (IF 1.807)
34. Liu H., Kelly M.S., Cook E.J., Black, K.D., Orr H., Zhu, J.X. and Dong, S.L. 200X. The effect of diet
type on growth and fatty-acid composition of sea urchin larvae, I Psammechinus miliaris (Gmelin). Aquaculture
35. Liu H., Kelly M.S., Cook E.J., Black, K.D., Orr H., Zhu, J.X. and Dong, S.L. 200X. The effect of diet
type on growth and fatty-acid composition of sea urchin larvae, II Paracentrotus lividus (Lamarck, 1816) (Echinodermata). Aquaculture
Network for Arctic Climate and Biological Diversity Studies ARCDIV.NET Shona Magill
Date of Birth: 13 9 1968 Support Scientist/Research Assistant Ecology QUALIFICATIONS (most recent first)
Year Qualification Awarding CAREER HISTORY (most recent first) Date Event Organisation
PUBLICATIONS Davenport, J. and Magill, S.H. (1996). Thermoregulation or osmotic control? Some preliminary observations of the function of emersion in the Diamondback terrapin Malaclemys terrapin (Latreille). Herpetological Journal, 6(1): 26-29. Davenport, J., Magill, S.H., and De Verteuille, N. (1997). The effects of current velocity and temperature upon swimming in juvenile green turtles Chelonia mydas (L). Herpetological Journal, 7:143-147. Field, R.H., Hills, J.M., Atkinson, R.J.A., Magill, S.H. and Shanks, A.M. (1998). Distribution and seasonal prevalence of Heamatodinium sp. infection of the Norway lobster (Nephrops norvegicus) around the west coast of Scotland. ICES Journal of Marine Science 55, 846-858. Davenport, J., Moore, P.G., Magill, S.H. and Fraser, L.A. (1998). Enhanced condition in dogwhelks, Nucella lapillus (L) living under mussel hummocks. Journal of Experimental Marine Biology and Ecology 230, 225-234. Magill, S.H. and Sayer, M.D.J. (2002). Seasonal and interannual variation in fish assemblages of northern temperate rocky subtidal habitats. Journal of Fish Biology 61, 1198-1216. Magill, S.H. and Sayer, M.D.J. (2004) The effect of reduced temperature and salinity on the blood physiology of juvenile Atlantic cod (Gadus morhua L.). Journal of Fish Biology 64: 1193-1205 Magill, S.H. and Sayer, M.D.J. (2004). Abundance of juvenile Atlantic cod (Gadus morhua) in the shallow rocky subtidal and the relationship to winter seawater temperature. Journal of the Marine Biological Association of the United Kingdom, 84: 439-442 Sayer, M.D.J., Magill, S.H., Pitcher, T.J., Morissette, L. and Ainsworth, C (2005). Using artificial habitats to restore or enhance inshore fisheries: an observation-based ecosystem simulation. Journal of Fish Biology 67 (Suppl B), 218-243 SH Magill, H Thetmeyer and CJ Cromey (2006). Settling velocity of faecal pellets of gilthead sea bream (Sparus aurata L.) and sea bass (Dicentrarchus labrax L.) and sensitivity analysis of measured data using a deposition model. Aquaculture 251, 295-305.
BOLHAS & COMICHÃO A varicela é uma infecção benigna da infância, causadapor um vírus – o varicella-zoster . É mais frequente noInverno e na Primavera, afectando sobretudo crianças atéFaz-se anunciar por febre ligeira, dores de cabeça, falta deapetite e mal-estar geral. Por vezes pode existir dor deA sua manifestação típica são pequenas manchas ver-melhas na pele,
Over-the-Counter Drug Eligibility: The recent Health Care Reform legislation introduced new plan provisions to Health Care Reimbursement Accounts. The first new provision to take effect is the change in eligibility of over-the-counter items. Effective January 1, 2011, over-the-counter (OTC) drugs and medicines (other than insulin) will require a prescription or Letter of Medical Necessity in ord