EPOCA WP 6

1. WP6 Performance: reproduction and growthPIs: Clemmesen, Catriona, IFM-GEOMARPiatkowski, Uwe, IFM-GEOMARPörtner, Hans, AWISommer, Frank, IFM-GEOMARWiddicombe, Steve, PML EPOCA WP 6

2. WP6 Performance: reproduction and growth • Objectives: • Identify critical stages in the life cycle (e.g. eggs, larvae) of functionally important marine organisms based on performance measures as indicators of sensitivity to ocean acidification • Analyse physiological mechanisms defining performance levels and sensitivity • Estimate acclimation capacity (gene expression capacity) for that mechanism as the background of physiological plasticity • Quantify impact and tolerance thresholds (tipping points) • Assess interaction between ocean acidification (OA) and ocean warming • Compare responses and mechanisms in different populations of a species (e.g. in a climate gradient) reflecting potential for evolutionary adaptation (genetic differences) • Available funding EPOCA (PhD student equ.): • 1 PhD student AWI • 1 PhD student IFM GEOMAR • 1 PhD student PML Application for complementary national funding underway: e.g. Bioacid, NERC …..as an integrating programme

3. WP6 Performance: reproduction and growth Tasks • T1: Sensitivity of critical and early life stages to OA will be assessed through analyses of rates and normality of development, growth, reproductive success, integrity of calcified structures, metabolic and acid-base regulation • T2: pH regulation capacity in different body compartments of the animal will be assessed as a potential mechanism shaping sensitivity • T3: Regulation capacity will be related to changes in gene expression • T4: Impact and tolerance thresholds will be quantified. • T5: CO2 effects on tolerance to temperature will be analysed. • T6: Responses to CO2of different populations, e.g. in a latitudinal cline, will be compared to investigate potential evolutionary adaptation. Work has started at various levels, EPOCA and additional funding. …..as an integrating programme

4. Impact of high CO2 on Early life Stages of fish and cephalopods Preliminary results (days) Experiments will be conducted with the Kiel CO2 manipulation system and during the 2009 Arctic experiment to determine the impact of high CO2 on fish and cephalopods eggs, larvae and juveniles. EPOCA WP 6 C. Clemmesen, U. Piatkowski et al.

5. Impact of high CO2 on early life stages of fish and cephalopods Histology (PML) SL pH regulation capacity (AWI) Homogenate Otolith micro-structure RNA/DNA ratio DW, Growth IFM-GEOMAR RNA/DNA ratio Decrease in condition and protein synthesis capacity with increase in pCO2 Clemmesen & Franke in prep. Performance Indicators EPOCA WP 6 C. Clemmesen, U. Piatkowski et al.

6. Impact of high CO2 on the physiology of key benthic bioturbators. EPOCA WP 6 Amphiura filiformis is an important benthic organism in terms of nutrient cycling and benthic-pelagic coupling. Recent work at PML has shown significant impacts of high CO2 on this species (Wood et al., 2008 Proc Roy Soc B). Exposure to low pH seawater causes: • increase in respiratory rate • increased growth • increased calcification • BUT at the cost of muscle wastage • Experiments will be conducted in the PML seawater acidification facility and during the 2009 Arctic experiment to determine the impact of high CO2 on otherkey benthic species. • A whole organism approach will be adopted to assess both the responses and the costs. S. Widdicombe et al.

7. Hyas araneus, North Sea vs Svalbard acceleration extension n=171-322 EPOCA WP 6 PhD project Kathleen Walther H. Pörtner et al.

8. Greenland smoothcockle (Serripes groenlandicus) modified after Ambrose et. al 2006 In situ enclosures • Goals • Develop a mechanistic understanding of the combined effects of ocean warming and acidification (OA) on biomineralization/ calcification in high latitude bivalves via whole organism physiology. • To evaluate the impact of simultaneous temperature rise and OA on “biorecording” properties of bivalve shells, i.e. growth increment formation and biogeochemical parameters • Field work - Spitzbergen (AWIPEV station, Ny-Alesund) • current status: individuals marked with calcein in 2006, 2007 • perspectives: 2008 - recapture specimens from 2006, 2007; mark new animals • 2009 - mesocosm experiments (EPOCA) EPOCA WP 6 NN, O. Heilmayer et al.

9. Na+K+-ATPase Na+HCO3-- Cotransporter Long term acclimation via gene expression of pH – regulation mechanisms in fish gills Expression (Real-time PCR) of essential gill transport proteins Eelpout (Z. viviparus) 6 weeks K. Deigweiher, M. Lucassen, H.O. Pörtner, unpubl. EPOCA WP 6

10. Impact of high CO2 and temperature on development and reproduction of mesozooplankton (copepods) To date, the sensitivity of juvenile stages to OA has not been assessed. In the lab we will assess : - growth and development - egg hatching success - respiratory costs and/or gene expression 2 temperatures 6 pCO2 levels (~380-4000 ppm) Results will be compared with experiments using constant pCO2 and variable food quality (microalgae grown under different CO2 levels; WP8) In the 2009 Arctic mesocosm we will assess : • copepod egg production, hatching success and naupliar survival F. Sommer et al. EPOCA WP 6

11. Shifts in: geographical distribution Species interactions ….foodweb structure CO2 and the concept of oxygen and capacity limited thermal tolerance CO2, hypoxia Effects on behaviour, growth, reproduction,….fitness CO2, hypoxia rate of aerobic perfor- mance 0 Temperature Tp Tp : Pejus T‘s: Limitation of aerobic performance 100 % oxygen limited aerobic scope : critical T‘s: Tc Tc Anaerobic metabolism 0 Pörtner et al, 2005, Metzger et al. 2007, Pörtner and Knust, Science 2007 EPOCA WP 6

12. D6.1: Methodologies used in long term CO2 incubations, presented at meeting (month 9; O, PU) • D6.2-3: Internal reports on sensitivities of life stages in individual species (months 12 and 24; R, PP) • D6.4: Concept of sensitivity in the life cycles of various groups of marine animals (invertebrates and fish), dissemination as a viewpoint article (month 36; O, PU) • D6.5: Conceptual model of mechanisms causing sensitivity and linking organismic responses to ecosystem change (linking to WP 7, 9), dissemination as a viewpoint article (month 36; R, PU) • D6.6: Conceptual model of mechanisms and potential for acclimation and adaptation (month 48; R; PU) • D6.7: Contribution to mechanism-based model of OA effects at ecosystem level (linking to WP7 and 9) (month 48; R; PU) • D6.8: Potentially dangerous thresholds in relevant species, naming uncertainties and probabilities and perspectives on new ecosystem states to PP, PU (linking to WP 13)

14. Addressing CO2 effects and sensitivities in warming oceans First linesof CO2sensitivity(withecologicalrelevance) likelydepend on CO2 effects on temperaturedependentperformance in rel. tocompensation capacity for extracellular acid-base status. Thisincludesdisturbanceofcalcificationthroughextracellularacidification. Implicationstobeconsidered: seasonalshifts in performancewindows climatedependentfunctionalspecialization temperaturedependentbiogeography climatedependentgrowth, fecundity synergisticinteractionswithfactors in additiontotemperature (hypoxia, pollutants, …) Germany United Kingdom

15. Principle considerations: Role of time scales and levels for lethal effects of CO2 exposure arbitrary units Mortality dependent on CO2 level and exposure time Zone of resistance Upper median lethal CO2 level (LD50) Mortality independent of exposure time Incipient lethal CO2 level (long term critical threshold) Zone of tolerance log exposure time (days, weeks, months, years) → †Acute asphyxiation: squid, fish No such complete data set exists Critical level and mechanism unknown? Tolerable organism and ecosystem (?) responses Pörtner et al., 2005

16. Impact of high CO2 on Early life Stages of fish and cephalopods Control system Kiel CO2 Manipulation system Valves and flowmeters for constant pCO2 air mixtures Alarm system EPOCA WP 6

17. Impact of high CO2 and temperature on development and reproduction of mesozooplankton Timetable:

18. Funding: DFG SPP 1158 (HE5753/1-1: 2008 - 2010) CO2 impact on calcification in marine bivalves: a key to understand past, present and future climate records of polar ecosystem • Objectives: • to develop a mechanistic understanding of the combined effects of temperature rise and ocean acidification (OA) on biomineralization/ calcification in high latitude bivalves • to evaluate the impact of simultaneous temperature rise and OA on “biorecording” properties of bivalve shells, i.e. growth increment formation and biogeochemical parameters • Greenland smoothcockle (Serripes groenlandicus) • circum-arctic distribution • subtidal down to 100m • 80 - 100 mm within ten years EPOCA WP 6