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Seasonal Patterns in the Physiology of Calanus Glacialis

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Seasonal Patterns in the Physiology of Calanus Glacialis Life history traits of copepods in a changing Arctic - Seasonal patterns in the physiology of Calanus glacialis DISSERTATION zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften - Dr. rer. nat. - vorgelegt dem Fachbereich 2 (Biologie/ Chemie) der Universität Bremen von DANIELA FREESE Februar 2015 ii 1. Gutachter: PD Dr. Barbara Niehoff (Alfred-Wegener-Institut, Helmholtz-Zentrum für Polar- und Meeresforschung) 2. Gutachter: Prof. Dr. Wilhelm Hagen (Universität Bremen, Fachbereich 2) 1. Prüfer: Dr. Franz Josef Sartoris (Alfred-Wegener-Institut, Helmholtz-Zentrum für Polar- und Meeresforschung) 2. Prüfer: Prof. Dr. Kai Bischof (Universität Bremen, Fachbereich 2) Tag des Promotionskolloquiums: 13.05.2015 iii iv Contents List of abbreviations .......................................................................................................... i Summary .......................................................................................................................... iii Zusammenfassung ........................................................................................................... vi 1 Introduction ................................................................................................................. 1 1.1 Scientific background ............................................................................................. 1 1.2 Dormancy ................................................................................................................ 5 1.3 Physiological characteristics of copepods ............................................................ 10 1.3.1 Biochemical composition ............................................................................... 10 1.3.2 Enzyme activities ........................................................................................... 11 1.4 Vertical migration - The role of buoyancy ........................................................... 13 1.5 Aims and outline of this thesis .............................................................................. 14 2 Material and methods ............................................................................................... 17 2.1 Field work ............................................................................................................. 17 2.2 Analytical work ..................................................................................................... 22 2.2.1 Biochemical composition ............................................................................... 22 2.2.2 Enzyme analyses ............................................................................................ 23 2.2.2.1 Digestive enzyme activities ........................................................................ 24 2.2.2.2 Metabolic enzyme activities ....................................................................... 25 2.2.2.3 Substrate SDS-PAGE .................................................................................. 26 2.2.3 Extracellular pH and cation concentrations .................................................. 28 2.3 Incubation experiments under different food and light conditions ...................... 28 2.4 Statistics ................................................................................................................ 30 v 3 Manuscripts ................................................................................................................ 31 Manuscript I: ............................................................................................................... 33 Seasonal patterns in extracellular ion concentrations and pH of the Arctic copepod Calanus glacialis ............................................................................................................ Manuscript II: ............................................................................................................. 53 Digestive enzyme activities in the Arctic copepod Calanus glacialis reflect its ontogenetic vertical migration ........................................................................................ Manuscript III: ............................................................................................................ 77 Metabolic enzyme activities and body composition during the ontogenetic vertical migration of the Arctic copepod Calanus glacialis ........................................................ 4 Results and synoptic discussion ................................................................................ 99 4.1 Physiological and biochemical adaptations during activity and diapause ............ 99 4.1.1 Seasonal study on the physiology of Calanus glacialis in Billefjorden ...... 100 4.1.2 Spatial variations in the physiology of Calanus glacialis ............................ 110 4.2 Effect of different food and light conditions on the physiology of Calanus glacialis ..................................................................................................................... 117 5 Conclusions and future perspectives ..................................................................... 123 6 References ................................................................................................................. 124 Acknowledgements ...................................................................................................... 143 Appendix ...................................................................................................................... 145 Erklärung ...................................................................................................................... 157 vi List of abbreviations AARS Aminoacyl-tRNA synthethase acetyl-CoA Acetyl-Coenzyme A ANOVA Analysis of Variance aqua dem. Distilled water AWI Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research C Carbon Ca 2+ Calcium CaCl 2 Calcium chloride CBB Commassie brilliant blue Chl a Chlorophyll a CI - CV Copepodite stage I - V CLEOPATRA II Climate effects on food quality and trophic transfer in the Arctic Marginal ice zone CS Citrate synthase CVIF Adult females DM Dry mass DMSO dimethyl sulfoxide DTNB 5,5'-dithiobis-(2-nitrobenzoic acid) EC Enzyme Commission number EDTA ethylene-diamineteraacetic acid FITC Casein fluorescein isothiocyanate from bovine milk HCl Hydrochloric acid HOAD 3-hydroxyacyl-CoA dehydrogenase HPTS 8-Hydroxypyrene-1,3,6-trisulfonic acid trisodium salt indv. Individual K+ Potassium KV Coast guard vessels Li + Lithium Mg 2+ Magnesium MDH Malate dehydrogenase MSA Methane sulfonic acid MUF Methylumbelliferyl i N Nitrogen Na + Sodium NADH nicotinamide adenine dinucleotide NH 3 Ammonia + NH 4 Ammonium pH e Extracellular pH PPi Pyrophosphate reagent POLMAR Helmholtz Graduate School for Polar and Marine Research RFU Relative fluorescence units RV Research vessel SD Standard deviation SE Standard error SDS-PAGE Sodium dodecylsulfate polyacrylamide gel electrophoresis spp. Species SR Spearman rank order correlation TAG Triacylglycerols TCA trichloroacetic acid Tris Tris(hydroxymethyl)aminomethan UNIS The University Centre in Svalbard WP-2 Working party 2 plankton sampling net WP-3 Working party 3 plankton sampling net ii Summary In the last few decades, the Arctic has experienced rapid changes in the physical and biological marine environment. The sea ice cover shrinks, sea surface temperatures increase and the ice-free season prolongs, which results in profound changes in the food and light regime. In Arctic shelf seas, zooplankton communities are dominated by the large calanoid copepod Calanus glacialis , which links primary production with higher trophic levels. C. glacialis accumulates energy reserves in surface waters during the productive season and overwinters in a state referred to as diapause in deep waters. Diapause is characterized by arrested development and metabolic depression. The physiology and metabolism and the factors that determine the duration and timing of diapause in C. glacialis are, however, poorly understood. With ongoing environmental changes it is most important to understand the physiology and timing of life cycle events of C. glacialis in order to predict the effects of climate change on the pelagic food web. Thus, in a comprehensive approach, this study aims to tackle seasonal patterns in the physiology of C. glacialis and elucidate if changes in the metabolic activity are related to external cues, i.e. light, food and temperature, or if they are internally regulated. Within the framework of the Norwegian research project CLEOPATRA II (Climate effects on food quality and trophic transfer in the Arctic marginal ice zone), C. glacialis was sampled monthly in Billefjorden, a high-Arctic sill fjord on the western coast of Svalbard. In order to investigate the influence of different environments on the physiology of C. glacialis , the copepods were also collected in Kongsfjorden and Rijpfjorden whenever logistically possible. In a combined field and experimental study, the biochemical composition, digestive and metabolic enzyme activities and pH and ion concentration in the haemolymph of the C. glacialis in different phases of diapause were related to depth distribution of the copepods and different food and light conditions. The present study showed a clear seasonal pattern in digestive and metabolic enzyme activities as well as acid-base regulation and extracellular ion concentrations in C. glacialis . The physiological patterns were similar between C. glacialis
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