Life in cold oceans: activity dependent on extracellular ion regulation? [Elektronische Ressource] = Die Rolle der extrazellulären Ionenregulation in der Kältetoleranz mariner Crustaceen / vorgelegt von Astrid C. Wittmann

Life in cold oceans: activity dependent on extracellular ion
regulation?


Die Rolle der extrazellulären Ionenregulation in der
Kältetoleranz mariner Crustaceen



Dissertation
zur Erlangung des akademischen Grades
– Dr. rer. nat –




dem Fachbereich 2 Biologie/Chemie
der Universität Bremen
vorgelegt von


Astrid C. Wittmann
Diplom-Biologin




Bremen 2010









































Gutachter:

1. Gutachter: Prof. Dr. Hans-Otto Pörtner
Alfred-Wegener-Institut für Polar- und Meeresforschung
Am Handelshafen 12, 27570 Bremerhaven


2. Gutachter: Prof. Dr. Ulrich Saint-Paul
Leibniz-Zentrum für Marine Tropenökologie
Fahrenheitstraße 6, 28359 Bremen


Prüfer: Dr. Franz J. Sartoris
Alfred-Wegener-Institut für Polar- und Meeresforschung
Am Handelshafen 12, 27570 Bremerhaven


Prüfer: PD Dr. Holger Auel
Universität Bremen, Fachbereich 2, Marine Zoologie
Leobener Straße NW2, 28359 Bremen


Tag des Promotionskolloquiums: 5. November 2010

TABLE OF CONTENTS

TABLE OF CONTENTS
LIST OF ABBREVIATIONS ....................................................................................................................................II
LIST OF FIGURES................................................................................................................................................... III
LIST OF TABLES......................................................................................................................................................IV
SUMMARY................................................................................................................................................................... V
ZUSAMMENFASSUNG..........................VII
1. INTRODUCTION.................................... 1
1.1. BIOGEOGRAPHY OF CRUSTACEANS AND THE SUB-ANTARCTIC AND ANTARCTIC THERMAL REGIMES............ 1
1.2. TEMPERATURE EFFECTS AND THERMAL TOLERANCE ........................................................................................ 2
1.3. THE MAGNESIUM HYPOTHESIS............................................................................................................................ 4
1.4. ION REGULATION AND THERMAL TOLERANCE ................................................................................................... 6
1.5. ONTOGENY AND THERMAL TOLERANCE............................................................................................................. 8
1.6. KEY QUESTIONS.................................................................................................................................................11
2. MATERIALS AND METHODS ..........................................................................................................................13
2.1. ACQUISITION AND MAINTENANCE OF ANIMALS...............................................................................................13
2.2. HAEMOLYMPH COLLECTION .............................................................................................................................14
2.3. ION CHROMATOGRAPHY.....14
2.4. EXPERIMENTS ON EARLY DEVELOPMENTAL STAGES OF PARALOMIS GRANULOSA...........................................15
2.4.1. CHN contents and dry mass.....................................................................................................................15
2.4.2. Oxygen consumption ................................................................................................................................15
2.4.3. Spontaneous swimming activity of zoeal stages at culture temperatures .............................................16
2.4.4. Activity of zoea I and crab I during acute cold exposure ......................................................................16
2.5. CARDIORESPIRATORY PHYSIOLOGY AND ACTIVITY OF ADULT P. GRANULOSA...............................................18
2.5.1. Preparation of animals and experimental protocol ...............................................................................18
2.5.2. Haemolymph PO .....................................................................................................................................20 2
2.5.3. Righting response......20
2.5.4. Analyses of heart and scaphognathite activity .......................................................................................21
2.6. EXPERIMENTS ON CARCINUS MAENAS ...............................................................................................................21
2.6.1. Spontaneous walking activity, food consumption and extracellular ion regulation ............................21
2.6.2. Cardiorespiratory physiology of C. maenas............22
2.7. STATISTICS.........................22
3. PUBLICATIONS....................................................................................................................................................24
PUBLICATION I..........................................................................................................................................................25
PUBLICATION II ........................................................................................................................................................49
PUBLICATION III ............................................................................................................................82
PUBLICATION IV........................111
4. DISCUSSION ........................................................................................................................................................132
4.1. HOW COLD TOLERANT ARE SUB-ANTARCTIC LITHODID CRABS? ..................................................................132
4.2. WHAT IS THE ROLE OF MAGNESIUM IN COLD TOLERANCE?...........................................................................137
4.3. WHY ARE “REPTANT” DECAPODS NOT PART OF THE ANTARCTIC SHELF FAUNA? ........................................143
4.4. CONCLUSIONS..................................................................................................................................................147
5. REFERENCES....................................150
6. APPENDIX ....................................................................................................................163
6.1. ADDITIONAL RESULTS......163
6.1.1. Effects of season, temperature and magnesium on Carcinus maenas ................................................163
6.1.1.1. Haemolymph ion composition, walking activity and food consumption ...................................................... 163
6.1.1.2. Haemolymph PO , ventilation and heart frequency in response to acute cold exposure ............................. 163 2
6.1.1.3. Seasonal effects on extracellular ion composition of C. maenas? ................................................................. 164
6.2. ADDITIONAL PUBLICATION .............................................................................................................................166
DANKSAGUNG........................................................................................................................................................169
ERKLÄRUNG GEM. § 5 (1) NR. 3 PROMO ......................................................................................................171


List of abbreviations
ASW artificial sea water
2+ASW –Mg artificial sea water with reduced magnesium concentration
-1 -1ßO oxygen solubility (mol L Torr ) 2
bpm beats per minute
CI crab I, first juvenile instar
CII crab II, second juvenile instar
C:N ratio Carbon to nitrogen ratio, is a proxy for the lipid:protein ratio
DW dry weight
frqfrequency
FW fresh weight
-1CO oxygen concentration (mol L ) 2
HLPO haemolymph oxygen partial pressure (kPa) 2
ind individual
M megalopa stage
-1 -1MO oxygen consumption, respiration, metabolic rate (molO h ind ) 2 2
Mya million years ago
n number of individuals
n.d. not determined
NSW natural sea water
2+ 2+NSW +Mg natural sea water of increased [Mg ]
P barometric pressure (Torr) B
PH O water vapour pressure (Torr) 2
Q temperature-velocity relationship, is a measure of thermal sensitivity 10
sc scaphognathite
s.d. standard deviation
s.e.standard error
t time
T temperature
T0 time point directly after hatching of larvae
V volume of the respiration chamber (L)
ZI zoea I, zoeal stage I
ZII zoea II, zoeal stage II
II




List of figures
Figure 1.1: Map of Antarctica and the Southern Ocean. .................................................................... 2
Figure 1.2: Oxygen- and capacity-limited thermal tolerance in ectothermic animals. ................ 4
Figure 1.3: Magnesium hypothesis for the biogeography of marine decapod crustaceans i