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Photo (by Elke S. Franke) shows fertilized 1-cell stage egg of E. chloroticus Egg diameter approx. 80 micrometers
Aspects of Fertilization Ecology in
Evechinus chloroticus and
Coscinasterias muricata
Elke Sabine Franke
Abstract
Broadcast spawning marine invertebrates have been widely used as model organisms to study processes of evolution. One of these is the study of various life history stages associated with reproduction. Fertilization ecology in broadcast spawning marine organisms, i.e. the process by which sperm and egg fusion occurs once released into the ocean, has been the subject of intensive study for roughly the last 20 years, and represents thus a recent field in ecological sciences. This growth in interest was sparked by studies that showed spawning events may take place predominantly under sperm limiting conditions. More recent findings however, suggest that the occurrence of sperm competition, and the risk of polyspermy (multiple sperm entry in to the egg, which is lethal in echinoderms) can also occur during reproductive events in marine invertebrates.
This has also been predicted on theoretical grounds. Even though polyspermy has been observed previously, particularly during aquaculture studies, evidence to assess the occurrence of polyspermy in situ, and the conditions under which it occurs is lacking.
Simulated field studies in Evechinus chloroticus as well as laboratory studies in E.
chloroticus and Coscinasterias muricata found high levels of polyspermy, even under the sperm limiting conditions that are naturally found in the field in other marine broadcast spawners. Furthermore, laboratory results in both Evechinus and Coscinasterias showed that polyspermy is most likely to increase when sperm concentrations are increased. In addition, even though increasing sperm concentrations increases monospermy to a certain extent, an increase in gamete contact times generates similar levels of monospermy.
Results from these studies confirmed the mathematical model predicting some of these events.
i
The timing of gamete release to achieve maximum fertilization success (monospermy) in a broadcast spawner is thought to be under strong selective pressures, because reproductive success is directly related to fitness. The spawning patterns in the sea urchin E. chloroticus observed in this study demonstrated that E. chloroticus spawns under varying environmental and ecological conditions. Interestingly, it appeared that highly synchronous and widespread spawning was found to occur when highly turbulent conditions existed. This may reduce the potential for polyspermy that may exist during mass spawnings under shallow and calm conditions.
The demonstration of polyspermy and the confirmation of the mathematical model, suggest that polyspermy is a common and frequent occurrence during broadcast spawning events. Thus polyspermy represents an evolutionary force that may shape the evolution of reproductive phenomena at levels ranging from the gamete to the population.
Table of Contents
Chapter 1 General Introduction………...…….…………...…….1-50
Chapter 2 “Sexual Conflict and Polyspermy under Sperm-limited Conditions: In situ evidence from Field simulations with the free-spawning marine echinoid Evechinus chloroticus”……….….……51-86
Chapter 3 “Fertilization Kinetics and the Importance of Polyspermy in Fertilization Ecology of the sea urchin Evechinus chloroticus”………...………87-132
Chapter 4 “Reproductive Pattern of the New Zealand sea urchin Evechinus chloroticus”
.………...……….. 133-191
Chapter 5 “Fertilization Kinetics and the Importance of Polyspermy in the Broadcast- Spawning starfish Coscinasterias muricata”………...………192-235
Chapter 6 General Discussion……….……..………..236-251
Acknowledgements………..……..…..252-254
Appendix………..…255
iii
Table of Figures, Illustrations and Tables
Chapter 1
Picture of Evechinus chloroticus………..……….14
Picture of Coscinasterias muricata………..………..17
Chapter 2 Fig.1……….………..……69
Fig. 2 ………...………..70
Fig. 3………..………71
Fig. 4………..72
Table 1………..…74-76 Chapter 3 Fig. 1………...……….114
Fig. 2………115
Fig. 3………...…….116
Fig. 4………..…..117
Fig. 5………118
Fig. 6………..………..119
Fig. 7………120
Fig.8……….121
Table 1……….123
Table 2………...124
Chapter 4
Fig. 1………..………..166
Fig. 2………167
Fig. 3………...……….168
Fig. 4………..…..169
Fig. 5………...…….170
Fig. 6………171
Fig. 7………172
Fig. 8………..……...173-174 Fig. 9………...…….175
Table 1……….179
Table 2……….179
Table 3………...……..179
Table 4………...…..180
Table 5………...………..181
Table 6………...…………..182
Chapter 5 Fig. 1………215
Fig. 2 A………...…….216
Fig. 2 B………...….217
Fig. 3………218
Fig. 4………219
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Fig. 5……….…….220 Fig. 6………..….…..221-225 Table 1……….227 Table 2……….228 Table 3……….…229