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Fertilization Success of an Arctic Sea Urchin Species

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Fertilization Success of an Arctic Sea Urchin Species EGU Journal Logos (RGB) Open Access Open Access Open Access Advances in Annales Nonlinear Processes Geosciences Geophysicae in Geophysics Open Access Open Access Natural Hazards Natural Hazards and Earth System and Earth System Sciences Sciences Discussions Open Access Open Access Atmospheric Atmospheric Chemistry Chemistry and Physics and Physics Discussions Open Access Open Access Atmospheric Atmospheric Measurement Measurement Techniques Techniques Discussions Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Open Access Biogeosciences Discuss., 10, 8027–8064, 2013 Open Access www.biogeosciences-discuss.net/10/8027/2013/ Biogeosciences Biogeosciences BGD doi:10.5194/bgd-10-8027-2013 Discussions © Author(s) 2013. CC Attribution 3.0 License. 10, 8027–8064, 2013 Open Access Open Access This discussion paper is/has been under review for the journal BiogeosciencesClimate (BG). Climate Fertilization success Please refer to the correspondingof finalthe Past paper in BG if available. of the Past Discussions of an arctic sea urchin species Open Access Open Access Earth System Earth System D. Bogner¨ et al. Fertilization successDynamics of an arcticDynamics sea Discussions urchin species, Strongylocentrotus Title Page Open Access Geoscientific Geoscientific Open Access droebachiensisInstrumentation(O. F. Muller,¨ 1776)Instrumentation under Abstract Introduction Methods and Methods and CO2-inducedData ocean Systems acidificationData Systems Conclusions References Discussions Open Access Open Access Tables Figures ¨ 1 1 ¨ 1,2 Geoscientific D. Bogner , U. BickmeyerGeoscientific, and A. Kohler Model Development 1 Model Development J I Alfred Wegener Institute Helmholtz Center for Polar and Marine Research, Bremerhaven,Discussions Germany 2 J I Open Access Jacobs University Bremen, Bremen, GermanyOpen Access Hydrology and Hydrology and Back Close Received: 20 March 2013 –Earth Accepted: System 21 March 2013 – Published: 14Earth May System 2013 Correspondence to: D. Bogner¨ ([email protected])Sciences Sciences Full Screen / Esc Discussions Open Access Published by Copernicus Publications on behalfOpen Access of the European Geosciences Union. Printer-friendly Version Ocean Science Ocean Science Discussions Interactive Discussion Open Access Open Access 8027 Solid Earth Solid Earth Discussions Open Access Open Access The Cryosphere The Cryosphere Discussions Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Abstract BGD Sea urchins as broadcasting spawners, release their gametes into open water for fer- tilization, thus being particularly vulnerable to ocean acidification. In this study, we as- 10, 8027–8064, 2013 sessed the effects of different pH scenarios on fertilization success of Strongylocen- 5 trotus droebachiensis, collected at Spitsbergen, Arctic. We achieved acidification by Fertilization success bubbling CO2 into filtered seawater using partial pressures (pCO2) of 180, 380, 980, of an arctic sea 1400 and 3000 µatm. Untreated filtered seawater was used as control. We recorded urchin species fertilization rates and diagnosed morphological aberrations after post-fertilization peri- ods of 1 h and 3 h under different exposure conditions in experiments with and without D. Bogner¨ et al. 10 pre-incubation of the eggs prior to fertilization. In parallel, we conducted measurements of intracellular pH changes using BCECF/AM in unfertilized eggs exposed to a range of acidified seawater. We observed increasing rates of polyspermy in relation to higher Title Page seawater pCO2, which might be due to failures in the formation of the fertilization en- Abstract Introduction velope. In addition, our experiments showed anomalies in fertilized eggs: incomplete 15 lifting-off of the fertilization envelope and blebs of the hyaline layer. Other drastic malfor- Conclusions References mations consisted of constriction, extrusion, vacuolization or degeneration (observed Tables Figures as a gradient from the cortex to the central region of the cell) of the egg cytoplasm, and irregular cell divisions until 2- to 4-cell stages. The intracellular pH (pHi) decreased significantly from 1400 µatm on. All results indicate a decreasing fertilization success J I 20 at CO2 concentrations from 1400 µatm upwards. Exposure time to low pH might be J I a threatening factor for the cellular buffer capacity, viability, and development after fer- tilization. Back Close Full Screen / Esc 1 Introduction Printer-friendly Version Ocean acidification (OA) is largely dependent on increasing carbon dioxide (CO2) emis- Interactive Discussion 25 sions into the atmosphere due to anthropogenic activities. The concentration of CO2 in the atmosphere has a direct effect on the carbonate system of the oceans and thus, 8028 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | elevated CO2 levels modify the distribution of carbonate species (Zeebe and Wolf- Gladrow, 2001), putatively compromising the life of calcifying animal groups. The IPCC- BGD Report (2007) states that by the middle of this century the CO level could reach more 2 10, 8027–8064, 2013 than 500 ppm and by 2100 could be over 800 ppm, which means in terms of pH, that 5 the ocean will be 100–150 % more acidic than at the beginning of the industrial revo- lution (around 280 ppm). Cao and Caldeira (2008) predicted a decrease in pH of 0.25 Fertilization success and 0.48 units when atmospheric CO2 reaches 550 and 1000 ppm, respectively. of an arctic sea Sea urchins and other broadcasting spawning species might be particularly vulner- urchin species able to ocean acidification, especially during critical phases of their life cycle, like fertil- D. Bogner¨ et al. 10 ization and posterior larval development. Therefore, it is of major relevance to analyze whether gametes released into the acidified environment are protected or not, in order to forecast the potential effects of predicted pH changes on key ecological species. Title Page Studies on OA effects on different species have already been done: in fishes (Crim et al., 2011; Melzner et al., 2009; Checkley et al., 2009; Ishimatsu et al., 2004), mol- Abstract Introduction 15 luscs (Navarro et al., 2013; Thiyagarajan and Ko, 2012; Dineshram et al., 2012; Parker et al., 2009; Gazeau et al., 2007), crustaceans (Pansch et al., 2012; Li and Gao, 2012; Conclusions References Fitzer et al., 2012; Findlay et al., 2010; Kurihara and Ishimatsu, 2008), corals (Movilla Tables Figures et al., 2012; Fine and Tchernov, 2007; Pelejero et al., 2005) and some sea urchin species (Evans et al., 2013; Albright et al., 2012; Foo et al., 2012; Reuter et al., 2011; J I 20 Martin et al., 2011; Stumpp et al., 2011b; Dupont et al., 2010; Byrne et al., 2010; Clark et al., 2009; Havenhand et al., 2008). J I Sea urchin gametes are widely used models in genomic, embryology and toxicol- Back Close ogy research (Dupont et al., 2010; Epel et al., 2006; Wong and Wessel, 2005; Wessel et al., 2000; Epel, 1998) and are of high value in studies on development and ocean Full Screen / Esc 25 acidification effects. To our knowledge, there are no studies on effects of OA at the cel- lular level upon fertilization and this study is the first in identifying the effects of external Printer-friendly Version pH changes, based on CO2, on intracellular pH (pHi) of eggs before fertilization. Until now published, results on fertilization experiment using different sea urchins species Interactive Discussion under acidified conditions are contradictory, mostly due to the variety of experimental 8029 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | approaches. The use of different fertilization protocols, limits the comparability of re- sults on OA effects. Therefore, the critical comparison of data should take into account BGD different methodologies and endpoints of effects measured (Reuter et al., 2011). 10, 8027–8064, 2013 Moreover, no data exists on the sea urchin species Strongylocentrotus droebachien- 5 sis from the Arctic. The study of arctic populations offers the possibility to identify ef- fects of OA on reproduction of animals living in a geographic area, where low temper- Fertilization success ature enhances dissolution of carbon dioxide into seawater. The green sea urchin S. of an arctic sea droebachiensis is an important key species that stabilizes community structure and is urchin species part of established food webs all along its life cycle (Dupont et al., 2010; Addison and D. Bogner¨ et al. 10 Hart, 2004; Scheibling and Hatcher, 2001). S. droebachiensis is widely distributed in northern circumpolar waters (Pacific and Atlantic coasts of North America and Arctic Ocean) (Scheibling and Hatcher, 2001; Levitan, 1998a) within a wide range of physi- Title Page cally different habitat conditions in terms of temperature and depth. Cyclic environmen- tal changes might influence the acclimation history of animal species within different Abstract Introduction 15 populations expanding their salinity and pH tolerance. In Kongsfjorden, Svalbard, the seasonal melting cycles produce changes in transparency of water masses and salin- Conclusions References ity (Svendsen et al., 2002) which may also determine seasonal pH changes due to Tables Figures photosynthetic activity. The spring bloom is the only predictable phytoplankton produc- tion peak of the Kongsfjorden (Hop et al., 2002). In Norwest Atlantic, spawning of S. J I 20 droebachiensis takes place in early spring and is triggered by phytoplankton blooms (Scheibling and Hatcher, 2001). J I Reproduction
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