Will krill fare well under Southern Ocean acidification?
So Kawaguchi
Haruko Kurihara
Robert King
Lillian Hale
Thomas Berli
James P. Robinson
Akio Ishida
Masahide Wakita
Patti Virtue
Stephen Nicol
Atsushi Ishimatsu
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Biol. Lett. (2011) 7, 288291
doi:10.1098/rsbl.2010.0777
Published online 13 October 2010
Global change biology
Will krill fare well under
Southern Ocean
acidification?
So Kawaguchi1,2,*, Haruko Kurihara3,
Robert King1, Lillian Hale4, Thomas Berli4,
James P. Robinson4, Akio Ishida5,
Masahide Wakita5,6, Patti Virtue4,
Stephen Nicol1,2 and Atsushi Ishimatsu7
1Australian Antarctic Division, Kingston, Tasmania 7050, Australia
2Antarctic Climate and Ecosystems Cooperative Research Centre,
Sandy Bay, Hobart, Tasmania 7001, Australia
3University of the Ryukyus, Okinawa 903-0213, Japan
4University of Tasmania, Sandy Bay, Tasmania 7005, Australia
5Research Institute for Global Change, JAMSTEC, Yokosuka
237-0061, Japan
6Mutsu Institute for Oceanography, JAMSTEC, Mutsu
035-0022, Japan
7Nagasaki University, Nagasaki 851-2213, Japan
*Author for correspondence ().
Antarctic krill embryos and larvae were
experimentally exposed to 380 (control), 1000 and
2000 matm pCO2 in order to assess the possible
impact of ocean acidification on early
development of krill. No significant effects were
detected on embryonic development or larval
behaviour at 1000 matm pCO2; however, at
2000 matm pCO2 development was disrupted
before gastrulation in 90 per cent of embryos,
and no larvae hatched successfully. Our model
projections demonstrated that Southern Ocean
sea water pCO2 could rise up to 1400 matm in
krills depth range under the IPCC IS92a
scenario by the year 2100 (atmospheric pCO2
788 matm). These results point out the urgent
need for understanding the pCO2-response
relationship for krill developmental and later
stages, in order to predict the possible fate of
this key species in the Southern Ocean.
1. INTRODUCTION
The ecosystems of the Southern Ocean are expected to
be most severely affected by ocean acidification (OA)
because of the higher solubilities of CO2 and CaCO3
in cold waters and because of regional upwelling of
hypercapnic deep sea water [1,2]. Moreover, a future
rise in surface water pCO2 may be augmented at
great depths [3], where sea water pCO2 is already
much higher than at the surface ([4]; figure 1).
Hence, vertically migrating animals in the Southern
Ocean will probably experience the most drastic
changes in carbonate chemistry in future oceans.
However, OA research has mainly dealt with tropical and
temperate shallow-water calcifying organisms [1], and
little attention has been paid to polar species [5].
Electronic supplementary material is available at http://dx.doi.org/
10.1098/rsbl.2010.0777 or via http://rsbl.royalsocietypublishing.org.
Antarctic krill (Euphausia superba, hereafter krill) is
the key species of the Southern Ocean ecosystem,
and is found in a range of water depths. Krill spawn
eggs at the surface which sink to 700 1000 m before
larvae hatch to swim back to the surface [6]. The
post-larval vertical distribution ranges from the surface
to at least 3500 m ([7]; figure 1). Thus, krill are already
exposed to high CO2 conditions at depth, which will
probably become far more hypercapnic than surface
waters (electronic supplementary material, S1).
The purpose of this study is to examine how
elevated CO2 conditions affect krill. We focused on early
developmental stages, since larvae and juveniles are
generally more vulnerable to environmental
perturbations, and their survival will largely determine
population abundance, distribution and community
structure [8].
2. MATERIAL AND METHODS
The stock population of krill was collected from the Indian Ocean
sector of the Southern Ocean between January and March in
the 20052006 field season [9]. The krill were maintained in the
Australian Antarctic Divisions marine research aquarium, where
they matured and spawned naturally [10].
(a) Experimental set-up
Experimental sea water was supplied from a 70 l header tank and
equilibrated with air (control) or CO2-enriched air before being
delivered to experimental jars (250 ml clear polycarbonate)
containing krill eggs (see electronic supplementary material, S2). The
CO2-enriched air was prepared with a mass flow controller
(Horiba STEC SEC-E-40) and by an air valve, to regulate flow
rates of pure CO2 and atmospheric air, respectively. The pCO2
levels of the CO2-enriched air and sea water were monitored by a
CO2 monitor (Telaire 7001) and indirectly from pH measurement
(Radiometer PHM 210 pH metre), respectively. Experimental
temperature was set at 0.58C. Effluent from each jar was drained into a
70 l sump, and recirculated through a degassing unit before
returning back to the header tank via a filtration and cooling system. For
details, see Kawaguchi et al. [10]. Total alkalinity was measured
through a two-stage, potentiometric, open-cell titration. The
carbonate chemistry of the experimental sea water is summarized in the
electronic supplementary material, S3.
(b) Hatching experiment
Fertilized eggs were obtained in January 2008 and 2009. In the 2008
experiments, three batches of eggs originating from three different
females were used. Each batch was randomly distributed into
experimental jars, with approximately 2030 eggs per jar. The embryos
were incubated at one of the three target CO2 levels: control
(380 matm), medium (1000 matm) or high (2000 matm). In the
2009 experiments, four batches of eggs were incubated as in 2008.
Hatch rates were determined for each jar after 710 days of
spawning. The number of jars in each treatment is summarized in table 1.
Embryonic stages were classified at the end of each 2009 experiment
after George [11].
(c) Observation of larval swimming activity
Three of the four batches in the hatching experiments in 2009 (S2,
W2 and Y2) were used for this observation. R2 was not used because
of the limited capacity of the set-up. CO2 exposure started within 1
day of spawning and continued throughout the experimental period.
Larval behaviour was observed on an average of 3 days after
hatching, when they were in the nauplius stage. Since almost no eggs
hatched in 2000 matm (table 1), observations were made only for
the 380 and 1000 matm groups.
(d) Statistical tests
Statistical tests were performed using SPLUS v. 8 software.
3. RESULTS
The hatch rates of control eggs used in our experiment
were highly variable (16.7 74.7%, table 1) but the
range is in fact comparable to the rates in field
experiments (0 89% [12,13]). There were significant
Krill under ocean acidification
S. Kawaguchi et al. 289
aNumber of replicates.
)
m
(
th 2000
p
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