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reported E. huxleyi to double its calcii cation in
response to p CO 2 increasing from 280 to 750 μatm.
However, the difference can be explained by a size
difference in the cells incubated in the different CO 2
treatments. Cells grown at high CO 2 had an initial
biomass two to three times greater than low CO 2 -
grown cells (possibly due to differences in pre-
cultures). Due to the difference in cell size, however,
a comparison between CO 2 treatments on a per cell
basis is meaningless, but can only be done on a per
biomass basis. In fact, when normalized to algal
biomass, the trends in calcii cation and primary
production with increasing p CO 2 disappear (see
Riebesell et al. 2008a). A re-analysis of the same
strain of E. huxleyi (NZEH) used by Iglesias-
Rodriguez et al. ( 2008 ) by Hoppe et al. (pers. comm.)
revealed no effect on growth rate and a moderate
decrease in calcii cation with increasing p CO 2 .
In a study by Shi et al. ( 2009 ) both growth rate
(cell division rate) and the cellular POC and PIC
content of E. huxleyi (strain NZEH) was higher at
pH T 7.8 compared with pH T 8.1, yielding higher
rates of organic carbon production and calcii cation
at elevated p CO 2 . The PIC:POC ratio was slightly
lower in cultures maintained at lower pH levels. As
discussed above, increased carbon cell quota and
cell size are frequently observed in coccolithophores
at elevated p CO 2 . However, the results reported by
Shi et al. (2009) differ from all other studies on coc-
colithophores in showing an increased cell division
rate at elevated p CO 2 . The signii cance of this i nd-
ing is difi cult to assess, partly because that study
was based on only two p CO 2 levels and was
observed in only two out of the three CO 2 manipu-
lation approaches, with the opposite trend when
carbonate chemistry manipulation was done
through CO 2 bubbling. Using the same strain of
E. huxleyi , Hoppe et al. (pers. comm.) found no
effect on growth rate.
In summary, although much of the work on
CO 2 /pH sensitivity focuses on coccolithophores,
evidence currently available suggests that ocean
acidii cation will cause a decline in CaCO 3 produc-
tion in most planktonic calcii ers. It is currently
unknown whether a decreased calcii cation rate will
affect the competitive i tness of calcifying organisms
relative to their non-calcifying competitors and to
what extent CO 2 -sensitive calcifying organisms will
6.3.4 Calcii cation
CaCO 3 is one of the most common building materials
used in the formation of skeletons, shells, and other
protective structures in the marine biota. Organisms
exploit the supersaturation with respect to CaCO 3 in
the surface ocean, which prevents crystallized CaCO 3
from dissolving. Calcii cation, the precipitation of
CaCO 3 , is facilitated by high pH and high carbonate
ion (CO 3 2- ) concentration (see Box 1.1 in Chapter 1). In
calcifying organisms these conditions are achieved at
the site of calcii cation through energy-consuming
ion transport processes (Mackinder et al. 2010 ). With
ocean acidii cation causing a decrease in pH and
[CO 3 2- ], the energetic cost of calcii cation is thought to
increase. The extra energy needed to compensate for
these changes in seawater carbonate chemistry
depends on the specii c pathways employed in
CaCO 3 precipitation, the details of which are cur-
rently poorly understood but which are likely to dif-
fer between taxonomic groups.
Most planktonic calcifying organisms tested so
far show a decrease in calcii cation in response to
elevated CO 2 /reduced pH ( Table 6.4 ), such as
foraminifera, pteropods, and planktonic larvae of
echinoderms. A wide range of responses to ocean
acidii cation was obtained for coccolithophores
( Table 6.4 ). Whereas calcii cation in E. huxleyi ,
G. oceanica , and Calcidiscus quadriperforatus decr-
eases to varying degrees with increasing p CO 2 ,
Calcidiscus leptoporus shows an optimum curve with
reduced calcii cation at p CO 2 levels below and
above present conditions and Coccolithus pelagicus/
braarudii appears to be insensitive to elevated p CO 2 .
In a comparison of different strains of E. huxleyi
Langer et al. (2009) observed either no change or a
decrease in calcii cation rate with increasing p CO 2 .
In all studies on coccolithophores the ratio of CaCO 3
to organic matter production (PIC:POC; PIC being
particulate inorganic carbon and POC particulate
organic carbon) decreases or remains unchanged
with elevated p CO 2 . No interacting effects of p CO 2
and temperature were observed on calcite produc-
tion, coccolith morphology, or on coccosphere size
by De Bodt et al. ( 2010 ).
Two recent studies on coccolithophores appear to
suggest a stimulating affect of ocean acidii cation
on calcii cation. Iglesias-Rodriguez et al.
( 2008 )
 
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