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JOURNAL OF GEOPHYSICAL RESEARCH, VOL. 117, G01020, doi:10.1029/2011JG001849, 2012

Carbon and oxygen cycles: Sensitivity to changes in environmental

forcing in a coastal upwelling system
L. Bianucci1,2 and K. L. Denman1,3,4
Received 31 August 2011; revised 6 December 2011; accepted 10 December 2011; published 22 February 2012.
[1] Biogeochemical cycles in the coastal ocean are changing and will continue to change
in response to a changing climate. Effects on the oxygen and carbon cycles are particularly
important, as either episodic or permanent shifts toward lower oxygen and/or higher
inorganic carbon conditions can impact coastal ecosystems negatively. Here we study the
sensitivity of these cycles to changes that may occur in the coastal ocean, focusing on a
summer wind-driven upwelling region off southern Vancouver Island shelf. We use a quasi
2-D configuration of the Regional Ocean Modeling System (ROMS) to perform six
sensitivity experiments. Results indicate that carbon and oxygen cycles in this region may
be significantly affected by an altered upwelling season, a shallower offshore Oxygen
Minimum Zone, and a carbon-enriched environment. Combinations of these scenarios
suggest a potentially increasing risk for the development of coastal hypoxia and corrosive
conditions in the region.
Citation: Bianucci, L., and K. L. Denman (2012), Carbon and oxygen cycles: Sensitivity to changes in environmental forcing in
a coastal upwelling system, J. Geophys. Res., 117, G01020, doi:10.1029/2011JG001849.

1. Introduction [3] Oxygen Minimum Zones (OMZ) are permanently

hypoxic regions of the open ocean (O2 < 60 mmol m3),
[2] Future climate change will affect every aspect of the
typically along the continental margins of the eastern Pacific,
ocean, from its physics to its chemistry and biology. In
eastern Atlantic, and Indian oceans [Helly and Levin, 2004;
particular, biogeochemical cycles will not only be altered
Stramma et al., 2008]. Time series of O2 show a persistent
directly by the rising partial pressure of carbon dioxide
decline of concentrations in different OMZs as well as a
(pCO2) in the atmosphere, but also by indirect effects, such
shoaling of their upper boundaries [Whitney et al., 2007;
as higher temperatures and changes in wind strength and
Stramma et al., 2008]. These changes could impact shallower
patterns [e.g., Bakun, 1990; Falkowski et al., 2000]. For
waters in regions where OMZs affect outer continental
instance, the increase in sea surface temperatures and the shelves and upper slopes [Helly and Levin, 2004], especially
strengthening of stratification as global climate warms is
where wind-driven upwelling brings deep waters closer to or
expected to be sufficient to intensify existing hypoxia and
onto the shelves (e.g., western North America). The shoaling
generate hypoxia in new areas [Rabalais et al., 2010]. In
of the OMZ in the northeast Pacific [Whitney et al., 2007]
addition, if the biotic carbon to nitrogen ratio (C:N) increa-
may be a cause of recently observed hypoxic events in the
ses under elevated CO2 conditions (as observed in a meso-
coastal waters off Oregon and California [Grantham et al.,
cosm experiment [Riebesell et al., 2007]), anthropogenic
2004; Bograd et al., 2008; Chan et al., 2008], which in
CO2 emissions may extend tropical oxygen (O2) deficient
some cases turned the shelves into “dead zones” due to the
zones [Oschlies et al., 2008]. Models project a decline in O2
widespread mortality of benthic animals.
concentrations in the oceans during the 21st century
[4] Cycling of carbon in the ocean is also expected to
[Frölicher et al., 2009] and over the next 100,000 years change as pCO2 increases in the atmosphere and climate
[Shaffer et al., 2009]. Recent warming in the 1990s has
changes. A decline in pH and an increase in surface pCO2
already produced an estimated global oceanic O2 outgassing
have already been observed at several oceanic time series
of 0.3
0.4 1014 mol-O2 yr1 [Keeling and Garcia, 2002]. sites: stations ALOHA in the central North Pacific [Dore
et al., 2009] and BATS, off Bermuda in the North Atlan-
1 tic (both time series longer than 20 years [Bates, 2007]), as
School of Earth and Ocean Sciences, University of Victoria, Victoria,
British Columbia, Canada. well as station ESTOC (100 km north off Gran Canaria Island
2
Now at Department of Oceanography, Dalhousie University, Halifax, in the North Atlantic [Santana-Casiano et al., 2007]), which
Nova Scotia, Canada.
3
has a 10 yearlong record. These stations show that surface
Canadian Centre for Climate Modeling and Analysis, Environment ocean pCO2 has increased at rates indistinguishable from the
Canada, Victoria, British Columbia, Canada.
4
Now at VENUS Project, University of Victoria, Victoria, British
atmospheric increase (1.5 to 1.9 matm yr1 [Bindoff et al.,
Columbia, Canada. 2007]). On the west coast of the US, model simulations indi-
cate a pH decrease of 0.1 since pre-industrial times [Hauri
Copyright 2012 by the American Geophysical Union. et al., 2009]. Moreover, corrosive waters have been observed
0148-0227/12/2011JG001849

G01020 1 of 13

G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING
Figure 1. Map of western North America showing the
location of the southern Vancouver Island shelf. The inset
shows the transect that the quasi 2-D model represents
(black line) and the meteorological buoy 46202 that pro-
vided winds to force the model (triangle). NCEP data used
to force surface net heat and shortwave fluxes are represen-
tative of a region of 1.9° latitude  2.4° longitude centered
on the solid black circle.
on shelves off western North America [Feely et al., 2008].
There, the undersaturation horizon of aragonite (the less
stable form of calcium carbonate, CaCO3, found in corals for
example) was observed at depths between 40 and 120 m,
even reaching the surface along one transect off northern
California.
[5] In contrast to other coastal regions of western North
America, wide spread hypoxic events and corrosive waters
have not yet been observed along the west coast of Van-
couver Island. This region represents the northern limit of
the California Current System (CCS) in Pacific Canada. The
outflow from the Juan de Fuca Strait, relatively fresh and
nutrient-rich due to intense tidal mixing in the Strait, gen-
erates a buoyancy-driven coastal current, known as the
Vancouver Island Coastal Current (VICC) [Freeland et al.,
1984; Thomson et al., 1989; Crawford and Dewey, 1989].
This O2-rich current and the relatively wide shelf protect the
shallower waters from developing hypoxia and aragonite
undersaturation [Bianucci et al., 2011]. Moreover, wind-
driven upwelling only occurs during summer [Strub et al.,
1987a, 1987b], such that the influence of carbon-rich and
O2-depleted deep waters from offshore is limited to that
season (winds are downwelling-favorable during winter).
However, it is uncertain if hypoxia and undersaturation may
develop in the future, given potential alterations to envi-
ronmental forcing with climate change. Projections for the
21st century from an earth system model have predicted a
decrease in O2 concentrations and increase in ocean acidi-
fication in the entire CCS [Rykaczewski and Dunne, 2010].
Therefore, we use a coastal circulation model to investigate
the sensitivity of the carbon and O2 cycles to different for-
cings in our region of interest. A quasi 2-D configuration
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G01020
represents summer upwelling over the southern Vancouver
Island shelf. Six experiments (details in section 2) investi-
gate the model sensitivity to changes in strength of upwell-
ing, depth of the OMZ, and inorganic carbon concentration.
We introduce to our analysis the concept of the Respiration
Index [Brewer and Peltzer, 2009a, section 3], which simul-
taneously considers the effects of changes in both O2 and
pCO2 on marine organisms. This index is calculated in the
different simulations and compared with the traditional
definitions of hypoxia.
2. Design of Sensitivity Experiments
[6] We have developed a quasi 2-D application of the
Regional Ocean Modeling System (ROMS, www.myroms.
org [Haidvogel et al., 2008]) to model upwelling on the
Vancouver Island shelf in response to time variable wind-
forcing. The domain is a transect perpendicular to the iso-
baths (i.e., cross-shore distance versus depth) at 49°N
(Figure 1) and the small alongshore dimension (5 km) has
uniform conditions. A summary of the model configuration
is provided in Table 1. This quasi 2-D approach precludes
the inclusion of an alongshore pressure gradient that varies
with cross-shelf distance and depth, preventing the modeling
of a dynamic VICC [Masson and Cummins, 1999]. Hence,
to model the VICC, water properties in the nearshore 6 km
are restored toward a fixed vertical profile representative of
the core of the VICC. The restoring is strongest at the
inshore boundary and decreases as exp(x2) toward off-
shore, where x is the distance from the inshore boundary (for
more details, see Bianucci et al. [2011, Appendix B]. The
model configuration and evaluation, as well as the bio-
logical and sediment modules coupled to the physical
model, are described in detail elsewhere [Bianucci, 2010;
Bianucci et al., 2011]. Vertical profiles represented cor-
rectly the observed vertical structure of temperature, nitrate,
and other variables. Moreover, the statistical properties of the
model and observations were in agreement. Modeled time
series of sea surface height captured some of the surface
dynamics observed at a meteorological buoy in the region.
The model also represented the observed cross-shore gradi-
ent of DIC [Ianson et al., 2003], such that VICC waters are
enriched in DIC with respect to shelf waters at comparable
depths.
[7] Coastal upwelling ecosystems are intrinsically 3-D
systems, where mesoscale phenomena play an important
role [e.g., Gruber et al., 2006; Lathuilière et al., 2010].
However, a 2-D model is able to represent locally forced
upwelling and facilitates extensive sensitivity analysis.
Table 1. Summary of Model Configuration
Configuration
Model dimensions Cross-shore (Lx) = 185 km.
Alongshore (Ly) = 5 km.
Vertical (H) = from 40 to 1500 m.
Grid resolution Cross-shore (dx) = 953 m
Alongshore (dy) = 1.67 km.
Vertical (dz) = from 1.3 to 72.8 m
(30 s-layers).
Time step dt = 320 sec.
Spin-up time 50 days.
Total length of experiments 125 days.
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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

Table 2. Description of Model Sensitivity Experiments

Experiment Description and Characteristic Valuesa
1 Control experiment with 1993 late spring and summer forcing from 27 May 1993 (day 0) to 29 September 1993 (day 125).
2 Like 1, but with 1993 wind stress during spin-up (day 0 to 50) and 2002 wind stress (increased upwelling) after day 50 (16 July).
Mean t y = 0.028 N m2 (0.018 N m2)
3 Like 1, but with 2002 wind stress from day 0.
Mean t y = 0.023 N m2 (0.018 N m2)
4 Like 1, but initial conditions for O2 with a shallower OMZ and lower O2 concentrations.
Min. initial O2 at 800 m = 1.3 mmol m3 (10.6 mmol m3)
OMZ hypoxic threshold depth = 280 m (380 m)
5 Like 4, but with 2002 wind stress from day 0.
Min. initial O2 at 800 m = 1.3 mmol m3 (10.6 mmol m3)
OMZ hypoxic threshold depth = 280 m (380 m)
Mean t y = 0.023 N m2 (0.018 N m2)
6 Like 1, but with higher pCO2atm and DIC initial conditions (expected 2000 to 2050 changes from CanESM 1.1 A2 simulation).
CanESM 1.1 is the Canadian Earth System Model version 1.1 [Arora et al., 2009; Christian et al., 2010].
Max. initial DIC = 2291 mmol m3 (2273 mmol m3)
pCO2atm = 513 ppmv (370 ppmv)
a
Values in parentheses represent conditions in control experiment 1.

Previous studies on the Oregon shelf have successfully used
2-D models [Allen et al., 1995; Federiuk and Allen, 1995;
Spitz et al., 2003]. Here, a set of model experiments helps to
determine the sensitivity of the carbon and O2 cycles to
different forcing and changing conditions. By changing one
aspect of the model at a time, we compare the differences
with respect to a control simulation (experiment 1). The
sensitivity experiments are described below and summarized
in Table 2. Surface incoming shortwave radiation and net
heat fluxes are specified in all experiments from NCEP
reanalysis as daily values for the period 27 May to 29
September 1993, at 48.57°N, 125.62°W. Wind stress is cal-
culated from observed hourly winds at meteorological
buoy 46206 (48.83°N, 126.00°W, Figure 1) following Smith
[1988], then filtered with a 6-hour low-pass Fast Fourier
Transform (FFT) filter. Atmospheric pCO2 concentration
(pCO2atm) is set as a constant boundary condition (370 ppmv,
except in experiment 6). To provide sufficient time for
spin-up, since sediments take more than a month to
approach equilibrium [Bianucci et al., 2011], the experi-
ments start on 27 May 1993 (day 0) and analyses begin on
16 July (day 50).
2.1. Control Experiment (Experiment 1)
[8] Wind and surface heat forcing corresponds to late
spring and summer 1993, a year representing a “normal
upwelling summer”: upwelling indices for July and August
(38 and 26 m3 s1 per meter of coastline, respectively)
were close to the climatological monthly averages (34 and
22 m3 s1 per meter of coastline; upwelling indices from
the Environmental Research Division, Pacific Fisheries
Environmental Laboratory). Observed deep ocean summer
profiles from the study region are used to create average
depth profiles to initialize scalar properties (i.e., horizon-
tally uniform distributions), and initial velocities are set to
zero.
2.2. Stronger Upwelling Experiments (Experiments 2
and 3)
[9] Increasing trends in upwelling intensity have been
observed in some major coastal upwelling systems of the
world during the 20th Century [e.g., Bakun, 1990; Schwing
and Mendelssohn, 1997; McGregor et al., 2007]. Intensified
summer upwelling due to increased atmospheric pCO2 is
predicted from a regional climate model with high resolution
over the coastal region of California [Snyder et al., 2003]. Off
the Vancouver Island shelf, an ensemble of 18 climate models
predicts increased upwelling summer winds in the 21st century
[Merryfield et al., 2009]. Therefore, we test the sensitivity of
the O2 and carbon cycles in the model to increased upwelling.
The summer of 2002 experienced higher than normal
upwelling-favorable winds off Vancouver Island. Mean
alongshore wind stress was 28 % greater in 2002 than in 1993
for the period 27 May to 29 September (0.018 N m2 in
1993 versus 0.023 N m2 in 2002; see Figure 2a). For the
period 16 July to 29 September (the period of analysis fol-
lowing 50 days of spin-up), mean alongshore wind stress
almost tripled in 2002 (0.026 N m2) relative to 1993
(0.009 N m2). The simulations with stronger upwelling are
forced with 2002 wind stress in two ways: experiment 2 has
the same spin-up as the control experiment (first 50 days with
1993 wind stress), while experiment 3 is forced from the start
with 2002 winds. The first approach allows a comparison of
results after a common forcing during the 50 day spin-up
period; the second allows examining the effect of using dif-
ferent forcing during spin-up.
2.3. Shallower OMZ Experiments (Experiments 4
and 5)
[10] Although not all species have the same tolerance to
low O2 concentrations [Vaquer-Sunyer and Duarte, 2008],
hypoxia is commonly defined as waters with O2 <
60 mmol m3 (= 60 mM  60 mmol kg1  1.4 mL L1)
[Gray et al., 2002; Whitney et al., 2007; Stramma et al.,
2008]. The hypoxic threshold at Ocean Station Papa
(OSP) has shoaled roughly by 100 m (from 400 to 300 m
depth) between 1956 and 2006 [Whitney et al., 2007]. The
southern CCS has experienced a shoaling of up to 90 m in
the period 1984–2006 [Bograd et al., 2008]. Moreover, O2
concentrations in the OMZ are decreasing with observed
rates of 0.18 mmol-O2 m3 yr1 at a depth of 800 m at
OSP and 0.15 mmol-O2 m3 yr1 at a depth of 500 m in
the southern CCS region [Whitney et al., 2007; Bograd et al.,
2008]. To analyze the effects of a shallower and more intense
OMZ on the southern Vancouver Island shelf system
(experiment 4), we modify the initial O2 field such that the

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

Figure 2. (a) Alongshore wind stress for experiments 1 (1993, black) and 3 (2002, red). Experiment 2 is
forced by 1993 winds up to day 50 (vertical dashed grey line) and 2002 winds afterwards. (b) Initial
O2 profiles (used over the whole domain) for experiments 1 (black) and 4 (red), which coincide in the
upper 130 m. The OMZ hypoxic threshold depth is defined as the depth where open ocean waters
have O2 = 60 mmol m3 (hypoxic threshold, vertical dashed lined). (c) Profiles of decadal means
for dissolved inorganic carbon (DIC) centered on years 2000 and 2050 from the Canadian Earth Sys-
tem Model (CanESM 1.1). (d) Profile of the DIC difference between decadal means, which is added to
initial conditions in experiment 6.

hypoxic threshold is 100 m shallower than in the control
simulation. Moreover, the minimum concentration is
changed to 1.3 mmol-O2 m3, 9 mmol-O2 m3 lower than
the minimun in the control experiment (Figure 2b). This O2
decrease would be achieved in 50 years at the observed rate at
800 m at OSP. In experiment 5, in addition to the shallower
OMZ, the model is forced with the stronger upwelling winds
from 2002.
2.4. Higher Carbon Scenario Experiment
(Experiment 6)
[11] As CO2 increases in the atmosphere due to anthropo-
genic activities, roughly one third of emissions are absorbed
by the ocean [Sabine et al., 2004; Sabine and Feely, 2007].
Experiment 6 examines the effect of increased pCO2atm and
ocean dissolved inorganic carbon (DIC) on biogeochemical
cycles in the model. The increments correspond to the
predicted changes in DIC and pCO2atm over the period 2000
to 2050 from a global climate model (Figures 2c and 2d). We
use the Canadian Earth System Model (CanESM 1.1), with
emission scenario A2 [Arora et al., 2009; Christian et al.,
2010] at the model location closest to the Vancouver Island
shelf (50°N, 130°W). DIC concentrations in the VICC were
assumed to increase by the same amount as those over the
shelf. pCO2atm, which is set as a constant boundary condi-
tion, was increased by 143 ppmv relative to the control
experiment (from 370 to 513 ppmv).
3. Respiration Index (RI) and Aragonite
Saturation State (WA)
[12] Recently, Brewer and Peltzer [2009a] argued that
elevated pCO2 may impose a physiological strain on higher
animals, and that the use of an O2 limit alone to define a

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

Figure 3. Across-shelf distribution of temporally and vertically averaged (a, c) O2 and (b, d) pCO2 for
experiments 1 (blue) and 2 (red). Temporal average is for days 50 to 125, after the spin-up period. Vertical
averages are shown for the upper 30 m of the water column (Figures 3a and 3b) and from 30 m to the sea-
floor (Figures 3c and 3d). (e, f) The bathymetry, with a magenta vertical line indicating the edge of the
shelf break. The dashed grey line in Figure 3b indicates atmospheric pCO2 (370 ppmv).

dead zone implicitly assumes that pCO2 levels are low and
inversely proportional to O2. They defined a Respiration
Index that is linearly related to available energy in basic oxic
respiration (RI = log(pO2/pCO2)). The RI reflects the ther-
modynamic energy yield of aerobic respiration as the con-
centration ratio of substrate and product changes. This
concept suggests that, as atmospheric pCO2 rises and more
carbon is absorbed by the ocean, dead zones could expand
even if O2 levels were not affected [Brewer and Peltzer,
2009a]. RI = 1 can be considered as a general boundary
for aerobic stress, although Brewer and Peltzer [2009a]
pointed out that the actual limits will be species-dependent.
[13] Some of the assumptions behind this index aroused
controversy. For instance, the calculation assumes a closed
thermodynamic system, while living organisms are essen-
tially open systems [Seibel et al., 2009]. These authors also
were concerned about the use of environmental gas partial
pressures, arguing that the intracellular concentrations are
regulated independently by kinetic and physiological
mechanisms. Despite these criticisms, to which Brewer and
Peltzer [2009b] responded, we use the RI in the context of
our modeling study to evaluate the combined effect of O2
and CO2 as a threshold for habitable shelf environments,
compared with the more typical O2 thresholds.
[14] Aragonite is the less stable form of CaCO3 in the
ocean and its degree of saturation can be approximated as
WA  [CO2 2 2
3 ]/[CO3 ]sat where [CO3 ] and [CO3 ]sat
2
represent the concentrations of the carbonate ions in
ambient seawater and at saturation. When WA > 1 (<1),
seawater is supersaturated (undersaturated) with respect to
aragonite and favors calcification (dissolution). We will
use WA as well as pCO2 to describe the carbon state of the
system.
[15] WA is calculated from modeled temperature (T),
salinity (S), DIC, and total alkalinity (TA) using the
CO2SYS software [Lewis and Wallace, 1998]. When com-
paring changes in WA between two simulations, we calculate
the total change as
DWATotal ¼ ðWAn  WA1 Þ=WA1  100% ð1Þ
where the subscripts 1 and n indicate the control experiment
and any of the sensitivity simulations, respectively. To
evaluate the role that DIC alone plays in DWATotal, we com-
pute DWADIC =(W∗A  WA1)/WA1  100%, where W∗A is calcu-
lated with T, S, and TA from experiment 1 and DIC from
experiment n. DWADIC allows us to quantify the effect of
changing only DIC on WA1. The same procedure can be
performed for every variable (e.g., computation of DWAT,
DWATA, etc) or for combinations of variables (e.g., the com-
bined role of TA and DIC leads to DWATA+DIC).
4. Results
4.1. Effect of Increased Upwelling
[16] Temporal and vertical averages of O2 and pCO2
across the shelf are compared for the control experiment
(experiment 1) and experiment 2, which has increased

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING
Figure 4. Temporal evolution (Hovmöller plots) for total
water column primary production in experiments (a) 1 and
(b) 2. The dashed magenta line represents the location of
the shelf break. On the right, alongshore wind stress for each
experiment (upwelling/downwelling in red/blue) is shown;
the horizontal dashed line denotes day 50, the end of spin-
up period with common 1993 forcing. Primary production
units are g-C m2 d1.
upwelling (Figure 3, blue and red lines, respectively). The
upper 30 m of the water column experiences higher O2
concentrations under intensified upwelling over most of the
shelf (Figure 3a), while below 30 m depth these wind con-
ditions decrease O2 concentrations (Figure 3c). The O2
changes are up to +7 and 11% in the upper and lower
region of the water column, respectively. The inner shelf
concentrations are similar in both simulations due to the
restoring to VICC properties. On average, pCO2 in the upper
ocean is lower than pCO2atm (dashed grey line in Figure 3b)
over the mid- and outer shelf in both experiments, so air-sea
CO2 fluxes are from the atmosphere to the ocean in those
regions (surface pCO2 has a similar pattern but lower values
than the average over the upper 30 m). However, the high
DIC concentrations specified in the VICC lead to pCO2
concentrations higher than pCO2atm over the inner shelf.
CO2 outgassing occurs on the inner 4 km of the model
domain in experiment 1 and within 6 km in experiment 2.
[17] In experiment 2, pCO2 is higher in the bottom layers
(Figure 3d) and lower in the upper 30 m over most of the
model domain (Figure 3b), with changes reaching up to +4
and 7% in each case. However, there is a region of the
inner shelf (10–25 km from inshore boundary) where
pCO2 is higher in the upper layer under intensified upwell-
ing; moreover, approximately in the same region (10–
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G01020
18 km) O2 is lower under stronger upwelling relative to the
control experiment 1. These changes respond to the position
of phytoplankton blooms in both simulations, since phyto-
plankton are advected offshore near the surface during
intense upwelling events (Figure 4). Regression analysis
predicts a 20 km displacement of the maximum primary
production from the inshore boundary for a mean alongshore
equatorward wind stress of 0.05 N m2 in the previous
9 days [Bianucci, 2010]. The bulk of the primary production
(which increases by 12%) occurs farther offshore in experi-
ment 2, over the mid- and outer shelf (between 10 and
40 km, Figure 4). Moreover, high primary production over
the mid-shelf leads to more organic matter reaching the
seafloor at those depths, contributing to enhanced exchange
of O2 and DIC with the sediments. Between 20 and 30 km
from the inshore boundary (i.e., at bottom depths between 90
and 115 m) O2 and DIC exchanges with the sediments
increase up to 50% (the increment is 8% on average).
[18] As the development of hypoxia and/or corrosive
conditions first occurs in the bottom waters over the shelf,
we focus on the spatial and temporal distributions of O2 and
WA in the near-bottom layer of the model (Figure 5). This
layer accurately represents the bottom boundary layer,
except within the shallowest 3 km of the domain where
restoring to the VICC is strongest [Bianucci et al., 2011].
With increased upwelling as in experiment 2, low O2 con-
centrations develop in the bottom layer inshore of the shelf
break, reaching a minimum of 34 mmol-O2 m3 (Figure 5c).
In particular, the hypoxic boundary (60 mmol-O2 m3, bold
black contour) migrates inshore across the shelf after the
intense upwelling-favorable wind event centered on day 80
(see Figure 2a). The onset of hypoxia in shallower waters is
triggered by advection of low oxygen during that event (red
line in Figure 6). Once upwelling intensity decreases, bio-
logical consumption (especially in the sediments; black line
in Figure 6) maintains the low O2 levels achieved by
advection. The enhanced advection of low O2 results partly
from the stronger upwelling circulation and partly from the
lower O2 content of upwelled waters (the depth of upwelling
increases 40 m in experiment 2). Since advection and local
biological consumption generate the lowest O2 concentra-
tions in bottom waters, vertical mixing generates a down-
ward flux of O2 to the bottom layers from the overlying,
more oxygenated waters (blue line in Figure 6).
[19] The wind-forcing used during the first 50 days of the
experiments with stronger upwelling affects the results. An
intense upwelling event between days 35 to 45 in 1993
(first week of July) advects the hypoxic threshold closer
to the shelf break. Therefore, if 1993 winds are used during
spin-up (experiment 2, Figure 5c), waters on the shelf
become more hypoxic than if 2002 winds are used from day 0
(experiment 3, Figure 5e). From day 50 to 125, hypoxia
covers 43% of the near-bottom waters over the shelf in
experiment 2 compared with 18% in experiment 3. More-
over, the minimum O2 concentration is lower in experiment 2
relative to experiment 3 (34 versus 41 mmol-O2 m3).
Hence, the timing of the onset of the upwelling season is
another factor determining both the timing and magnitude of
hypoxia (and analogously, of events with lower aragonite
saturation state).
[20] Dissolved carbon concentrations in the near-bottom
layer over the shelf respond to the same processes as
 21562202g, 2012, G1, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2011JG001849 by Cochrane Philippines, Wiley Online Library on [04/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

Figure 5. Hovmöller plots for near-bottom (left) O2 and (right) WA for experiments (a, b) 1, (c, d) 2, (e) 3,
(f) 6, (g) 4 and (h) 5. Spin-up period (first 50 days) not shown. The bold black contours represent either the
hypoxic threshold (60 mmol-O2 m3) or the limit for aragonite dissolution (WA = 1). The bold yellow con-
tour is RI = 1 (area with RI < 1 is indicated) and the dashed magenta line represents the location of the
shelf break. The black dashed line in Figure 5c indicates the 90 m isobath; the dash-dotted yellow lines
in Figure 5g show the 130 and 90 m isobaths.

dissolved O2: DIC increases due to advection of high con- experiment 2, WA decreases in the near-bottom layer over the
centrations from offshore, while sediment remineralization shelf near day 90 (Figure 5d) relative to the control experi-
maintains high DIC over the mid-shelf during relaxation and ment 1 (Figure 5b). As WA depends on T, S, DIC and TA, we
periods with weak winds (DIC budget terms not shown). In evaluate the contribution of each variable to the total change

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING
Figure 6. (top) Time series of O2 fluxes from advection
(red), vertical mixing (blue), and biological O2 sinks (black,
remineralization within the sediments; gray, remineralization
plus nitrification in the water column) for experiment 2 in
the bottom 10 m of the water column at the 90 m isobath
(dashed black line in Figure 5c). Positive (negative) fluxes
indicate a gain (loss) of O2. (bottom) Alongshore wind
stress: upwelling occurs when t y < 0.
in WA between both simulations as explained in section 3
(Figure 7). The increase in DIC by upwelling is primarily
responsible for the drop in WA (Figure 7a), while the increase
in TA cancels out about 40% of the DIC effect (Figure 7b).
Upwelling also brings colder waters from offshore that
reduce the temperature near the bottom over the shelf by
up to 3°C (the mean cooling over the near-bottom shelf is
0.63°C), which tends to decrease WA by increasing
3 ]sat (section 3). The reduction in WA due to cooling
[CO2
(Figure 7c) is small compared with the change due to
increasing DIC (Figure 7a). The changes in salinity due to
stronger upwelling do not significantly affect WA over the
near-bottom of the shelf outside the VICC region (Figure 7c).
The combined effect of DIC and TA (Figure 7d) accounts for
essentially all of the change in near-bottom WA between
experiments 1 and 2 (black bars in Figure 7).
4.2. Effect of a Shallower Offshore OMZ
[21] On average (from day 50 to 125), the bottom waters
over the shelf have lower O2 in the experiment with a shal-
lower OMZ (experiment 4) compared with the control
experiment 1 (Figure 8). Most of the shelf has bottom con-
centrations below 80 mmol-O2 m3 in experiment 4, while
in experiment 1 shelf waters are mainly above that level
(waters between 60 and 80 mmol-O2 m3 are shaded in
Figure 8). The upper layers of both simulations remain
similar, since their initialization is the same (Figure 2b) and
the changes in deeper waters do not greatly modify biology
in the upper ocean or air-sea O2 exchange. The mean hyp-
oxic threshold (60 mmol-O2 m3, bold black line in
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G01020
Figures 8a and 8b) reaches the shelf break in experiment 4,
penetrating up to depths of 135 m. During the 75 days of
analysis, the near-bottom hypoxic threshold in experiment 4
is either near the shelf break or over the outer shelf (bold
black contour in Figure 5g). The threshold penetrates
onshore after strong upwelling events (after days 40 and
108, see Figure 2a). The lower O2 concentrations with
respect to the control experiment 1 respond to two factors:
(1) the initial conditions, which are already lower for depths
greater than 130 m (Figure 2b), and (2) the enhanced
advection of lower O2 from offshore onto the mid- and outer
shelf (see O2 budget in Figure 9). Biological sinks (both in
the water column and the sediments) remain essentially
unchanged over the shelf between experiments 1 and 4
(Figure 9), which differ only in the initial profiles of O2
concentrations. Since the lower bottom O2 concentrations
increase the vertical gradient of O2, vertical mixing is
enhanced in experiment 4 and partly compensates the
decrease in O2 by advection.
[22] When the modified initial conditions with a shallower
OMZ are used in combination with stronger upwelling-
favorable winds (experiment 5), O2 concentrations decrease
even more in the near-bottom layer over the shelf
(Figure 5h). Most of the bottom waters over the shelf
become hypoxic after the strong upwelling event around day
80, except for the area influenced by the O2-rich VICC. O2
concentrations drop to 21 mmol-O2 m3 near the shelf
break, the lowest over the shelf for all the experiments pre-
sented here. Concentrations lower than 20 mmol-O2 m3 (or
0.5 mL L1) are usually considered to represent “severe
hypoxia” [e.g., Monteiro et al., 2006; Chan et al., 2008;
Diaz and Rosenberg, 2008].
[23] In experiments 4 and 5 (Figures 5g and 5h), the
region with RI ≤ 1 expands greatly compared with experi-
ments 1 to 3 (Figures 5a, 5c, and 5e; bold yellow contours
indicate the RI = 1 contour). The decrease in RI is mainly
due to the lower O2 concentrations in the near-bottom layer
of the open ocean, since pCO2 does not change significantly
in those waters. In particular, the shoaling of the OMZ does
not significantly affect the carbon cycle, e.g., differences
between experiments 1 and 4 for pCO2 and WA are < 0.6%
and 0.4%, respectively (not shown).
4.3. Effect of Higher Inorganic Carbon
[24] Higher DIC initial conditions (experiment 6, see
Figures 2c and 2d) increase pCO2 throughout the model
domain (Figure 10). The bold black contours in the averaged
pCO2 vertical sections correspond to the atmospheric
pCO2atm values in experiments 1 (370 ppmv, Figure 10a)
and 6 (513 ppmv, Figure 10b). Over most of the shelf, the
surface ocean is on average undersaturated in both simula-
tions and absorbs CO2 from the atmosphere. However,
surface VICC waters within 4 km from the inshore
boundary are oversaturated in the control experiment 1
(releasing CO2 to the atmosphere), in contrast to experi-
ment 6. Aragonite undersaturation expands considerably
over the shelf in the latter scenario. The average saturation
horizon (WA = 1) coincides with averaged pCO2 values of
710 and 800 ppmv in experiments 1 and 6, respectively
(Figure 10). The sensitivity of WA to pCO2 varies between
these experiments due to the different effect of DIC on WA
and pCO2. We can approximate the former using the
 21562202g, 2012, G1, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2011JG001849 by Cochrane Philippines, Wiley Online Library on [04/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

Figure 7. Histograms of WA change (DWA) in the near-bottom layer of experiment 2 relative to experi-
ment 1 due to the contributions of individual and combined variables: (a) dissolved inorganic carbon
(DIC), (b) total alkalinity (TA), (c) temperature (T) and salinity (S), and (d) the combination of TA and
DIC (TA + DIC). Black histograms are the same in all plots and show the total change in near-bottom
layer WA between experiments (DWATotal). The histograms do not include the spin-up period or the area with
strong VICC restoring on the inner 3 km.

carbonate ion concentration in seawater and at saturation

[Sarmiento and Gruber, 2006]: WA  [CO2 3 ]/[CO3 ]sat 
2
2
(TA-DIC)/[CO3 ]sat. Hence, an increase in DIC leads to lower
WA. However, pCO2 can be approximated by (2  DIC 
TA)2/(TA-DIC), such that higher DIC increases pCO2.
[25] In experiment 1, the saturation horizon in the open
ocean is below 250 m (not visible in Figure 10a). However,
in experiment 6, the saturation horizon shoals 250 m in the
open ocean with respect to experiment 1 and extends over
the shelf up to 30 m above the seafloor (Figure 10b).
Moreover, the saturation state in experiment 6 is below unity
(reaching down to 0.78) after day 50 in the near-bottom
layer over the whole shelf seaward of 8 km from the inshore
boundary (Figure 5f). There, the VICC carries waters with
lower DIC relative to near-bottom waters over the shelf
[Bianucci et al., 2011, and references therein]. However, in
the control experiment 1, near-bottom waters with WA ≤ 1 are
found only offshore from 45 km from the inshore bound-
ary (Figure 5b). Figure 8. Mean O2 vertical sections down to 250 m
[26] The area with RI < 1 expands modestly in experiment 6 (average for days 50 to 125) for experiments (a) 1 and
compared with the control experiment 1 (yellow bold contour (b) 4. Bold black contours indicate the hypoxic threshold
in Figure 5f). In waters deeper than 1000 m, the modified (60 mmol-O2 m3); the areas shaded in yellow emphasize
initial conditions do not show a large increase in DIC waters between 60 and 80 mmol-O2 m3. The dashed
(Figure 2f). Consequently, pCO2 increases only moderately in magenta line indicates the position of the shelf break.

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING G01020

the near-bottom waters of the slope (where RI < 1, at 60 km

from the inshore boundary). If RI is calculated using pO2 from
experiment 4 instead, the area with indices values <1 is almost
identical to that of experiment 4 (only slightly larger). The
combination of pO2 from experiment 4 and pCO2 from
experiment 6 is consistent with an scenario of low-O2 and
high-DIC, since T and S are the same in both simulations and
changes in TA are minimal.

5. Discussion and Conclusions

Figure 9. Source and sink fluxes of O2 in the bottom 10 m
of the water column at locations where the bottom depth is
(top) 90 m and (bottom) 130 m (dash-dotted yellow lines in
Figure 5g). Fluxes are integrated from day 50 to 125. Experi-
ments 1 and 4 shown in black and white bars, respectively.
Abbreviations on the horizontal axis represent advection
(horizontal plus vertical), vertical mixing, exchanges with
the sediments, water column remineralization of semilabile
dissolved organic matter and detritus, and water column
nitrification.
[27] The experiments described here provide insight into
the sensitivity of the carbon and O2 cycles to changes in
forcing or initial conditions in a seasonal upwelling region.
Both cycles respond to changes in upwelling in our model
experiments. However, only O2 concentrations are affected
by the shoaling of the OMZ and only the carbon cycle
responds to changes in the oceanic DIC inventory. None-
theless, changes in the depth of the OMZ and carbon content
of the ocean will not be independent of each other, as both
are expected to respond to the increase in atmospheric CO2
from human activities. Moreover, these changes will be
coupled to modifications in the cycling of other nutrients.
Given the complexity of the feedbacks between biogeo-
chemical cycles, the introduction of a single change per
experiment allows a clearer understanding of the responses
to any given change.
[28] This study extends that of Ianson and Allen [2002],
which described results from a compartment model. Our
quasi 2-D model [Bianucci et al., 2011] can resolve the
spatiotemporal response to wind-forcing in both the vertical
and across-shelf directions. Moreover, the current work
focuses on sensitivities to climate-related change of inter-
actions between the carbon system and O2, which was not
included in the model of Ianson and Allen [2002].

Figure 10. Mean pCO2 vertical sections down to 250 m (average for days 50 to 125) for experiments
(a) 1 and (b) 6. Bold black contours show where the pCO2 is equal to the surface atmospheric pCO2:
(Figure 10a) 370 ppmv; (Figure 10b) 513 ppmv. The bold white contour in Figure 10b shows the sat-
uration horizon (WA = 1), which lies below 250 m depth in Figure 10a. The dashed magenta line indi-
cates the position of the shelf break.

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G01020 BIANUCCI AND DENMAN: CARBON AND O2 SENSITIVITY TO FORCING
[29] Strong upwelling events (such as the one at day 80
in experiment 2) result in onshore advection of low O2 and
high DIC from offshore that triggers low O2 and WA events
in the near-bottom layers over the shelf. The WA decrease is
partially compensated by the onshore advection of high TA
(advection of cooler waters has a much smaller effect). In
addition to the intensified onshore advection under increased
upwelling scenarios, O2 consumption and DIC production in
the sediments are enhanced over the mid-shelf due to off-
shore transport of near-surface phytoplankton blooms. The
latter increases deposition of organic matter to the seafloor
over the mid-shelf, enhancing the exchange of O2 and DIC
between the water column and the sediments. Therefore, low
O2 and WA values are maintained after the upwelling events
by remineralization within the sediments. O2 concentrations
in particular are demonstrated to be sensitive to intensified
upwelling, since hypoxia develops in the near-bottom waters
on the shelf in both simulations with stronger upwelling
(experiments 2 and 3).
[30] Differences between experiments 2 and 3 (which
differ only in the wind conditions during the spin-up period
of 50 days) demonstrate the effect of the timing of the onset
of the upwelling season. If summer upwelling is stronger
and starts earlier (experiment 2), O2-poor and DIC-rich near-
bottom waters are advected upwards closer to the shelf break
earlier in the season. Thus, subsequent upwelling events
further reduce O2 and WA in the near-bottom layers over the
shelf. A regional climate model of the northern California
shelf, with 40 km horizontal resolution, projects a 1-month
delay of the onset of seasonal upwelling for increased
atmospheric CO2 (560 to 686 ppmv), as well as intensified
upwelling [Snyder et al., 2003]. In the context of the present
study, a delayed upwelling season would reduce the poten-
tial for hypoxia and acidification on the Vancouver Island
shelf. In contrast, Barth et al. [2007] reported on the nega-
tive consequences of a one-month delay in the 2005 transi-
tion to upwelling-favorable wind stress on the California and
Oregon shelves (warm waters, low nutrient levels, low pri-
mary productivity, and low recruitment of rocky intertidal
organisms).
[31] The effect of a shallower OMZ (experiment 4) is
straightforward: it moves the hypoxic threshold closer to the
shelf break, so upwelling events are more likely to advect
O2-poor waters onto the shelf. Therefore, the combination of
stronger upwelling and a shallower OMZ (experiment 5)
would further reduce O2 concentrations on the shelf, repre-
senting a potential increasing stress on ecosystems. The
present model represents the shelf off Vancouver Island:
despite the considerable shelf width, the influence of a
shallower OMZ reaches the inner shelf (except for the region
influenced by the O2-rich VICC). The effect could be more
severe on narrower shelves, where upwelling can transport
offshore waters to even shallower depths onshore. The
expected overall decrease in O2 concentration in the sub-
surface open ocean would lead to an expansion of the region
with RI < 1, implying a larger area where aerobic marine life
would be under stress [Brewer and Peltzer, 2009a]. In none
of the experiments performed did RI reach values below
unity over the shelf (although O2 levels did decline below
the 60 mmol-O2 m3 hypoxic threshold). These results
suggest that RI is an indicator of more severe aerobic stress
than the 60 mmol-O2 m3 threshold.
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G01020
[32] RI calculated for a hypothetical scenario with lower
O2 and higher DIC showed a higher sensitivity of this index
to O2 than to CO2 changes. This difference results from the
changes in the initial conditions in experiments 4 and 6,
since the fractional decrease of initial O2 is larger than the
fractional increase of DIC (both changes aim to represent a
projection 50 years into the future).
[33] Under elevated inorganic carbon conditions (experi-
ment 6), near-bottom waters over the shelf (beyond the area
of influence of the VICC) become corrosive after 50 days,
i.e., WA drops below unity. In the current model configu-
ration, no biological process depends on DIC concentra-
tion (e.g., calcification is not included), so the increase of
DIC in the ocean does not affect the O2 or nitrogen cycles
directly. Decreased calcification in a carbon-rich ocean
may affect photosynthesis and phytoplankton community
structure [Sikes et al., 1980; Paasche, 2001], providing a
potential link between O2 and carbon. Furthermore, the
model is missing a negative feedback in the carbon cycle
that could potentially reduce acidification: a reduction of
calcification (increase of CaCO3 dissolution) reduces the
production (enhances the consumption) of CO2 in seawater
through the reaction 2HCO 3 + Ca
2 +
⇌ CO2 + CaCO3 +
H2O. Most published works agree on the reduction of
calcification rates with higher pCO2, although Iglesias-
Rodriguez et al. [2008] reported the opposite and
aroused some controversy [see Riebesell et al., 2008].
Blooms of calcifiers (e.g., coccolithophorids) are not fre-
quent over the Vancouver Island shelf, although they have
been observed (D. Ianson, personal communication, 2010).
[34] We need to understand how biogeochemical cycles
will respond to climate change in the coastal ocean, since we
depend on its resources. The present work contributes
toward that goal by focusing on the sensitivities of biogeo-
chemical cycling to environmental factors, individually and
collectively, that may change (in most cases, certainly will)
as the anthropocene evolves. The perturbations analyzed
here (increased upwelling, shallower OMZ, and higher DIC
and pCO2atm) consistently drive the system toward lower O2,
pH, and WA states. These results emphasize the potential
negative impact that these perturbations may have on ben-
thic ecosystems, along the lines of the consequences first
hypothesized by Bakun [1990] for increased upwelling.
Therefore, this sensitivity analysis suggests that a region
such as the Vancouver Island shelf could develop hypoxia
and corrosive conditions (WA < 1) in the future. Full 3-D
biogeochemical modeling as well as continuous and simul-
taneous observations of carbon and O2 are needed to assess
the current state of these cycles in the coastal ocean and how
they vary in time and space.
[35] Acknowledgments. We thank K. Fennel for her constructive
comments on an earlier version of this manuscript, as well as the valuable
input from two anonymous reviewers. L.B. acknowledges several sources
of graduate support: a UVic Fellowship, a Maritime Award Society of
Canada Scholarship, and a Bob Wright Fellowship. Computing support
was provided by the Canadian Centre for Climate Modeling and Analysis,
Environment Canada. K.D. acknowledges funding from an NSERC Dis-
covery Grant.
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L. Bianucci, Department of Oceanography, Dalhousie University, 1355
Oxford St., Halifax, NS B3H 4J1, Canada. (laura.bianucci@dal.ca)
K. L. Denman, VENUS Project, University of Victoria, PO Box 1700,
STN CSC, Victoria, BC V8W 2Y2, Canada. (denmank@uvic.ca)