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Lit Review Charlotte Horn
Lit Review Charlotte Horn
Charlotte Horn
Dr. Wilfong
Fall 2023
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Introduction
Ocean Acidification refers to the decrease in pH of the ocean due to the increase in Carbon
Dioxide (CO2) uptake from the atmosphere. As CO2 dissolves in seawater, it forms a carbonic acid,
leading to a decline in ocean pH. During this time in the Anthropocene, CO2 has increased because of the
burning of fossil fuels, cement production, and land use changes (Heinze et al.,2015). A decrease in the
pH of the ocean can have profound implications for coral reefs and other marine animals. Acidification of
seawater poses severe threats to coral reefs, shell-forming organisms, and various marine species that rely
on stable pH levels for their survival and growth. My review will examine the causes, consequences, and
potential ocean acidification solutions, emphasizing the urgent need for collective action to mitigate its
Since the Industrial Revolution, humans are producing Carbon dioxide much faster than in the
past 20,000 years (Heinze et al., 2015). As atmospheric CO2 levels increase in the atmosphere, so do the
levels in the ocean. CO2 levels are rising due to fossil fuels, land-use change, and agriculture. CO 2 levels
were at 278 ppm (pre-industrial), 315 ppm (beginning industrial), and now 400 ppm (Heinze et al., 2015).
As the carbon dioxide levels in the atmosphere increase, so does the global temperature and the pressure
of CO2 on surface waters (Ivanina et al., 2013). An increase in CO2 is then absorbed by the ocean,
absorbed through plant photosynthesis, and/or stays in the atmosphere as a greenhouse gas.
Increased CO2 levels mean higher levels of carbon, which lowers the pH of the ocean (Noor &
Das, 2019). The ocean absorbs CO2 through physical and biological activity (Heinze et al.,2015). In
physical activity, water (H2O) and CO2 combine to form carbonic acid (H2CO3). When carbon dioxide and
carbonic acid (H2CO3) combine with water (through turbulence and currents) they dissociate into
bicarbonate (HCO3-) and carbonate (CO32-), and a minimal amount of CO2 is kept in its dissolved state
(Heinze et al.,2015). The sum of HCO3-, CO32-, and CO2 is called total dissolved inorganic carbon (DIC).
Inorganic carbon cycling leads to a slight increase in DIC with depth (Heinze et al.,2015). The slower the
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ocean circulation due to climate change, the less efficient the ocean will become for absorbing further
In a more simplified sense, water and carbon dioxide combine to form carbonic acid (H 2CO3).
Weak acids then dissolve into H+ and bicarbonate ions (HCO3-). However, the available carbonate ions
(CO32-) bond with excess hydrogen (H+), resulting in fewer carbonate ions available for calcifying
organisms to build their shell, skeletons, and other calcium carbonate structures. Figure 1 visually
explains how lower pH favors the formation of bicarbonate ions (HCO3-) over carbonate ions (CO32-),
Figure 1
Biological carbon pumps are responsible for turning DIC into particulate organic carbon (POC)
and dissolved organic carbon (DOC) (Heinze et al.,2015). Biological carbon binding occurs on the
surface level of the water column where phytoplankton go through photosynthesis (Heinze et al.,2015).
Phytoplankton absorb CO2 and water (H2O) and use the sun’s energy to produce glucose (C₆H₁₂O₆) and
oxygen (O2). Microscopic marine plankton are primary producers, and responsible for distributing their
energy down the food chain. All living organisms consume CO2 directly or indirectly. Once plants and
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animals die, they sink, storing the carbon in the sediment and reducing the amount of carbon in the
atmosphere. The ocean is responsible for absorbing 25% of anthropogenic CO2 emissions over the past
several decades, and may lead to a reduction in the ability to absorb CO 2 in the future (DeVries, 2022).
Impact on Mollusks
Ocean acidification affects the calcium carbonate structures of mollusks and corals. Many studies
describe the negative impacts of rising temperatures and ocean acidification on calcification, energy
metabolism, reproduction, development, and growth of marine calcifiers (Goncalves, 2017). Harvey et al.
(2016) studied the effect of decreased pH on hexaplex trunculus, a widespread Mediterranean mollusk.
Through their experiment, the increase in CO2 levels in the ocean caused a decrease in shell size but an
Furthermore, mollusks can exhibit altered growth and strength of the exoskeletons (Amaral et al.,
2017). Amaral and other’s study examined the effects of estuarine acidification on predator-prey
interactions. They tested how acidification sustained from runoff from acid sulfate soils (or soils that
contain iron sulfides) affected the strength of saccostrea glomerata (prey-oyster), bembicium auratum
exoskeletons in the mangroves of Australia (Amaral et al., 2017). They found the shells of S. glomerata,
B. auratum, and M. marginalba were weakened by the abundance of CO2 while H. cordiformis
exoskeletons showed no structural difference. With M. marginalba having greater rates of predation
overcompensates for the shell strength, as the gastropod’s capacity to consume prey items is driven by its
size but not its shell strength (Amaral et al., 2017 and Cotton et al., 2004). While B. auratum in acidified
sites leads to greater mortality, although increase in predator avoidance (Cotton et al. 2004). The effects
of acidification on mollusks allow oysters and other prey gastropods to become more susceptible to
predation (Amaral et al., 2017). Climate change resilience can differ from species to species and even
The third and final mollusk-based study took place in 2013 and examined eastern oysters
(crassostrea virginica) and hard clams (mercenaria mercenaria) (Ivanina et al., 2013). Both species are
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found on the eastern shores of the United States. The study exposed the two bivalves to varying PCO 2
(~400 and 800μatm) and temperature (22 and 27°C) levels for 15 weeks (Ivanina et al., 2013). They
found a combination of high temperature and elevated PCO2 led to a significant decrease in shell hardness
in both species (Ivanina et al., 2013). Rising temperatures and ocean acidification can have serious
implications on the calcification, energy metabolism, reproduction, development, and growth of estuarine
Impact on fish
Ocean acidification can affect the development of fish, behavior, hearing, physiological
functioning, and novel species interactions (Radford et al., 2021; Mitchell et al., 2023). Ocean
acidification and climate change directly and indirectly modify species responses to novel environmental
conditions (Mitchell et al., 2023). Mitchell et al. (2023) used laboratory experiments to explore how
ocean acidification, future summer vs winter temperatures, and novel species could affect coral reef fish.
The future winter experiment consisted of 20 degrees Celsius and elevated pCO 2 levels. Coral fish
showed reduced physiological performance which included lower body conditions, cellular defense, and
high oxidative damage (Mitchell et al., 2023). Contrastingly, shoaling temperate fish showed higher
oxidative damage, reduced short-term energy storage, and cellular deference in future summers compared
to future winters (Mitchell et al., 2023). Temperate fish were found to benefit (higher body condition and
short-term energy storage) from shoaling with coral reef fish compared to same-species fish (Mitchell et
al., 2023). In conclusion, this study found that coral reef fish will benefit from ocean warming and have
contrasting benefits in winter conditions, and therefore slow their establishment at high latitudes (Mitchell
et al., 2023). Temperate fish species benefited from co-shoaling with smaller-sized tropical fish, but these
benefits dissipate due to their reduced physiological function under future summer temperatures (Mitchell
et al., 2023).
However, increased CO2 and lower pH levels cause sensory deficits and altered behavior of
marine fish (Radford et al., 2021). In fish, changes in pH can alter the effectiveness of their hearing. Fish
that matured in water that had elevated CO2 levels suffered from low-frequency hearing loss due to either
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otolith asymmetry or larger otoliths (Radford et al., 2021). Otoliths aid in balance and hearing and are
calcium carbonate structures found inside the heads of bony fish. Fish use low frequencies (less than 500
Hz) to communicate and to find mates (Radford et al., 2021). Hearing loss could have important
consequences for the maintenance of fish populations globally (Radford et al., 2021).
Impact on corals
Ocean acidification mainly affects the calcifiers of the ocean including mollusks and coral reefs
(Edmunds et al., 2013). Corals make their calcium carbonate structure by forming aragonite crystals on
top of each other (Stewart et al., 2022). Coral reefs provide habitat, feeding, spawning, and coastal
protection against storms, floods, and tsunamis (Guannel et al., 2016). Coral reefs are extremely
important to the health of marine ecosystems and biodiversity (Sebens, 1994). Ocean acidification can
affect coral reefs' direct physiological function as well as indirect effects of compromised satisfaction of
reef-building corals (Hill & Hoogenboom, 2022). Coral direct physiological functions can relate to the
upward growth (extension) and lateral thickening (densification) of coral (Mollica et al., 2018). Mollica et
al. (2018) found that the skeletal density of porites corals could decline up to 20.3% over the 21st century
solely due to the acidification of the ocean (Mollica et al., 2018). Porites corals are a type of stony corals
that are the main building blocks of the reef structure which provide food, shelter, and substrate for reef
ecosystems (Mollica et al., 2018). In this specific study, they chose 6 species of porites off the coast of
Puala (Mollica et al., 2018). Ocean acidification puts more pressure on coral reefs and may cause them to
grow more slowly and unable to overcome typical pressure (reef dwellers breaking apart reef skeletons
for food or habitat). Coral reefs don’t need another added pressure, ocean acidification already threatens
their future by reducing the carbonate ions needed to construct their skeletons (Mollica et al., 2018). This
study did not consider temperature, sea level, and nutrient concentrations that may impact corals in the
future.
Another large group of corals affected by ocean acidification is cold-water corals (CWC). Maier
et al. (2012) examined calcification rates and the effect of ocean acidification on Mediterranean cold-
water corals. The study examined three species of Mediterranean cold-water scleractinian corals including
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lophelia pertusa, madrepora oculata, and desmophyllum dianthus (Maier et al., 2012). Cold water corals'
abundance is determined by the temperature of the water, preferring temperatures between 4 and 12
degrees Celsius (Roberts et al., 2006). CWCs will be highly affected by ocean acidification because they
predominantly live at relatively high latitudes and/or deep water (Maier et al., 2012). Both of these
environments are the first to be affected by the lowering pH and shallowing of the aragonite saturation
(Turley et al., 2007). This study found that elevated pCO2 levels (867 μatm) had no negative response to
calcification rates. However, calcification doubled at the pre-industrial pCO 2 levels (285 μatm) for both L.
pertusa and M. oculata. This reveals that M. oculata had a reduction in calcification by 50 percent since
pre-industrial times (Maier et al., 2012). Suggesting that M. oculata has already hit its highest temperature
My literature review examines how ocean acidification and Anthropogenic factors impacts
mollusks, fish, and corals. The complexities of these environmental changes alter organisms' behavior and
physiological function (Mitchell et al., 2023). Some studies examine the future impacts of increased pCO 2
levels, and others explore the impacts already seen in today's elevated CO 2 levels. Temperature rise
heightens the impact of ocean acidification and stresses marine ecosystems. Ecosystems all over the
world, from the coast of Austalia to the eastern shores of the United States are effected. Now the question
is where do we go from here, and how do we stay optimistic about the degradation of marine ecosystems?
For now, we hope that certain species can evolve quickly enough to adjust to the changing pH and
temperature changes (Kang et al., 2022). Additionally, we must make deep cuts to our carbon emissions if
we want to save the already impacted ecosystems. One way to do this is to become a climate citizen, vote
for climate action, and prioritize policies to reduce and restrict excessive carbon emissions. One person
can’t turn back climate change, but if we all work together we can accomplish great things.
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References
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Cotton, P. A., Rundle, S. D., & Smith, K. E. (2004). Trait Compensation in Marine Gastropods: Shell
DeVries, T. (2022). The Ocean Carbon Cycle. Annual Review of Environment & Resources, 47(1), 317–
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Edmunds, P. J., Carpenter, R. C., & Comeau, S. (2013). Understanding the Threats of Ocean
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