Alyssa Faye R.

Mateo
2011-30731

MS 102

Effect of Ocean Acidification on Marine Microorganisms

1.

Introduction
Ocean acidification refers to the decrease of pH and the increase in the concentration of dissolved

carbon dioxide in the worlds oceans. Rising atmospheric carbon dioxide levels due to anthropogenic
emission from fossil fuels causes the reduction of oceanic pH as well as changes in seawater carbonate
chemistry. The acidification of oceans has altered the potential seawater chemical speciation as well
biogeochemical cycles of many elements and compounds. One well-known effect is the significant
alteration of calcium carbonate saturation states, which affects the formation of calcium carbonate shells
and skeletons of organisms ranging to plankton, echinoderms, molluscs, and corals. The rate of ocean
acidification will continue to accelerate for the next 100 years unless future of emissions of carbon
dioxide to the atmosphere would be curbed dramatically. Figure 1 shows a graph highlighting the increase
of concentration of atmospheric carbon dioxide, as well as the gradual decline of average oceanic pH.
Research has shown that the oceanic pH has already declined by at least 0.1 unit. Unmitigated CO 2
emissions will cause pH to decrease by as much as 0.4 by 2100 and 0.77 by 2300. These changes are a
threat to important biological processes (productivity, calcification, internal physiology, fertilization,
embryo development, larval settlement, and communication) and ultimately to the survival of many
marine organisms.

Figure 1. Graph showing the historical and projected decline of oceanic pH alongside the increase in the
concentration of dissolved carbon dioxide in seawater.
1.1

Seawater Carbonate Chemistry

As said previously, ocean acidification has serious consequences in chemical reactions of

dissolved carbon dioxide and its conversion to protons and carbonate ions. Figure 2 shows a diagram
summarizing ocean carbonate chemistry.

Figure 2. Diagram showing the alteration of ocean carbonate chemistry in response to the dramatic
increase of dissolved carbon dioxide in the oceans.
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The diagram shows the addition of dissolved carbon dioxide and water forms carbonic acid
(H2CO3). Carbonic acid is then ionized into bicarbonate ions (HCO 3-) and hydrogen ions, or protons (H+).
The increase in protons indicates the increase in the acidity of seawater. Bicarbonate ions are then
converted ions (CO32-). These bicarbonate ions impede in the formation and cause deformation of shells
and skeletons of various marine organisms.
1.2

Marine Microorganisms

Marine microorganisms constitute of the 98% of the biomass present in world oceans. They
encompass all microorganisms present in seawater and they are distributed across three taxonomical
domains. (Eukarya, Archaea, Bacteria). Most marine microorganisms are acellular. They can be
considered as a very diverse classification as they are further divided into different kingdoms: viruses,
protists, and prokaryotes. Marine microorganisms are involved in primary production, which involves the
creation of organic matter, through either photosynthesis or chemosynthesis. They also contribute to
second and tertiary production. Additionally, marine microorganisms are vital to marine life as they are
the drivers of chemical reactions and processes involved in the biogeochemistry of global ocean
ecosystems.
3.

Objectives
The term paper primarily discusses the various consequences of ocean acidification on various

species of marine microorganisms. Since marine microorganisms are invisible to the naked eye, the effect
of ocean acidification on these organisms cannot be directly observed visually, unlike in macroorganisms
such as coral reefs. Therefore, the paper will account for changes in physiology, behavior and function of
several species of marine microorganisms through different processes.
4.

Biogeochemical Processes of Marine Microorganisms affected by Ocean Acidification

The journal article authored by Das and Mangwani gives an overview showing the possible

effects of the decline in pH and increased dissolved carbon dioxide concentration on marine
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microorganisms. The biogeochemical processes of marine microorganisms that would most likely be
affected are given as follows: microbial based primary productivity, nitrogen fixation, trace gas emission,
extracellular enzyme activity and degradation of organic matter, microbial diversity and composition,
impact on microbial biofilms and quorum sensing.
4.1

Microbial-based Primary Productivity

Primary productivity refers to the manufacture of organic matter from CO 2 through the action of
photosynthesis. Photosynthetic marine organisms are responsible for the production of more than 90% of
total organic carbon produced in the sea through CO 2 fixation. Among photosynthetic marine organisms,
cyanobacteria has been shown to have the most impact on the ocean carbon cycle, as they are the most
abundant photosynthetic marine prokaryotes. In fact, they have contribute to 50% of total carbon fixation.
Marine photoautotrophs have developed specialized mechanisms for carbon fixation. One of
these mechanisms is referred to as Inorganic Carbon Concentrating Mechanism (CCM). In CCM,
photosynthetic organisms take up HCO3- ions to counter the limited dissolved CO2 concentration. CCM is
also found to assist in the maintenance of rapid growth under low peripheral dissolve inorganic carbon
conditions. (Badger, et. al, 2006). Research on the effect of elevated CO 2 on cyanobacteria seem to
indicate the increase in growth rate of species such as Synechococcus sp., Trichodesmium sp.,
Crocosphaera watsonii and Prochlorococcus sp. (Barcelos e Ramos et al., 2007; Fu et al., 2007; Kranz et
al., 2009, 2010) On the other hand, decrease in growth rate has been observed in Nodularia spumigena.
(Czerny et al., 2009). Research conducted by Coffin, et. al in 2004 have shown that longer incubation
times and warmer temperatures reduced bacterial metabolic production in samples exposed to the lower
pH.
Increased pCO2 levels have significantly affected cell division of marine microorganisms, in such
a way that it affects cellular elemental C:N and P:N ratio. A study conducted by Thomas, et. al in 2012
among cyanobacteria in the subtropical North Atlantic found that the cyanobacteria were able to

acclimatize rapidly to the changing pH and CO 2 levels. Variations in the results indicate that the response
of marine photosynthetic microorganisms to ocean acidification is species-specific. Elevated CO 2 levels
and reduced pH levels can interact with ultraviolet radiation from sunlight which can definitely affect
marine producers. Ocean warming and freshening cause the shoaling of the upper layer of oceans, which
may lead to additional photosynthetically active radiation (PAR) and ultraviolet (UV) exposure. This may
cause the increase in the rate of photosynthesis in both phytoplankton and macroalgae. Other experiments
have shown the small but significant increase in oceanic primary production in response to elevated pCO 2
concentrations. However, increased rate of primary production may have profound effects as it may lead
to massive eutrophication and anoxia in ocean surface waters, which can affect the nutrient balance in the
oceans.
4.2

Nitrogen fixation

Nitrogen fixation is a biogeochemical process that involves the conversion of atmospheric

nitrogen (N2) into biologically useful forms such as NH 4+, NO22-, and NO3-. These biologically useful
forms aid in other processes such as biological carbon export and sequestration.

Figure 3. Nitrogen fixation in the marine environment.

Diazotrophic cyanobacteria are one of the marine microorganisms that are able to process
substantial amounts of atmospheric nitrogen. Trichodesmium spp. contribute to more than 50% of
nitrogen fixation of all of marine environment. Research has indicated that the effect of elevated pCO 2
levels on N2 fixing bacteria on some groups of diazotrophic cyanobacteria still remain largely unknown.
Other short-term experiments conducted on Trichodesmium spp. have shown that ocean acidification
(OA) and elevated pCO2 levels may have the capacity to increase N 2 fixation rate at substantial amounts.
Enhanced N2 fixation is probably due to an increase in the availability of energetic resources for cellular
processes. On the other hand, Czerny, et. al in 2009 reported a decrease of N 2 fixation in other nitrogen
fixing species such as Nodularia spumigena.at increased pCO2 levels. Another research conducted by
Hayashik, et. al has shown that the ammonium oxidation activity among the nitrification process was
suppressed by the elevated pCO2 levels. However, the quantity and the diversity of amoA gene, which is
the specific coding of the ammonium oxidation, were not significantly affected by elevated pCO 2.

Figure 4. A graph showing the results of the study conducted by Hayashik, et. al on the effect of elevated
dissolved carbon dioxide levels on the nitrogen fixing activity of marine bacteria located on marine
sediment.
The changes to the nitrogen fixing activity of marine microorganisms brought about by OA may
also have effects in the iron uptake of marine microorganisms due to limited bioavailability of iron. A
study conducted by Shi, et.al has reported that the acidification of media containing various Fe
compounds decreases the Fe uptake rate of diatoms and coccolithophores to an extent predicted by the
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changes in Fe chemistry. A similar study has also demonstrated a slower Fe uptake by a model diatom
with decreasing pH as observed in the Atlantic surface water.
4.3

Trace gas emission

Many biological and photochemical processes contribute to the formation of small amounts of
volatile gases in the oceans such as dimethyl sulfide, organohalogens, ammonia, nonmethane, oxygenated
hydrocarbons, volatile oxygenated organics and nitrous oxides. These volatile organic gases are emitted
by marine microbes. These compounds play a role in both marine and atmospheric processes.
Changes in the ocean water chemistry brought about by ocean acidification may have effects on
the emission of trace gases in marine environments. If the atmospheric carbon dioxide rises to two times
of its current amount, the emission of trace gases will significantly decline. This may have serious
repercussions on atmospheric chemistry and global climate.
4.4

Extracellular enzyme activity and degradation of organic matter

Most marine microorganisms are involved in the decomposition, mineralization and recycling of
organic matter. Marine microorganisms decompose larger macromolecules for nutrition. Cell bound or
extracellular enzymes aid in the decomposition of polymers and other organic compounds. In the marine
environment, organic matter is chiefly composed of high molecular weight dissolved organic
matter (DOM). The activity of heterotrophic bacteria is important for the flux of DOM across the ocean.
The extracellular enzyme activity in marine bacteria is important as it aids in the formation or organic
nutrient for themselves for other organisms. It remains in question as to whether ocean acidification has a
profound effect on the activity of extracellular enzymes. A study conducted by Yamada, et. al in 2013 has
reported that there was little to no effect of increased CO 2 concentration on the short-term decomposition
of labile DOM. On the other hand, microbial activities producing new DOM were apparently enhanced by
some acidification conditions (2000 and 5000 ppm CO 2). These conditions could also influence the

processes involved in organic aggregate formation. Research needs to be conducted on the long-term
effects of increased pCO2 levels and ocean acidification on DOM formation and decomposition.
It is known that enzyme activity is limited to a narrow pH range. Small changes in pH can
dramatically affect the activity of enzymes, thus the existence of cytoplasmic buffers which resists
extreme changes in the pH. Grossart, et. al in 2006 has reported that OA has the capacity to affect many
microbial cellular processes, including extracellular enzymes. They also have reported the increase in the
activity of bacterial protease, alpha-glucosidase and beta-glucosidase in elevated pCO 2 levels. Similar
results have also been obtained by Piontek, et. al on acidification studies on microbial polysaccharide
degradation in 2010. On the other hand, Yamada and Suzumura in 2011 have reported decreased leucine
aminopeptidase and lipase activities under acidification conditions. Protease and beta-glucosidase activity
did not show changes in activity as the pH was decreased from 8.2 to 7.8. Increased extracellular activity
accompanied by the increased in dissolved carbon dioxide levels have consequences on the increased
availability of carbon sources. This may have significant effects on the primary production in the oceans.
4.5

Microbial diversity and composition

A study conducted on the response of microbial diversity to ocean acidification have shown
considerable impact. Krause, et. al In 2012 has reported that small pH shifts can induce drastic changes in
the bacterial composition in the North Sea. They have cited that Gammaproteobacteria,
Flavobacteriaceae, Rhodobacteraceae and Campylobacteraceae as phylogenetic groups responding
remarkably to differences in pH. Another research conducted by Meron, et.al in 2011 has reported
increase in growth of many pathogenic microorganisms such as Vibrionaceae and Alteromonadaceae in
corals. Karvonen, et.al in 2010 has reported that bacterial composition is also significantly affected by
other factor such as climate and growth conditions. The difficulty of assessing the effects of OA on
microbial community diversity has been limited by insufficient information.
4.6

Microbial biofilms

Biofilms are defined by the International Union of Pure and Applied Chemistry (IUPAC) as the
Aggregate of microorganisms in which cells that are frequently embedded within a self-produced matrix
of extracellular polymeric substance (EPS) adhere to each other and/or to a surface. Extracellular
polymeric substances (EPS) are defined as a polymeric conglomeration generally composed of
extracellular biopolymers in various structural forms. The common culturable bacteria studied for
marine biofilm are Exiguobacterium, Bacillus, Halotalea, Micrococcus, Halomonas, Arthrobacter,
Jeotgalibacillus, Pseudoalteromonas, Pseudomonas, Paracoccus and Vibrio, among others. The impacts
of ocean acidification on coastal biofilms remain poorly understood. Witt, et.al in 2011 has reported
changes in microbial composition in marine biofilms in the Great Barrier Reef under acidification
conditions. Changes in the community structure of biofilms can influence the settlement of invertebrate
larvae in some cases. Marine biofilms also serve as substrates for both microorganisms and eukaryotes.
Therefore, the changes in the microbial diversity of marine biofilms may have profound effects on the
current ecosystems.
4.7

Quorum Sensing

Quorum sensing (QS) is referred to as regulation of gene expression in response to fluctuations

in cell-population density. Quorum sensing bacteria produce and release chemical signal molecules
called autoinducers that increase in concentration as a function of cell density. The detection of a minimal
threshold stimulatory concentration of an autoinducer leads to an alteration in gene expression. (Miller
and Bassler, 2001). Quorum sensing is regulated by compounds such as autoinducing peptides (AIPs)
and acyl homoserine lactones (AHLs) in Grampositive and Gram-negative bacteria respectively. In
marine bacteria, QS is mainly governed by AHLs.
Quorum sensing regulates many processes in marine bacteria, which includes bioluminescence,
symbiosis, competence, virulence, secondary metabolites, extracellular enzymes and biofilm formation.
(De Kievit, 2009; Mangwani et al., 2012; Miller and Bassler, 2001; Weber et al., 2009).
Gammaproteobacteria, which produces AHLs typically live in corals, biofilms, sponges, algae and
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animals in marine environments. Krause, et.al has reported that OA plays a role in shifting of
microbial
community from Gammaproteobacteria to Flavobacteriaceae. This may have
profound effects on higher organisms (corals, algae, sponges) which live near these
microbes. The decline in pH due to OA favors the growth of pathogenic microbes in
the nearby corals. As a defense mechanism, corals and few coral associated
microbes synthesize number of QS inhibitors to protect themselves from these
pathogenic microbes.
5.

Challenges
Studying the effect of ocean acidification on marine microorganisms has been

described as a difficult, arduous process. Most of the studies on microorganisms and

the effect of decreased pH are limited to selected species and short-term
experiments. The results of the experiments are also of ambiguous nature, as
marine microorganisms tend to display acclimatization to slightly enhanced
dissolved carbon dioxide levels. It is also important to note that ocean acidification
is a very slow, gradual process. Long-term experiments that may range from years
to decades are better suited to assess the impacts of OA on marine microorganisms.
There is also a need to develop new methods and approaches to investigate the
effects of ocean acidification.
6.

Personal Insight
The topic of ocean acidification is both interesting and overwhelming for me.

Ocean acidification is a massive and a very serious problem, and it is something

that is inevitable given the amount of carbon dioxide humans have been emitting to
the atmosphere since the advent of Industrial Revolution. In class, I saw the effects
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of OA in corals and molluscs, particularly the coral bleaching phenomenon. I also

saw that the effects are not limited to isolated areas, but are also felt across the
globe. This piqued my curiosity about the topic which led me to writing this paper.
Learning about ocean acidification has shown me the importance of the
ecosystem of these marine microorganisms. Its amazing that some marine
microorganisms are resilient to the increase in dissolved carbon dioxide levels. In
fact, some species even acclimatized to these environmental changes. But for how
long? Many studies regarding global oceanic pH has projected the continued
decrease in global ocean pH levels and at some point some of these species may
not be able to handle such degrees of physico-chemical stresses. Whats alarming
about this fact is that this phenomenon is something that will continue even if
anthropogenic carbon dioxide emissions into the atmosphere are cut in half, unless
an atmospheric carbon dioxide chelating technology can be developed in the near
future. This goes that it is only after decades of carbon dioxide accumulation we can
begin to observe its serious consequences.
It is important to note that marine microorganisms comprise the biggest
percentage of ocean biomass and partake in very important biogeochemical
processes. Many higher organisms depend on them for nutrition and survival. Their
contributions to the marine ecosystem is massive, and we can say that marine
microorganisms form the backbone of ocean life. As such, the effects of ocean
acidification on marine microorganisms cannot be underestimated. The effects and
consequences of OA is an interesting topic for research and further discovery.
7.

Conclusions

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The effects of ocean acidification on marine microorganisms remains an

active area of research as information regarding the long-term effects of OA remain
understudied due to the extreme diversity of marine microorganisms, as well as
temporal and spatial limitations. Previous researches on changing ocean carbonate
chemistry and its consequences on marine ecosystems have shed light to
understanding of the effects on marine microbes. However, much remains to be
seen regarding the understanding of the consequences at the level of functional
genomics. Ocean acidification has also highlighted the valuable contributions of
these microorganisms, as they are the main drivers of biogeochemical processes in
the ocean and are the primary producers of organic matter. Therefore, the decline in
oceanic pH not only has a serious consequence on the microorganisms themselves,
but also it has ramifications in the entire marine ecosystem. Novel approaches and
methods in bacterial physiology and genomics are still being developed and are
seen as one of possible mitigations to combat this environmental problem and also
the condition of the oceans in the future.
8.

References

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2.

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