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d head: CORAL EVOLUTION AND OCEAN5 .2 ndiACIDIFICATION 1 5 .2 ndi
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Coral evolution in the face of increasing ocean acidification: A review

Mandi T. Jebe

Brigham Young University, Provo UT

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9 .1 f M Author
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As climate change and its various factors, specifically ocean acidification, change oceans
worldwide, the coral reef health is in decline. Amidst this decline, researchers have investigated
the evolutionary capabilities of these ecosystem giants to better understand potential for coral
survival as ocean acidification worsens. While the science community varies on some
perspectives, the possible for coral evolution seems promising, both in genetic and non-genetic
adaptation and evolution passed throughout generations. It is still undetermined whether these
mechanisms of evolution will function if ocean acidification continues to rise at its current rate
considering adaptation takes time, and a consistently increasing rate of acidification could
overwhelm coral systems, damaging potential for adaptation and evolution. However, many
experiments have proven the adaptability of corals in the face of ocean acidification, adding
promise to the idea of coral restoration efforts for the time being.

Keywords: evolution, coral evolution, coral reefs, ocean acidification, calcifiers,

calcification, coral skeletons, reef deterioration, climate change

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As climate change continues to affect many facets of the natural world, oceans are
continually impacted by various factors. Not only have oceans experienced rising sea levels and
temperatures due to climate change factors, but they have also become more acidic as carbon
emissions increase. Hoegh-Guldberg et. al. (2007) state the carbon emissions of the 1900’s not
only affected areas such as sea level and temperature, but also depleted carbonate concentrations,
therefore increasing the acidity of the seawater. They continued to explain that currently
approximately 25% of the anthropogenic CO2 emissions enter the ocean, producing carbonic
acid and limiting the amount of carbon available to other marine organisms. (Hoegh-Guldberg et.
al., 2007). Kleypas et. al. (2006) add their opinion on the future of the issue, stating “Model
projections suggest that on millennial timescales the ocean will be the ultimate sink for about
90% of the anthropogenic carbon released to the atmosphere.” On this point, Knoll et. al. (2008)
further this information by concluding that is the rate at which atmospheric CO2 concentrations
are rising, not the absolute values of them that make ocean acidification an issue. Based on this
shared scientific knowledge one may logically conclude that if an issue of ocean acidity currently
exists with the ocean absorbing 25% of anthropogenic CO2 emissions, an increase of absorption
to 90% of anthropogenic CO2 emissions in a biologically and geologically short period of time
would e further and intensify the ocean acidification process. e e
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5 .2 ndi With such a scientific outlook on the future 5 .2ofndocean acidification, it is important to 5 .2 ndi
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Researchers have begun investigations on rthe evolutionary capabilities of corals, the results of r
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which will be the focus of this review. Corals appear to be adaptable and to evolve over
generations, however a critical remaining question is if corals will be able to adapt at the
unprecedented rate at which the ocean is gaining acidity. Research remains unfocused on this
area, leaving an opening for future researchers to begin experimentation. Researchers have
discovered a promising amount of evidence in support of corals’ ability to evolve to a changing
climate, indicating the rapid rate of change is an obstacle that could be overcome by these
resilient organisms.

Coral Reefs and Ocean Acidification

Amidst this shift in ocean acidity, one area of increasing interest is coral reefs. Coral
reefs are among the most diverse ecosystems on the planet, making them valuable is retaining
diversity. Not only this, but coral reefs hold ecological significance for the thousands of species
they provide habitat for. Therefore, the health of coral reefs in the face of climate change factors
has become of interest to many scientists.

Coral reefs are particularly sensitive to ocean acidification levels due to the fact they are
8 e be skeletons. In order to do this, they 8 ebe
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acidification could negatively affect marine
approximately 40 years ago (Kleypas et. al., 2006). Since the emergence of the idea, corals have
been studied thoroughly to determine how rising ocean acidity will affect their livelihood, the
ecosystems they create, and therefore the other species they provide for.

Before continuing with this discussion about ocean acidification, it is worthwhile to

discuss how this may relate to other climate change factors. Rising sea temperature is large topic
of discussion in the face of climate change, and the factor of temperature is not completely
independent of the factor of ocean acidification. These two factors can affect each other and
further exacerbate those effects. Kleypas et. al. (2006) explain this effect by stating “human
activities are causing changes in all of these factors and light, temperature, and nutrients have all
been demonstrated to affect calcification rates of corals, either singly or in combination with
saturation state.” However, viewing the effects separately is beneficial in order to break down the
amount of information about each subject. So, while this review focuses purely on the effects
seen in coral due to ocean acidification, many of the sources referenced here also discuss the
effects on coral from rising temperatures since the two factors often appear together.

As for ocean acidification, although the hypotheseseand reasoning may differ, it appears
8 ebe
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(Hoegh-Guldberg et. al.,
2007). Van Oppen et. al.
express concern that this
effect on corals may
jeopardize their placement
within an ecosystem as
Figure 1| Hoegh-Guldberg et. al. (2007) illustrate depleting aragonite
engineers that support
concentrations. Aragonite is a form of carbonate that corals and other ecosystem services. The
8 eb e
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8 eb e rapid deterioration of reefs in 8 ebe
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op ro
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dissolution rates. However, the researchers hypothesize there is a critical threshold of CO2 intake
from the ocean at which calcification rates are less than dissolution rates. This would lead to a
net loss of corals biomass under high CO2 conditions (Fine & Tchernov, 2007). Thus,
investigations on the potential responses of corals and coral reefs in the face of climate change
are relevant and have been undertaken by many scientists around the world.

While scientists may agree on the dire state of our coral reefs, there is still debate on the
mechanisms involved in this deterioration, and more importantly, how corals may be able to
respond to these issues. Hoegh-Guldberg et. al. (2007) state that corals have three natural
responses they can undertake:

1) A decreased linear extension rate and skeletal density of coral colonies.

2) A maintenance of extension and growth rate through a reduction in skeletal
density.
3) A maintenance of both extension and skeletal density amidst reduced carbonate
saturation through a greater energy investment in calcification.
e 8 eberesponse leads to a net loss of coral 8 ebe
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Coral Evolution

Many researchers have commented on this idea of coral evolution or adaptation. The goal
of reef restoration is to use the science behind evolution and adaptation and applying such
science to threatened reefs to prevent deterioration. Van Oppen et. al. (2015) explain that one of
the goals of such restoration is to increase the ability of corals to accelerate their recovery after
acute impacts. To do so, researchers use widely accepted techniques to attempt to accelerate the
evolutionary and adaptive processes that occur naturally. Albright & Cooley (2019) add to this
concept by explaining the goal of reef restoration is not a simple replanting, but to develop
natural recovery through re-establishing populations with genotypic diversity, making them able
promote sexual reproduction and natural recovery. This concept is key in reef restoration.
Afterall, a simple replanting of reefs doesn’t address the core issue leading to reef deterioration.
If reef restoration attempts didn’t address the need for corals to adapt to fluctuating climate and
environment, the attempts would be wasted. However, researchers believe that an increase in
genetic diversity through various avenues can encourage adaptation to our changing climate.
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evolvability. Van Oppen et. al. (2015) offers

“1) The common occurrence of asexual reproduction in addition to sexual reproduction

2) A lack of segregation of the germ cell from the somatic cell line
3) The existence of symbiosis with a range of potentially fast-evolving microbes
4) Naturally occurring high levels of genetic diversity and the occurrence of interspecific
hybridization in some taxa.”

These traits together indicate that corals could respond well to assisted evolution efforts
and many researchers agree. Jury et. al. (2019) discuss the high heritability values reported for
corals, concluding “that a large portion of the observed variation in calcification rates among
unrelated coral colonies can be explained by heritable genetic factors. The high heritability
values combined with considerable variation among individual colonies and very large
population sizes indicates high potential for selection to act on calcification rates….” And lastly,
Cornwall et. al. (2020) provide more evidence for the evolvability of corals through diurnal
variability. Shallow water reefs experience daily increases in acidity during the night, and
Cornwall et. al. propose that this variability may impart tolerance to a low pH in corals through
selection
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P Not only this, but they additionally statedPthat corals have long generation times coupled with Pa
low genetic diversity, meaning they will cannot adapt quickly. While these statements are noted,
most researchers are of the persuasion that corals can evolve to meet these conditions. Hoegh-
Guldberg et al. performed their study almost thirteen years ago, and while their study is a still
valuable and was a forerunner in such experimentation, more recent studies seem to negate these
findings. Those negating such findings have undergone many projects to determine the
evolvability of corals.

Simple observation done by Jury et. al. (2019) has shown that corals can adapt to
changing climate relatively quickly. In an experiment done on corals from varying environments
in Hawaii, researchers replicated an experiment done in 1970 and illustrated that corals’
threshold for high temperatures had increased by approximately 2° Celsius. While this study
doesn’t directly relate to ocean acidification, it does illustrate a coral’s ability to adapt and
change in just a matter of decades—a relatively short period of time when discussing evolution.
Experimentation in coral evolution has varied as researchers began studying how to approach
assisted evolution. Among these efforts are assisted migration, selective breeding, and even
developing
e genetically modified organisms and most of thee experiments and efforts have been e
8
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based on a usage of asexually produced coral (van Oppen et. al., 2006). Albright & IP ert
op op restoration, something they define as “the Pro
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Cooley (2019) advocate an approach of ecological
process of assisting the recovery of an ecosystem that has been degraded, damaged, or
destroyed.” They suggest using all the experimental processes mentioned above by van Oppen
et. al., along with coral gardening and nurseries since these techniques have been widely
acknowledged as possible reservoirs of resilience to climate change (Albright & Cooley, 2019).
However, Albright & Cooley are quick to state that a single intervention strategy isn’t going to
fit every reef system due each system’s unique factors. This is a point Cornwall et. al. (2020)
agree with, stating that predicting beforehand the winners and losers of ocean acidification is too
difficult with our limited knowledge. Comments such as these illustrate why there is value in
varieties of experimentation. As more researchers determine what can work for coral ecosystems,
more options will be available to managers of reef ecosystems.

Throughout this experimentation, Jury et. al. (2019) are quick to state their own
hypothesis: that corals experience broad sense heritability. They state that is includes genetic,
maternal, epigenetic, and other heritable sources of variation. Albright & Cooley (2020) agree
with this statement, stating that even though immediate evidence for heritability is difficult to
find, examples of genotype-dependent responses to temperature and ocean acidification are
e
abundant. This hypothesis and the support for it is invaluable e because it supports the line of e
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6 experimentation occurring: it shows that corals can26adapt and evolve through heritable traits, and 6
5 .2 ndi 5 . ndi 5 .2 ndi
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evolution, also known as adaptation, and non-genetic evolution, also known as acclimatization.
To provide a better outlook on how coral evolution can occur and the scope of experimentation
that has occurred, both will be discussed below.

Genetic Evolution

Van Oppen et. al. (2015) define genetic adaptation as “change in the phenotype from one
generation to the next through natural selection and involves a genetic change in the form of
allele frequency changes between generations.” In summary, genetic evolution has to do with a
shift in the actual genes of the organism. It is also known as hard inheritance. Several
experiments have documented this kind of evolution at work. Kleypas et. al. (2006) explain in
their research that some species adapt to lower calcification rates by changing morphology or
growth strategies. There is prehistoric evidence to support this idea, offered by Hoegh-Guldberg
et. al. (2007). They state that their studies suggest that corals survived the Permian-Triassic
extinction event through forms lacking calcified skeletons. Due to this hypothesis, Fine &
Tchernov (2007) conducted an experiment to test the idea. They tested the adaptability of corals
to restrain
e to a soft bodied system in highly acidic conditions. e After 12 months, all corals that e
8
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transitioned to a skeleton-free system survived, IP ert
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relies on whether the tolerance action is heritable, and a trait that selection can act upon. Fine &
Tchernov’s experiment demonstrates that the tolerance response of a soft bodied coral is
heritable. When these soft bodied corals were returned to ambient pH conditions, they calcified
and reformed colonies. Fine and Tchernov called these soft bodied corals “ecophenotypes,”
playing on the idea of ecotypes common in botany. Ecotypes are organisms within the same
species that are genetically different. This idea supports the idea of genetic adaptation occurring
to provide better survival for corals. Fine & Tchernov did, however, find it important to state that
while this genetic adaptation could increase the survival of corals, the loss of structures in coral
reefs due to a lack of calcification would have massive ecological ramifications.

Van Oppen et. al. (2015) also speak on the idea of genetic adaptation in corals, although
they discuss theory of the process as opposed to experimentation. They state that genetic
adaptation could be constrained in corals due to tradeoffs between competing functions. In
various breeding programs there had been a lack of trait improvement even in the face of
artificial selection. Such behavior indicates tradeoffs present. Van Oppen et. al. express a desire
for more information about selective breeding in coral reef conservation. As more scientists
begin eto research selective breeding, the researchers point e
about tradeoffs should be well minded. e
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Non-Genetic Evolution

Van Oppen et. al. (2015) define non-genetic evolution as “a phenotypic response to
variation in the natural environment that alters performance and possibly enhances fitness but
does not involve genetic change.” This process is also known as acclimatization. According to
van Oppen et. al., until recently acclimatization had been considered non-heritable and to only
occur within the life span of a specific organism. One example of such non-genetic evolution is
evident in the experiments that have shown that exposure to mild stress conditions leads to an
increase in tolerance to more severe stress later in life. Jury et. al. preformed an experiment on
Hawaiian coral that support this idea. The determined after a cloning of corals from two varying
environments and reciprocal transplantation they found that the clonal corals from a low pH site
were less sensitive to ocean acidification than clonal corals from a high pH site. This provides
support for the hypothesis from van Oppen et. al. Acclimatization does appear to occur in corals,
and its apparently heritable since the asexual offspring retained those capabilities.

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Pr evolution as well. Experimentation focused Pr specifically on the symbiotic relationship almost all
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corals posses with coralline algae. They studied these algae in correlation with ocean
acidification because these algae often provide the source of energy a coral needs to calcify its
skeleton. In an experiment that spanned six generations, the researchers determined that the
corals were initially highly sensitive to ocean acidification, but that after six generations of
exposure to ocean acidity, the effects seemed to disappear. In a profound statement by the
researchers, the support the idea of non-genetic evolution. They state:

The trends observed here were also rapid in the context of evolutionary adaptation. The rapid yet
incremental, nature of these changes and the predominately asexual reproduction indicate that
transgenerational acclimatory processes could have played a key role in inducing the changes
observed.

Cornwall et. al. concluded that the corals had acquired tolerance to acidic ocean
conditions and that those results demonstrate that reef building taxa could maintain their
ecological role. While these results do appear promising, the experiment did fail to take into
account continuously rising acidification as opposed to an increase followed by a plateau,
meaning the research may be less applicable to the climate change we are actually experiencing.
e be 8 ebe
.58 Jeb way, we can’t belittle the significance of results
Either . 58 showcasing
e non-genetic evolution in . 5
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Jury et. al. (2019) also performed an: experiment
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that highlight non-genetic evolution in : 6 y o
IP ert corals. Through studying corals from varyingI environments off the coast of O’ahu, Jury et. al. I pe
P rt
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Pr Pr
studied the corals control over their calcifying fluid. It had become a well-established fact thatPr
corals could exert control over the fluid, upregulating its pH even as seawater pH decreases. Jury
et. al. had observed that some corals upregulate the pH of their calcifying fluid more than others
or maintain pH despite changes in seawater pH. They hypothesized that this ability to regulate
calcifying fluid plays a role in coral tolerance to ocean acidity. They further hypothesized that
corals from a reef with higher ocean acidity would upregulate more than corals from a reef with
lower ocean acidity. To test this, they cloned corals from each environment and then preformed a
reciprocal transplantation. Their hypothesis was correct; corals from higher ocean acidity sites
did upregulate more. However, all corals upregulated in ocean acidity, indicating a reaction
regardless of previous conditions. Since these were clonal organisms reacting to this
transplantation, it indicates that the acclimatization wasn’t restricted to the single, contained
organism, meaning this response could be a heritable form of acclimatization.

Albright & Cooley (2019) add a simple comment to the debate of non-genetic evolution:
one that helps sum up the research done so far. They conclude that transgenerational
acclimatization could lead to resistance to ocean acidification that may last for several
generations. This idea makes the study of non-genetic evolution or transgenerational
e 8 ebeof our coral reefs in the face of 8 ebe
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acclimatization very valuable considering the current.5state . 5
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important role transgenerational acclimatization Pr
can have on corals in regard to temperature and
ocean acidity. In support of that statement, there should be more research done on this topic. The
studies referenced here are a great start to understanding transgenerational acclimatization, but
it’s an area that would benefit from continual study.

Conclusion

The issue of coral deterioration is not new, but in the face of a continuously changing
climate and environment, the threat is now being addressed. Research has been conducted
throughout the decade, indicating that corals do have potential to evolve and adapt to changing
conditions. However, as was pointed out by one scientist, corals may not be able to adapt in a
climate where ocean acidification continuously increases. In all the research, no scientist was
able to refute this claim. It is unlikely that atmospheric carbon concentrations will continue to
rise in the future as they have in the recent past due to increased regulation efforts. Therefore, the
findings from various scientists that show not only plasticity amongst coral populations, but
genetic and non-genetic evolution throughout generations hold promise for the future of coral
reefs. It is likely that through reef restoration efforts, assisted evolution can occur within corals,
increasing the populations’ genetic diversity and providinge avenues for adaptation. There are still
8 ebe
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As research in this area continues, more 5 n
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p
Pr P
such research justifies the anthropogenic increase in CO2 concentrations. While this avenue of P
science is necessary and interesting, better care for our planet would have postponed or even rid
us of the current issues and deterioration within coral reefs. While reef restoration efforts are
promising, we should not loose sight of the fact that we, as humans, effect these valuable and
irreplaceable ecosystems. As governments regulate CO2 emissions, we see a decrease in the
drastic consequences of ocean acidification and as “green” practices continue, coral reefs have a
higher chance of surviving longer. Therefore, along with all the best science has to offer to
rescue our coral reefs, human efforts cannot be understated in their importance as well. A
combination of such efforts may lengthen the life of coral reefs and provide science time to
better understand the evolutionary capabilities of our ocean corals.

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op op op
Pr Pr References Pr
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