Tulioc 1

Mitigation of Coral Bleaching through Shading

Techniques Under Conditions of Thermal Stress in
Acropora aspera

By
Serafina Tulioc
Submitted in partial fulfillment of the requirements of the
Department of Natural Sciences and Mathematics and the Honors
Program
Dominican University of California
2015

First Reader: Dr. Vania Coelho

Sciences and

Department of Natural
Mathematics

Tulioc 2
Second Reader: Dr. Kenneth Frost

Department of Natural
Sciences and Mathematics

Honors Director: Dr. Diara Spain

Department of Natural Sciences

and Mathematics

Table of Contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Background Information . . . . . . . . . . . . . . . . . 6
Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Preparing Coral Environment . . . . . . . . . . . . . 10
Addition of Corals . . . . . . . . . . . . . . . . . . . . . . 12
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Tulioc 3

List of Tables & Figures

Table 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Table 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Figure A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Figure B . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Figure C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Figure D . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Figure E . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Tulioc 4
Abstract
The global temperature has been increasing and causing climate
change over the last century. This rise in temperature causes the sea surface
temperature to increase which in turn causes environmental stress to corals.
When exposed to environmental stress, coral bleaching occurs. Coral
bleaching is the expulsion of a corals symbiotic organisms; potentially
leading to host death. This phenomenon is caused by thermal stress which is
a result of increased water temperature and overexposure to light. In this
study we examine the impact of decreasing stress by reducing the light
exposure of the species Acropora aspera in environments with high thermal
stress. Corals were separated into six treatment tanks with water
temperature of 31 C, which represented high thermal stress. The light
exposure for the treatment tanks varied from 75% shade, 50% shade, or no
shade. Four control tanks with water at either 26 C or 29.3 C without
shade represented corals in an environment ideal for survival. Twenty-six
degrees Celsius was chosen as a control tank because it is the optimal
temperature for the survival of Acropora aspera and well below the mean
maximum sea surface temperature (MSST). The second group of control
tanks was established as 29.3 C due to the fact that it is at the highest
temperature of this species thermal tolerance range; however it is warm
enough to trigger moderate levels of stress. Coral survival was measured by
analysis of mean coral shade color over the length of the experiment as well
as monthly coral growth. Higher mean coral shade was correlated to darker

Tulioc 5
coral pigmentation as well as zooxanthallae density, while decreased mean
coral shade indicated loss of pigmentation or coral bleaching. Monthly coral
growth per tank indicated the positive growth of corals; specimen that
experienced bleaching would have decreased growth rates or no growth at
all. Environmental stress was measured in terms of degree heating weeks
(DHW), or the accumulation of heat stress over time. Analysis of corals
exposed to thermal stress in tanks with no shade indicated bleaching which
began as early as DHW 01. The results showed corals with 50% shade had
dark mean coral shade color up through DHW 05 whereas corals under 75%
shade exhibited healthy mean coral shade up through degree heating week
03. Shading did offer some protection to corals at the start of the
experiment, however once DHW 05 had been reached, the cumulative stress
resulted in bleaching of corals in all experimental tanks. Examination of
monthly coral growth data for corals exposed to high thermal stress of 31 C
with no shade exhibited decreased growth indicative of bleaching. Monthly
coral growth data revealed growth of corals under 50% and 75% shaded
experimental tanks at 31 C to have coral growth statistically similar to
growth of corals in both 26 C and 29.3 C. These results indicate positive
growth of corals experiencing thermal stress was possible and instances of
bleaching were decreased through the use of shade protection. Therefore
results show that shading of corals offers protection from thermal stress and
can help attenuate coral bleaching in terms of coral pigmentation and coral

Tulioc 6
growth, but once corals reach their cumulative stress threshold, bleaching is
inevitable.

Introduction
Background Information
According to the Intergovernmental Panel on Climate Change (IPCC),
global climate change is defined as any significant change in the measures
of climate lasting for an extended period of time. These changes are seen in
the land and ocean surface temperatures, which have increased 0.85 C from
1880 to 2012. Climate change is caused by an increase in greenhouse gases,
most significantly carbon dioxide, that cause heat to be trapped in our
atmosphere. Carbon dioxide concentrations have gone up about 40% since
1750 due to the burning of fossil fuels and deforestation and are increasing
by a rate of 2.0 0.1 ppm/yr (IPCC, 2014). Land and ocean temperature
measurements are taken around the world every day and analyzed to
determine the global average temperature change; it is this temperature
change that indicates the presence of a warming trend. Retreating snow
cover in the northern hemisphere, shrinking glacier volume, and the rising
global surface temperature contribute to rising sea level and rising upper
ocean heat (IPCC, 2014). According to a review in global change, the tropical
sea-surface temperature will increase by 1-3 C (Smith, 1992) and the global
sea level will rise 8 to 16 mm/year over the next century (IPCC, 2014). Each
of these factors has consequences ecologically, which are particularly
evident in the examination of coral reefs.

The relationship between global climate change and its effects on

corals are most obvious when examining ocean acidification and sea surface
temperature (SST). Twenty-five percent of the carbon dioxide released into
the atmosphere is absorbed into the ocean resulting in acidification that
decreases the calcification rates of corals. This inhibits growth in a species
already known for slow growth. In an examination of coral coverage
throughout the world, the impact of a 1% rise in CO2, where carbon dioxide
levels start off between 360 and 380 parts per million, leads to a decrease of
0.61% coverage (Chen, Chen, Chu & McCarl, 2015). Another significant factor
on coral coverage is the rising global surface temperature and rising upper
ocean heat that lead to SST increases and result in thermal stress which
induces coral bleaching. The effect of hikes in the annual mean SST directly
correlates to the sustained water temperature of corals over time. When the
temperature of the water is above the highest temperature corals can
sustain before escalations in SST, thermal stress of corals accumulates early
and continues accumulating, therefore the effects of increases in SST are
compounded due to cumulative thermal stress. In fact, a 1% increase in SST
when temperature is above 26.85 C leads to a 2.26% decrease in coral
coverage (Chen, Chen, Chu & McCarl, 2015). These are clear illustrations of
the consequences of climate change on the survival of corals.
Stony corals are colonial organisms found throughout the ocean, often
in warm climates. A reef is built when a calcium carbonate skeleton forms
and individual corals become connected by thin layers of tissue. The

skeletons that corals form are merely structural support for their over-laying
living tissues. It takes millions of years for corals to grow into the large reefs
that we see around the world today ("NOAA CoRIS - What Are Coral Reefs).
On average, corals can grow from 0.5 to 2 cm per year, however under
extremely favorable circumstances some species have been known to grow
4.5 cm in a year. Most corals survive in water between 23 and 29C. There
are known exceptions, with certain corals surviving regular drops as low as
11C or as high as 40C ("NOAA CoRIS - What Are Coral Reefs). Corals
maintain a facultative mutualistic relationship with algae known as
zooxanthallae. It is this microorganism that provides the corals with their rich
colors and produces organic compounds and oxygen as a result of
photosynthesis which the coral then utilizes. Through respiration, the coral
produces carbon dioxide and inorganic nutrients that the algae use for
photosynthesis ("NOAA CoRIS - What Are Coral Reefs). According to a review
in the evolution of coral reefs, this photosymbiosis enables stronger growth
and calcification rates of corals and is reliant on high light intensities.
Colonies of corals contain multiple types of zooxanthallae which enables a
broader range of physiological capabilities (Baker, 2003); recombination of
these symbiont types change over time and may be related to the stress
response of coral bleaching (Wood, 1998).
Light penetration is essential to the photosymbiosis of corals and
zooxanthallae; therefore corals can only survive in very clear water.
Irradiance that corals are exposed to due to ultraviolet light (UV) or

photosynthetically active radiation (PAR) is affected by the depth of the

water. High light levels enhance coral sensitivity to environmental stress
factors such thermally and salinity-induced stress (Smith, 1992). During
times of environmental stress corals expel zooxanthallae from their skeleton,
an organismal response known as coral bleaching. Corals may recover from
episodes of bleaching through the reestablishment of symbiont distribution
(Baker, 2003), which can take up to a year because loss of zooxanthallae
results in decreased respiration and calcification (Smith, 1992). When
recovery is not possible coral death occurs; repeated bleaching incidents
over a short time increase mortality rates. Thermal stress due to increased
water temperatures is a major cause of coral bleaching and related coral
death and has the potential for extensive damage to corals (Smith, 1992).

Experiment
The coral species Acropora aspera is a branching coral that is highly
sensitive to environmental stress which makes it more susceptible to
bleaching. Due to its low tolerance and inability to recover from stress, it is
classified as a vulnerable species on the International Union for Conservation
of Nature (IUCN) Red List of threatened species. This organism thrives in
warm shallow waters from the northern Indian Ocean to the oceanic west
Pacific (Aeby, Delbeek, Lovell, et. Al, 2014) and of all the reef-building corals,
this genus is the largest surviving in terms of global coverage (Wallace,
1994). We examined the possibility of preventing coral bleaching through

shading from light in a high thermal stress environment on Acropora aspera.

The ideal temperature range of this particular coral is 25.605 - 28.617 C
("Descriptions and Articles about Acropora Aspera"). High thermal stress for
this species of coral was established as 31 C. Shading of tanks was an
attempt to reduce the amount of stress placed on the organisms. By
reducing stress, we had hoped to improve corals ability to cope with thermal
stress and therefore minimize bleaching. Experimental controls were
established by four separate tanks which were not placed under thermal
stress. All tanks were then observed over a six week period. It was
hypothesized that shading would offer protection from bleaching and corals
with the highest shade coverage of 75% would have the highest survival
rates during thermal stress.

Methodology
Preparing Coral Environment
The experiment was confined to ten twenty gallon aquaria. Before
introducing the corals into the tanks, each aquaria was set up with a heater
and a chiller which was connected to a temperature controller. These
temperature controllers were set to 80 F (equivalent to 26 C) and allowed
for a fluctuation of 1 F. Protein skimmers were added to eliminate protein
and filters were added to keep the water clean and prevent algae build up.
Filters were changed every two weeks. Each tank contained two power heads
to maintain circulation and aeration of water. Temperature loggers were put
in tanks to record temperature every ten minutes; these readings were
uploaded to the computer daily and daily averages and standard deviation of
these readings was calculated in excel to ensure water was maintained
between 26 and 26.5 C. Salt and nutrients were added to the tanks to
provide nutrients, salinity, calcium, and pH levels that corresponded to the
natural habitat of the corals.
Each day we took readings for salinity, calcium and pH to ensure water
quality in each tank was compatible with normal oceanic conditions in
America Samoa. Salinity was measured using a refractometer that was
cleaned and calibrated before each use. Refractometers take measurements
of salinity from two to three drops of water from a tank. This water is
dropped onto the glass lens and then covered with a clear plastic flap.

Salinity levels can be read by holding the refractometer up to the light and
focusing the eyepiece. After reading salinity, the refractometer is cleaned
with drops of distilled water and dried with kim wipes. The ideal salinity level
is 35 parts per thousand (ppt). Calcium readings were taken with a Salifert
calcium proficiency test. We used 2 mL of water from one tank at a time and
added Ca-1 and Ca-2 solutions provided in the kit to obtain a pink solution. A
third reagent was then added to the mixture from a syringe one drop at a
time. After each drop the mixture had to be swirled. Once the mixture
changed from pink to clear blue, the amount of reagent left in the syringe
was read. This amount corresponds with the calcium level of the tank.
Calcium must be kept between 420 and 450 parts per million (ppm). If the
calcium level was too low, additional 5 mL of nutrients was added. pH was
measured with a pH meter and must be 8.0 and 8.3. If pH levels were either
too high or too low, pH up or pH down solutions were be added.
Lights were installed above the tanks, and readings were taken to
measure the output of ultraviolet light (UV) and photosynthetically active
radiation (PAR). The measurements for UV were taken with a single reading
taken at the bottom center of the tank. Three separate measurements were
taken: without any shade, with 50% shade and with 75% shade. The PAR
measurements were also taken from the bottom of the tank in three
categories: without any shade, with 50% shade and with 75% shade. We
measured PAR from nine different areas for each category within the tank;
left front, center front, right front, left center, center center, right center, left

back, center back and right back. Once our light readings had been taken,
we set up photoperiods to regulate the time intervals that the lights would
turn on and off throughout the day. These photoperiods ensured that each
tank received the required amount of light necessary for corals to survive
without exceeding them.

Addition of Corals
The corals arrived from America Samoa on March 2, 2013. Thirty corals
were allotted for each tank and locked into egg crate stands, two spaces
apart. The corals in each tank were numbered as looked at from above.
Corals were numbered left to right, starting from the row at the front of the
tank and moving backwards. The coral closest to the front of the tank and at
the far left was coral number one. A total of 314 corals were used. The water
at this time was 26 C. For the first week, corals acclimated to their tanks
with the application of 75% shade and an adjusted the photoperiod of the
lights. By reducing the intensity of the light, the possibility of environmental
stress was decreased. Tanks were changed to 50% shade and photoperiods
were adjusted on March 8th. We allowed the corals to acclimate to this

environment for one week and then took off all shade and shortened
photoperiods on March 13th. After a week of light exposure without shade, we
then adjusted the temperature and photoperiod of each tank individually. On
March 12th we started to adjust the temperature in each tank. Each of the
twelve tanks had their own controlled temperature and light exposure: two
26 C no shade (26 C NS), two 29.3 C no shade (29.3 C NS), two 31C no
shade (31 C NS), two 31 C 50% shade (31 C 50%), and two 31 C 75%
shade (31 C 75%). All of the tanks at 26 C and 29.3 C were control tanks.
The 26 C tanks represented the lowest temperature range in which this
species could survive. The 29.3 C tanks represented the highest
temperature range corals could survive. Corals in control tanks received no
shade because they were placed in conditions ideal for survival.
Experimental tanks were those tanks with temperatures set at 31 C, which
signified thermal stress. By March 19th each tank was at its required
temperature. The experiment began on March 21st. With the launch of the
experiment, we took daily readings of salinity and temperature loggers to
ensure the correct levels were maintained. Even slight changes in salinity
and temperature can lead to heightened levels of coral stress so daily
readings ensured control of these factors. Calcium and pH readings were

taken every other day because calcium and pH levels fluctuated more
gradually. Analysis of temperature logger data was done daily in order to
keep track of degree heating weeks. The National Oceanic and Atmospheric
Administration (NOAA) defines a degree heating week as a cumulative
measurement of the intensity and duration of thermal stress, and is
expressed in the unit C-weeks. A degree heating week was calculated by
subtraction of 29.3 (which signify the highest temperature before the
threshold of stress) from the calculated daily average temperature; the sum
of this number was then divided by seven. When the calculated number is
1, a new DHW has been reached. Once the DHW 01 had been reached,
designated experimental tanks were shaded with 50% and 75% mesh
covers. When tanks reached a degree heating week, pictures of the corals
were taken from the top, front, and sides with a color scale included in the
shot. These pictures were renamed and later analyzed for coral shade.
Control tanks did not experience DHWs due to the fact that they were not
placed under thermal stress. Therefore, every time an experimental tank
reached a DHW, pictures of control tanks were also taken. These pictures are
the record of the bleaching progress in each tank.

Table 1. Light measurements per aquarium (tank).

Treatment

Tank

Mean PAR
NS

UV

50S

75S

NS

50S

75S

Control 26C

1255.6

371.6

177.8

0.95

0.21

0.04

Control 26C

11

969.9

281.9

135.0

0.40

0.09

-0.03

Control 29.3C

1203.8

365.5

186.9

0.91

0.22

0.05

Control 29.3C

1003.9

314.2

150.4

1.00

0.23

0.05

No Shade 31C

1012.2

312.8

125.5

0.20

0.02

-0.01

No Shade 31C

1196.2

409.5

170.3

0.85

0.18

0.02

50% Shade 31C

12

1143.6

365.0

167.9

0.13

0.02

-0.01

50% Shade 31C

1096.1

328.7

159.6

0.49

-0.11

-0.01

75% Shade 31C

1161.0

369.4

173.8

0.80

0.18

0.03

75% Shade 31C

10

1087.6

327.0

155.4

0.89

0.20

0.04

50S: 50% shade; 75S: 75% shade; ND: no data available; NS: not shaded; PAR:
Photosynthetic Active Radiation measured in mol m-2 s-1, values are means
calculated from 9 measurements in each tank; UV: ultraviolet radiation reading
taken at the center of the tank, measured in mol m-2 s-1

Table 2. Photoperiod and total amount of photosynthetic active radiation received

per aquarium (tank); at the beginning of the experiment and after shading was
placed in some of the aquaria in the initial stage of cumulative thermal stress
(degree heating week 01).
Beginning
Treatment
Control 26C
Control 26C

After DHW 01

Tank

Photoperiod
Total PAR
(h)
4
4.4
20

Photoperiod
Total PAR
(h)
4.4
20

11

5.7

20

5.7

20

Control 29.3C

4.6

20

4.6

20

Control 29.3C

5.5

20

5.5

20

No Shade 31C

5.5

20

5.5

20

No Shade 31C

4.6

20

4.6

20

50% Shade 31C

12

4.9

20

7.5

10

50% Shade 31C

5.1

20

8.5

10

75% Shade 31C

4.8

20

8.0

75% Shade 31C

10

5.1

20

9.0

DHW: Degree Heating Week, h: hours, PAR: Photosynthetic Active Radiation

measured in mol m-2 day-1

Control 26

Control 29

Figure A: The average temperature for every day of the experiment in the
control tanks and the standard deviation of that data. MSSST: Mean Summer
Sea Surface Temperature

HT NS

HT NS

HT 50S

HT 50S

HT 75S

HT 75S

Figure B: The mean average temperature for every day of the experiment in
the experimental tanks and the time the degree heating weeks were met.
Bars on graphs A, C and E illustrate standard deviation. MSSST: Mean
Summer Sea Surface Temperature, HT NS: High Temperature No Shade, HT
50S: High Temperature 50% Shade, HT 75S: High Temperature 75% Shade

Control 26C

n = 62
corals

Control 29.3C

n = 63
corals

HT NS

n = 64 corals

HT 50S

n = 62 corals

HT 75S

n = 63
corals

Figure C: Average coral color during each degree heating week in each of the
treatments.

n = sample size of treatment, 2<x1 = bleached, 3<x2 = bleaching, x3

= not bleached
Results
5.0

4.0

Mean Coral Color Shade

Control 26C

3.0

Control 29C

NS 31C

50S 31C

75S 31C

2.0

1.0
0.0

1.0

2.0

3.0

4.0

5.0

6.0

Degree Heating Week

* = Significant

7.0

8.0

Figure D: Mean coral shade of the different treatment tanks in relation to the
degree heating week. The chart shows the Kruskal-Wallis (KW) test and the
Dunn's Multiple Comparisons (DMC) Test and significant data. DHW: Degree
Heating Week, P: P-value
The results gathered indicate that there was no statistically significant
difference of mean coral color shade between control 26 C, control 29.3 C,
and the tank with no shade at 31 C at the start of the experiment (DHW 00).
From DHW 01 to DHW 08 the corals in treatment tanks of high thermal stress
(31 C) without any shade were statistically different from the corals in
control tanks. This shows that the specimen in treatment tanks started with
mean coral color shades comparable to those in control tanks, however once
the first DHW had been reached, experimental corals without shade quickly
bleached. Due to the fact that treatment corals were not shaded until after
DHW 01 was reached, the expected results would have been for control
corals and treatment corals to be similar up through DHW 01. Treatment
tanks with the 50% shade were comparable to the control tanks at 26 C up
to DHW 03 and for control tanks at 29.3 C up to DHW 02. Once these DHW
passed, corals under the 50% shade show statistically significant differences
indicating that the corals in the 50% shade treatment tanks bleached
steadily for the remainder of the experiment. Treatment tanks under 75%
shade were statistically different from the corals in control tanks at both 26
C and 29.3 C from DHW 03 to DHW 08. Therefore they avoided bleaching
significantly only through DHW 02. Comparisons of corals from the treatment
tanks with no shade with those under 50% shade show specimen in the two
treatment tanks to be statistically different from DHW 01 through DHW 05.

This shows that corals in both treatments started off with similar coral color;
however from DHW 01 to 05, corals under 50% shade had lower instances of
bleaching. Corals in treatment tanks under 75% shade were statistically
different from those with no shade only from DHW 02 to DHW 03. Corals in
the tank with 75% shade fared slightly better than those with no shade only
through DHW 03. Therefore, both the 75% and 50% shades offer adequate
protection through DHW 03. The 50% shade provided protection longer than
the 75% with better results up through DHW 05. These results indicate that
the 50% shade offers better protection than the 75%, however neither types
of shade will prevent coral bleaching once corals experience thermal stress
up to six DHWs.
0.4

0.3

0.2

Growth Per Month (Buoyant weight g)

0.1

* = Significant
Figure E: Mean coral growth per month based on the sample size in each
tank treatment. The chart shows the Kruskal-Wallis (KW) test and the Dunn's
Multiple Comparisons (DMC) Test and significant data. Numbers above bars
indicate sample size. P: P-value
The monthly coral growth data indicates that the corals in control tanks
at 26 C and 29.3 C had growth that was not statistically different which
shows positive growth for all corals in control tanks. The corals in treatment
tanks under conditions of thermal stress (31 C) with no shade had a
statistically significant difference from corals in control tanks at 26 C,
however they did not have this difference when compared to organisms in
control tanks at 29.3 C. Further examination of control tanks at 29.3 C
revealed that the temperature in tank 06 fluctuated up to 30 C for two days,
down to 28 C for another two days and then back up to 30 C for four more
days; this was the last fluctuation, the temperature was kept at 29.3 for the
remainder of the experiment. These increases in temperature could have
caused enough thermal stress to the corals to inhibit coral growth or the
temperature of 29.3 C may be high enough to induce stress levels that halt
growth. The only clear significance that can be drawn from the temperature
fluctuations is the lack of statistical difference in growth from the corals in
the control tanks with the corals in experimental no shade tanks. It was

expected that corals exposed to the treatment with no shade would have no
growth while corals in both control tanks were expected to have positive
coral growth. This indicates that the corals in control tank 26 C grew more
than in control tank 29.3 C; this difference is not statistically significant, but
it accounts for the lack of significant difference in the growth comparison of
the corals under no shade and 31 C with controls in the 29.3 C
environment. The monthly coral growth of treatment tanks under both 50%
shade and 75% shade were not significantly statistically different from
control tanks 26 C and 29.3 C. This implies that the growth of shaded
corals exposed to thermal stress was similar to the positive growth of corals
in control tanks. There was a difference in the monthly coral growth of the
treatment corals under no shade compared to treatment corals under 50%
shade and 75% shade that was statistically different. This difference reveals
that corals experiencing high thermal stress of 31 C were protected from
light stress that inhibited growth through both 50% and 75% shading
techniques. Comparison of corals in treatment tanks with 50% shade and
75% shade revealed no statistically significant difference in monthly coral
growth. Therefore, both shade treatments provided comparable protection
from light stress for corals.
Analysis of the mean coral color shade data and the monthly coral
growth data indicates that shading of corals exposed to high thermal stress
of 31 C offers some degree of protection from high irradiance and a certain
extent of protection from bleaching. Shading of both 50% and 75% offers

minimal overall protection because once corals have reached a certain stress
threshold, indicated by the data to be DHW 03, bleaching is inevitable. The
50% shade protected corals for a longer duration, as indicated in Figure D,
but it still did not provide full protection from bleaching. Under conditions of
high thermal stress and high light stress, corals exhibited increased rates of
bleaching and lack of coral growth. Corals kept under low light levels (50%
and 75% shade) and increased thermal stress did eventually bleach,
however they also exhibited positive growth which indicates photosynthetic
activity. The bleaching of these corals was a result of thermal stress however
it is possible that the remaining presence of zooxanthallae on the host
continued photosynthesis which created coral growth. The parameters of this
experiment did not account for the rates of photosynthesis, therefore the
relationship between growth and photosynthetic capability cannot be
established.

Discussion
In each of the aquaria the results were near what we expected in
comparison to each other. The control tanks bleached a lot later in time than
the tanks under thermal stress, even when tanks were shaded. In a study
done by Bhagooli and Hidaka it was found that light and photosynthetic
symbionts dramatically change the way different corals respond to
bleaching. Their experiment indicated that thermal stress reduced corals

ability to withstand light intensity (Bhagooli & Hidaka, 2003). Based off of
our results, thermal stress is much more significant than stress from light. In
a study done by Anthony, Connolly and Hoegh-Guldberg, the relationship
between thermal stress, light stress and sedimentation was observed. In this
particular study, it was found that under conditions of high thermal stress,
high light intensity, and high water sedimentation, corals exhibited positive
growth and darker pigmentation whereas corals kept in conditions of high
thermal stress, high light intensity and low levels of sedimentation exhibited
no growth and loss of pigment. The growth and pigment of specimen in the
tanks with high sedimentation was due to protection from environmental
stress provided by the cloudy water. This protection from light is similar to
the shading technique used in our experiment, with protection from light
stemming from suspended sediment in the water instead of mesh covers.
This is an approach to decreasing irradiance that could be cross-examined
with overhead shading to find further coral protection. This research revealed
that the importance of light was relevant only to corals under thermal stress
and effects became more apparent over time (Anthony, Connolly, & HoeghGuldberg, 2007). Similarly to the results obtained in our experiment, these
results revealed that high light intensity increased coral mortality when
corals were exposed to thermal stress. It was also found that bleaching
occurred in higher instances later in the experiment which implies the buildup of stress factors. At 31 degrees Celsius the corals began to bleach earlier
than our control tanks no matter what shade they were under. In fact, our

50% shade tank had only slightly less bleached corals as time passed. The
parameters of our experiment did not account for the possibility of
photoinhibition, a factor that may have contributed to incidence of bleaching
and the inhibition of growth due to the photosynthetic nature of the
organism. Photoinhibition is the reduction of photosynthetic capability of
corals due to an abundance of light and it is found through the comparison of
the rate of photodamage to photosystem II (PS II) and the rate of PS II repair.
In a review of environmental stress and photoinhibition done by Takahasi and
Murata, it was found that environmental stress coupled with high light
intensities increases photoinhibition because of an inability to repair PSII, the
reaction center that controls oxygenic photosynthesis. During times of
thermal stress, photosynthetic fixation of carbon dioxide is limited, which in
turn limits the photosynthetic capabilities of the corals (Takahashi & Murata,
2008). Future repetitions of this experiment could include photoinhibition
factors and measurements of photosynthetic rates which would provide more
all-inclusive results. Although thermal stress was the main cause for our
corals to bleach, reducing light stress did show some signs of benefitting the
coral growth. In conclusion one thing we can state as fact is that higher
temperatures cause corals to bleach faster no matter what protection from
light is added into the equation.

Acknowledgements
Supported by Dr. Vania Coelho alongside the Department of Natural Sciences
at Dominican University of California.

I would like to thank Dr. Vania Coelho for allowing me to work with research
she is so passionate and invested in, as well as for providing me with
guidance support throughout the entirety of this research project. Thanks are
also extended to Dr. Kenneth Frost for working with me during the editing of
this thesis and providing an outside ecological perspective on data analysis. I
would also like to thank Jacob Steele, Elizabeth Rice, Connor Bickler, Devan
Klein, Leeanne Obilor, Mary Barana, and Kathryn Que for their hard work and
gathering of data in the coral lab. Lastly, I would like to thank Dr. Diara Spain
for her guidance through this Honors thesis and her support and
encouragement throughout my career in the Honors Program.

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