Coral bleaching is the loss of intracellular endosymbionts (Symbiodinium, also known as zooxanthellae) through either expulsion or loss of algal pigmentation. The corals that form the structure of the great reef ecosystems of tropical seas depend upon a symbiotic relationship with algae-like unicellular flagellate protozoa that are photosynthetic and live within their tissues. Zooxanthellae give coral its coloration, with the specific color depending on the particular clade. Some scientists consider bleaching a poorly-understood type of "stress" related to high irradiance; environmental factors like sediments, harmful chemicals and freshwater; and high or low water temperatures. This "stress" causes corals to expel their zooxanthellae, which leads to a lighter or completely white appearance, hence the term "bleached". Bleaching has been attributed to a defense mechanism in corals; this is called the "adaptive bleaching hypothesis," from a 1993 paper by Robert Buddemeier and Daphne Fautin. Bleached corals continue to live, but growth is limited until the protozoa return.
- 1 Causes
- 2 Triggers
- 3 Effects
- 4 Mass bleaching events
- 5 Impact
- 6 Coral adaptation
- 7 See also
- 8 Notes
- 9 References
- 10 External links
Bleaching occurs when the conditions necessary to sustain the coral's zooxanthellae cannot be maintained. Any environmental trigger that affects the coral's ability to supply the zooxanthellae with nutrients for photosynthesis (carbon dioxide, ammonium) will lead to expulsion. This process is a "downward spiral", whereby the coral's failure to prevent the division of zooxanthellae leads to ever-greater amounts of the photosynthesis-derived carbon to be diverted into the algae rather than the coral. This makes the energy balance required for the coral to continue sustaining its algae more fragile, and hence the coral loses the ability to maintain its parasitic control on its zooxanthellae.
Physiologically the lipid composition of the symbiont thylakoid membrane affects their structural integrity when there is a change in temperature, which combined with increased nitric acid results in damage to photosystem II. As a result of accumulated oxidative stress and the damage to the thylakoid of chloroplasts there is an increase in degradation of the symbiosis and the symbionts will eventually abandon their host. Not only does the change in temperature in the water increase the chances of bleaching, but there are other factors that play a role. Other factors include an increase in solar radiation (UV and visible light), regional weather conditions, and for intertidal corals, exposure to cold winds.
Coral bleaching is theorized to be a generalized stress response of corals that may be caused by a number of biotic and abiotic factors, including:
- increased (most commonly), or reduced water temperatures
- oxygen starvation caused by an increase in zooplankton levels as a result of overfishing
- increased solar irradiance (photosynthetically active radiation and ultraviolet light)
- changes in water chemistry (in particular acidification)
- increased sedimentation (due to silt runoff)
- bacterial infections
- changes in salinity
- low tide and exposure
- cyanide fishing
- elevated sea levels due to global warming (Watson)
- mineral dust from African dust storms caused by drought
- four common sunscreen ingredients, that are nonbiodegradable, and can wash off from tourists swimming, snorkeling, or diving
While most of these triggers may result in localized bleaching events (tens to hundreds of kilometers), mass coral bleaching events occur at a regional or global scale and are triggered by periods of elevated thermal stress resulting from increased sea surface temperatures. The coral reefs that are more subject to continued bleaching threats are the ones located in warm and shallow water with low water flow. Physical factors that can prevent or reduce the severity of bleaching are available for the reefs located under conditions that include low light, cloud cover, high water flow and higher nutrient availability.
The color of a coral depends largely on the species of symbiont. A reduction in concentration of zooxanthellae causes paling and an increase results in deepening of color. Stony corals have calcium carbonate skeletons and most have transparent tissues, so expulsion of the zooxanthellae causes them to lose their color and become white. Although the coral polyps feed on zooplankton and other food particles, the majority of reef-forming corals rely for a large proportion of their nutritional requirements on their zooxanthellae. This means that without them they are liable to starve. Coral growth and reproduction are reduced and the coral becomes increasingly susceptible to disease. If stress factors reduce and the zooxanthellae return, the coral can recover, but prolonged bleaching causes death of the coral.
Ejection increases the polyp's chance of surviving short-term stress. It can regain symbionts, possibly of a different species, at a later time. If the stressful conditions persist, the polyp eventually dies.
Mass bleaching events
Most evidence indicates that elevated temperature is the cause of mass bleaching events. Sixty major episodes of coral bleaching have occurred between 1979 and 1990, with the associated coral mortality affecting reefs in every part of the world. Correlative field studies have pointed to warmer-than normal conditions as being responsible for triggering mass bleaching events. These studies show a tight association between warmer-than-normal conditions (at least 1 °C higher than the summer maximum) and the incidence of coral bleaching.
Factors that influence the outcome of a bleaching event include stress-resistance which reduces bleaching, tolerance to the absence of zooxanthellae, and how quickly new coral grows to replace the dead. Due to the patchy nature of bleaching, local climatic conditions such as shade or a stream of cooler water can reduce bleaching incidence. Coral and zooxanthellae health and genetics also influence bleaching.
Large coral colonies such as Porites are able to withstand extreme temperature shocks, while fragile branching corals such Acropora are far more susceptible to stress following a temperature change. Corals consistently exposed to low stress levels may be more resistant to bleaching.
Monitoring reef sea surface temperature
The US National Oceanic and Atmospheric Administration (NOAA) monitors for bleaching "hot spots", areas where sea surface temperature rises 1 °C or more above the long-term monthly average. This system detected the worldwide 1998 bleaching event, that corresponded to an El Niño. NOAA also uses a nighttime-only satellite; these observations are taken at night to avoid the increase in temperature due to daily warming caused by solar heating at the sea surface during the day. This is also a precaution to avoid glare from the sun.
Changes in ocean chemistry
Increasing ocean acidification due to rises in carbon dioxide levels exacerbates the bleaching effects of thermal stress. Acidification affects the corals' ability to create calcareous skeletons, essential to their survival. A recent study from the Atkinson Center for a Sustainable Future found that with the combination of acidification and temperature rises, the levels of CO2 could become too high for coral to survive in as little as 50 years.
Infectious bacteria of the species Vibrio shiloi are the bleaching agent of Oculina patagonica in the Mediterranean Sea, causing this effect by attacking the zooxanthellae. V. shiloi is infectious only during warm periods. Elevated temperature increases the virulence of V. shiloi, which then become able to adhere to a beta-galactoside-containing receptor in the surface mucus of the host coral. V. shiloi then penetrates the coral's epidermis, multiplies, and produces both heat-stable and heat-sensitive toxins, which affect zooxanthellae by inhibiting photosynthesis and causing lysis.
During the summer of 2003, coral reefs in the Mediterranean Sea appeared to gain resistance to the pathogen, and further infection was not observed. The main hypothesis for the emerged resistance is the presence of symbiotic communities of protective bacteria living in the corals. The bacterial species capable of lysing V. shiloi had not been identified as of 2011.
In the 2012-2040 period, coral reefs are expected to experience more frequent bleaching events. The Intergovernmental Panel on Climate Change (IPCC) sees this as the greatest threat to the world's reef systems.
Great Barrier Reef
The Great Barrier Reef along the coast of Australia experienced bleaching events in 1980, 1982, 1992, 1994, 1998, 2002, and 2006. Some locations suffered severe damage, with up to 90% mortality. The most widespread and intense events occurred in the summers of 1998 and 2002, with 42% and 54% respectively of reefs bleached to some extent, and 18% strongly bleached. However coral losses on the reef between 1995 and 2009 were largely offset by growth of new corals. An overall analysis of coral loss found that coral populations on the Great Barrier Reef had declined by 50.7% from 1985 to 2012, but with only about 10% of that decline attributable to bleaching, and the remaining 90% caused about equally by tropical cyclones and by predation by crown-of-thorns starfishes.
The IPCC's moderate warming scenarios (B1 to A1T, 2 °C by 2100, IPCC, 2007, Table SPM.3, p. 13) forecast that corals on the Great Barrier Reef are very likely to regularly experience summer temperatures high enough to induce bleaching.
Other coral reef provinces have been permanently damaged by warm sea temperatures, most severely in the Indian Ocean. Up to 90% of coral cover has been lost in the Maldives, Sri Lanka, Kenya and Tanzania and in the Seychelles.
Evidence of thermal tolerance in Hawaiian corals and of oceanic warming from research in the 1970s led researchers in 1990 to predict mass occurrences of coral bleaching throughout Hawaii. Major bleaching occurred in 1996 and in 2002. Biologists from the University of Queensland observed the first mass bleaching event for Hawaiian coral reefs in 2014, and attributed it to The Blob.
Economic and political impact
According to Brian Skoloff of The Christian Science Monitor, "If the reefs vanished, experts say, hunger, poverty and political instability could ensue." Since countless sea life depends on the reefs for shelter and protection from predators, the extinction of the reefs would ultimately create a domino effect that would trickle down to the many human societies that depend on those fish for food and livelihood. There has been a 44% decline over the last 20 years in the Florida Keys, and up to 80% in the Caribbean alone.
In 2010, researchers at Penn State discovered corals that were thriving while utilizing an unusual species of symbiotic algae in the warm waters of the Andaman Sea located in the Indian Ocean. Normal zooxanthellae cannot withstand temperatures as high as in that location, so this finding was unexpected. This gives researchers hope that with rising temperatures due to global warming, coral reefs will develop tolerance for different species of symbiotic algae that are resistant to high temperature, and can live within the reefs.
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