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Pacific in Peril - Greenpeace

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Figure 3. Bleached corals<br />

photographed <strong>in</strong> French<br />

Polynesia dur<strong>in</strong>g the 1994<br />

<strong>Pacific</strong> bleach<strong>in</strong>g event.<br />

Photography: Roger<br />

Grace/<strong>Greenpeace</strong><br />

International<br />

THERMAL STRESS AND CORAL BLEACHING<br />

Coral bleach<strong>in</strong>g is one of the pr<strong>in</strong>cipal responses of<br />

corals and their symbiotic d<strong>in</strong>oflagellates to sudden<br />

changes <strong>in</strong> a range of conditions (eg, sal<strong>in</strong>ity,<br />

temperature, light; Brown, 1997b; Hoegh-Guldberg,<br />

1999). Normally the population densities of<br />

zooxanthellae <strong>in</strong> reef-build<strong>in</strong>g corals range between 0.5<br />

and 5 x 106 cell.cm -2 (Drew, 1972; Porter et al, 1984;<br />

Hoegh-Guldberg and Smith, 1989). Zooxanthellae<br />

typically show very low rates of migration or expulsion<br />

to the water column under normal conditions (Hoegh-<br />

Guldberg et al, 1987). Populations of zooxanthellae may<br />

be adjusted, however, as corals acclimatise to different<br />

environmental or seasonal conditions (Jones, 1995;<br />

Fagoonee et al, 1999; Fitt et al, 1999). Sudden<br />

reductions <strong>in</strong> the number of zooxanthellae will occur<br />

when conditions such as water temperature and light<br />

change around them. Normally zooxanthellae impart a<br />

brown colour to the tissues of corals. Consequently,<br />

when the zooxanthellae are expelled, corals turn from<br />

brown to white, the process termed bleach<strong>in</strong>g (Figure 3).<br />

Coral bleach<strong>in</strong>g may occur at local reef scales (ie,<br />

over hundreds of square metres) <strong>in</strong> response to sudden<br />

changes <strong>in</strong> sal<strong>in</strong>ity, temperature or light (Egana and<br />

DiSalvo, 1982; Goreau, 1964; Hoegh-Guldberg, 1999)<br />

or at geographic scales that may <strong>in</strong>volve entire reef<br />

systems and geographic realms (Glynn, 1984; Goreau,<br />

1990; Williams and Williams, 1990; Hoegh-Guldberg<br />

and Salvat, 1995; Brown, 1997b; Hoegh-Guldberg,<br />

1999).<br />

The latter has been referred to as mass coral<br />

bleach<strong>in</strong>g. Its scale and <strong>in</strong>tensity, as well as its <strong>in</strong>creas<strong>in</strong>g<br />

frequency, has many scientists conv<strong>in</strong>ced that it<br />

represents a serious challenge to the health of the world’s<br />

coral reefs. This concern has been heightened by recent<br />

evidence that the specific thermal trigger required to<br />

cause coral bleach<strong>in</strong>g will be exceeded on an annual<br />

basis <strong>in</strong> the next few decades.<br />

Mass coral bleach<strong>in</strong>g events have occurred <strong>in</strong> six<br />

major periods that have generally co<strong>in</strong>cided with<br />

disturbances to the El Niño Southern Oscillation<br />

(ENSO) cycle (Glynn, 1993; Hoegh-Guldberg, 1999).<br />

While reduced sal<strong>in</strong>ity (Egana and DiSalvo, 1982;<br />

Goreau, 1964), <strong>in</strong>creased or decreased light (Vaughan,<br />

1914; Yonge and Nicholls, 1931; Hoegh-Guldberg and<br />

Smith, 1989b; Gleason and Well<strong>in</strong>gton, 1993; Lesser et<br />

al, 1990) and chemical factors such as copper<br />

contam<strong>in</strong>ation (Jones, 1997a), cyanide (Jones and<br />

Steven, 1997; Jones and Hoegh-Guldberg, 1999),<br />

herbicides, pesticides and biological factors (eg, bacteria,<br />

Kushmaro and Loya, 1996) can also evoke the loss of<br />

algal pigments from symbiotic <strong>in</strong>vertebrates, the<br />

primary cause of mass coral bleach<strong>in</strong>g is temperature<br />

(Hoegh-Guldberg and Smith, 1989; Glynn and D’Croz,<br />

1991; Glynn, 1993; Hoegh-Guldberg, 1999). The<br />

widespread bleach<strong>in</strong>g events of 1998 have added<br />

further weight to the argument that elevated<br />

temperature is the primary variable trigger<strong>in</strong>g coral<br />

bleach<strong>in</strong>g. The accuracy and predictive capability of the<br />

“HotSpot” program (Goreau and Hayes, 1994; Strong et<br />

al, 1996) has further strengthened the case for<br />

temperature be<strong>in</strong>g the primary variable that expla<strong>in</strong>s the<br />

recent spate of mass bleach<strong>in</strong>g events <strong>in</strong> the 1980s and<br />

1990s.<br />

The advent of thermal stress <strong>in</strong> corald<strong>in</strong>oflagellate<br />

symbioses is very similar to the<br />

mechanism of heat stress <strong>in</strong> higher plants (Jones et al,<br />

1998; Hoegh-Guldberg, 1999) and results <strong>in</strong><br />

hypersensitivity to light of the symbiotic d<strong>in</strong>oflagellates<br />

of corals. Whereas normal light levels are essential for<br />

the survival of corals (through the power<strong>in</strong>g of the<br />

photosynthetic processes of the symbiotic<br />

d<strong>in</strong>oflagellates), light <strong>in</strong>tensities after heat stress are a<br />

liability. This leads to chronic photo<strong>in</strong>hibition of the<br />

symbiotic d<strong>in</strong>oflagellates and the light-dependent<br />

destruction of the symbiotic d<strong>in</strong>oflagellates. After they<br />

become non-functional, the damaged symbionts are<br />

expelled from (or beg<strong>in</strong> to leave) the coral host. A<br />

complete review of the biochemical and physiological<br />

basis of coral bleach<strong>in</strong>g can be found <strong>in</strong> Hoegh-<br />

Guldberg (1999).<br />

The fate of corals follow<strong>in</strong>g coral bleach<strong>in</strong>g is of<br />

crucial importance. The severity of the thermal stress<br />

largely determ<strong>in</strong>es the outcome of any particular<br />

thermal event. In large and long-lived thermal<br />

anomalies, reef-build<strong>in</strong>g corals will die <strong>in</strong> large numbers<br />

(Brown and Suharsono, 1990; Glynn, 1990). In 1998,<br />

almost total mortality of corals was seen on part of the<br />

Great Barrier Reef <strong>in</strong> Queensland (Marshall and Baird,<br />

2000), on Scott Reef of north Western Australia (Smith<br />

and Heywood, pers comm), <strong>in</strong> the Seychelles and<br />

Maldives (Spencer et al, 2000), <strong>in</strong> Ok<strong>in</strong>awa (Y Loya, W<br />

Loh and K Sakai, pers comm) and <strong>in</strong> Palau (J Bruno,<br />

pers comm).<br />

Not all corals are equally susceptible. Branch<strong>in</strong>g<br />

corals (eg, Acropora) tend to be more sensitive than more<br />

massive types of corals (eg, Porites). In milder events,<br />

death rates may be m<strong>in</strong>imal (Harriott, 1985) and corals<br />

may make almost complete recoveries. This may take<br />

weeks to months and may result <strong>in</strong> corals that appear<br />

unaffected by the previous thermal <strong>in</strong>sult (Hoegh-<br />

Guldberg and Smith, 1989). Recent evidence, however,<br />

reveals that while corals may not die, their physiologies<br />

can be severely disrupted by thermal stress (Ward et al,<br />

1998; Hoegh-Guldberg et al, 2000).<br />

Corals that have bleached and recovered may<br />

have reduced growth, as well as depressed calcification<br />

and repair capabilities (Goreau and Macfarlane, 1990;<br />

Glynn, 1993; Meesters and Bak, 1993). These impacts<br />

are probably a direct result of the expulsion of the<br />

13 | <strong>Pacific</strong> <strong>in</strong> <strong>Peril</strong>

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