Pacific in Peril - Greenpeace
Pacific in Peril - Greenpeace
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>