Chapter Coral reef disturbances This chapter covers the principal disturbances at various levels occurring or2 seen as a problem in coral reefs (including non-reefal coral communities) in Japan. The disturbances seen on natural environments are often caused by combination of anthropogenic and natural causes. On coral reefs, dredging by con- struction works and terrestrial red soil inflows are included in the former cause to a large extent, and world-wide mass bleaching event due to modern human activities can also be included as well with high possibilities. While outbreak of Acanthaster planci and widespread of disease-causing sponges on cor- als are generally considered as the later cause, causal link with human activities still cannot be denied. A new problem has brought about accompanied by the modern economic framework, turning some rural peoples’ simple, traditional lives to destructive modern living styles. In addition, a problem of garbage outflow on the sea has become a cross-border debate. We have to take these situations seriously, under- stand each disturbing factors, and implement concrete countermeasures toward improvement. Furthermore, it is necessary to understand these disparate-looking phenomena as total, interdependent phenomena, including social and economic aspects, and establish a comprehensive philosophy on our behavior. We should strongly be aware that the most ‘threatening disturbance’ is our ‘resignation and dis- interest’ to the facing problems. It must be our duty, extending to the future, to exert an effort to achieve a good relationship with the blessing nature. 0201_03（英） 04.9.3 3:03 PM ページ 42 02 Coral Reefs of Japan -1 Global environmental change and coral bleaching 2 Yoshikatsu Nakano Milleporina in the Class Hydrozoa (Nishihira and Veron 1 Introduction 1995). Many coral reef researchers around the world recorded The symbiotic algae in corals are called zooxanthellae, the large-scale coral bleaching event that occurred from and are dinoflagellates (~10 µm in diameter) belonging to 1997 to 1998. Wilkinson (1998) used the Internet to moni- the genus Symbiodinium (Photo. 1), which lives in the tor bleaching all over the world, and provided a record of gastrodermal cells of the coral polyp (Kawaguti 1944; the extent of the event. In Japan, information about the Muscatine 1980). To summarize the symbiotic relation- coral bleaching was communicated to the public via the ship, the zooxanthellae use some of the carbon dioxide website of the Japanese Coral Reef Society*1 (Kayanne generated by coral respiration for photosynthesis, and the 2002). coral utilizes glycerol etc. produced by algal photosynthe- sis as an energy source (Grant et al. 1999). The algae also The rapid communication of this event symbolized the use coral metabolic waste products, such as ammoniums age of the Internet, indicating that our use of technology and phosphates, as nutrients for growth. This symbiosis to disseminate information was advancing in parallel with allows corals to flourish in oligotrophic tropical seas. The global environmental change. Unlike local disturbances presence of zooxanthellae in coral tissue also promotes in coral reefs, this bleaching event occurred on a large coral calcification (Hidaka 2002); therefore, this symbio- scale, within a short period. Research on the bleaching in sis is important for reef formation by these hermatypic or hermatypic corals was accelerated worldwide. Surveys ‘reef-building’ corals. and research conducted in Japan resulted in many pub- lished reports on coral bleaching in that region (Research There are millions of zooxanthellae in 1 cm2 of the tissue Institute for Subtropics 1999; Nature Conservation surface of a coral colony. Zooxanthellae undergo daily Bureau, Ministry of the Environment 2002). cell divisions in the coral tissue and cells, and are dis- charged when they cease to function. This cycle is con- This section is an overview of the status of coral reef trolled by the interaction of corals and zooxanthellae, and ecosystems, one of the oldest existing ecosystems on the density of zooxanthellae is adjusted according to Earth (Veron 1995), the hermatypic corals that are the external environment parameters (Smith and Muscatine keystone species of these habitats, and the current state 1999). of Japanese corals. This information has been collected from recent reports by academic societies, the media, and other sources. 2 Symbiosis between corals and zooxanthellae Hermatypic coral (hereafter, coral) is the general name applied to those species that belong to several different groups of the phylum Cnidaria, and which contain unicel- lular algae in their tissues. This includes most species of the order Scleractinia, all the species of the orders Photo. 1. Zooxanthellae (diameter ~10 µm), with some Coenothecalia, Stolonifera in the Class Anthozoa, and undergoing cell division. 42 0201_03（英） 04.9.3 3:03 PM ページ 43 A BC Photo. 2. A: Healthy Acropora hyacinthus colony on the reef flat at 2m depth (April 1998) in front of the Sesoko Station, Tropical Biosphere Research Center, University of the Ryukyus. B: Same colony as in 2A, showing bleaching (August 1998). C: Dead colony, the surface of which is covered with algae and silt (October 1998). alone; therefore, they will become energy deficient. If this 3 What is coral bleaching? condition lasts for an extended period, corals exhaust stored energy supplies and mortality occurs (Photo. 2C). Rapid environmental change causes coral stress, making it difficult to maintain the symbiotic relationship. When There is a boundary between survival and mortality in the symbiotic relationship collapses, zooxanthellae are bleached corals. Jones (1997) has reported that bleached eliminated from the host coral, causing ‘bleaching’. corals are recoverable if the density of zooxanthellae has Because zooxanthellae are brown in color and very abun- decreased; they are not recoverable, however, if photo- dant, a healthy coral appears to have a brown coloration synthetic pigment has been lost in conjunction with the (Photo. 2A). Coral tissues are thin and almost transpar- decrease in number. While coral diseases have also been ent, over a white calcareous skeleton, therefore when reported as potential bleaching factors, the underlying zooxanthellae are expelled, the colonies appear white and mechanisms have not yet been clarified (Kushmaro et al. the corals are referred to as ‘bleached’ (Photo. 2B). This 1996; Nakano and Yamashiro 2002). bleaching is a symptom of stress, but coral tissues can survive under bleached conditions for a limited period of time. 4 Response of corals to high water temperature Extremes of environmental factors such as temperature, light, and salinity can induce bleaching (Nakano 2002a). There is a consensus of opinion that an abnormal rise in When these factors become higher or lower than the tol- seawater temperatures around the world was the main erance range of the coral-zooxanthellae association, the cause of the large-scale bleaching of coral reefs in 1997- resultant stress can lead to bleaching. Patterns of zooxan- 98 (Wilkinson 1998). Jokiel and Coles (1990) reported thellae expulsion vary, according to the fluctuation level that a water temperature rise of as little as 1°C in summer of environmental stress: 1) When the change is rapid and can cause bleaching, eventually leading to coral death. large, coral tissues are damaged first and then zooxan- Subsequent bleaching studies have indicated that the thellae will be discharged (Muscatine et al. 1992). This is main cause of the worldwide coral bleaching in 1997-1998 considered acute bleaching, and is a severe threat to was a sustained temperature that was 1-2°C higher than coral health. In such cases, most of the discharged zoox- the average water temperature in summer. anthellae are healthy. 2) When the change is compara- tively gradual, there is no remarkable damage to coral tis- High water temperature increases the production of toxic sues, however, zooxanthellae are gradually discharged. substances such as active oxygen within zooxanthellae Such a discharge is considered to occur due to abnormali- (Lesser et al. 1990). Oxygen free radicals has an unstable ty of cells (ie., loss of pigment and/or cell contraction) charge and is in a high-energy state, which causes dam- (Kuroki and van Woesik 1999). This condition is known age to cell organelles as well as to Photo System II, thus as chronic bleaching and, although corals can survive for obstructing photosynthesis (Lesser 1997). Corals may short periods in the bleached state, they cannot supply discharge zooxanthellae in self-defense. their energy requirements from heterotrophic sources Chapter-2 ■ Coral reef disturbances 43 0201_03（英） 04.9.3 3:03 PM ページ 44 02 Coral Reefs of Japan High water temperature reduces the amount of photosyn- thesis and increases the amount of respiration, which 5 Impacts of bleaching on corals and increases the carbon dioxide concentration in cells and coral reefs creates a situation where bleaching readily occurs (Pecheux 1998). After an improvement in environmental conditions, the coral colonies that survived the bleaching showed The response of corals to high water temperature differs increased levels of infection by diseases during the recov- according to optical conditions, including penetration of ery process (Williams and Williams 1990). In some ultraviolet rays (Glynn et al. 1992; Hough-Guldberg and corals, the effects of bleaching were manifested in the fol- Smith 1989). Takahashi et al. (2001) reported that lowing reproductive season. In Pocillopora verrucosa Acropora digitifera, which is a dominant species on many colonies that recovered after bleaching in 1998, both the reefs, showed photoinhibition by 250 µmol/m2/s or more number of reproductive colonies and their fertilization of radiation at a water temperature of 28°C, and by 100 rates decreased in 1999. Similarly, colonies of Montipora µmol/m2/s at 32°C. The recovery speed from photoinhi- digitata that recovered from bleaching showed decreased bition rapidly decreases at temperatures above 30°C, as numbers of oocytes and smaller testes in the reproduc- compared to lower temperatures.