Coles S L, Brown Barbara E
Department of Natural Sciences, Bishop Museum, 1525 Bernice St., Honolulu, HI 96734, USA.
Adv Mar Biol. 2003;46:183-223. doi: 10.1016/s0065-2881(03)46004-5.
Coral bleaching, i.e., loss of most of the symbiotic zooxanthellae normally found within coral tissue, has occurred with increasing frequency on coral reefs throughout the world in the last 20 years, mostly during periods of El Nino Southern Oscillation (ENSO). Experiments and observations indicate that coral bleaching results primarily from elevated seawater temperatures under high light conditions, which increases rates of biochemical reactions associated with zooxanthellar photosynthesis, producing toxic forms of oxygen that interfere with cellular processes. Published projections of a baseline of increasing ocean temperature resulting from global warming have suggested that annual temperature maxima within 30 years may be at levels that will cause frequent coral bleaching and widespread mortality leading to decline of corals as dominant organisms on reefs. However, these projections have not considered the high variability in bleaching response that occurs among corals both within and among species. There is information that corals and their symbionts may be capable of acclimatization and selective adaptation to elevated temperatures that have already resulted in bleaching resistant coral populations, both locally and regionally, in various areas of the world. There are possible mechanisms that might provide resistance and protection to increased temperature and light. These include inducible heat shock proteins that act in refolding denatured cellular and structural proteins, production of oxidative enzymes that inactivate harmful oxygen radicals, fluorescent coral pigments that both reflect and dissipate light energy, and phenotypic adaptations of zooxanthellae and adaptive shifts in their populations at higher temperatures. Such mechanisms, when considered in conjunction with experimental and observational evidence for coral recovery in areas that have undergone coral bleaching, suggest an as yet undefined capacity in corals and zooxanthellae to adapt to conditions that have induced coral bleaching. Clearly, there are limits to acclimatory processes that can counter coral bleaching resulting from elevated sea temperatures, but scientific models will not accurately predict the fate of reef corals until we have a better understanding of coral-algal acclimatization/adaptation potential. Research is particularly needed with respect to the molecular and physiological mechanisms that promote thermal tolerance in corals and zooxanthellae and identification of genetic characteristics responsible for the variety of responses that occur in a coral bleaching event. Only then will we have some idea of the nature of likely responses, the timescales involved and the role of 'experience' in modifying bleaching impact.
珊瑚白化,即珊瑚组织内通常存在的大部分共生虫黄藻的丧失,在过去20年里,在世界各地的珊瑚礁上出现的频率越来越高,主要发生在厄尔尼诺南方涛动(ENSO)期间。实验和观察表明,珊瑚白化主要是由高光条件下海水温度升高导致的,这会加快与虫黄藻光合作用相关的生化反应速率,产生有毒形式的氧,干扰细胞过程。已发表的关于全球变暖导致海洋温度上升基线的预测表明,30年内的年度最高温度可能达到会导致频繁珊瑚白化和广泛死亡的水平,从而导致珊瑚作为珊瑚礁上的优势生物数量减少。然而,这些预测没有考虑到珊瑚种内和种间白化反应的高度变异性。有信息表明,珊瑚及其共生体可能能够适应并选择性地适应已经导致具有抗白化能力的珊瑚种群出现的升高温度,这种现象在世界不同地区的局部和区域都有发生。可能存在一些机制可以为温度和光照升高提供抗性和保护。这些机制包括可诱导的热休克蛋白,其作用是重新折叠变性的细胞和结构蛋白;产生氧化酶,使有害的氧自由基失活;荧光珊瑚色素,既能反射又能消散光能;虫黄藻的表型适应以及它们在较高温度下种群的适应性变化。当这些机制与经历过珊瑚白化地区珊瑚恢复的实验和观察证据结合起来考虑时,表明珊瑚和虫黄藻具有尚未明确的适应导致珊瑚白化条件的能力。显然,能够对抗海水温度升高导致的珊瑚白化的适应过程是有限的,但在我们更好地了解珊瑚 - 藻类的适应/适应潜力之前,科学模型无法准确预测珊瑚礁珊瑚的命运。尤其需要开展关于促进珊瑚和虫黄藻耐热性的分子和生理机制以及确定导致珊瑚白化事件中各种反应的遗传特征的研究。只有到那时,我们才能对可能的反应性质、所涉及的时间尺度以及“经验”在减轻白化影响方面的作用有一些了解。