珊瑚礁白化研究进展

珊瑚礁白化是由于珊瑚失去体内共生的虫黄藻和（或）共生的虫黄藻失去体内色素而导致五彩缤纷的珊瑚礁变白的生态现象。近年来，频繁发生的珊瑚礁白化导致了珊瑚礁生态系统严重退化，并已经影响到全球珊瑚礁生态系统的平衡，受到了人们的高度重视。研究认为：（1）大范围珊瑚礁白化主要是全球环境变化引起的，尤其是全球变暖和紫外辐射增强；（2）导致珊瑚礁白化的机制主要在于细胞机制和光抑制机制；（3）珊瑚礁白化后的恢复与白化程度有关，大范围白化的珊瑚礁完全恢复需要几年到几十年；（4）珊瑚礁白化的后果在于降低珊瑚繁殖能力、减缓珊瑚礁生长、改变礁栖生物的群落结构，导致大面积珊瑚死亡和改变珊瑚礁生态类型，如变为海藻型等；（5）与珊瑚共生的D系群虫黄藻更适应高温环境，珊瑚礁有可能通过D系群逐渐取代C系群的方式适应全球环境变化。     

珊瑚礁生态系统病毒研究进展

病毒对珊瑚礁生态系统中的生物进化、生物地球化学循环、珊瑚疾病等方面具有重要的生态影响。随着珊瑚礁的全球性退化，病毒在珊瑚礁生态系统中的功能与危害日益显现。综述了珊瑚礁生态系统中病毒的研究现状与进展，包括：（1）珊瑚礁病毒的多样性与分布特征（水体、宿主、核心病毒组）；（2）珊瑚礁病毒的生态功能（感染方式、促进生物进化、生物地球化学循环）；（3）珊瑚礁病毒对全球气候变化的响应（热压力、珊瑚疾病）。总体而言，珊瑚礁生态系统具有极高的病毒多样性，所发现的60个科占已知所有病毒科数量的58%。珊瑚的核心病毒组主要由双链DNA病毒、单链DNA病毒、单链逆转录病毒所组成，珊瑚黏液层对病毒具有富集作用。"Piggyback-the-Winner"（依附-胜利）是病毒在珊瑚礁中主要的生物动力学模式，其可通过水平基因迁移的方式促进礁区生物进化。病毒可通过裂解细菌与浮游藻类的途径参与珊瑚礁的生物地球化学循环，尤其是碳循环与氮循环过程。此外，病毒还具有介导珊瑚热白化与直接引发珊瑚疾病的能力，这会影响珊瑚礁生态系统应对气候变化的适应性与恢复力。基于国际上的研究进展综述，结合南海珊瑚礁生态现状提出以下研究方向，以期促进我国珊瑚礁病毒学的发展：（1）开展南海珊瑚礁中病毒多样性的识别及其时-空分布特征研究；（2）探索病毒对南海珊瑚热白化、珊瑚疾病的介导作用及其与气候变化的关系；（3）揭示病毒对南海珊瑚礁生物地球化学循环的贡献。     

造礁石珊瑚与其共生藻(Symbiodinium)共生研究进展

对造礁石珊瑚与其共生藻共生研究现状及其在全球变化下的适应能力进行较全面的综述.造礁石珊瑚与遗传和生理功能独特的共生藻组成内共生关系是成功演化的范例.近年来对珊瑚共生体的分子系统学研究表明共生藻遗传多样性极为丰富,当前认为共生藻属至少包括8个(A-H)各自包含亚系群的世系或系群.珊瑚-共生藻共生功能体对诸如全球变化引起的海水温度上升等环境变化十分敏感.由于珊瑚以及珊瑚礁面临气候变化的严峻挑战,对珊瑚与其共生藻共生关系和共生功体适应能力的研究将是未来重要的研究领域之一.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

卫星遥感珊瑚礁白化概述

珊瑚礁白化是由于珊瑚失去体内共生的虫黄藻或者共生的虫黄藻失去体内色素而导致五彩缤纷的珊瑚礁变白的现象,严重的白化可以带来珊瑚礁的死亡.国内外研究表明海水温度升高和珊瑚礁白化关系最为紧密.卫星遥感能够提供大范围、同步与连续的海洋数据,如海水表层温度和海色数据,从而能够及时监测和预测珊瑚礁的白化.基于AVHRR (Advanced Very High Resolution Radiometer),NOAA(National Oceanic and Atmospheric Administration,US)开发了全球监测珊瑚礁白化的方法,热点(HotSpot)和周热度(DHW)两种主要指数.目前,我国珊瑚礁白化现象的监测和研究明显滞后于国际动态,迫切需要发展和利用卫星遥感的方法监测南海珊瑚礁白化状况.     

运用PCR-RFLP方法研究三亚鹿回头岸礁造礁石珊瑚共生藻的组成

造礁石珊瑚共生藻的系统分类研究对于理解珊瑚礁生态系统对全球变化的响应具有十分重要的意义.本研究利用PCR技术扩增核糖体基因人亚基片段以及限制性片段长度多态性(RFLP)的方法,对海南三亚鹿回头岸礁的8科14属22种造礁石珊瑚的共生藻组成进行了研究.结果表明鹿回头岸礁造礁石珊瑚共生藻以C系群为优势系群,偶尔发现D系群与鹿角杯形珊瑚(Pocilolpora damieornis)和黄癣蜂巢珊瑚(Favia favus)共生:另外丑鹿角珊瑚(Acropora horrida)和丛生盔形珊瑚(Galaxea fascicularis)可以同时与C系群和D系群共生.共生多型性的发现暗示珊瑚与共生藻的共生关系具有灵活性.研究结果同样显示共生藻的核糖体基因人片段的DNA多态性偏低.未来应该结合其他的分子标记对鹿回头岸礁造礁石珊瑚共生藻的DNA多态性进行更深入的研究.     

珊瑚白化及温度相关基因的研究进展

近年来,由于海洋酸化、海洋污染、海洋表面温度升高以及人类活动的影响,珊瑚礁遭受到了严重的白化袭击,导致珊瑚礁处于退化状态。相关功能基因在珊瑚白化过程中可能起到增强珊瑚的抗逆性和恢复能力等作用,探讨这些基因在珊瑚白化过程中的表达变化及调控作用也已经成为了当今的研究热点,通过分析其生理调控作用,可以深入了解珊瑚白化机制与恢复机制。本综述总结了珊瑚白化与温度的关系和珊瑚温度相关基因的研究进展,并重点介绍了细胞响应机制中的细胞免疫相关基因、细胞生长调控基因、钙离子相关基因、核酸稳定与修复相关基因以及其它重要基因的研究。     

微生物在珊瑚礁生态系统中的作用与功能

珊瑚礁是由珊瑚、鱼类、底栖生物、藻类以及微生物等多种生命形式组成的聚集体,代表着一类典型的海洋生态系统.珊瑚礁存在于热带和亚热带的寡营养环境,拥有极高的初级生产力和生产效率,被誉为“海底热带雨林”.微生物在珊瑚礁生态系统的生物地球化学循环、物质转化以及健康维护上具有重要作用.随着分子生态学的发展,微生物在珊瑚中的作用和功能日益凸显.本文总结了微生物生态学的研究现状,包括珊瑚生态系统中微生物的定植方式,共生微生物的特性(专一性、可塑性、协同进化),共生微生物与珊瑚疾病的关系与信号调节,以及微生物应对全球变化(气温升高、海水酸化、富营养化)的响应.从“珊瑚 微生物”共生体的发生、共生微生物的特性与生态功能,以及全球环境变化下微生物的衍生效应来梳理最新理论与成果,明确珊瑚微生物生态学机制,为更好地保护珊瑚资源、维护海洋生物多样性提供理论借鉴.     

不同生长状态珊瑚光谱特征

珊瑚礁生态系统迅速退化是目前重要的生态环境问题之一,应用遥感技术监测大范围珊瑚礁的结构组成和变迁有很大的潜力。珊瑚光谱响应特征受珊瑚生态习性影响,在光学上相似而容易造成混淆误判。采集了西沙群岛大量石珊瑚样品的光谱,对其光谱特征进行分析及成因探讨。通过导数光谱、主成分分析研究了不同生长状态珊瑚的光谱差异,并建立珊瑚生长状态高光谱遥感判别准则。结果表明,珊瑚的光谱特性及其变化均较为复杂,受珊瑚种类和生长环境影响,光谱形状主要由共生藻色素吸收决定的。结合520—530 nm、564—574 nm和600—605 nm的导数光谱可以区分健康珊瑚、白化珊瑚和藻类覆盖的死珊瑚。总体判定准确度优于80%,误判的主要来源是种内珊瑚反射率差异。研究表明珊瑚礁环境高光谱遥感可以定量评估珊瑚状态的变化。     

海南三亚后海海域珊瑚礁生态系统的健康状况及其影响因素

分别采用鹞式调查法和断面监测法调查了海南三亚后海海域珊瑚的物种多样性、覆盖率、病害和补充量等指标,利用健康指数(CI)评估了后海珊瑚礁生态系统的健康状况,并初步分析了影响该区域珊瑚礁健康状况的主要因素。调查发现:后海海域造礁石珊瑚54种,覆盖率达到50%以上,珊瑚病害及死亡率低,珊瑚补充量高达4.5个·m-2,CI值介于1.87~2.27,表明后海海域的珊瑚礁生态系统非常健康。分析认为:后海海域浅水区域存在的海草床和大型藻类,以及珊瑚礁区高密度的植食性动物和夏季上升流的存在是该区域珊瑚礁生态系统健康的主要原因。海草和海藻将陆源污染物过滤吸收,确保进入珊瑚礁生态系统的水质良好;数量众多的植食性动物(如魔鬼海胆Diadema setosum等)调控了大型藻类和珊瑚之间的竞争关系,保证大型海藻不会威胁到珊瑚的健康生长;而后海海域夏季上升流的出现使得该海域珊瑚礁生态系统不会受到高温的影响,不会产生热白化现象。这样,多个因素的共同作用保证了后海珊瑚礁生态系统的健康。     

Local Stressors Reduce Coral Resilience to Bleaching

Coral bleaching, during which corals lose their symbiotic dinoflagellates, typically corresponds with periods of intense heat stress, and appears to be increasing in frequency and geographic extent as the climate warms. A fundamental question in coral reef ecology is whether chronic local stress reduces coral resistance and resilience from episodic stress such as bleaching, or alternatively promotes acclimatization, potentially increasing resistance and resilience. Here we show that following a major bleaching event, Montastraea faveolata coral growth rates at sites with higher local anthropogenic stressors remained suppressed for at least 8 years, while coral growth rates at sites with lower stress recovered in 2–3 years. Instead of promoting acclimatization, our data indicate that background stress reduces coral fitness and resilience to episodic events. We also suggest that reducing chronic stress through local coral reef management efforts may increase coral resilience to global climate change.     

Heterotrophic Compensation: A Possible Mechanism for Resilience of Coral Reefs to Global Warming or a Sign of Prolonged Stress?

Thermally induced bleaching has caused a global decline in corals and the frequency of such bleaching events will increase. Thermal bleaching severely disrupts the trophic behaviour of the coral holobiont, reducing the photosynthetically derived energy available to the coral host. In the short term this reduction in energy transfer from endosymbiotic algae results in an energy deficit for the coral host. If the bleaching event is short-lived then the coral may survive this energy deficit by depleting its lipid reserves, or by increasing heterotrophic energy acquisition. We show for the first time that the coral animal is capable of increasing the amount of heterotrophic carbon incorporated into its tissues for almost a year following bleaching. This prolonged heterotrophic compensation could be a sign of resilience or prolonged stress. If the heterotrophic compensation is in fact an acclimatization response, then this physiological response could act as a buffer from future bleaching by providing sufficient heterotrophic energy to compensate for photoautotrophic energy losses during bleaching, and potentially minimizing the effect of subsequent elevated temperature stresses. However, if the elevated incorporation of zooplankton is a sign that the effects of bleaching continue to be stressful on the holobiont, even after 11 months of recovery, then this physiological response would indicate that complete coral recovery requires more than 11 months to achieve. If coral bleaching becomes an annual global phenomenon by mid-century, then present temporal refugia will not be sufficient to allow coral colonies to recover between bleaching events and coral reefs will become increasingly less resilient to future climate change. If, however, increasing their sequestration of zooplankton-derived nutrition into their tissues over prolonged periods of time is a compensating mechanism, the impacts of annual bleaching may be reduced. Thus, some coral species may be better equipped to face repeated bleaching stress than previously thought.     

Resistance to thermal stress in corals without changes in symbiont composition

Discovering how corals can adjust their thermal sensitivity in the context of global climate change is important in understanding the long-term persistence of coral reefs. In this study, we showed that short-term preconditioning to higher temperatures, 3°C below the experimentally determined bleaching threshold, for a period of 10 days provides thermal tolerance for the symbiosis stability between the scleractinian coral, Acropora millepora and Symbiodinium. Based on genotypic analysis, our results indicate that the acclimatization of this coral species to thermal stress does not come down to simple changes in Symbiodinium and/or the bacterial communities that associate with reef-building corals. This suggests that the physiological plasticity of the host and/or symbiotic components appears to play an important role in responding to ocean warming. The further study of host and symbiont physiology, both of Symbiodinium and prokaryotes, is of paramount importance in the context of global climate change, as mechanisms for rapid holobiont acclimatization will become increasingly important to the long-standing persistence of coral reefs.     

Identity and diversity of coral endosymbionts (zooxanthellae) from three Palauan reefs with contrasting bleaching, temperature and shading histories

The potential of corals to associate with more temperature-tolerant strains of algae (zooxanthellae, Symbiodinium) can have important implications for the future of coral reefs in an era of global climate change. In this study, the genetic identity and diversity of zooxanthellae was investigated at three reefs with contrasting histories of bleaching mortality, water temperature and shading, in the Republic of Palau (Micronesia). Single-stranded conformation polymorphism and sequence analysis of the ribosomal DNA internal transcribed spacer (ITS)1 region was used for genotyping. A chronically warm but partly shaded coral reef in a marine lake that is hydrographically well connected to the surrounding waters harboured only two single-stranded conformation polymorphism profiles (i.e. zooxanthella communities). It consisted only of Symbiodinium D in all 13 nonporitid species and two Porites species investigated, with the remaining five Porites harbouring C*. Despite the high temperature in this lake (> 0.5 degrees above ambient), this reef did not suffer coral mortality during the (1998) bleaching event, however, no bleaching-sensitive coral families and genera occur in the coral community. This setting contrasts strongly with two other reefs with generally lower temperatures, in which 10 and 12 zooxanthella communities with moderate to low proportions of clade D zooxanthellae were found. The data indicate that whole coral assemblages, when growing in elevated seawater temperatures and at reduced irradiance, can be composed of colonies associated with the more thermo-tolerant clade D zooxanthellae. Future increases in seawater temperature might, therefore, result in an increasing prevalence of Symbiodinium phylotype D in scleractinian corals, possibly associated with a loss of diversity in both zooxanthellae and corals.     

How much time can herbivore protection buy for coral reefs under realistic regimes of hurricanes and coral bleaching?

Coral reefs have been more severely impacted by recent climate instability than any other ecosystem on Earth. Corals tolerate a narrow range of physical environmental stress, and increases in sea temperature of just 1 °C over several weeks can result in mass coral mortality, often exceeding 95% of individuals over hundreds of square kilometres. Even conservative climate models predict that mass coral bleaching events could occur annually by 2050. Unfortunately, managers of coral‐reef resources have few options available to meet this challenge. Here, we investigate the role that fisheries conservation tools, including the designation of marine reserves, can play in altering future trajectories of Caribbean coral reefs. We use an individual‐based model of the ecological dynamics to test the influence of spatially realistic regimes of disturbance on coral populations. Two major sources of disturbance, hurricanes and coral bleaching, are simulated in contrasting regions of the Caribbean: Belize, Bonaire, and the Bahamas. Simulations are extended to 2099 using the HadGEM1 climate model. We find that coral populations can maintain themselves under all levels of hurricane disturbance providing that grazing levels are high. Regional differences in hurricane frequency are found to cause strikingly different spatial patterns of reef health with greater patchiness occurring in Belize, which has less frequent disturbance, than the Bahamas. The addition of coral bleaching led to a much more homogenous reef state over the seascape. Moreover, in the presence of bleaching, all reefs exhibited a decline in health over time, though with substantial variation among regions. Although the protection of herbivores does not prevent reef degradation it does delay rates of coral loss even under the most severe thermal and hurricane regimes. Thus, we can estimate the degree to which local conservation can help buy time for reefs with values ranging between 18 years in the Bahamas and over 50 years in Bonaire, compared with heavily fished systems. Ultimately, we demonstrate that local conservation measures can benefit reef ecosystem services but that their impact will vary spatially and temporally. Recognizing where such management interventions will either help or fail is an important step towards both achieving sustainable use of coral‐reef resources and maximizing resource management investments.     

Symbiophagy as a cellular mechanism for coral bleaching

Coral bleaching is a major contributor to the global declines of coral reefs. This phenomenon is characterized by the loss of symbiotic algae, their pigments or both. Despite wide scientific interest, the mechanisms by which bleaching occurs is still poorly understood. Here we report that the removal of the symbiont during light and temperature stress is achieved using the host's cellular autophagic-associated machinery. Host cellular and sub-cellular morphologies showed increased vacuolization and appearance of autophagic membranes surrounding a variety of organelles and surrounding the symbiotic algae. Markers of autophagy (Rab 7 and LAS) corroborate these observations. Results showed that during stress the symbiont vacuolar membrane is transformed from a conduit of nutrient exchange to a digestive organelle resulting in the consumption of the symbiont, a process we term symbiophagy. We posit that during a stress event, the mechanism maintaining symbiosis is destabilized and symbiophagy is activated, ultimately resulting in the phenomenon of bleaching. Symbiophagy may have evolved from a more general primordial innate intracellular protective pathway termed xenophagy.     

Detriments to post-bleaching recovery of corals

Predicting the response of coral reefs to large-scale mortality induced by climate change will depend greatly on the factors that influence recovery after bleaching events. We experimentally transplanted hard corals from a shallow reef with highly variable seawater temperature (23–36°C) to three unfished marine parks and three fished reefs with variable coral predator abundance and benthic cover. The transplanted corals were fragmented colonies collected from a reef that was relatively undisturbed by the 1997–1998 warm-water temperature anomaly, one of the most extreme thermal events of the past century, and it was assumed that they would represent corals likely to succeed in the future temperature environment. We examined the effects of four taxa, two fragment sizes, an acclimation period, benthic cover components, predators and tourists on the survival of the coral fragments. We found the lowest survival of transplants occurred in the unfished marine parks and this could be attributed to predation and not tourist damage. The density of small coral recruits approximately 6 months after the spawning season was generally moderate (~40–60/m²), and not different on fished and unfished reefs. Coral recovery between 1998 and 2002 was variable (0–25%), low (mean of 6.5%), and not different between fished and unfished reefs. There was high variability in coral mortality among the three unfished areas despite low variation in estimates of predator biomass, with the highest predation occurring in the Malindi MNP, a site with high coralline algal cover. Stepwise multiple regression analysis with 14 variables of coral predators and substratum showed that coralline algae was positively, and turf algae negatively associated with mortality of the transplants, with all other variables being statistically insignificant. This suggests that alternate food resources and predator choices are more important than predator biomass in determining coral survival. Nonetheless, large predatory fish in areas dominated by coralline algae may considerably retard recovery of eurythermal corals. This will not necessarily retard total hard coral recovery, as other more predator-tolerant taxa can recover. Based on the results, global climate change will not necessarily favor eurythermal over stenothermal coral taxa in remote or unfished reefs, where predation is a major cause of coral mortality.     

Coral reef bleaching: ecological perspectives

Coral reef bleaching, the whitening of diverse invertebrate taxa, results from the loss of symbiotic zooxanthellae and/or a reduction in photosynthetic pigment concentrations in zooxanthellae residing within the gastrodermal tissues of host animals. Of particular concern are the consequences of bleaching of large numbers of reef-building scleractinian corals and hydrocorals. Published records of coral reef bleaching events from 1870 to the present suggest that the frequency (60 major events from 1979 to 1990), scale (co-occurrence in many coral reef regions and often over the bathymetric depth range of corals) and severity (>95% mortality in some areas) of recent bleaching disturbances are unprecedented in the scientific literature. The causes of small scale, isolated bleaching events can often be explained by particular stressors (e.g., temperature, salinity, light, sedimentation, aerial exposure and pollutants), but attempts to explain large scale bleaching events in terms of possible global change (e.g., greenhouse warming, increased UV radiation flux, deteriorating ecosystem health, or some combination of the above) have not been convincing. Attempts to relate the severity and extent of large scale coral reef bleaching events to particular causes have been hampered by a lack of (a) standardized methods to assess bleaching and (b) continuous, long-term data bases of environmental conditions over the periods of interest. An effort must be made to understand the impact of bleaching on the remainder of the reef community and the long-term effects on competition, predation, symbioses, bioerosion and substrate condition, all factors that can influence coral recruitment and reef recovery. If projected rates of sea warming are realized by mid to late AD 2000, i.e. a 2°C increase in high latitude coral seas, the upper thermal tolerance limits of many reef-building corals could be exceeded. Present evidence suggests that many corals would be unable to adapt physiologically or genetically to such marked and rapid temperature increases.