微生物参与海草床元素循环及其生态修复功能

海草是分布在全球海岸带的沉水被子植物,与周围环境共同形成的海草床生态系统是三大典型海洋生态系统之一,具有十分重要的生态功能。20世纪以来,全球海草床衰退严重,研究海草床的生态修复迫在眉睫,现有修复方法未能足够重视微生物在海草床中的重要作用。本文综合阐述了微生物在海草床生态系统有机物矿化和营养流动过程中起到的作用,分析了微生物驱动下的海草床水体与沉积物之间的元素循环,提出了人类活动引起海草床退化的原因,总结了海草床微生物的系统研究方法,并在此基础上提出从微生物生态的角度修复海草床的新思路。     

海草生态学研究进展

海草床生态系统是生物圈中最具生产力的水生生态系统之一,具有重要的生态系统服务功能。作者根据海草生态学及相关领域的最新研究进展,对世界范围内海草床的空间分布、海草床的生态系统服务功能以及外界因素对海草床的影响等研究进展进行了综述。海草床生态系统服务功能主要包括净化水质、护堤减灾、提供栖息地和生态系统营养循环等。对海草床影响较大的外界环境因素包括盐度、温度、营养盐、光照、其他动物摄食、人类活动和气候变化等。海草普查、海草生态功能研究,影响海草床的主要环境因素,海草修复研究等将是我国海草研究的主要方向。     

海草生态系统的固碳机理及贡献

由海草、红树林、盐沼草等植被组成的滨海和海洋生态系统是地球中高效的碳汇热点,它们所固定的碳被称为"蓝碳".作为全球生态服务功能价值最高的生态系统之一,海草生态系统所固定的碳是蓝碳里的重要组成部分.高生产力、高效过滤及高稳定性造就了海草生态系统巨大的固碳能力,进而对全球碳循环具有深刻影响.然而,人为影响以及全球气候变化使全球海草床加速衰退,成为地球生物圈中退化速度最快的生态系统之一.当前,国内外对海草床等滨海生态系统固碳能力的关注、研究深度与广度仍远远不足,对全球海草固碳的评估仍存在诸多不确定性.为了能更准确地评估全球海草床的碳埋存,一些基础性的科学问题应优先考虑:1)全国和全球海草的准确分布面积;2)不同海草优势种类或不同地域的海草床碳汇能力的差异;3)人为干扰和全球气候变化对海草生态系统碳捕获和碳埋存的影响.     

海草床生态系统的退化及其恢复

海草床是生产力和生物多样性最高的典型海洋生态系统之一,目前全球海草床呈现退化趋势,自然干扰和人类活动的负面影响是其退化原因,以后者为主.海草退化的生理生态机制主要是光合作用速率、光合色素荧光强度和含量、酶活性等生理生态指标在胁迫(高温、光照、盐度、污染物等)下发生显著变化.海草床监测和保护计划已经在局部区域取得成效,使部分海草床得到恢复.中国的海草床恢复研究重点应放在生物多样性保护、海草移植和海草床生态系统服务价值评估.     

海草床育幼功能及其机理

海草床是近岸海域中生产力极高的生态系统,是许多海洋水生动物的重要育幼场所。从生物幼体的密度、生长率、存活率和生境迁移4个方面阐述海草床育幼功能,并从食源和捕食压力两个方面探讨海草床育幼功能机理。许多生物幼体在海草床都呈现出较高的密度、生长率和存活率,并且在个体发育到一定阶段从海草床向成体栖息环境迁移。丰富的食物来源或较低的捕食压力可能是海草床具有育幼功能的主要原因,但不同的生物幼体对海草床的利用有差异,海草床育幼功能的机理在不同环境条件下也存在差异。提出未来海草床育幼功能的重点研究方向:(1)量化海草床对成体栖息环境贡献量;(2)全球气候变化和人类活动对海草床育幼功能的影响;(3)海草床育幼功能对海草床斑块效应和边缘效应的响应,以期为促进我国海草床育幼研究和海草床生态系统保护提供依据。     

营养盐富集和全球温度升高对海草的影响

海草床具有稳定沉积物、净化水质和碳储存等重要的生态系统服务功能。近几十年来,世界范围内海草床衰退严重。在全球近岸水体富营养化和气温升高的背景下,本文对营养盐富集和全球温度升高对海草的影响进行了分析。硝态氮和铵态氮作为海草重要的营养来源,贫营养环境下,会促进海草的生长。由于硝态氮需要转化为亚硝态氮,再通过一系列新陈代谢过程转化为氨基酸,硝态氮富集会影响海草组织的碳平衡,从而对海草床造成负面影响。较高浓度的铵态氮会对海草产生毒性,引起海草床的衰退。有机氮作为海草可选择吸收的氮源,是对海草氮吸收的有效补充。温度是控制全球海草分布和生长的主要因素,春季一定程度的营养盐富集可以提高海草的生产力,而夏季高温和营养盐富集对海草的生长具有抑制作用。营养盐富集和夏季高温还可以通过促进大型海藻爆发性生长,导致光衰减,从而引起海草床向大型海藻生态系统的逆向演替。本文提出了未来海草主要研究方向,主要包括:长时间序列海草床野外观测;有机氮对海草的影响机制;营养盐富集和全球温度升高对海草的协同影响机制;热带海草呼吸作用对全球温度升高的响应。     

非共生生物固氮微生物分子生态学研究进展

氮是限制生态系统生产力的主要元素,生物固氮是自然生态系统中氮的主要来源.生物固氮包括共生、联合和自生固氮3种类型,其中联合固氮和自生固氮统称为非共生固氮.相对于共生固氮而言,非共生固氮速率虽然较低,但其不需要与其他生物形成共生体系就可以生存并进行固氮,在时空分布上更加广泛,因此对生态系统氮循环特别是素输入具有重要贡献.本文对近年有关非共生固氮微生物的多样性、土壤和叶际固氮微生物的分布特征及影响因素等研究进展进行了综述,并在此基础上阐述了现有研究中存在的问题和发展前景.     

海草与其附生藻类之间的相互作用

海草床是滨海湿地中重要的生态系统,近年来世界范围内出现大面积的海草床衰退,水体富营养化引发附生藻类的大量繁殖是其中的一个主要原因。本文综述了近20年来关于海草与其附生藻类之间相互作用的研究进展,包括海草床中附生藻类的地位及其积极作用,附生藻类与海草的物质交换关系,附生藻类对海草的影响（主要是光利用限制和营养物质利用限制）。对海草床中海草-附生藻类群落复杂的生态关系提出了研究展望,包括海草与其附生藻类的物质交换机制,海草与其附生藻类对营养物质利用的竞争关系,附生藻类生物量分别对海草可利用光和光合作用能力的削弱关系,海草与附生藻类对海草床初级生产力的贡献随环境因子的变化关系,通过原位观测数据的积累建立海草生长与生物作用的模型等。     

固氮类植物在陆地生态系统中的作用研究进展

固氮类植物是陆地生态系统中的一个重要功能群,它们广泛存在于陆地生态系统中,是陆地生态系统最重要的氮源,大量研究表明,固氮类植物在陆地生态系统中的生态功能表现出明显的复杂性和多样性,对于生态过程有很强的主导性和控制性.通过与固氮微生物形成不同的共生固氮类型,它们可以改善土壤结构和营养组成以及地表微生境,直接影响碳氮磷循环、种子萌发和植株生长、植被演替、凋落物构成及其分解4个关键生态过程,进而提高群落结构复杂性、生物多样性和初级生产力,促进植被恢复和减少水土流失.未来研究中,应该加强对陆地生态系统固氮过程的定量化分析,确定影响共生固氮的主导因子和生态机制从而为陆地系统的生态恢复提供依据.     

一株海草沉积物菲降解菌的筛选、鉴定和降解特性

【背景】多环芳烃(Polycyclic Aromatic Hydrocarbons,PAHs)是一类高毒性的有机污染物,在海洋环境尤其是沿海环境中广泛分布。海草床生态系统作为沿海环境的重要组成部分,深受环境污染等人类活动的影响而处于严重衰退的状态。微生物修复是修复环境中多环芳烃污染的重要途径,具有经济简便、环境友好和无二次污染等特点。【目的】从深圳市大亚湾的海草床沉积物中筛选获得高效多环芳烃降解菌,并分析其降解特性,从而探究海草床生态系统中多环芳烃污染物的微生物修复可行性。【方法】以多环芳烃菲为唯一碳源从海草床沉积物样品中筛选菌株,再通过形态学观察、生理生化实验和16SrRNA基因序列对筛选的菌株进行鉴定,并利用特定引物扩增多环芳烃降解的功能基因——双加氧酶(nidA)基因,最后通过培养实验分析该菌株对菲的降解特性。【结果】筛选出一株高效降解菲的菌株SCSIO 43702,经鉴定为玫瑰杆菌属(Roseovarius)的潜在新菌,并成功扩增得到双加氧酶相似(nidA like)基因;培养实验结果表明,玫瑰杆菌SCSIO 43702在10 d内对100 mg/L菲的降解率最高可达96%,而且其对菲的最适降解条件为:温度30°C、pH值7.5和8.0、盐度3%。【结论】玫瑰杆菌SCSIO 43702凭借其良好的菲降解能力和较强的环境适应性,具有进一步被开发为微生物菌剂以用于多环芳烃污染修复的巨大潜力,为海草床生态系统中多环芳烃污染的微生物修复研究提供了理论依据和可利用的微生物资源。     

Nitrogen fixation in sediments and the rhizosphere of the seagrassZostera capricorni

Rates of nitrogen fixation in seagrass beds (Zostera capricorni) were determined with¹⁵N and reduction of acetylene in intact cores of sediment and seagrass. There was good agreement in the results from the two techniques, with a molar ratio of 3∶1.9 ethylene: ammonia produced. Fixed nitrogen was rapidly utilized by the plants, with significant amounts of¹⁵N found in the roots and rhizomes and 50% of fixed¹⁵N apparently translocated to the leaves. Rates of fixation were high in summer (25 to 40 mg N m⁻² day⁻¹) and lower in winter (10 mg N m⁻² day⁻¹) and were estimated to supply between one-third and one-half of the nitrogen requirements of the seagrass. Rates of nitrogen fixation were greater in the light than in the dark, and in cores of intact seagrass than in defoliated cores, indicating that the bacteria were dependent on organic compounds secreted by the plants.     

Tiny Is Mighty: Seagrass Beds Have a Large Role in the Export of Organic Material in the Tropical Coastal Zone

Ecosystems in the tropical coastal zone exchange particulate organic matter (POM) with adjacent systems, but differences in this function among ecosystems remain poorly quantified. Seagrass beds are often a relatively small section of this coastal zone, but have a potentially much larger ecological influence than suggested by their surface area. Using stable isotopes as tracers of oceanic, terrestrial, mangrove and seagrass sources, we investigated the origin of particulate organic matter in nine mangrove bays around the island of Phuket (Thailand). We used a linear mixing model based on bulk organic carbon, total nitrogen and δ¹³C and δ¹⁵N and found that oceanic sources dominated suspended particulate organic matter samples along the mangrove-seagrass-ocean gradient. Sediment trap samples showed contributions from four sources oceanic, mangrove forest/terrestrial and seagrass beds where oceanic had the strongest contribution and seagrass beds the smallest. Based on ecosystem area, however, the contribution of suspended particulate organic matter derived from seagrass beds was disproportionally high, relative to the entire area occupied by mangrove forests, the catchment area (terrestrial) and seagrass beds. The contribution from mangrove forests was approximately equal to their surface area, whereas terrestrial contributions to suspended organic matter under contributed compared to their relative catchment area. Interestingly, mangrove forest contribution at 0 m on the transects showed a positive relationship with the exposed frontal width of the mangrove, indicating that mangrove forest exposure to hydrodynamic energy may be a controlling factor in mangrove outwelling. However we found no relationship between seagrass bed contribution and any physical factors, which we measured. Our results indicate that although seagrass beds occupy a relatively small area of the coastal zone, their role in the export of organic matter is disproportional and should be considered in coastal management especially with respect to their importance as a nutrient source for other ecosystems and organisms.     

Decomposition and nitrogen dynamics of turtle grass (<Emphasis Type="Italic">Thalassia testudinum</Emphasis>) in a subtropical estuarine system

Seagrass beds are pivotal in the functioning of coastal ecosystems in terms of productivity, organic matter turnover and nutrient cycling. Aiming to document decay and nitrogen (N) dynamics of turtle grass (Thalassia testudinum) in a subtropical estuarine system, decomposition patterns of leaves and rhizomes were characterized and compared. Nitrogen usage during decomposition of tissues, and of live tissues and epiphytes growing on live leaves, was also quantified and compared. Stable isotope ratios allowed tracing N within the seagrass bed, following N incorporation into seagrass tissues from the surrounding media (water, sediment). Leaves had a higher N content and decomposed at a faster rate (~6.4 times) compared to rhizomes. Leaching of soluble materials explain the rapid (0–3 days) initial mass loss of leaves (20%) and rhizomes (18%); with a loss of 85 and 56%, respectively, by the end of the study (77 days). Overall, leaves released N while rhizomes immobilized it. Nitrogen concentration was significantly different among live tissues. The main source of N for both seagrass tissues was the sediment, and water column for epiphytes. Differences in decomposition rates among seagrass tissues can be explained by the quality of the substrate and its susceptibility to microbial use. Seagrass leaves and rhizomes are equally important in taking up nutrients from either the water column or the sediments. This study provides a platform to study energy and matter transfers through detrital foodwebs linked to seagrass meadows.     

Introduced species in seagrass ecosystems: Status and concerns

A literature review revealed that at least 56 non-native species, primarily invertebrates and seaweeds, have been introduced to seagrass beds, largely through shipping/boating activities and aquaculture. Four seagrass species also have been introduced. The introductions of the seaweeds Caulerpa taxifolia, C. racemosa v. cylindracea, Codium fragile ssp. tomentosoides, Sargassum muticum, the Asian mussel, Musculista senhousia, and the seagrass, Zostera japonica, are the best-known examples in seagrass beds. The ecological effects on seagrasses and associated communities have been examined for slightly less than half of the introduced species, which have predominantly negative effects. There is a paucity of experimental data for ecological effects, particularly for seagrass community structure and function. The exception to this finding is the introduction of the seagrass Z. japonica with oyster aquaculture to native eelgrass beds on the Pacific coasts of Canada and the USA. Recent experiments in several different seagrass ecosystems confirmed that disturbance contributes to the invasibility of seagrass beds. More definitive studies are required to elucidate the relative effects of nutrient pollution and introduced species in causing seagrass decline, particularly where reduced herbivory and boating activity also covary. Seagrass beds often are subject to multiple introduced species, but their cumulative effect has been virtually unstudied. The potential for compounded negative effects merits serious attention. Heightened attention to the issue of introduced species in seagrass beds is called for given the evidence that introduced species can contribute to seagrass decline, to biodiversity changes that could affect seagrass ecosystem functions, and that they can compromise seagrass restoration. Comprehensive surveys in seagrass beds, complemented by more stringent experimental and mensurative sampling designs, are needed. In the interim, conserving seagrass density and bed size can offer resistance to introduced species. Managing to prevent the introductions, including restricting transplantations of non-native biota during seagrass restorations, is likely to bear positive benefits for seagrass ecosystems.     

Material exchange and food web of seagrass beds in the Sylt-Rømø Bight: how significant are community changes at the ecosystem level?

Material exchange, biodiversity and trophic transfer within the food web were investigated in two different types of intertidal seagrass beds: a sheltered, dense Zostera marina bed and a more exposed, sparse Z. noltii bed, in the Northern Wadden Sea. Both types of Zostera beds show a seasonal development of above-ground biomass, and therefore measurements were carried out during the vegetation period in summer. The exchange of particles and nutrients between seagrass beds and the overlying water was measured directly using an in situ flume. Particle sedimentation [carbon (C), nitrogen (N) and phosphorus (P) constituents] from the water column prevailed in dense seagrass beds. In the sheltered, dense seagrass bed, a net particle uptake was found even on windy days (7–8 Beaufort). Dissolved inorganic N and orthophosphate were mainly taken up by the dense seagrass bed. At times of strong winds, nutrients were released from the benthic community to tidal waters. In a budget calculation of total N and total P, the dense seagrass beds were characterised as a material sink. The seagrass beds with sparse Z. noltii were a source of particles even during calm weather. The uptake of dissolved inorganic N in the sparse seagrass bed was low but significant, while the uptake of inorganic phosphate and silicate by seagrasses and their epiphytes was exceeded by release processes from the sediment into the overlying water. Estimates at the ecosystem level showed that material fluxes of seagrass beds in the Sylt-Rømø Bight are dominated by the dense type of Zostera beds. Therefore, seagrass beds act as a sink for particles and for dissolved inorganic nutrients. During storms, seagrass beds are distinct sources for inorganic nutrients. The total intertidal area of the Sylt-Rømø Bight could be described as a sink for particles and a source for dissolved nutrients. This balance of the material budget was estimated by either including or excluding seagrass beds. Including the subtidal part, the function of the ecosystem as a source for particles increased, supposing that all seagrass beds were lost from the area. During the vegetation period, seagrass beds act as a storage compartment for material accumulated in the living biomass of the community. There was great biodiversity among the plant and animal groups found in intertidal seagrass beds of the Sylt-Rømø Bay, representing 50–86% of the total number of species investigated, depending on the particular group. Since most species are not exclusively seagrass residents, the loss of intertidal seagrass beds would be of minor importance for biodiversity at the ecosystem level. Food web structure in seagrass beds is different from other intertidal communities. Primary production and detritus input is high, but secondary production is similar to that of unvegetated areas, although the relative importance of the trophic guilds is different. The loss of seagrass beds leads to profound alterations in the food web of the total ecosystem. Historical as well as recent changes in material fluxes and energy flow due to man-made alterations to the ecosystem are discussed.     

In situ nitrogen fixation associated with seagrasses in the Gulf of Elat (Red Sea)

In situ nitrogen fixation associated with the seagrass Halophila stipulacea, at the northern Gulf of Elat (Red Sea), is eight to ten times higher than that of nearby plant-free areas. A daily cycle of nitrogen fixation is evident, with rates during the day being seven times greater than during the night. Removal of seagrass leaves only from a patch within a seagrass bed gradually decreases nitrogen fixation activity, reaching the rates of plant-free areas after ten hours. A method devised for the in situ measurement of nitrogen fixation rates using belljars is described in detail. Nitrogen fixation rates in situ are higher than in the laboratory and lack the lag period typical to laboratory measurements. In laboratory experiments using intact plant samples, glucose enhances nitrogen fixation rates both in light and dark. Photosystem II inhibitor (3-3,4-dichloro-phenyl-1,1-dimethylurea) doubles nitrogen fixation rates in light. Both field and laboratory results indicate that light is essential for nitrogen fixation activity in the H. stipulacea bed possibly through its effect on cyanobacterial population that occupy the aerobic niches of the phyllosphere and on photosynthetic Rhodospirillacean bacteria that inhabit the anaerobic ones. Nitrogen fixation rates evident in H. stipulacea beds in situ account for a considerable portion of the biomass production by the seagrasses. The dependence of high nitrogenase activity by the diazotrophs on the presence of the seagrasses indicates the great importance of the seagrass community to the nitrogen cycle in its highly oligotrophic surroundings of the Gulf of Elat.     

Nutrient content of the seagrass Thalassia testudinum reveals regional patterns of relative availability of nitrogen and phosphorus in the Florida Keys USA

Between 1992 and 2000, we sampled 504 randomly chosen locations in theFlorida Keys, Florida, USA, for the elemental content of green leaves of theseagrass Thalassia testudinum. Carbon content ranged from29.4–43.3% (dry weight), nitrogen content from 0.88–3.96%, andphosphorus content from 0.048–0.243%. N and P content of the samples werenot correlated, suggesting that the relative availability of N and P variedacross the sampling region. Spatial pattern in C:N indicated a decrease in Navailability from inshore waters to the reef tract 10 km offshore;in contrast, the pattern in C:P indicated an increase in P availability frominshore waters to the reef tract. The spatial pattern in N:P was used to definea P-limited region of seagrass beds in Florida Bay and near shore, and anN-limited region of seagrass beds offshore. The close juxtaposition ofN–and P-limited regions allows the possibility that N loading from thesuburban Florida Keys could influence the offshore, N-limited seagrass bedswithout impacting the more nearshore, P-limited seagrass beds.     

Azoarcus sp. strain BH72 as a model for nitrogen-fixing grass endophytes

The availability of nitrogen often limits plant growth in terrestrial ecosystems. The only biological reaction counterbalancing the loss of N from soils or ecosystems is biological nitrogen fixation, the enzymatic reduction of N2 to ammonia. Some gramineous crops such as certain Brazilian sugar cane cultivars or Kallar grass can derive a substantial part of the plant nitrogen from biological nitrogen fixation. Our research on grass-associated diazotrophs focuses on endophytic bacteria, microorganisms that multiply and spread inside plants without causing damage of the host plants or conferring an ecological threat to the plant. This review summarizes the current knowledge on the diazotrophic endophyte Azoarcus sp. BH72, which is capable of colonizing the interior of rice roots, one of the globally most important crops.     

Potential roles of Labyrinthula spp. in global seagrass population declines

Overwhelming evidence suggests that seagrass ecosystems are declining around the world. Pathogens from the genus Labyrinthula have repeatedly been found to cause disease in a variety of seagrass species. For example, the ‘wasting disease’ of Zostera marina has been attributed to Labyrinthula infection. Although poorly characterized taxonomically, species of Labyrinthula are very common in marine ecosystems, virulence of genotypes/phylotypes is known to be variable, and highly virulent species are able to cause ecologically significant diseases of protists, plants and animals. Here, the pathosystem model is applied to host–parasite relationships in seagrass ecosystems. Known physical and biological stressors of seagrass are reviewed. Finally, we make the case that it is time to expand research on this poorly studied microorganism in order to quantify the role of disease in seagrass populations world-wide.