中国沿海污损性海鞘生态特点及研究展望

海鞘是海洋污损生物群落的重要组成部分, 其附着会给水产养殖业带来严重危害。在中国沿海, 引发生物污损的海鞘共8 科20 属40 种, 其中渤海海域7 种, 黄海17 种, 东海23 种和南海27 种。北方海域以柄瘤海鞘(Styela clava)、乳突皮海鞘(Molgula manhattensis)和米氏小叶鞘(Diplosoma listerianum)为优势种, 南方则以冠瘤海鞘(Styela canopus)、皱瘤海鞘(S.plicata)和大洋纵列海鞘(Symplegma oceania)为优势种。海鞘污损特点与设施所处地理位置、浸海时间、环境状况等因子密切有关。今后除了继续对沿岸代表性海域开展深入研究外, 还需结合海洋经济发展和科学研究的需要, 加强离岸设施(尤其深海环境)污损生物群落的研究, 以期更好地掌握污损性海鞘的组成分布、种群特点及发展趋势。     

污损性海鞘的生态特点研究展望

海鞘生长快,繁殖迅速,能产生大量在短时间内附着的幼虫,是海洋污损生物群落中的重要成员,对海上人工设施会产生严重危害。污损性海鞘主要由悉尼海鞘(Ascidia sydneiensis)、史氏菊海鞘(Botryllus schlosseri)、米氏小叶鞘(Diplosoma listerianum)、柄瘤海鞘(Styela clava)、红贺海鞘(Herdmania momus)等9科29属103种组成,其中在太平洋海域64种、印度洋23种、大西洋44种,而北冰洋海域仅3种;另外,其附着污损具有明显的地域性和季节特点,并与深度有关。今后应加强污损性海鞘的生态调查和分类研究,阐明深海及两极海域附着污损特点,揭示幼虫附着变态过程的分子调控机理,完善幼虫采集培养技术,以期更好地掌握海鞘生物学特性与生态特点,丰富和发展海洋生态学内容,并为海洋污损生物的防除奠定基础,促进海洋经济产业的发展。     

中国沿海海鞘的物种多样性

至今,中国海已经记录了66种海鞘。其中,渤海有5种,黄海21种,东海24种,南海53种。种类分布从北往南逐渐递增,柄瘤海鞘在黄海、渤海数量很大(8100个/m²),但往南分布,未发现超过罗源湾。皱瘤海鞘在南海数量很大(2225个/m²),往北也未发现超过罗源湾。玻璃海鞘和乳突皮海鞘在全国沿海都有分布。     

黄、渤海沿岸污损生物中的海鞘类

本文根据1974—1985在黄渤海进行污损生物调查的资料,报道该海域的海鞘类14种。同时对这些种类的附着季节、数量及具在生物污损中的意义作初步研究。并对优势种柄瘤海鞘(Styela clava)的种群进行了初步分析。     

桑沟湾栉孔扇贝生物沉积的现场测定

运用沉积物捕集器于桑沟湾对栉孔扇贝 (Chlamysfarreri)的生物沉积进行了现场测定 ,以评价贝类养殖对近岸生态环境的影响。结果表明 :桑沟湾栉孔扇贝具有相当高的生物沉积速率。壳高 68 1～77 9mm、软体干重 2 75～ 3 91g的栉孔扇贝 ,其生物沉积速率为 0 93～ 6 97g ind·d,平均 3 99g ind·d ;而壳高 40 8～ 67 4mm、软体干重 0 5 9～ 1 85g的栉孔扇贝生物沉积速率的变化范围为 0 5 2～ 6 42g ind·d,平均为 2 3 8g ind·d ;另外 ,壳高为 2 5 8～ 2 8 3mm、软体干重 0 1 2～ 0 1 7g的栉孔扇贝 ,其生物沉积速率在2 0 0 2年的 1 1和 1 2月以及 2 0 0 3年的 1月分别为 0 74、1 1 1和 0 1 5g ind·d。在桑沟湾 ,影响栉孔扇贝生物沉积的主要因素包括水温、悬浮颗粒物、扇贝个体大小和年龄。高密度、大规模的近岸浅海贝类养殖所产生的大量生物沉积物可能会对海区的物理、化学和生物环境产生影响。     

汕头南澳养殖区表层沉积物中生源要素的分布及其污染评价

于2010年9月采集南澳海域鱼类养殖区、贝类养殖区、大型藻类栽培区和对照区4个功能区表层沉积物样品,分析沉积物中生物硅(BSi)、总有机碳(TOC)、总氮(TN)和总磷(TP)4种生源要素的含量。结果表明,南澳海域生源要素含量与国内外养殖区相比属于中等水平。鱼类养殖区TOC、TN、TP的含量最高,而BSi的最高含量出现在藻类栽培区(平均含量为0.30%)。鱼类养殖区BSi、TOC、TN和TP的平均含量分别为0.24%、0.89%、0.13%和0.097%。根据沉积物中TOC/TN比值分析,发现鱼类养殖区的TOC主要来源于水生,贝类、藻类和对照区TOC则主要来源于陆源。生源要素污染评价表明,4个功能区的TN均属于Ⅱ类污染,鱼类和贝类养殖区的TP属于Ⅱ类轻度污染。     

庄河海域菲律宾蛤仔底播增殖区自身污染

采用生物沉积物捕集器和封闭式代谢瓶,周年现场研究了庄河海域菲律宾蛤仔的生物沉积速率、排氨率和排磷率.结果表明:菲律宾蛤仔的生物沉积速率、排氨率和排磷率均具有明显的季节变化.生物沉积速率为0.15～1.47 g·ind-1·d-1(年均0.61 g·ind-1·d-1);其排氨率及排磷率分别为0.02～0.40 mg·ind-1·d-1(年均0.17 mg·ind-1·d-1)和0.01～0.39 mg·ind-1·d-1(年均0.13 mg·ind-1·d-1).根据以上结果,估算庄河海域底播增殖菲律宾蛤仔每年产生的生物沉积物达到5.46×107t(干质量),折合有机物9.07×106t、有机碳1.00x106t和有机氮1.18xlO5t;而氨氮和磷酸盐分别为1.49x104t和1.15x104t.表明浅海高密度、规模化菲律宾蛤仔增养殖区自身污染严重,其对环境的影响不可忽视.     

我国海洋大型底栖生物多样性研究及展望:以黄海为例

本文综述了我国海洋大型底栖生物生态学和生物多样性研究进展,指出在该领域已有的研究中以胶州湾、长江口、福建和浙江沿岸等海域研究较多,其他海域还需要加强.介绍了常规大型底栖生物群落研究(包括种类组成、栖息密度(丰度)、生物量、生物多样性、次级生产力、能量级等)中的研究方法,环境变化、环境污染、海洋工程建设等对大型底栖生物的影响等方面的主要研究成果和进展,并重点以黄海为例,列举了已有的主要调查项目和研究成果.我国已有的研究表明,渤海海域底栖动物区系简单,大型底栖生物共413种,种数在四个海区中最少,优势种主要是低温、广盐暖水种;年总平均生物量为19.83 g/m2,以软体动物占有绝对优势;年总平均栖息密度474个/m2,多毛类和软体动物为主要贡献类群.黄海大型底栖生物共853种,优势种为狭盐性北温带种;北黄海年总平均生物量为99.66 g/m2,以棘皮动物最高;南黄海年总平均生物量为27.69 g/m2,多毛类贡献量最高;北黄海年总平均密度为2,017.40个/m2,而南黄海只有88.67个/m2差异较大;北黄海大型底栖生物的总平均密度和生物量远高于其他海域.南黄海年平均次级生产力为4.98 g(去灰干重)/m2,两个高生产力区位于黄海冷水团两侧.东海共发现大型底栖生物l,300种,主要优势种类有45种,与黄海优势种生物之间有很大差别;东海平均生物量的季节变化不显著,基本趋势为春季＞冬季＞夏季＞秋季;东海各海域的年总平均生物量浙江海域(28.22 g/m2)＞长江口海域(15.55 g/m2)＞台湾海峡(8.98 g/m2);年总平均栖息密度为164个/m2,呈现由北向南逐渐增加的趋势.南海大型底栖生物共发现1,661种,明显高于渤、黄、东海,部分优势种类与东海相同,与黄海、渤海差别较大;年总平均生物量为20.06 g/m2,基本趋势为春季＞冬季＞秋季＞夏季,软体动物和棘皮动物为主要贡献类群;年总平均栖息密度为198个/m2,夏季＞春季＞冬季＞秋季,多毛类生物为主要贡献者.指出随着人类活动和全球气候变化对海洋环境的影响越来越剧烈,对海洋生物资源和生物多样性的保护显得尤为急迫,而对海洋大型底栖生物长期的变化规律、变化机理、未来变化趋势预测的研究,是探究海洋大型底栖生物生态服务功能,提出资源有效保护和合理开发建议,使之永续利用的基础和迫切任务.     

西藏达则错盐湖沉积背景与有机沉积结构

以西藏拟溞(Daphniopsis tibetana Sars)为优势浮游动物物种的低盐度盐湖是西藏湖泊的一个重要类型,以达则错为代表,分析了其沉积背景及沉积物组成。结果如下:(1)湖泊敞水区无机沉积以内生化学沉积为主,可代表深水盐湖无机沉积物的自然沉积过程。(2)达则错盐湖浮游植物以蓝藻、硅藻、裸藻、绿藻为主,总生物量11.35 mg/L;浮游动物生物量为4.92 mg/L,其中西藏拟溞占 82.30%;浮游植物残体受盐梯度影响在盐梯度层之上聚集,而浮游动物残体及粪粒(Fecal pellets)因外表有碳酸盐附着可穿过盐梯度层沉积湖底,生物残体与浮游动物代谢产物构成了沉积有机物的物质基础。(3)表层沉积物平均含水量为66.70%,粒径0.004-0.02 mm范围内的颗粒物含量最大,占20.42%,其次为<0.004 mm的粘土,占4.53%。(4)表层沉积物总有机碳(TOC)平均含量为27.99 mg/g(干重),其中颗粒有机碳(POC)约为18.11 mg/g,占TOC的64.70%;在POC中,西藏拟溞粪粒贡献最大,约占POC的60.48%,占TOC的39.06%,占沉积物总量的1.12%,其次为西藏拟溞残体,占POC的38.85%。分析结果表明盐湖因其独特的水化学和生物学特征具有较强的沉积能力,以化学沉积为主的无机沉积及以西藏拟溞粪粒和残肢碎屑为主的有机沉积构成了该类型盐湖颗粒物沉降及沉积的主要过程。     

黄东海陆架区沉积物中磷的形态分布及生物可利用性

采用1992年Ruttenberg连续提取法(SEDEX)将黄东海陆架区沉积物中的磷分为交换态磷(Ex-P),Fe结合态磷(Fe-P),自生磷(Au-P),碎屑磷(De-P),有机磷(Or-P),分析了各形态磷的平面和垂直分布特征;利用沉积物年代序列测定的结果,探讨了柱状沉积物中不同形态磷的含量变化,并进一步分析了该区域磷形态的生物可利用性.结果表明,黄东海陆架区表层沉积物各形态磷平均含量为:Au-P(140.72 μg/g)＞De-P(59.23 μg/g)＞Or-P(32.69 μg/g)＞Fe-P(29.91 μg/g)＞Ex-P(5.92 μg/g);各形态磷在沉积时间序列上分布不同,反映了不同时期人类活动和气候环境等因子对磷埋藏量影响的不同,其中Au-P在长江口H1-18站位含量比南黄海中部3个站位要低得多;调查区表层潜在生物有效磷为13.55％左右,仅仅占沉积磷中的一小部分.     

Preparation, properties and crystal structure of two isomers of R,R,S,S- and R,S,R,S-[chloro(1,3,10,12,16,19-hexaazatetracyclo[17,3,1,1,0]tetracosane)copper(II)]

Two isomers (R,S,R,S- and R,R,S,S-) of five coordinate complex [Cu(L)Cl]⁺ have been separated and characterised. These two isomers have significantly different spectrochemical and electrochemical properties. Absorption maximum of R,S,R,S-[Cu(L)Cl]⁺ shifts to longer wavelength and its reduction potential shifts to more positive direction comparing those of R,R,S,S-[Cu(L)Cl]⁺. R,S,R,S-[Cu(L)Cl]⁺ is significantly distorted to trigonal-bipyramidal structure, whereas R,R,S,S-[Cu(L)Cl]⁺ retains almost square-planar geometry. The average bond distance of Cu-N in basal plane of R,S,R,S-[Cu(L)Cl]⁺ is longer by 0.024 Å than that of R,R,S,S-[Cu(L)Cl]⁺, whereas the bond distance of Cu-Cl in former is shorter by 0.200 Å than that in latter. The isolated square-planar complexes of R,R,S,S- and R,S,R,S-[Cu(L)](ClO₄)₂ are converted to the R,R,S,S- and R,S,R,S-[Cu(L)Cl]⁺ by the addition of Cl⁻ in nitromethane solution with the rate constants, k=1.70 (±0.02) and 8.31 (±0.07) M⁻¹ s⁻¹, respectively.     

Temporal variation in zebra mussel (Dreissena polymorpha) density structure the benthic food web and community composition on hard substrates in Lake Constance, Germany

Invasive species often influence existing biocenoses by altering their environment or facilitating the ecology of other species. Here we combined stable isotope analysis with quantitative benthic community sampling to investigate temporal variation in the influence of biodeposition of organic material (biodeposits) by the zebra mussel (Dreissena polymorpha) on the benthic food web in hard substrate habitats of Lake Constance, Germany. The accumulation of organic material excreted by zebra mussels (faeces and pseudofaeces) is hypothesised to implement a biodeposition based food web. By means of stable isotope analyses, we found that the feeding strategy of amphipods was dependent on the availability of zebra mussel biodeposits. The proportion of pelagic resources contributing to the diet (by ??¹³C) and trophic position (by ??¹⁵N) of amphipods were significantly and positively correlated to the production of zebra mussel biodeposits. Also, the benthic community in Lower Lake Constance showed a shift towards higher densities of the mayfly Caenis spp., Chironominae, and the caddisfly Ecnomus tenellus, which might act as links to higher trophic levels when more zebra mussel biodeposits were available. These results demonstrate that temporal variation in zebra mussel density, and thus in organic biodeposition, have a strong impact on the benthic communities and food web structure associated with hard substrates to the extent that there may be dependence on zebra mussel occurrence in such habitats in Lake Constance.     

Experimental test of biodeposition and ammonium excretion from blue mussels (Mytilus edulis) on eelgrass (Zostera marina) performance

Blue mussels and eelgrass have been found to coexist in many locations. However, knowledge of the interactions between these species is limited. Two experiments were conducted in the laboratory, a “Deposit” and an “Epiphyte” experiment. The Deposit experiment examined possible effects of increasing load of blue mussel (Mytilus edulis) biodeposits on sediment biogeochemistry and eelgrass (Zostera marina) performance. Z. marina mesocosms received normal or high loads of mussel biodeposits (Normal and High), while no biodeposits were added to the Control. High dosage had overall negative effects on Z. marina, which was reflected as lower leaf numbers and biomass and accumulation of elemental sulphur in rhizomes. The sediment biogeochemical conditions were altered, as the mussel biodeposits enhanced sulphate reduction rates and increased sulphide concentrations in the porewater, which may result in sulphide invasion and reduced growth of Z. marina.In the Epiphyte experiment effects of mussel excretion, with particular emphasis on ammonium, on the growth of Z. marina and their epiphytes were examined. A thick cover of epiphytes developed on Z. marina growing together with M. edulis, and the relative growth rate was reduced with 20% compared to plants from control without mussels. Overall the experiments showed negative effects on Z. marina growing together with M. edulis, thereby supporting a preceding field study by Vinther et al. [Vinther, H.F., Laursen, J.S., Holmer, M. 2008. Negative effects of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds in Flensborg fjord, Denmark. Est. Coast Shelf. Sci. 77, 91-103.].     

Biofouling ascidians on aquaculture gear as potential vectors of harmful algal introductions

Biofouling ascidians are ubiquitous in coastal ecosystems and are among the main colonizers of aquaculture gear. Our study tested the hypothesis that the transport, removal, and transfer of fouling ascidian species by aquaculturists provide a mechanism for concentration and distribution of harmful-algal cells to new areas. Wild-caught specimens of common, biofouling ascidian species (Styela clava, Ciona intestinalis, Molgula manhattensis, Botrylloides violaceus, Didemnum vexillum, and Botryllus schlosseri) were exposed individually to cultured strains of co-occurring harmful algae (Prorocentrum minimum, Alexandrium fundyense, Alexandrium monilatum, Karenia brevis, Aureococcus anophagefferens, or Heterosigma akashiwo) at simulated bloom cell densities of each HAB species. After feeding, ascidians were transferred to ultrafiltered seawater. Immediately after exposure, and after 24 and 48 h in ultrafiltered seawater, biodeposits were collected and observed microscopically for the presence of intact, potentially viable cells. Subsamples of biodeposits were transferred into culture tubes with ultrafiltered seawater and monitored for algal growth during 8 weeks. Cells of all HAB species were found to pass intact through the ascidian digestive system, remained viable, and in many cases were capable of re-establishing populations at least 48 h post-ingestion. The results of our study will inform industry and managers of the potential threat and ecological impact of spreading biofouling ascidians, and practices to mitigate adverse impacts. Additionally, these management practices have been formally incorporated into a new cost-share program developed to help shellfish producers prevent the further spread of ascidians and associated HAB species.     

Variation in the ability of Didemnum sp. to invade established communities

Over the past 30 years southern New England, USA has been invaded by several species of ascidians, including Botrylloides violaceus, Diplosoma listerianum, Styela clava, and Ascidiella aspersa. These species have become dominate in coastal embayments and marinas but are usually absent from more open water coastal areas. A colonial ascidian, Didemnum sp. has invaded southern New England during the past 10 years and we first observed this species in eastern Long Island Sound in 2000. It has become the dominant at several field sites while remaining in low abundance at others. We conducted an experiment at two places, a protected marina and an open coast site, to examine its ability to compete with the established fouling community. Small colonies of Didemnum were transplanted onto panels with communities that varied in age from one to four weeks old and these treatments along with controls with only Didemnum were exposed at both sites. In most treatments Didemnum became a dominant species in the communities at both sites but it reached higher abundances at the open coast site. Potential causes of the observed differences are predation on other species of ascidians at the open coast site reducing recruitment of these species and competition, lower tolerance for elevated temperatures at the marina site, or other environmental parameters that might affect growth rates.     

The non-native solitary ascidian Ciona intestinalis (L.) depresses species richness

Non-native ascidians are a dominant feature of many sessile marine communities throughout the world and may have negative effects on species diversity. We tested effects of the non-native Ciona intestinalis on the sessile invertebrate community in San Francisco Bay, where it occurs in dense aggregations. In particular, we compared species richness between PVC panels from which C. intestinalis were experimentally removed to panels with naturally dense C. intestinalis growth, using fouling panels of four sizes (between 49 cm² and 1177 cm²) to measure the effect of C. intestinalis recruitment on species-area relationships. We initially deployed 120 fouling panels (30 of each size) at a site known to have dense populations of C. intestinalis, assigning these to three different treatments: (1) Experimental removal, whereby new recruits of C. intestinalis were removed on a weekly basis, pulling panels out of the water for a short time period to do so; (2) Manipulated control, whereby panels were removed from the water each week (as in the experimental removal) but without C. intestinalis removal; and (3) Unmanipulated control, which remained in the water throughout the experiment. After 4 months, all of the panels were collected and analyzed to estimate species richness and relative abundance (percent cover) of sessile invertebrates and of C. intestinalis. Across all panels, species richness was negatively correlated with C. intestinalis abundance. The removal of C. intestinalis produced communities with significantly higher species richness than the controls. The overall species composition of treated and control panels was also distinctly different, with many species occurring more often in the absence of C. intestinalis, while others occurred more often on C. intestinalis-dominated panels. These data suggest that C. intestinalis both depress local species diversity and alter community assembly processes to fundamentally change sessile community composition.     

Feeding preference of Strongylocentrotus droebachiensis (Echinoidea) for a dominant native ascidian, Aplidium glabrum, relative to the invasive ascidian Botrylloides violaceus

Subtidal benthic communities show distinct patterns of community structure related to substrate angle. Suspension-feeding invertebrate communities often dominate vertical and undercut rock surfaces, while macroalgae dominate horizontal surfaces. One factor that may shape this pattern is sea urchin grazing, which is often more intense on horizontal surfaces. The native Gulf of Maine ascidian Aplidium glabrum, like other native ascidians, is generally restricted to vertical and undercut rock walls, whereas the introduced ascidian Botrylloides violaceous is often abundant on horizontal surfaces. We tested the hypothesis that this pattern could be due to differing predation intensity on these two ascidians by Strongylocentrotus droebachiensis, a dominant omnivore in the Gulf of Maine. Feeding preference of S. droebachiensis on the native A. glabrum vs. B. violaceous was estimated in the laboratory and in field experiments. Laboratory results show that S. droebachiensis prefers to feed on the native ascidian A. glabrum over B. violaceous. In the field, potential differences in the impact of S. droebachiensis on the two species were masked by the much greater growth rate of B. violaceus compared to A. glabrum. These results may help explain observed patterns in ascidian distribution in the Gulf of Maine, and ultimately the overall success of B. violaceus as a major invader throughout New England.     

Continuous culture of the microalgae Schizochytrium limacinum on biodiesel-derived crude glycerol for producing docosahexaenoic acid

Crude glycerol is a major byproduct of the biodiesel industry; previous research has proved the feasibility of producing docosahexaenoic acid (DHA, 22:6 n − 3) through fermentation of the algae Schizochytrium limacinum on crude glycerol. The objective of this work is to investigate the cell growth kinetics, substrate utilization efficiency, and DHA production of the algae through a continuous culture. Steady-state biomass yield, biomass productivity, growth yield on glycerol, specific glycerol consumption rate, and fatty acid composition were investigated within the range of dilution rate (D) from 0.2 to 0.6 day⁻¹, and the range of feed crude glycerol concentration (S₀) from 15 to 120 g/L. The maximum specific growth rate was determined as 0.692 day⁻¹. The cells had a true growth yield of 0.283 g/g but with a relatively high maintenance coefficient (0.2216 day⁻¹). The highest biomass productivity of 3.88 g/L-day was obtained at D = 0.3 day⁻¹ and S₀ = 60 g/L, while the highest DHA productivity (0.52 g/L-day) was obtained at D = 0.3 day⁻¹ and S₀ = 90 g/L due to the higher DHA content at S₀ = 90 g/L. The biomass and DHA productivity of the continuous culture was comparable to those of batch culture, while lower than the fed-batch culture, mainly because of the lower DHA content obtained by the continuous culture. Overall, the results show that continuous culture is a powerful tool to investigate the cell growth kinetics and physiological behaviors of the algae growing on biodiesel-derived crude glycerol.     

Antimalarial sesquiterpene lactones from Distephanus angulifolius

Combined use of bioassay-guided fractionation based on in vitro antiplasmodial assay and dereplication based on HPLC-PDA-MS-SPE-NMR led to isolation of (6S,7R,8S)-14-acetoxy-8-[2-hydroxymethylacrylat]-15-helianga-1(10),4,11(13)-trien-15-al-6,12-olid and (5R,6R,7R,8S,10S)-14-acetoxy-8-[2-hydroxymethylacrylat]-elema-1,3,11(13)-trien-15-al-6,12-olid, along with vernodalol, vernodalin, and 11,13β-dihydroxyvernodalin from extract of Distephanus angulifolius. All compounds were identified by spectroscopic methods, including 1D and 2D homo- and heteronuclear NMR experiments. The isolated compounds showed IC₅₀ values in the range 1.6-3.8 μM and 2.1-4.9 μM against chloroquine sensitive D10 and chloroquine resistant W2 Plasmodium falciparum strains, respectively.     

Metabolic Activity and Functional Diversity Changes in Sediment Prokaryotic Communities Organically Enriched with Mussel Biodeposits

This experimental microcosm study reports the influence of organic enrichments by mussel biodeposits on the metabolic activity and functional diversity of benthic prokaryotic communities. The different biodeposit enrichment regimes created, which mimicked the quantity of faeces and pseudo-faeces potentially deposited below mussel farms, show a clear stimulatory effect of this organic enrichment on prokaryotic metabolic activity. This effect was detected once a certain level of biodeposition was attained with a tipping point estimated between 3.25 and 10 g day^-1 m^-2. Prokaryotic communities recovered their initial metabolic activity by 11 days after the cessation of biodeposit additions. However, their functional diversity remained greater than prior to the disturbance suggesting that mussel biodeposit enrichment may disturb the functioning and perhaps the role of prokaryotic communities in benthic ecosystems. This manipulative approach provided new information on the influence of mussel biodeposition on benthic prokaryotic communities and dose-response relationships and may support the development of carrying capacity models for bivalve culture.