A mathematical model for the uptake of heavy metals in benthic algae

A mathematical model for the uptake of heavy metals in the benthic algae Ascophyllum nodosum has been developed. The model allows assignment of age-dependent growth parameters which also may be a function of external factors. Mortality curves for A. nodosum corresponding to a highly polluted and a nearly unpolluted area have been used in the study of metal uptake. Uptake has been simulated for different values of growth parameters and different concentrations of heavy metals in seawater. Emphasis has been on the uptake of zinc. The simulations showed that the concentration of zinc in the algae changed much less (6%) than the biomass (30%) with changes in intrinsic growth rate, mortality and carrying capacity.The concentration in the algae was found to be an approximately linear function of the mean concentration in seawater up to about 100 ppb. At very high concentrations, associated with high mortality, the deviation from linearity may be significant (about 13% at 162 ppb at one locality). The calculation indicates further that variations in the zinc concentration in the seawater are considerably damped in the algae.Simulations have also been performed to study the effects of sampling different parts of the algae, and some initial simulations have been performed to study the effects of heavy metal exchange between algae and sea water.     

A mathematical model for the distribution and abundance of benthic algae in a Norwegian fjord

A mathematical model has been developed to study the distribution and abundance of benthic algae species in a Norwegian fjord. The main forcing functions are the physical factors which have well defined gradients in this area, but biotic interactions are also accounted for in the model.The distribution of algae along vertical transects in the intermediate fjord area has been simulated, and the observed distribution is reproduced fairly well. This indicates that the major factors determining the equilibrium elevation between intertidal and subtidal vegetations in this area are adequately represented in the model.Based on observations in a fjord branch with a high level of freshwater influx and polluted by heavy metals, assumptions are made of the combined influence of these contaminants on growth characteristics of the species considered. The resulting effects on community development and community patterns are simulated. For a certain level of contamination, the decrease in abundance, community development rate and species separation have been determined. It is also shown that the fluctuations in biomass during a year are more pronounced than under unaffected conditions.The relative importance of external growth regulating factors and self-shading is demonstrated for different degrees of community development, and we have indicated that this may be an important factor in the assessment of transient and long-term responses to “natural” and man-made influences on the shore-line vegetation.One of the main problems during model development was the formulation of the interaction function for factors influencing growth parameters. Several alternatives were examined.The differences between the most frequently used forms were small with respect to abundance and not very large with respect to possible changes in distribution. It was noted that even by including only the most limiting factor at each instant, (Liebig's form) a high percentage of the potentially limiting factors did actually limit growth during periods of the year.The model is intended as a working tool for the studies of changes in the shore-line vegetation caused by man-made interferences like heavy metal and oil pollution, and we believe that the results presented demonstrate that it can successfully be used for this purpose.     

Alkanes and alkenes in marine benthic algae

Saturated and olefinic hydrocarbons were determined in additional species of benthic marine algae from the Cape Cod (Massachusetts, USA) area (see: Youngblood et al., 1971). The distribution of homologous and isomeric olefins was studied in plants of different age and in morphologically different parts of the same specimen. With two minor exceptions, only normal alkanes and alkenes are present. The methylene-interrupted C₁₉- and C₂₁-polyolefins are particularly abundant; ¹-heneicosahexaene and the corresponding pentaene are common to all brown algae, while the corresponding ³-isomers occur in green algae. The hydrocarbon concentration, the alkene-to-alkane ratio and the polyolefin content are highest in young plants or in rapidly growing tissues of older plants. This suggests a deeper involvement in cell biochemistry of straight-chain hydrocarbons than previously considered. The biosynthesis of the plant polyolefins remains to be explored; no immediately obvious precursors of the ¹-polyolefins were found among the algal fatty acids. The hydrocarbon composition of these benthic algae differs greatly from that of fossil fuels in its simplicity and predominately unsaturated nature. The separation of the isomers by gas chromatography and their structural elucidation by mass spectrometry, alone and in combination with hydrogenation and ozonolysis, are discussed.Contribution No. 3155 of the Woods Hole Oceanographic Institution.     

Global biodiversity patterns of benthic marine algae

Species richness patterns are remarkably similar across many marine taxa, yet explanations of how such patterns are generated and maintained are conflicting. I use published occurrence data to identify previously masked latitudinal and longitudinal diversity gradients for all genera of benthic marine macroalgae and for species in the Order Bryopsidales. I also quantify the size, location, and overlap of macroalgal geographic ranges to determine how the observed richness patterns are generated. Algal genera exhibit an inverse latitudinal gradient, with biodiversity hotspots in temperate regions, while bryopsidalean species reach peak diversity in the tropics. The geographic distribution of range locations results in distinct clusters of range mid-points. In particular, widespread taxa are centered within tight latitudinal and longitudinal bands in the middle of the Indo-Pacific and Atlantic Oceans while small-ranged taxa are clustered in peripheral locations, suggesting that variation in speciation and extinction are important drivers of algal diversity patterns. Hypotheses about factors that regulate diversity contain underlying assumptions about the size and location of geographic ranges, in addition to predictions as to why species numbers will differ among regions. Yet these assumptions are rarely considered in assessing the validity of the prevailing hypotheses. I assess a suite of hypotheses, suggested to explain patterns of marine diversity, by comparing algal-richness patterns in combination with the size and location of algal geographic ranges, to the richness and range locations predicted by these hypotheses. In particular, the results implicate habitat areas and ocean currents as the most plausible drivers of observed diversity patterns.     

Exosmosis and “free space” in marine benthic algae

In distilled water, marine benthic algae immediately lose ions. This can be demonstrated by means of chloride titration or conductivity measurements. The rapidity and the complete reversibility of this process show that ion movements out of and into the free space are involved. When Laminaria saccharina thalli, exposed to a series of increasing NaCl-concentrations, are subsequently transferred into distilled water, the external concentration increases proportionally to rate of ion loss. Through its free space, the alga establishes an ionic equilibrium with its external medium. If the alga thallus is killed (20 sec boiling in isosmotic sea water), the extent of chloride loss is much higher. Because of thermal destruction of the osmotic space, the chloride can then escape from the entirc thallus. Since sublittoral algae die upon drying, chloride loss from dry thalli is much higher than that from wet thalli: this difference is small in littoral algae, which tolerate short periods of dryness. A close relationship exists between extent of chloride loss and degree of resistance to drying.     

Relative and interactive effects of plant and grazer richness in a benthic marine community

The interactive effects of changing biodiversity of consumers and their prey are poorly understood but are likely to be important under realistic scenarios of biodiversity loss and gain. We performed two factorial manipulations of macroalgal group (greens, reds, and browns) and herbivore species (amphipods, sea urchin, and fish) composition and richness in outdoor mesocosms simulating a subtidal, hard-substratum estuarine community in North Carolina, U.S.A. In the experiment where grazer richness treatments were substitutive, there were no significant effects of algal or herbivore richness on final algal biomass. However, in the experiment in which grazer treatments were additive (i.e., species-specific densities were held constant across richness treatments), we found strong independent and interactive effects of algal and herbivore richness. Herbivore polycultures reduced algal biomass to a greater degree than the sum of the three herbivore monocultures, indicating that the measured grazer richness effects were not due solely to increased herbivore density in the polycultures. Taking grazer density into account also revealed that increasing algal richness dampened grazer richness effects. Additionally, the effect of algal richness on algal biomass accumulation was far stronger when herbivores were absent, suggesting that grazers can utilize the increased productivity and mask the positive effects of plant biodiversity on primary production. Our results highlight the complex independent and interactive effects of biodiversity between adjacent trophic levels and emphasize the importance of performing biodiversity-ecosystem functioning experiments in a realistic multi-trophic context.     

Photosynthetic rates of benthic marine algae in relation to light intensity and seasonal variations

The effect of light intensity on rate of photosynthesis was measured at irregular intervals over a 12-month period for 22 benthic marine algae in the western Baltic Sea. In most species photosynthesis (mg O₂·g dry weight^-1·h^-1) was highest in spring and summer, corresponding to the seasonal growth pattern of the algae. In winter all the species showed adaptation of the light compensation point. Highest productivity was shown by algae which are: short-lived annual species rather than perennials, eulittoral rather than sublittoral, and which possess sheet-like or filamentous thalli rather than coarsely branched forms. These factors are clearly inter-related.This research was carried out while one of us (R.J.K.) was an Alexander von Humboldt fellow at the University of Kiel, and is part of the programme Sonderforschungsbereich 95, Wechselwirkung Meer-Meeresboden, Universität Kiel.     

A trait specific model of competition in a spatially structured plant community

The non-random dispersal of plant propagules is thought to counter competitive exclusion and thus promote the survival of competitively inferior species. We investigated this process by modelling the outcome of interactions between species with competitive ability defined as a function of both life-history traits and the environment with both random and clustered dispersal strategies and in environmentally homogeneous and heterogeneous environments. Four main results emerged: (1) environmental heterogeneity was seen to promote co-existence in conjunction with associated trait variation for tolerance to the environmental variable and where this trait variation was effectively limited by ‘trade-off’ such that no single species had an overall competitive advantage, (2) consistent with theory, random dispersal appeared to enhance the likelihood of competitive exclusion, whereas clustering favoured co-existence, (3) the ecological outcome of interactions between dispersal and competitive relationships varied as a function of the trait determining the competitive advantage within a particular environment, and (4) promotion of co-existence by clustered dispersal was most marked when associated with environmental heterogeneity. It is argued that these results suggest that current ecological models of species interactions may need to be modified to incorporate a more realistic understanding of competitive ability if we are to better understand the factors effecting species co-existence.     

Calorific values of benthic marine algae and their postulated relation to invertebrate food preference

Data are given on the kcal/g dry weight, percent ash, and kcal/ash-free g of dry weight for 1 marine diatom species, 70 macroscopic benthic algae, and 1 marine tracheophyte species. For 41 of these, the data have also been converted to kcal/g wet weight. Calorific values, though relatively homogeneous within genera, appear to be influenced by phyletic affinity, water purity, or depth of immersion, and such ecological properties as growth form, generation time, and relative susceptibility to herbivory. Seasonal factors, portion of the plant combusted and growth rate appear to have little effect. The food preferences of various invertebrate herbivores seem to have evolved more in response to an availability factor than to absolute food value.     

RQ of benthic marine invertebrates

This study investigated an incubation method which employed simultaneous measurement of CO₂ production and O₂ consumption rates to calculate the RQ (respiratory quotient; CO₂ production rate: O₂ consumption rate) of individual benthic marine invertebrates. Carbon dioxide production rates were calculated from changes in CO₂ concentration determined using seawater pH. O₂ consumption rates were calculated from changes in O₂ concentration with a correction applied for O₂ flux across the air/water interface due to gaseous exchange. Species examined were Triphyllozoon sp. cf. moniliferum (MacGillivray 1860), a bryozoan; Herdmania momus (Savigny), a solitary ascidian; Poneroplax albida (Blainville 1825), a chiton; and Haliotis roei (Gray 1826), an abalone. Six individuals of each were collected on 14 November 1985 from the limestone walls of a cave in a nearshore reef off Marmion, Western Australia. After acclimation for 6 h in experimental conditions, rates of CO₂ production and O₂ consumption were measured. A minimum period of 4 h was required to obtain consistent RQ values for each species. The standard error (SE) of the (calculated) RQ ratio was 14 to 33% of the mean in incubations of 4 h, and less than 14% in incubations of 4 to 12 h. The RQ is commonly used as an indicator of unknown catabolic substrates by comparing it with biochemically determined limits for known substrates. This study provides a strong argument against using the RQ of individual animals to draw any conclusions about catabolic substrates. Unexplained variation in the components of the RQ of an individual, measured over short time periods, and the potential involvement of stored reserves in catabolism, over longer time periods, obscure the relationship between the RQ of individual animals and the ratio's biochemically determined limits.     

Deep-water benthic algae of the Florida Middle Ground

The composition and seasonality of the benthic algae of the Florida Middle Ground, an offshore area of extensive reef outcroppings, 25 to 60 m deep in the eastern Gulf of Mexico, are described. Collections and observations were made by SCUBA diving during June, July, and September, 1971, and January 1972. Ninety-one algal species (92 taxa) were obtained, including 6 species newly reported for Florida and the eastern Gulf of Mexico. The flora is predominantly tropical, with Caribbean affinities. Marked seasonal differences in species diversity and abundance were present. An extensive or well-anchored holdfast system was a common feature of those species which appeared to be perennial.     

Environmental grain and polychaete species' diversity in a bathyal benthic community

Patterns of polychaete species' dispersion in the San Diego Trough, Southern California Continental Borderland, North Pacific Ocean, at a depth of 1230 m are analyzed. Samples consist of 4 replicate 0.25-m² box cores, each partitioned into twenty-five 0.01-m² subcores, and of 1 unpartitioned core. The sampler is biased but, when this bias is taken into account, few species show strong aggregation either between or within cores. If all species are considered together as replicates, intraspecific aggregation is detected between cores, but uniform dispersion dominates within cores. Patchiness or habitat partitioning on the assayed scales can thus account for little of the extremely high species' diversity observed. Apparently due to an environmental structure having a “grain” smaller than 0.01-m², the diversity of species assumed to be sedentary is higher than the diversity of species assumed to be mobile. One evidence of such structure is a negative covariance between paraonid abundance and density of “mudballs” constructed by a cirratulid. Consideration of these results suggests that either grain specialization or simultaneous, biogenic disequilibrium or both act on spatial scales smaller than 0.01-m² to maintain high polychaete species' diversity in the San Diego Trough.     

Hydrocarbons in benthic algae from the eastern Gulf of Mexico

Thirty-six samples of benthic algae were collected from the continental shelf along the eastern Gulf of Mexico. The algae contain 82.8±143 ppm aliphatic hydrocarbons by dry weight and 11.8±22.7 ppm aromatic and polyolefinic hydrocarbons. The aliphatic constituents of red algae are composed almost entirely of n-C₁₇ (70 to 95% of aliphatic weight); green algae have varying amounts of n-alkanes in the range of n-C₁₅ to n-C₁₉, with homologous series of odd carbon number n-alkenes. Phytadienes occur as high as 740 ppm dry weight in the green algae and do not appear to be related to time or place of collection or to presence of any petroleum pollutants. The single brown algal species contains predominantly n-C₁₅. About 30% of the samples have hydrocarbons resembling degraded petroleum residues, but no petroleum pollution of recent origin was detected in any specimen.     

Surface area in ecological analysis: Quantification of benthic coral-reef algae

The surface area of organisms and substrata is shown to be a significant ecological parameter because of its functional importance in the system. Quantification of surface area can be of particular value in morphologically complex environments such as coral reefs. The amount of surface in a reef habitat can be estimated by direct measurements and theoretical approximations, using a surface index (SI) for the amount of surface increase over that of a similarly bounded plane. SI values for a section of the British Honduran barrier reef ranged up to 15 in the reef-crest area at scales significant to macro-organisms. By combining substrate-area measurements with estimates of percent coverage of the major benthic algal components, a reef transect with a horizontal area of 300 m² was shown to have over 300 m² covered by benthic macroalgae. The potential for the further development of surface estimation techniques is discussed.     

CO2 fixation and translocation in benthic marine algae. III. Rates and ecological significance of translocation in Laminaria hyperborea and L. saccharina

Translocation of ¹⁴C-labelled assimilates in Laminaria hyperborea and L. saccharina is strictly basipetal within the thallus. The lowermost 20 cm of the frond accumulates 71% (L. hyperborea) or 63% (L. saccharina) of the basipetallymoving translocate after a period of 96 h from initial feeding with NaH¹⁴CO₃. Whereas translocation was found to occur at all seasons in L. saccharina, which grows throughout the year, no translocation of newly synthesized photosynthate occurred in L. hyperborea in October, when no frond enlargement takes place, nor in January, when the new fond occurs as a tiny outgrowth. In L. hyperborea, organic material, assimilated by the old frond during the period of rapid growth, is exported at the rate of 0.5 mg C · dm^-2 · h^-1. In L. saccharina, 14 to 18 cm frond length are necessary for an elongation rate of the growing zone, situated in lowermost 10 cm of the frond, of 1 cm per week. The significance of translocation in the two Laminaria species is demonstrated by comparison of export rates with growth rates measured in situ and in the laboratory.Dedicated to Professor Dr. Otto Stocker on his 85th birthday.     

Biosorption of lead and cadmium using marine algae

The use of algae (Ulva fasciata, green and Sargassum sp., brown) to reduce lead and cadmium levels from mono-metal solutions was investigated. The brown algae showed higher efficiency for the accumulation of lead (～1.5 times) and cadmium (～2 times) than green algae. The optimum pH value is found to be between 4 and 5.5. Regarding biomass concentration, an increase in metals percentage removal and a decrease in metal uptake capacity coincided with the increase in biomass concentration. All light metals (Ca, Mg and Na) showed a suppressive effect on biosorption capacity. The enhancement of biosorption in the case of NaOH was obvious. The biosorption process (65–90%) occurred within 3?min. Experimental data were in high agreement with the pseudo-second-order kinetic model and Freundlich model for lead and cadmium biosorption using different biosorbents. In the desorption study, 0.2?mol?L^?1 HCl recorded the best concentration for the elution of metals from the biomass. The biosorption capacity decreased over the four operational cycles for both lead and cadmium. Infrared analysis showed that amino, hydroxyl and carboxyl functional groups provide the major biosorption sites for metal binding. Use of the above-mentioned algae for cheap metal absorbance is considered as one water treatment criterion.     

Effects of enhanced UVB radiation on a marine benthic diatom mat

In order to study the ecophysiological response of a natural diatom-dominated microbial mat to an enhanced level of ultraviolet-B radiation (UVBR), intact sediment cores from a shallow microtidal bay on the Swedish west coast were incubated in an outdoor flow-through system and exposed to either no UVBR or to an artificially enhanced level of UVBR. The microbial mat was cohesive and dominated by the large (≈ 400 μm) motile diatom Gyrosigma balticum. Functional response to UVBR was assessed by measuring carbon fixation (¹⁴C) and allocation of photosynthetic products. Sediment oxygen microprofiles were measured as indicators of the balance between the photosynthetic and respiratory activity of the community. Structural variables included biomass and composition of microalgae, pigment composition (HPLC), content of UV-absorbing compounds, and composition of fatty acids. An enhanced level of UVBR resulted in significant functional changes in the microalgal community. Significantly decreased carbon fixation and lower net oxygen production (as calculated from oxygen microprofiles) suggest that primary productivity in the type of microbial mat studied is potentially sensitive to an enhanced level of UVBR. The pattern of carbon allocation hinted a change as to the photosynthetically active part of the community when exposed to UVBR, indicating a differential sensitivity among algal cells. Despite significant functional effects, pigment or algal composition were not significantly affected by enhanced UVBR. The concentration of UV-absorbing compounds was low and did not increase at UVBR exposure. Due to the short duration of the experiment (4 d), a change in algal biomass or composition was not to be expected. Moreover, the thickness of the sampled sediment layer, may have contributed to the lack of observed structural effects of enhanced UVBR. Visual observations suggested that UVBR affected the motility of G. balticum. This study, as well as other experiments with a similar diatom mat, suggest that vertical migration is a key mechanism to be further studied in relation to UVBR exposure of diatom-dominated microbial mats. Received: 7 September 1996 / Accepted: 8 October 1996     

Immunological methods for food web analysis in a soft-bottom benthic community

Basic to nearly any prediction concerning the behavior and structure of entire communities or of their components is knowledge of trophic connections among species. A major impediment to such understanding of soft-bottom benthos is methodological. Because none of the routinely available methods of food web analysis (e.g. visual gut content analysis, direct observation of feeding, tracer techniques) is generally suitable for examining all trophic interactions of benthic infauna, we rought to evaluate the potential of immunological methods for identifying predatorprey relationships in one typical, estuarine, intertidal sand flat. Whole-organism extracts of individual macro- and meiofaunal taxa were injected into rabbits to produce antisera of varying specificity. Double immunodiffusion precipitin tests of antiserum specificity revealed both phyletic and trophic relationships among 20 taxa. Using relatively unspecific antisera, preliminary analysis of the stomach contents of a few surface deposit-feeders and particle browsers was successful, giving positive identification of several trophic links which would otherwise have gone undetected. The production of taxon-specific antisera is expected to provide the methodological tool necessary to document the breadth of trophic connections in a marine benthic food web.     

非稳态条件下藻类种间非生物资源竞争理论及研究进展

从理论及实验研究的角度论述了藻类非稳态种间资源竞争研究进展。实验室内稳态条件下的竞争理论得到了大量实验验证,由于自然水体处于非稳状态,非稳态条件下藻类种间资源竞争日益成为藻类水华形成机制研究的重点。主要理论包括Tilman的R*法则和资源比假说,Grover的拾遗者-机会主义者交替竞争假说(VIS),中度干扰假说(IDH)及关键光强(CLIH)假说。这些假说的提出丰富了藻类资源竞争的理论体系,为研究自然状态下藻类增殖和竞争行为奠定理论基础。今后的藻类种间资源竞争的研究重点为非稳态下物理条件变化以及物理、化学和生物资源共同变化下藻类种间关系及生物多样性的研究。     

A mathematical model of PCB bioaccumulation in plankton

In batch experiments exposing individual plankton constituents to Aroclor 1254 PCB, the rate at which the organism approaches partitioning equilibrium appears to be partly size-dependent while the extent of PCB accumulation is species-specific. The sorptive desorptive kinetics of PCB in these experiments can be described mathematically by a first-order expression. Employing this expression in a model plankton food web permits examination of the role of feeding and sorptive processes in determining PCB body burden under various environmental conditions. When ingestion rates exceed desorption and excretion rates, a consuming organism accumulates PCB above levels predicted by equilibrium partitioning relationships. Feeding-induced oscillations in PCB body burden could thus obscure the reduction of soluble PCB concentration which determine a “baseline” PCB body burden. Unless referenced to a specific set of biological and environmental conditions, the importance of direct partitioning from water vs. food uptake appears to be a moot topic.