Close and delayed benthic–pelagic coupling in coastal ecosystems: the role of physical constraints

We aimed to analyse the temporal scales of the variability of benthic ammonium flux using data from an estuarine bay (Alfacs Bay, N. W. Mediterranean). Several nitrogen (N) pools in the sediment, their reactivities and their associated fluxes were estimated. Decomposition of labile buried N (4.5 mol N m^-2) was found to cause an ammonium flux of 0.1 mmol N m^-2 d ^-1, referred to as background flux. The fluxes measured from bell-jar incubations were usually higher, between 2 and 6 mmol N m^-2 d ^-1, which reflects mineralization of recent sedimentation. A typical sedimentation event was found to account for 25 – 75 mmol m^-2 of freshly settled N, which should bring on an ammonium flux of about 1.7–5.0 mmol N m^-2 d ^-1, referred to as fast flux. The concordance between measured and computed benthic fluxes is associated with the coupling of benthic fluxes to production and sedimentation. Close benthic–pelagic coupling was observed in winter and early spring, while a delayed flux response to sedimentation, with transient variations of pore water ammonium profiles, showing surface peaks and decreasing concentrations with depth, was found in autumn. Structures, such as viscous layers, which develop over the sediment–water interface, were found to be essential to the regulation of benthic processes and to lead to transient variations of pore water nutrient concentrations and associated fluxes. The temporal scales of the benthic flux response to sedimentation were discussed in terms of the physical structures involved in decomposition (the bulk sediment, the viscous layers or the fresh settled layer), the processes controlling kinetics and diffusion laws. Several scenarios for the benthic- pelagic coupling in Alfacs Bay, in which local (estuarine circulation) and climatic components combine to yield the variability observed, were examined.     

Responses of benthic–pelagic coupling to climate change in a temperate estuary

This article reports the first demonstration of the impact of climate change on benthic–pelagic coupling and the biogeochemical cycles of a coastal marine system. Over the last 30 years Narragansett Bay, a 328-km² temperate estuary on the east coast of the United States, has undergone a variety of ecological changes. Building on a robust data set that spans three decades, we present a link between warming (+1.7°C in annual mean water temperature) in the bay and a marked decrease in sediment oxygen consumption, in the fluxes of ammonium and phosphate from sediments to the overlying water, and in sediment denitrification. We attribute this reduction in biogeochemical exchange to a dramatic drop in the standing crop of water-column chlorophyll as the system has shifted from one characterized by a dominant winter–spring bloom to one supported by more ephemeral and less intense summer–autumn blooms. The recent climate-induced oligotrophication of the bay will be further exacerbated by forthcoming nitrogen reductions due to tertiary sewage treatment. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Plant–animal interactions in suburban environments: implications for floral evolution

Plant interactions with mutualists and antagonists vary remarkably across space, and have played key roles in the ecology and evolution of flowering plants. One dominant form of spatial variation is human modification of the landscape, including urbanization and suburbanization. Our goal was to assess how suburbanization affected plant–animal interactions in Gelsemium sempervirens in the southeastern United States, including interactions with mutualists (pollination) and antagonists (nectar robbing and florivory). Based on differences in plant–animal interactions measured in multiple replicate sites, we then developed predictions for how these differences would affect patterns of natural selection, and we explored the patterns using measurements of floral and defensive traits in the field and in a common garden. We found that Gelsemium growing in suburban sites experienced more robbing and florivory as well as more heterospecific but not conspecific pollen transfer. Floral traits, particularly corolla length and width, influenced the susceptibility of plants to particular interactors. Observational data of floral traits measured in the field and in a common garden provided some supporting but also some conflicting evidence for the hypothesis that floral traits evolved in response to differences in species interactions in suburban vs. wild sites. However, the degree to which plants can respond to any one interactor may be constrained by correlations among floral morphological traits. Taken together, consideration of the broader geographic context in which organisms interact, in both suburban and wild areas, is fundamental to our understanding of the forces that shape contemporary plant–animal interactions and selection pressures in native species.     

The native–invasive balance: implications for nutrient cycling in ecosystems

We conducted single- and mixed-litter experiments in a hardwood forest in Long Island, New York, using leaf litter from phylogenetically paired native and invasive species. We selected long-established, abundant invasive species with wide-ranging distributions in the eastern United States that likely make substantial contributions to the litter pool of invaded areas. Overall, leaf litter from invasive species differed from native litter, though differences varied by phylogenetic grouping. Invasive litter had lower carbon:nitrogen ratios (30.9 ± 1.96 SE vs. 32.8 ± 1.36, P = 0.034) and invasive species lost 0.03 ± 0.007 g of nitrogen and had 23.4 ± 4.9 % of their starting mass remaining at the end of 1 year compared with a loss of 0.02 ± 0.003 g nitrogen and 31.1 ± 2.6 % mass remaining for native species. Mixing litter from two species did not alter decomposition rates when native species were mixed with other native species, or when invasive species were mixed with other invasive species. However, mixing litter of native and invasive species resulted in significantly less mass and nitrogen loss than was seen in unmixed invasive litter. Mixtures of native and invasive litter lost all but 47 ± 2.2 % of initial mass, compared to 37 ± 5.8 % for invasive litter and 50 ± 5.1 % for native litter. This non-additive effect of mixing native and invasive litter suggests that an additive model of metabolic characteristics may not suffice for predicting invasion impacts in a community context, particularly as invasion proceeds over time. Because the more rapid decomposition of invasive litter tends to slow to rates typical of native species when native and invasive litters are mixed together, there may be little impact of invasive species on nutrient cycling early in an invasion, when native leaf litter is abundant (providing litter deposition is the dominant control on nutrient cycling).     

Metazoan meiofauna dynamics and pelagic–benthic coupling in the Southeastern Beaufort Sea,Arctic Ocean

Pelagic–benthic coupling is relatively well studied in the marginal seas of the Arctic Ocean. Responses of meiofauna with regard to seasonal pulses of particulate organic matter are, however, rarely investigated. We examined the dynamics of metazoan meiofauna and assessed the strength of pelagic–benthic coupling in the Southeastern Beaufort Sea, during autumn 2003 and spring–summer 2004. Meiofauna abundance varied largely (range: 2.3 × 10⁵ to 5 × 10⁶ ind m⁻²), both spatially and temporally, and decreased with increasing depth (range: 24–549 m). Total meiofauna biomass exhibited similar temporal as well as spatial patterns as abundance and varied from 25 to 914 mg C m⁻². Significant relationships between sediment photopigments and various representatives of meiofauna in summer and autumn likely indicate the use of sediment phytodetritus as food source for meiofauna. A carbon-based grazing model provided estimates of potential daily ingestion rates ranging from 32 to 723 mg C m⁻². Estimated potential ingestion rates showed that meiofauna consumed from 11 to 477% of the sediment phytodetritus and that meiofauna were likely not food-restricted during spring and autumn. These results show that factors governing the distribution and abundance of metazoan meiofauna need to be better elucidated if we are to estimate the benthic carbon fluxes in marginal seas of the Arctic Ocean. This paper is dedicated to the memory of our dear friend and colleague Gaston Desrosiers who contributed so much to benthic ecology. We will continue in his spirit.     

Biogeochemical responses following coral mass spawning on the Great Barrier Reef: pelagic–benthic coupling

This study quantified how the pulse of organic matter from the release of coral gametes triggered a chain of pelagic and benthic processes during an annual mass spawning event on the Australian Great Barrier Reef. Particulate organic matter (POM) concentrations in reef waters increased by threefold to 11-fold the day after spawning and resulted in a stimulation of pelagic oxygen consumption rates that lasted for at least 1 week. Water column microbial communities degraded the organic carbon of gametes of the broadcast-spawning coral Acropora millepora at a rate of >15% h⁻¹, which is about three times faster than the degradation rate measured for larvae of the brooding coral Stylophora pistillata. Stable isotope signatures of POM in the water column reflected the fast transfer of organic matter from coral gametes into higher levels of the food chain, and the amount of POM reaching the seafloor immediately increased after coral spawning and then tailed-off in the next 2 weeks. Short-lasting phytoplankton blooms developed within a few days after the spawning event, indicating a prompt recycling of nutrients released through the degradation of spawning products. These data show the profound effects of coral mass spawning on the reef community and demonstrate the tight recycling of nutrients in this oligotrophic ecosystem.     

A method for measuring depth-integrated community metabolism in experimental planktonic–benthic ecosystems

We developed incubation chambers of clear acrylic extending the full depth of the water column to measure in situ depth-integrated community metabolism in experimental planktonic–benthic ecosystems. A magnetic stirring apparatus at mid-depth in each chamber prevented stratification and maintained a vertical turnover time that roughly matched turnover in the experimental ecosystems. We used the chambers in light–dark experiments to partition the depth-integrated photosynthesis and respiration occurring in the experimental ecosystems among water column, wall periphyton, and benthic communities. Community metabolism measured with the chambers was positively related to the chlorophyll-a concentrations of the water column and wall periphyton communities. We also found close agreement between the summation of partitioned community metabolism measured in the chambers and an independent measure of total ecosystem metabolism in the experimental ecosystems. Incubations in chambers and in light–dark bottles revealed similar patterns of water column and wall periphyton community metabolism, but magnitudes differed. In addition to application in experimental ecosystems, this incubation chamber technique may provide an efficient and realistic measure of water column and benthic metabolism and nutrient uptake in lakes and estuaries.     

Upwelling-like bottom intrusion enhances the pelagic–benthic coupling by a fish predator in a coastal food web

Upwelling regions where nutrients are transported from deep to surface waters are among the most productive in the oceans. Although it is well known that the upwelling affects fishery production through bottom-up trophic cascading, it remains unexplored how temporal variation in its intensity alters overall trophic energy flows within a focal food web. In the present study, we demonstrate that inter-annual variation in the intensity of upwelling-like bottom intrusion alters food web properties in coastal waters of the Uwa Sea by focusing on the levels of δ¹³C and δ¹⁵N for a demersal fish predator, Acropoma japonicum. This approach integrates information on prey–predator interactions. In the season following a stratification period when pelagic productivity is limited by nutrient availability, A. japonicum showed lower levels of δ¹³C in years with high bottom intrusion intensity than in those with low intensity. One possible cause for this isotopic depletion is that the bottom intrusion-induced nutrient supply enhances pelagic productivity and consequently facilitates a foraging shift by A. japonicum from ordinary benthic prey to supplementary pelagic prey with a lower δ¹³C. In conclusion, the increased intensity of bottom intrusion results in coupling of two major trophic energy flows, pelagic and benthic food chains, through the demersal predator’s foraging shift.     

Patterns in benthic food webs: a role for omnivorous crayfish?

1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.     

Temperature effects on herbivory for an Indo-Pacific parrotfish in Panamá: implications for coral–algal competition

An assumption that a positive relationship exists between temperature and herbivorous fish grazing rates in coral reef environments has been used to explain seasonal and latitudinal trends in herbivore pressure. However, this assumption has not been systematically quantified over short-term (hours–days) changes in temperature, avoiding the confounding influences that can occur on seasonal or latitudinal scales. This study measured grazing activities of the pan Indo-Pacific parrotfish Scarus ghobban over short-term changes in temperature in upwelling and non-upwelling environments on the Pacific coast of Panamá. Individual juvenile fish were followed over naturally varying temperatures to determine their bite rates (bites min⁻¹), the foray frequency (forays min⁻¹) and bites per foray. In the upwelling environment, there was a significant positive correlation between temperature and bite rate (R _Partial = 0.63, P < 0.0001) and there was a marked fourfold change in bite rates over the range of temperatures encountered in the study (21.2–29.4°C). Bites per foray were also positively correlated to temperature (R _partial = 0.27, P < 0.0001), and tide height (R _partial = 0.26, P < 0.001), whilst foray frequency was positively correlated to temperature (R _partial = 0.63, P < 0.0001), but negatively to tide height (R _partial = −0.31, P < 0.0001). The effect of temperature on grazing rates may explain differences in herbivore pressure across different thermal environments and may be a factor contributing to algal biomass increases in low-temperature coral reef environments.     

Boundary forcing-induced oscillations in a phytoplankton–zooplankton patch

We consider one- and two-dimensional minimal models of a phytoplankton–zooplankton dynamical system subject to boundary forced oscillations in the prey. As a consequence, waves propagate in the space dimensions in both species. The difference between the peaks and the troughs are at times of orders of magnitude.     

Evidence for the participation of calmodulin in stimulus–secretion coupling in the pancreatic β-cell

1. The ability of a range of phenothiazines to inhibit activation of brain phosphodiesterase by purified calmodulin was studied. Trifluoperazine, prochlorperazine and 8-hydroxyprochlorperazine produced equipotent dose-dependent inhibition with half-maximum inhibition at 12mum. When tested at 10 or 50mum, 7-hydroxyprochlorperazine was a similarly potent inhibitor. However, trifluoperazine-5-oxide and N-methyl-2-(trifluoromethyl)phenothiazine were ineffective at concentrations up to 50mum, and produced only a modest inhibition at 100mum. 2. The same phenothiazines were tested for their ability to inhibit activation of brain phosphodiesterase by boiled extracts of rat islets of Langerhans. At a concentration of 20mum, 70-80% inhibition was observed with trifluoperazine, prochlorperazine, 7-hydroxyprochlorperazine or 8-hydroxyprochlorperazine, whereas trifluoperazine-5-oxide and N-methyl-2-(trifluoromethyl)phenothiazine were less effective. 3. The effect of these phenothiazines on insulin release from pancreatic islets was studied in batch-type incubations. Insulin release stimulated by glucose (20mm) was markedly inhibited by 10mum-trifluoperazine or -prochlorperazine and further inhibited at a concentration of 20mum. 8-Hydroxyprochlorperazine (20mum) was also a potent inhibitor but 7-hydroxyprochlorperazine (20mum) elicited only a modest inhibition of glucose-stimulated insulin release; no inhibition was observed with trifluoperazine-5-oxide or N-methyl-2-(trifluoromethyl)phenothiazine. 4. Trifluoperazine (20mum) markedly inhibited insulin release stimulated by leucine or 4-methyl-2-oxopentanoate in the absence of glucose, and both trifluoperazine and prochlorperazine (20mum) decreased insulin release stimulated by glibenclamide in the presence of 3.3mm-glucose. 5. None of the phenothiazines affected basal insulin release in the presence of 2mm-glucose. 6. Trifluoperazine (20mum) did not inhibit islet glucose utilization nor the incorporation of [(3)H]leucine into (pro)insulin or total islet protein. 7. Islet extracts catalysed the incorporation of (32)P from [gamma-(32)P]ATP into endogenous protein substrates. Sodium dodecyl sulphate/polyacrylamide-gel electrophoresis resolved several phosphorylated bands, but incorporation was slight. However, calmodulin in the presence of Ca(2+) greatly enhanced incorporation: the predominant phosphorylated band had an estimated mol.wt. of 55000. This enhanced incorporation was abolished by trifluoperazine, but not by cyclic AMP-dependent protein kinase inhibitor protein. 8. These results suggest that islet phosphodiesterase-stimulating activity is similar to, although not necessarily identical with, calmodulin from skeletal muscle; that islet calmodulin may play an important role in Ca(2+)-dependent stimulus-secretion coupling in the beta-cell; and that calmodulin may exert part at least of its effect on secretion via phosphorylation of endogenous islet proteins.     

Transcomplementation and synergism in plants: implications for viral transgenes?

In plants, viral synergisms occur when one virus enhances infection by a distinct or unrelated virus. Such synergisms may be unidirectional or mutualistic but, in either case, synergism implies that protein(s) from one virus can enhance infection by another. A mechanistically related phenomenon is transcomplementation, in which a viral protein, usually expressed from a transgene, enhances or supports the infection of a virus from a distinct species. To gain an insight into the characteristics and limitations of these helper functions of individual viral genes, and to assess their effects on the plant-pathogen relationship, reports of successful synergism and transcomplementation were compiled from the peer-reviewed literature and combined with data from successful viral gene exchange experiments. Results from these experiments were tabulated to highlight the phylogenetic relationship between the helper and dependent viruses and, where possible, to identify the protein responsible for the altered infection process. The analysis of more than 150 publications, each containing one or more reports of successful exchanges, transcomplementation or synergism, revealed the following: (i) diverse viral traits can be enhanced by synergism and transcomplementation; these include the expansion of host range, acquisition of mechanical transmission, enhanced specific infectivity, enhanced cell-to-cell and long-distance movement, elevated or novel vector transmission, elevated viral titre and enhanced seed transmission; (ii) transcomplementation and synergism are mediated by many viral proteins, including inhibitors of gene silencing, replicases, coat proteins and movement proteins; (iii) although more frequent between closely related viruses, transcomplementation and synergism can occur between viruses that are phylogenetically highly divergent. As indicators of the interoperability of viral genes, these results are of general interest, but they can also be applied to the risk assessment of transgenic crops expressing viral proteins. In particular, they can contribute to the identification of potential hazards, and can be used to identify data gaps and limitations in predicting the likelihood of transgene-mediated transcomplementation.     

The ecological role of moss in a polar desert: implications for aboveground–belowground and terrestrial–aquatic linkages

The McMurdo Dry Valleys of Antarctica is one of the coldest and driest habitats on the planet. As vascular plants are absent in this region, moss is the main form of aboveground primary production with a potentially important contribution to biogeochemical cycling, yet little is known about their ecological role. To determine the relationship between moss and soil properties relevant to biogeochemistry, we sampled both from a variety of locations in the Dry Valleys. Moss presence was compared to soil properties, and we measured the plasticity of moss stoichiometry (C:N:P) across gradients in nutrient availability. Results demonstrate that many soil properties significantly differed with moss presence, particularly conductivity and pH, but there is no strong evidence that this is caused by the moss presence and not the conditions inherent to the microsites where moss was found. There is great variability in moss stoichiometry, with some significant differences between sites, but generally variability within sites is larger than variation among sites. Results suggest that the main source of moss nutrients is from the soil, rather than water, but correlations with any one nutrient source are weak, suggesting a great deal of plasticity in moss stoichiometry and nutrient uptake.     

Perception–action coupling model for human locomotor pointing

How do humans achieve the precise positioning of the feet during walking, for example, to reach the first step of a stairway? We addressed this question at the visuomotor integration level. Based on the optical specification of the required adaptation, a dynamical system model of the visuomotor control of human locomotor pointing was devised for the positioning of a foot on a visible target on the floor during walking. Visuomotor integration consists of directly linking optical information to a motor command that specifically modulates step length in accordance with the ongoing dynamics of locomotor pattern generation. The adaptation of locomotion emerges from a perception-action coupling type of control based on temporal information rather than on feedforward planning of movements. The proposed model reproduces experimental results obtained for human locomotor pointing.     

Quantitative structure–activity relationships in enzymatic single-electron reduction of nitroaromatic explosives: implications for their cytotoxicity

The mechanisms of cytotoxicity of polynitroaromatic explosives, an important group of environmental pollutants, remain insufficiently studied so far. We have found that the rate constants of single-electron enzymatic reduction, and the enthalpies of single-electron reduction of nitroaromatic compounds (ΔHf(ArNO₂^−⋅)), obtained by quantum mechanical calculation, may serve as useful tools for the analysis of cytotoxicity of nitroaromatic explosives with respect to the possible involvement of oxidative stress. The single-electron reduction rate constants of a number of explosives including 2,4,6-trinitrotoluene (TNT) and 2,4,6-trinitrophenyl-N-methylnitramine (tetryl), and model nitroaromatic compounds by ferredoxin:NADP⁺ reductase (FNR, EC 1.18.1.2) and NADPH:cytochrome P-450 reductase (P-450R, EC 1.6.2.4) increased with a decrease in ΔHf(ArNO₂^−⋅). This indicates that the reduction rates are determined by the electron transfer energetics, but not by the particular structure of the explosives. The cytotoxicity of explosives to bovine leukemia virus-transformed lamb kidney fibroblasts (line FLK) increased with a corresponding increase in their reduction rate constant by P-450R and FNR, or with a decrease in their ΔHf(ArNO₂^−⋅). This points to an importance of oxidative stress in the toxicity of explosives in this cell line, which was further evidenced by the protective effects of desferrioxamine and the antioxidant N,N′-diphenyl-p-phenylene diamine, and an increase in lipid peroxidation. DT-diaphorase (EC 1.6.99.2) exerted a minor and equivocal role in the cytotoxicity of explosives to FLK cells.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.