Multiple functions increase the importance of biodiversity for overall ecosystem functioning

Biodiversity is proposed to be important for the rate of ecosystem functions. Most biodiversity-ecosystem function studies, however, consider only one response variable at a time, and even when multiple variables are examined they are analyzed separately. This means that a very important aspect of biodiversity is overlooked: the possibility for different species to carry out different functions at any one time. We propose a conceptual model to explore the effects of species loss on overall ecosystem functioning, where overall functioning is defined as the joint effect of many ecosystem functions. We show that, due to multifunctional complementarity among species, overall functioning is more susceptible to species loss than are single functions. Modeled relationships between species richness and overall ecosystem functioning using five empirical data sets on monocultures reflected the range of effects of species loss on multiple functions predicted by the model. Furthermore, an exploration of the correlations across functions and the degree of redundancy within functions revealed that multifunctional redundancy was generally lower than single-function redundancy in these empirical data sets. We suggest that by shifting the focus to the variety of functions maintained by a diversity of species, the full importance of biodiversity for the functioning of ecosystems can be uncovered. Our results are thus important for conservation and management of biota and ecosystem services.     

Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship?

The relationship between plant diversity and productivity has largely been attributed to niche complementarity, assuming that plant species are complementary in their resource use. In this context, we conducted an 15N field study in three different grasslands, testing complementarity nitrogen (N) uptake patterns in terms of space, time, and chemical form as well as N strategies such as soil N use, symbiotic N fixation, or internal N recycling for different plant species. The relative contribution of different spatial, temporal, and chemical soil N pools to total soil N uptake of plants varied significantly among the investigated plant species, within and across functional groups. This suggests that plants occupy distinct niches with respect to their relative N uptake. However, when the absolute N uptake from the different soil N pools was analyzed, no spatial, temporal, or chemical variability was detected, but plants, and in particular functional groups, differed significantly with respect to their total soil N uptake irrespective of treatment. Consequently, our data suggest that absolute N exploitation on the ecosystem level is determined by species or functional group identity and thus by community composition rather than by complementary biodiversity effects. Across functional groups, total N uptake from the soil was negatively correlated with leaf N concentrations, suggesting that these functional groups follow different N use strategies to meet their N demands. While our findings give no evidence for a biodiversity effect on the quantitative exploitation of different soil N pools, there is evidence for different and complementary N strategies and thus a potentially beneficial effect of functional group diversity on ecosystem functioning.     

Herbivores, tidal elevation, and species richness simultaneously mediate nitrate uptake by seaweed assemblages

In order for research into the consequences of biodiversity changes to be more applicable to real-world ecosystems, experiments must be conducted in the field, where a variety of factors other than diversity can affect the rates of key biogeochemical and physiological processes. Here, we experimentally evaluate the effects of two factors known to affect the diversity and composition of intertidal seaweed assemblages--tidal elevation and herbivory--on nitrate uptake by those assemblages. Based on surveys of community composition at the end of a 1.5-year press experiment, we found that both tide height and herbivores affected seaweed community structure. Not surprisingly, seaweed species richness was greater at lower tidal elevations. Herbivores did not affect richness, but they altered the types of species that were present; seaweed species characterized by higher rates of nitrate uptake were more abundant in herbivore-removal plots. Both tide height and herbivores affected nitrate uptake by seaweed assemblages. Individual seaweed species, as well as entire seaweed assemblages, living higher on the shore had greater rates of biomass-specific nitrate uptake, particularly at high ambient nitrate concentrations. Grazed seaweed assemblages exhibited reduced nitrate uptake, but only at low nitrate concentrations. We evaluated the effect of seaweed richness on nitrate uptake, both alone and after accounting for effects of tidal elevation and herbivores. When only richness was considered, we found no effect on uptake. However, when simultaneous effects of richness, tide height, and herbivores on uptake were evaluated, we found that all three had relatively large and comparable effects on nitrate uptake coefficients and that there was a negative relationship between seaweed richness and nitrate uptake. Particularly because effects of richness on uptake were not apparent unless the effects of tide height and herbivory were also considered, these results highlight the importance of considering the effects of environmental context when evaluating the consequences of biodiversity change in more realistic systems.     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Short-term interaction between nitrate and ammonium uptake inThalassiosira pseudonana: Effect of preconditioning nitrogen source and growth rate

The effect of preconditioning nitrogen source and growth rate on the interaction between nitrate and ammonium uptake was determined inThalassiosira pseudonana (Clone 3H). A new method, using cells on a filter (Parslow et al. 1985), allowed continuous measurement of uptake from 0.5 to 9 min after the addition of nitrate, ammonium, or both, with no variation in concentration during the course of the experiment. For each preconditioning N source and growth rate, a series of uptake experiments was conducted, including controls with only nitrate or only ammonium, and others with different combinations of concentrations of nitrate and ammonium. For the first time, preference for ammonium was separated from inhibition of nitrate uptake by ammonium. Ammonium was the preferred N source, i.e. if nitrate and ammonium were presented separately, ammonium uptake rates exceeded nitrate uptake rates. Preference for ammonium varied with both preconditioning N source and growth rate. Inhibition of nitrate uptake by ammonium, determined by comparing nitrate uptake in the presence and absence of ammonium, was observed at ammonium concentrations > 1µM, but was rarely complete. Inhibition of nitrate uptake by ammonium was less in the ammonium-limited culture than in the cultures growing on nitrate, but invariant with growth rate in the nitrate-grown cultures. Below 1µM ammonium, nitrate uptake was often stimulated and rates exceeded those in the controls without ammonium. Ammonium uptake was not inhibited by the presence of nitrate.T. pseudonana fits the classical view of the interaction between nitrate and ammonium uptake in some respects, such as preference for ammonium, and inhibition of nitrate uptake by ammonium concentrations > 1µM. However, at ammonium concentrations typical of most marine environments, nitrate uptake occurs at rapid rates. In other respects, N uptake inT. pseudonana deviates from the classical view in the following ways: (1) stimulation of nitrate uptake by low concentrations of ammonium; (2) lack of inhibition of nitrate uptake by ammonium at low nitrate concentrations; and (3) variation in preference and inhibition with preconditioning, which is markedly different for other species. Because of the apparent enormous species variation in the interaction between nitrate and ammonium uptake and the lack of detailed information for a variety of species, it is difficult to generalize about the effect of ammonium on nitrate uptake, especially in the field, where prior N availability and species composition are not usually addressed.     

CO2 enrichment inhibits shoot nitrate assimilation in C3 but not C4 plants and slows growth under nitrate in C3 plants

The CO2 concentration in Earth's atmosphere may double during this century. Plant responses to such an increase depend strongly on their nitrogen status, but the reasons have been uncertain. Here, we assessed shoot nitrate assimilation into amino acids via the shift in shoot CO2 and O2 fluxes when plants received nitrate instead of ammonium as a nitrogen source (deltaAQ). Shoot nitrate assimilation became negligible with increasing CO2 in a taxonomically diverse group of eight C3 plant species, was relatively insensitive to CO2 in three C4 species, and showed an intermediate sensitivity in two C3-C4 intermediate species. We then examined the influence of CO2 level and ammonium vs. nitrate nutrition on growth, assessed in terms of changes in fresh mass, of several C3 species and a Crassulacean acid metabolism (CAM) species. Elevated CO2 (720 micromol CO2/mol of all gases present) stimulated growth or had no effect in the five C3 species tested when they received ammonium as a nitrogen source but inhibited growth or had no effect if they received nitrate. Under nitrate, two C3 species grew faster at sub-ambient (approximately 310 micromol/mol) than elevated CO2. A CAM species grew faster at ambient than elevated or sub-ambient CO2 under either ammonium or nitrate nutrition. This study establishes that CO2 enrichment inhibits shoot nitrate assimilation in a wide variety of C3 plants and that this phenomenon can have a profound effect on their growth. This indicates that shoot nitrate assimilation provides an important contribution to the nitrate assimilation of an entire C3 plant. Thus, rising CO2 and its effects on shoot nitrate assimilation may influence the distribution of C3 plant species.     

Synergistic effects of nitrate and ammonium ion on the growth and uptake kinetics of Monochrysis lutheri in continuous culture

Continuous-culture results for Monochrysis lutheri grown on 12 h light-12 h dark cycles with a spectrum of ratios of nitrate and ammonium serving as limiting nutrients are compared with continuous light, exclusively nitrate, and exclusively ammonium-limited data for this species. The diel effects of the light regime on the maximum specific uptake rate are examined for both nitrate and ammonium. Synergistic effects on uptake by various initial concentrations of these two nutrients are presented. Preconditioning with light-dark cycles did not affect maximum uptake rate, but preconditioning on a combination of nitrate and ammonium gave much lower uptake rates than those observed for populations preconditioned on either nutrient exclusively. The implications of high maximum specific-uptake rates compared to maximum specific-growth rates in terms of the range of nitrate and ammonium ion concentrations associated with nutrient limitation are reviewed.Hawaii Institute of Marine Biology Contribution No. 477.     

不同氮肥对油菜B与Mo吸收的影响

利用盆栽试验研究了潮土中不同氮肥对油菜硼钼吸收的影响，结果表明，前期(5周)施氮并没有促进油菜的生长，而且大部分氮肥对油菜生长表现抑制作用，氯化铵处理的作用最大，硫酸铵和尿素处理次之，硝酸铵处理再次之；不同氮肥均有抑制油菜生长前期(5周)硼吸收和累积的作用，其中以氯化铵、硝酸铵和尿素的作用为甚；不同氮肥同样抑制油菜生长前期(5周)对钼的吸收和累积，以硫酸铵处理作用最突出，氯化铵处理次之。但随着时间的推移，10、15周不同氮肥大多促进油菜的生长，增加油菜对硼钼的吸收和累积，且各氮肥处理间的差异逐步减小。     

Interactions between nitrate and ammonium uptake: variation with growth rate,nitrogen source and species

The interaction between nitrate and ammonium uptake was examined as a function of preconditioning growth rate and nitrogen source by adding nitrate, ammonium, or both to nitrogen-sufficient,-deficient, and-starvedSkeletonema costatum (Grev.) Cleve and nitrogen-deficientChaetoceros debilis Cleve. By simultaneously measuring the internal accumulation of intermediates of nitrogen assimilation and the rates of nitrogen assimilation, the metabolic control of nitrogen uptake could be assessed. After the simultaneous addition of nitrate and ammonium to culture, both nitrate and ammonium uptake rates were decreased in comparison with the rates observed when each was added alone, although nitrate uptake was usually decreased more than ammonium uptake. Since both nitrate and ammonium uptake rates vary with time, preconditioning growth conditions, nitrogen sources present, and species, it was necessary to use several different indices to quantify inhibition. In general, ammonium inhibition of nitrate uptake inS. costatum was greatest in cultures preconditioned to ammonium and those at low growth rates, whereas ammonium uptake was inhibited most in cultures preconditioned to nitrate. In nitrogen-deficientC. debilis, nitrate uptake was more inhibited by ammonium, but uptake returned to normal rates more quickly than inS. costatum, whereas inhibition of ammonium uptake was similar. These results explain why the interaction between nitrate and ammonium uptake in the field can be so variable. Inhibition of uptake is not controlled by internal ammonium or total amino acids, nor is it related to the inability to reduce nitrate. Instead, inhibition must be determined in part by the external concentration of nitrogen compounds and in part by some intermediate(s) of nitrogen assimilation present inside the cell.Bigelow Laboratory Contribution No. 82022     

Incorporating Surrogate Species and Seascape Connectivity to Improve Marine Conservation Outcomes

Conservation focuses on maintaining biodiversity and ecosystem functioning, but gaps in our knowledge of species biology and ecological processes often impede progress. For this reason, focal species and habitats are used as surrogates for multispecies conservation, but species‐based approaches are not widely adopted in marine ecosystems. Reserves in the Solomon Islands were designed on the basis of local ecological knowledge to conserve bumphead parrotfish (Bolbometopon muricatum) and to protect food security and ecosystem functioning. Bumphead parrotfish are an iconic threatened species and may be a useful surrogate for multispecies conservation. They move across tropical seascapes throughout their life history, in a pattern of habitat use that is shared with many other species. We examined their value as a conservation surrogate and assessed the importance of seascape connectivity (i.e., the physical connectedness of patches in the seascape) among reefs, mangroves, and seagrass to marine reserve performance. Reserves were designed for bumphead parrotfish, but also enhanced the abundance of other species. Integration of local ecological knowledge and seascape connectivity enhanced the abundance of 17 other harvested fish species in local reserves. This result has important implications for ecosystem functioning and local villagers because many of these species perform important ecological processes and provide the foundation for extensive subsistence fisheries. Our findings suggest greater success in maintaining and restoring marine ecosystems may be achieved when they are managed to conserve surrogate species and preserve functional seascape connections. Incorporación de Especies Sustitutas y de Conectividad Marina para Mejorar los Resultados de Conservación     

Rare Species and Ecosystem Functioning

Abstract:  The role of diversity in the maintenance of ecosystems has been studied widely in the past decade. By correlating richness and diversity with basic ecosystem processes, these investigations lend support to the hypothesis that species diversity significantly influences ecosystem functioning and, in turn, provide support for the conservation of biodiversity. Nonetheless, the majority of these investigations demonstrate that conservation of a relatively small number of generally dominant species is sufficient to maintain most processes. Indeed, there is remarkably little evidence to support the contention that less common species, those likely of highest conservation concern, are important in the maintenance of ecosystem functioning. Here we summarize studies, most employing alternative methodological strategies, wherein less common and rare species are demonstrated to make significant contributions to ecosystem functioning. Evidence exists among studies of keystone species, aggregate effects of less common species, and species turnover. Our findings suggest that (1) less common species can make significant ecosystem contributions; (2) further investigation into the effects of rare and less common species on ecosystem maintenance is sorely needed; (3) further investigation should embrace a variety of approaches; and (4) until further research is conducted a prudent conservation approach is warranted wherein the contribution of less common species to ecosystem functioning is assumed.     

Negative selection effects suppress relationships between bacterial diversity and ecosystem functioning

I assembled bacterial communities to explore the effects of bacterial diversity on multiple ecosystem functions, including bacterial community biovolume, decomposition of particulate organic matter, and biomass transfer to the next trophic level. The experiment used a two-way factorial design with four levels of bacterial diversity (one to four species) and the absence/presence of a bacterivorous ciliated protist Tetrahymena pyriformis as two main factors, and all possible combinations of the four bacterial taxa nested within each diversity level. Bacterial community biovolume increased as bacterial diversity increased, a result due largely to positive selection effects. Decomposition and consumer abundance, however, were unaffected by bacterial diversity, though both varied among bacterial composition treatments. Negative selection effects, the dominance of species that do not contribute significantly to ecosystem functioning, accounted for the lack of diversity effects on decomposition and consumer abundance. The presence of Tetrahymena reduced bacterial community biovolume but increased decomposition, without altering the diversity-functioning relationships. Decomposition was strongly linked with consumer abundance such that communities supporting larger consumer biomass exhibited higher decomposition rates. This study suggests that if the negative selection effect is common, as it might be when examining ecosystem variables other than biomass (due to the presence of keystone species that can contribute disproportionably to ecosystem functioning relative to their abundances), basic bacteria-mediated ecosystem processes, such as decomposition and energy transfer to the next trophic level, may not increase with bacterial diversity.     

Relating effect and response traits in submersed aquatic macrophytes.

Reliably predicting the consequences of short- or long-term changes in the environment is important as anthropogenic pressures are increasingly stressing the world's ecosystems. One approach is to examine the manner in which biota respond to changes in the environment ("response traits") and how biota, in turn, affect ecosystem processes ("effect traits"). I compared the response and effect traits of four submersed aquatic macrophytes to understand how water level management may affect wetland plant populations and ecosystem processes. I measured resource properties (nutrients in sediment and water), non-resource properties (pH, alkalinity, sediment temperature, oxygen production), and biotic properties (periphyton biomass) in replicated outdoor monocultures of Stuckenia pectinata, Potamogeton nodosus, P. crispus, and Zannichellia palustris. After seven weeks, three of eight replicates of each species treatment were subjected to a temporary water draw-down that desiccated aboveground plant parts. The four species differed in their effects on ecosystem properties associated with nutrient uptake and photosynthetic activity. Shoot growth rate was negatively correlated with light transmittance to the sediment surface whereas root growth rate and root:shoot ratio were correlated with a species' ability to deplete nutrients in sediment interstitial water. Occupation of space in the water column was correlated with water alkalinity and pH and with sediment temperature. Root growth rate was related simultaneously to species effects on sediment nutrient dynamics and recovery of ecosystem properties after water draw-down. This suggests that this morphological trait may be used to predict the effects of environmental change on ecosystem functioning within the context of water level management. Expanding these analyses to more species, different environmental stressors, and across aquatic and terrestrial ecosystems should enhance predictions of the complex effects of global environmental change on ecosystem functioning.     

The rapid response of the marine diatom Skeletonema costatum to changes in external and internal nutrient concentration

A nitrogen-deficient batch culture of the marine diatom Skeletonema costatum, when resupplied with a mixture of nitrate and ammonium, showed an initial enhanced nitrate uptake rate leading to a large internal concentration (pool) of nitrate. Following this initial nitrate uptake event, nitrate uptake ceased, and nitrate assimilation was inhibited until the ammonium present was used. At this point, nitrate uptake resumed and nitrate assimilation began. No internal ammonium pool was observed during nitrate utilization, but a large nitrate pool remained throughout the utilization of external nitrate. The internal nitrate pool decreased rapidly after exhaustion of nitrate from the culture medium, but growth of cellular particulate nitrogen continued for about 24 h. A mathematical simulation model was developed from these data. The model cell consisted of a nitrate pool, ammonium pool, dissolved organic nitrogen pool, and particulate nitrogen. It was found that simple Michaelis-Menten functions for uptake and assimilation gave inadequate fit to the data. Michaelis-Menten functions were modified by inclusion of inhibitory and stimulatory feedback from the internal pools to more accurately represent the observed nutrient utilization.     

Incubation time, functional litter diversity, and habitat characteristics predict litter-mixing effects on decomposition

Plant diversity influences many fundamental ecosystem functions, including carbon and nutrient dynamics, during litter breakdown. Mixing different litter species causes litter mixtures to lose mass at different rates than expected from component species incubated in isolation. Such nonadditive litter-mixing effects on breakdown processes often occur idiosyncratically because their direction and magnitude change with incubation time, litter species composition, and ecosystem characteristics. Taking advantage of results from 18 litter mixture experiments in streams, we examined whether the direction and magnitude of nonadditive mixing effects are randomly determined. Across 171 tested litter mixtures and 510 incubation time-by-mixture combinations, nonadditive effects on breakdown were common and on average resulted in slightly faster decomposition than expected. In addition, we found that the magnitude of nonadditive effects and the relative balance of positive and negative responses in mixtures change predictably over time, and both were related to an index of functional litter diversity and selected environmental characteristics. Based on these, it should be expected that nonadditive effects are stronger for litter mixtures made of functionally dissimilar species especially in smaller streams. Our findings demonstrate that effects of litter diversity on plant mixture breakdown are more predictable than generally thought. We further argue that the consequences of current worldwide homogenization in the composition of plant traits on carbon and nutrient dynamics could be better inferred from long-duration experiments that manipulate both functional litter diversity and ecosystem characteristics in "hotspots of biodiversity effects," such as small streams.     

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.     

Species phylogenetic relatedness, priority effects, and ecosystem functioning

Species immigration history can structure ecological communities through priority effects, which are often mediated by competition. As competition tends to be stronger between species with more similar niches, we hypothesize that species phylogenetic relatedness, under niche conservatism, may be a reasonable surrogate of niche similarity between species, and thus influence the strength of priority effects. We tested this hypothesis using a laboratory microcosm experiment in which we established bacterial species pools with different levels of phylogenetic relatedness and manipulated the immigration history of species from each pool into microcosms. Our results showed that strong priority effects, and hence multiple community states, only emerged for the species pool with the greatest phylogenetic relatedness. Community assembly also resulted in a significant positive relationship between bacterial phylogenetic diversity and ecosystem functions. Interestingly, these results emerged despite a lack of phylogenetic conservatism for most of the bacterial functional traits considered. Our results highlight the utility of phylogenetic information for understanding the structure and functioning of ecological communities, even when phylogenetically conserved functional traits are not identified or measured.     

From individuals to ecosystem function: toward an integration of evolutionary and ecosystem ecology

An important goal in ecology is developing general theory on how the species composition of ecosystems is related to ecosystem properties and functions. Progress on this front is limited partly because of the need to identify mechanisms controlling functions that are common to a wide range of ecosystem types. We propose that one general mechanism, rooted in the evolutionary ecology of all species, is adaptive foraging behavior in response to predation risk. To support our claim, we present two kinds of empirical evidence from plant-based and detritus-based food chains of terrestrial and aquatic ecosystems. The first kind comes from experiments that explicitly trace how adaptive foraging influences ecosystem properties and functions. The second kind comes from a synthesis of studies that individually examine complementary components of particular ecosystems that together provide an integrated perspective on the link between adaptive foraging and ecosystem function. We show that the indirect effects of predators on plant diversity, plant productivity, nutrient cycling, trophic transfer efficiencies, and energy flux caused by consumer foraging shifts in response to risk are qualitatively different from effects caused by reductions in prey density due to direct predation. We argue that a perspective of ecosystem function that considers effects of consumer behavior in response to predation risk will broaden our capacity to explain the range of outcomes and contingencies in trophic control of ecosystems. This perspective also provides an operational way to integrate evolutionary and ecosystem ecology, which is an important challenge in ecology.     

Type of disturbance and ecological history determine structural stability.

This study aims to reveal whether complexity, namely, community and trophic structure, of chronically stressed soil systems is at increased risk or remains stable when confronted with a subsequent disturbance. Therefore, we focused on a grassland with a history of four centuries of patchy contamination. Nematodes were used as model organisms because they are an abundant and trophically diverse group and representative of the soil food web and ecosystem complexity. In a field survey, a relationship between contaminants and community structures was established. Following, two groups of soil mesocosms from the field that differed in contamination level were exposed to different disturbance regimes, namely, to the contaminant zinc and a heat shock. The zinc treatment revealed that community structure is stable, irrespective of soil contamination levels. This implies that centuries of exposure to contamination led to adaptation of the soil nematode community irrespective of the patchy distribution of contaminants. In contrast, the heat shock had adverse effects on species richness in the highly contaminated soils only. The total nematode biomass was lower in the highly contaminated field samples; however, the biomass was not affected by zinc and heat treatments of the mesocosms. This means that density compensation occurred rapidly, i.e., tolerant species quickly replaced sensitive species. Our results support the hypothesis that the history of contamination and the type of disturbance determine the response of communities. Despite that ecosystems may be exposed for centuries to contamination and communities show adaptation, biodiversity in highly contaminated sites is at increased risk when exposed to a different disturbance regime. We discuss how the loss of higher trophic levels from the entire system, such as represented by carnivorous nematodes after the heat shock, accompanied by local biodiversity loss at highly contaminated sites, may result in detrimental effects on ecosystem functions.     

Nitrogen uptake kinetics in three year-classes of Laminaria groenlandica (Laminariales: Phaeophyta)

Nitrate and ammonium uptake rates were measured for three year-classes of the perennial macrophyte Laminaria groenlandica Rosenvinge, collected from nitrogen-depleted waters in Barkley Sound, British Columbia, Canada, in summer 1981. A time course of uptake rate revealed that ammonium uptake was high during the first hour and then decreased for all three year-classes; the opposite pattern was exhibited for the time course of nitrate uptake rate. Nitrate uptake rate increased linearly with nitrate concentration up to the highest level tested (60 M). The nitrate uptake rate of first-year plants was three times higher than second- and third-year plants; ammonium uptake rates showed similar patterns to those for nitrate. The interaction between nitrate and ammonium was examined for first-year plants. Nitrate and ammonium were taken up simultaneously and uptake rates were identical and equal to uptake rates when only nitrate or ammonium was present in the medium. Therefore, first-year plants are able to take up twice as much inorganic nitrogen per unit time when both nitrate and ammonium are present. First-year plants showed significant diel periodicity in ammonium uptake rates, whereas second- and third-year plants showed no periodicity in nitrate or ammonium uptake rates.