Tipping point arises earlier under a multiple-stressor scenario.

Anthropogenic impacts and global changes have profound implications for natural ecosystems and may lead to their modification, degradation or collapse. Increases in the intensity of single stressors may create abrupt shifts in biotic responses (i.e. thresholds). The effects of multiple interacting stressors may create non-additive responses, known as synergistic or antagonistic interactions. Here we combine both concepts-ecological thresholds and interactions between multiple stressors-to understand the effects of multiple interacting stressors along environmental gradients, and how this can affect the occurrence of thresholds. Using an experimental approach to investigate the effect of nutrient enrichment and saltwater intrusion on mortality in the zebra mussel, Dreissena polymorpha, we show that multiple stressors can create thresholds at lower levels of an environmental gradient. Our results reveal a major shortcoming in how we currently investigate these two ecological concepts, as considering them separately may be causing underestimation of thresholds and stressor-interaction impacts.

Photoacclimation and Light Thresholds for Cold Temperate Seagrasses.

Water quality deterioration is expected to worsen the light conditions in shallow coastal waters with increasing human activities. Temperate seagrasses are known to tolerate a highly fluctuating light environment. However, depending on their ability to adjust to some decline in light conditions, decreases in daily light quantity and quality could affect seagrass physiology, productivity, and, eventually, survival if the Minimum Quantum Requirements (MQR) are not reached. To better understand if, how, and to what extent photosynthetic adjustments contribute to light acclimation, eelgrass (Zostera marina L.) shoots from the cold temperate St. Lawrence marine estuary (Rimouski, QC, Canada) were exposed to seven light intensity treatments (6, 36, 74, 133, 355, 503, and 860 µmol photons m-2 s-1, 14:10 light:dark photoperiod). Photosynthetic capacity and efficiency were quantified after five and 25 days of light exposure by Pulse Amplitude Modulated (PAM) fluorometry to assess the rapid response of the photosynthetic apparatus and its acclimation potential. Photoacclimation was also studied through physiological responses of leaves and shoots (gross and net primary production, pigment content, and light absorption). Shoots showed proof of photosynthetic adjustments at irradiances below 200 µmol photons m-2 s-1, which was identified as the threshold between limiting and saturating irradiances. Rapid Light Curves (RLC) and net primary production (NPP) rates revealed sustained maximal photosynthetic rates from the highest light treatments down to 74 µmol photons m-2 s-1, while a compensation point (NPP = 0) of 13.7 µmol photons m-2 s-1 was identified. In addition, an important package effect was observed, since an almost three-fold increase in chlorophyll content in the lowest compared to the highest light treatment did not change the leaves' light absorption. These results shed new light on photosynthetic and physiological processes, triggering light acclimation in cold temperate eelgrass. Our study documents an MQR value for eelgrass in the St. Lawrence estuary, which is highly pertinent in the context of conservation and restoration of eelgrass meadows.

Benthos response to nutrient enrichment and functional consequences in coastal ecosystems.

As land use intensifies, many coastal waters are becoming enriched with otherwise limiting nutrients, leading to eutrophication. While the extreme effects of eutrophication on benthic communities are well documented, there is still a lack of knowledge about how nutrient enrichment alters biogeochemical interactions occurring at the sediment-water interface. Using ex-situ experiments, this study explores the consequences of nutrient enrichment on sediment characteristics, macrofauna community and benthic fluxes. The quantity of sedimentary organic matter and porewater concentration of NH4+, NOx and PO43- increased in enriched treatments. These changes did not affect the macrobenthic community structure. However, macroinfauna buried less deep and increased their ventilation activity. As consequences, nutrient efflux increased, thereby favouring eutrophication processes. These effects were reduced in presence of seagrass, thus illustrating the buffering capacity of seagrass in the context of environmental changes, and particularly, of eutrophication. Overall, this study highlights that the functional consequences of nutrient enrichment involve interconnected processes that are variable in space and time.

Individual and population dietary specialization decline in fin whales during a period of ecosystem shift.

This study sought to estimate the effect of an anthropogenic and climate-driven change in prey availability on the degree of individual and population specialization of a large marine predator, the fin whale (Balaenoptera physalus). We examined skin biopsies from 99 fin whales sampled in the St. Lawrence Estuary (Canada) over a nine year period (1998-2006) during which environmental change was documented. We analyzed stable isotope ratios in skin and fatty acid signatures in blubber samples of whales, as well as in seven potential prey species, and diet was quantitatively assessed using Bayesian isotopic models. An abrupt change in fin whale dietary niche coincided with a decrease in biomass of their predominant prey, Arctic krill (Thysanoessa spp.). This dietary niche widening toward generalist diets occurred in nearly 60% of sampled individuals. The fin whale population, typically composed of specialists of either krill or lipid-rich pelagic fishes, shifted toward one composed either of krill specialists or true generalists feeding on various zooplankton and fish prey. This change likely reduced intraspecific competition. In the context of the current "Atlantification" of northern water masses, our findings emphasize the importance of considering individual-specific foraging tactics and not only population or group average responses when assessing population resilience or when implementing conservation measures.

Poor prey quality is compensated by higher provisioning effort in passerine birds.

In altricial avian species, nutrition can significantly impact nestling fitness by increasing their survival and recruitment chances after fledging. Therefore, the effort invested by parents towards provisioning nestlings is crucial and represents a critical link between habitat resources and reproductive success. Recent studies suggest that the provisioning rate has little or no effect on the nestling growth rate. However, these studies do not consider prey quality, which may force breeding pairs to adjust provisioning rates to account for variation in prey nutritional value. In this 8-year study using black-capped (Poecile atricapillus) and boreal (Poecile hudsonicus) chickadees, we hypothesized that provisioning rates would negatively correlate with prey quality (i.e., energy content) across years if parents adjust their effort to maintain nestling growth rates. The mean daily growth rate was consistent across years in both species. However, prey energy content differed among years, and our results showed that parents brought more food to the nest and fed at a higher rate in years of low prey quality. This compensatory effect likely explains the lack of relationship between provisioning rate and growth rate reported in this and other studies. Therefore, our data support the hypothesis that parents increase provisioning efforts to compensate for poor prey quality and maintain offspring growth rates.

From Africa to Antarctica: Exploring the Metabolism of Fish Heart Mitochondria Across a Wide Thermal Range.

The thermal sensitivity of ectotherms is largely dictated by the impact of temperature on cellular bioenergetics, particularly on mitochondrial functions. As the thermal sensitivity of bioenergetic pathways depends on the structural and kinetic properties of its component enzymes, optimization of their collective function to different thermal niches is expected to have occurred through selection. In the present study, we sought to characterize mitochondrial phenotypic adjustments to thermal niches in eight ray-finned fish species occupying a wide range of thermal habitats by comparing the activities of key mitochondrial enzymes in their hearts. We measured the activity of four enzymes that control substrate entrance into the tricarboxylic acid (TCA) cycle: pyruvate kinase (PK), pyruvate dehydrogenase complex (PDHc), carnitine palmitoyltransferase (CPT), and hydroxyacyl-CoA dehydrogenase (HOAD). We also assayed enzymes of the electron transport system (ETS): complexes I, II, I + III, and IV. Enzymes were assayed at five temperatures (5, 10, 15, 20, and 25°C). Our results showed that the activity of CPT, a gatekeeper of the fatty acid pathway, was higher in the cold-water fish than in the warmer-adapted fish relative to the ETS (complexes I and III) when measured close to the species optimal temperatures. The activity of HOAD showed a similar pattern relative to CI + III and thermal environment. By contrast, PDHc and PK did not show the similar patterns with respect to CI + III and temperature. Cold-adapted species had high CIV activities compared to those of upstream complexes (I, II, I + III) whereas the converse was true for warm-adapted species. Our findings reveal a significant variability of heart mitochondrial organization among species that can be linked to temperature adaptation. Cold-adapted fish do not appear to compensate for PDHc activity but likely adjust fatty acids oxidation through higher activities of CPT and HOAD relative to complexes I + III.

Bioturbation activity of three macrofaunal species and the presence of meiofauna affect the abundance and composition of benthic bacterial communities.

Given concerns of increasing rates of species extinctions, the relationship between biodiversity and ecosystem functioning has become a major research focus over the past two decades. Many studies have shown that biodiversity per se (e.g. species richness) or species-specific traits may be good predictors of changes in ecosystem function. Although numerous studies on this subject have focused on terrestrial systems, few have evaluated benthic marine systems. We used the Limecola balthica community as a model to test whether the number or identity of three well-studied macrofaunal species influence the sediment bacterial compartment, which drives important biogeochemical processes and influence ecosystem functioning. We also investigated the poorly known role of meiofauna in the interactions between macrofauna and bacteria. Eight combinations of 0-3 species were maintained in microcosms for 34 days in the presence or absence of meiofauna. The abundance and composition of the bacterial community, defined by the relative percentage of cells with a high (HNA) vs low (LNA) nucleic acid content, were measured. Species identity of macrofauna was a better indicator of changes in the microbial compartment than was species richness per se. In particular, the gallery-diffuser behaviour of the polychaete Alitta virens likely induced strong changes in sediment physical and geochemical properties with a major impact on the bacterial compartment. Moreover, the presence of meiofauna modulated the influence of macrofauna on bacterial communities. This study provides evidence that species identity provides greater explanatory power than species richness to predict changes in the bacterial compartment. We propose that multi-compartment approaches to describe interactions amongst different size classes of organisms and their ecological roles should be further developed to improve our understanding of benthic ecosystem functioning.

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.

Impact of forest harvesting on trophic structure of eastern Canadian Boreal Shield lakes: insights from stable isotope analyses.

Perturbations on ecosystems can have profound immediate effects and can, accordingly, greatly alter the natural community. Land-use such as forestry activities in the Canadian Boreal region have increased in the last decades, raising concerns about their potential impact on aquatic ecosystems. The objective of this study was to evaluate the impact of forest harvesting on trophic structure in eastern Canadian Boreal Shield lakes. We measured carbon and nitrogen stable isotopes values for aquatic primary producers, terrestrial detritus, benthic macroinvertebrates, zooplankton and brook trout (Salvelinus fontinalis) over a three-year period in eight eastern Boreal Shield lakes. Four lakes were studied before, one and two years after forest harvesting (perturbed lakes) and compared with four undisturbed reference lakes (unperturbed lakes) sampled at the same time. Stable isotope mixing models showed leaf-litter to be the main food source for benthic primary consumers in both perturbed and unperturbed lakes, suggesting no logging impact on allochthonous subsidies to the littoral food web. Brook trout derived their food mainly from benthic predatory macroinvertebrates in unperturbed lakes. However, in perturbed lakes one year after harvesting, zooplankton appeared to be the main contributor to brook trout diet. This change in brook trout diet was mitigated two years after harvesting. Size-related diet shift were also observed for brook trout, indicating a diet shift related to size. Our study suggests that carbon from terrestrial habitat may be a significant contribution to the food web of oligotrophic Canadian Boreal Shield lakes. Forest harvesting did not have an impact on the diet of benthic primary consumers. On the other hand, brook trout diet composition was affected by logging with greater zooplankton contribution in perturbed lakes, possibly induced by darker-colored environment in these lakes one year after logging.

Body size as a predictor of species loss effect on ecosystem functioning.

There is an urgent need to develop predictive indicators of the effect of species loss on ecosystem functioning. Body size is often considered as a good indicator because of its relationship to extinction risk and several functional traits. Here, we examined the predictive capacity of species body size in marine and freshwater multitrophic systems. We found a significant, but weak, effect of body size on functioning. The effect was much stronger when considering the effect of body size within trophic position levels. Compared to extinctions ordered by body size, random extinction sequences had lower multiple species loss effects on functioning. Our study is the first to show experimentally, in multitrophic systems, a more negative impact of ordered extinction sequences on ecosystem functioning than random losses. Our results suggest apparent ease in predicting species loss effect on functioning based on easily measured ecological traits that are body size and trophic position.

Identity effects dominate the impacts of multiple species extinctions on the functioning of complex food webs.

Understanding the impacts of species extinctions on the functioning of food webs is a challenging task because of the complexity of ecological interactions. We report the impacts of experimental species extinctions on the functioning of two food webs of freshwater and marine systems. We used a linear model to partition the variance among the multiple components of the diversity effect (linear group richness, nonlinear group richness, and identity). The identity of each functional group was the best explaining variable of ecosystem functioning for both systems. We assessed the contribution of each functional group in multifunctional space and found that, although the effect of functional group varied across ecosystem functions, some functional groups shared common effects on functions. This study is the first experimental demonstration that functional identity dominates the effects of extinctions on ecosystem functioning, suggesting that generalizations are possible despite the inherent complexity of interactions.

Ecodynamics of PAHs at a peri-urban site of the French Mediterranean Sea.

The PAH contamination level and biochemical composition of sinking particles and surficial sediments (0-0.5 cm layer) were assessed at a rural coastal site in the northwestern Mediterranean Sea. Surficial sediment contamination (≈20 ng g(-1)) was considerably lower than at other Mediterranean sites, yet particles collected in sediment traps had 6-8 times more PAH. Contaminated particles were mostly marine in origin. Temporal variation of contamination levels correlated with organic content of the particles, but some of the observed variability could be attributed to seasonal changes in pyrolytic PAH production. Sinking organic particles were potentially as readily digestible as surficial sediments for prospective consumers however, transfer of PAHs along the benthic food chain is probably enhanced because of the particles' higher nutritional value.

Genome size differences in Hyalella cryptic species.

The Hyalella azteca (Saussure) complex includes numerous amphipod cryptic species in freshwater habitats in America as revealed by DNA barcoding surveys. Two ecomorphs (small and large) have evolved numerous times in this complex. Few phenotypic criteria have been found to differentiate between the numerous species of this complex. The present study aims to explore genome size differences between some species of the H. azteca complex co-occurring in a Canadian boreal lake using flow cytometry. Nuclear DNA content was estimated for 50 individuals belonging to six COI haplotypes corresponding to four provisional species of the H. azteca complex. Species from the large ecomorph had C-values significantly larger than species from the small ecomorph, whereas slight differences were found among species of the small ecomorph. These differences in genome sizes might be linked to ecological and physiological differences among species of the H. azteca complex.

DIEL VARIATIONS IN OPTICAL PROPERTIES OF IMANTONIA ROTUNDA (HAPTOPHYCEAE) AND THALASSIOSIRA PSEUDONANA (BACILLARIOPHYCEAE) EXPOSED TO DIFFERENT IRRADIANCE LEVELS(1).

Diel variations of cellular optical properties were examined for cultures of the haptophyte Imantonia rotunda N. Reynolds and the diatom Thalassiosira pseudonana (Hust.) Hasle et Heimdal grown under a 14:10 light:dark (L:D) cycle and transferred from 100 µmol photons · m(-2) · s(-1) to higher irradiances of 250 and 500 µmol photons · m(-2) · s(-1) . Cell volume and abundance, phytoplankton absorption coefficients, flow-cytometric light scattering and chl fluorescence, and pigment composition were measured every 2 h over a 24 h period. Results showed that cell division was more synchronous for I. rotunda than for T. pseudonana. Several variables exhibited diel variability with an amplitude >100%, notably mean cell volume for the haptophyte and photoprotective carotenoids for both species, while optical properties such as flow-cytometric scattering and chl a-specific phytoplankton absorption generally showed <50% diel variability. Increased irradiance induced changes in pigments (both species) and mean cell volume (for the diatom) and amplified diel variability for most variables. This increase in amplitude is larger for pigments (factor of 2 or more, notably for cellular photoprotective carotenoid content in I. rotunda and for photosynthetic pigments in T. pseudonana) than for optical properties (a factor of 1.5 for chl a-specific absorption, at 440 nm, in I. rotunda and a factor of 2 for the absorption cross-section and the chl a-specific scattering in T. pseudonana). Consequently, diel changes in optical properties and pigmentation associated with the L:D cycle and amplified by concurrent changes in irradiance likely contribute significantly to the variability in optical properties observed in biooptical field studies.

UV effects on marine planktonic food webs: A synthesis of results from mesocosm studies.

UV irradiance has a broad range of effects on marine planktonic organisms. Direct and indirect effects on individual organisms have complex impacts on food-web structure and dynamics, with implications for carbon and nutrient cycling. Mesocosm experiments are well suited for the study of such complex interrelationships. Mesocosms offer the possibility to conduct well-controlled experiments with intact planktonic communities in physical, chemical and light conditions mimicking those of the natural environment. In allowing the manipulation of UV intensities and light spectral composition, the experimental mesocosm approach has proven to be especially useful in assessing the impacts at the community level. This review of mesocosm studies shows that, although a UV increase even well above natural intensities often has subtle effects on bulk biomass (carbon and chlorophyll), it can significantly impact the food-web structure because of different sensitivity to UV among planktonic organisms. Given the complexity of UV impacts, as evidenced by results of mesocosm studies, interactions between UV and changing environmental conditions (e.g. eutrophication and climate change) are likely to have significant effects on the function of marine ecosystems.