Extreme stresses, niches, and positive species interactions along stress gradients.

Since proposed two decades ago, the stress-gradient hypothesis (SGH), suggesting that species interactions shift from competition to facilitation with stress, has been widely examined. Despite broad support across species and ecosystems, ecologists debate whether the SGH applies to extreme environments, arguing that species interactions switch to competition or collapse under extreme stress. We show that facilitation often expands distributions on species borders. SGH exceptions occur when weak stress gradients or stresses outside of species' niches are examined, multiple stresses co-occur canceling out their effects, temporally dependent effects are involved, or results are improperly analyzed. We suggest that ecologists resolve debates by standardizing key SGH terms, such as fundamental and realized niche, stress gradients vs. environmental gradients, by quantitatively defining extreme stress, and by critically evaluating the functionality of stress gradients. We also suggest that new research examine the breadth and relevance of the SGH. More rigor needs to be applied to SGH tests to identify actual exceptions rather than those due to failures to meet its underlying assumptions, so that the general principles of the SGH and its exceptions can be incorporated into ecological theory, conservation strategies, and environmental change predictions.

Transcriptional and metabolomic analysis of Ascophyllum nodosum mediated freezing tolerance in Arabidopsis thaliana.

BACKGROUND: We have previously shown that lipophilic components (LPC) of the brown seaweed Ascophyllum nodosum (ANE) improved freezing tolerance in Arabidopsis thaliana. However, the mechanism(s) of this induced freezing stress tolerance is largely unknown. Here, we investigated LPC induced changes in the transcriptome and metabolome of A. thaliana undergoing freezing stress. RESULTS: Gene expression studies revealed that the accumulation of proline was mediated by an increase in the expression of the proline synthesis genes P5CS1 and P5CS2 and a marginal reduction in the expression of the proline dehydrogenase (ProDH) gene. Moreover, LPC application significantly increased the concentration of total soluble sugars in the cytosol in response to freezing stress. Arabidopsis sfr4 mutant plants, defective in the accumulation of free sugars, treated with LPC, exhibited freezing sensitivity similar to that of untreated controls. The 1H NMR metabolite profile of LPC-treated Arabidopsis plants exposed to freezing stress revealed a spectrum dominated by chemical shifts (δ) representing soluble sugars, sugar alcohols, organic acids and lipophilic components like fatty acids, as compared to control plants. Additionally, 2D NMR spectra suggested an increase in the degree of unsaturation of fatty acids in LPC treated plants under freezing stress. These results were supported by global transcriptome analysis. Transcriptome analysis revealed that LPC treatment altered the expression of 1113 genes (5%) in comparison with untreated plants. A total of 463 genes (2%) were up regulated while 650 genes (3%) were down regulated. CONCLUSION: Taken together, the results of the experiments presented in this paper provide evidence to support LPC mediated freezing tolerance enhancement through a combination of the priming of plants for the increased accumulation of osmoprotectants and alteration of cellular fatty acid composition.

Dynamics of species interaction strength in space, time and with developmental stage.

Quantifying species interaction strengths enhances prediction of community dynamics, but variability in the strength of species interactions in space and time complicates accurate prediction. Interaction strengths can vary in response to density, indirect effects, priority effects or a changing environment, but the mechanism(s) causing direction and magnitudes of change are often unclear. We designed an experiment to characterize how environmental factors influence the direction and the strength of priority effects between sessile species. We estimated per capita non-trophic effects of barnacles (Semibalanus balanoides) on newly settled germlings of the fucoid, Ascophyllum nodosum, in the presence and absence of consumers in experiments on rocky shores throughout the Gulf of Maine, USA. Per capita effects on germlings varied among environments and barnacle life stages, and these interaction strengths were largely unaltered by changing consumer abundance. Whereas previous evidence shows adult barnacles facilitate fucoids, here, we show that recent settlers and established juveniles initially compete with germlings. As barnacles mature, they switch to become facilitators of fucoids. Consumers caused variable mortality of germlings through time comparable to that from competition. Temporally variable effects of interactors (e.g. S. balanoides), or spatial variation in their population structure, in different regions differentially affect target populations (e.g. A. nodosum). This may affect abundance of critical stages and the resilience of target species to environmental change in different geographical regions.

Seaweed attachment to bedrock: biophysical evidence for a new geophycology paradigm.

Seaweeds are amongst the most obvious and ecologically important components of rocky shore communities worldwide but until now little has been known about the processes involved in their attachment. This multidisciplinary study investigated for the first time the interactions between marine macroalgal holdfasts and their underlying substrata, requiring the development of specialized sample preparation techniques to maintain the structural integrity of the holdfast-bedrock interface. Transmitted plane polarized light microscopy, scanning electron microscopy with energy dispersive spectroscopy and structured light illumination microscopy were used in the examination of the interface between Ascophyllum nodosum (Fucales, Heterokontophyta) and crustose red algae Lithothamnion sp. (Corallinales, Rhodophyta) on granite and limestone substrates. The new evidence presented here represents a paradigm shift in the way we view seaweed attachment because results show that the holdfasts exploit the physical characteristics of the rock-forming minerals in order to penetrate the bedrock and thus facilitate the attachment process. Mineral cleavage planes together with intercrystalline and intracrystalline boundaries and fractures provide penetration pathways for the holdfast tissue. This process causes disaggregation of rock-forming minerals to depths <10 mm and therefore assists in the bioerosion of coastal bedrock. It is concluded that seaweeds are able to cause weathering of natural rock and the term 'geophycology' is introduced to describe seaweed-bedrock interactions, including seaweed-induced weathering.

Modeling variation in interaction strength between barnacles and fucoids.

The strength by which species interact can vary throughout their ontogeny, as environments vary in space and time, and with the density of their populations. Characterizing strengths of interaction in situ for even a small number of species is logistically difficult and may apply only to those conditions under which the estimates were derived. We sought to combine data from field experiments estimating interaction strength of life stages of the barnacle, Semibalanus balanoides, on germlings of Ascophyllum nodosum, with a model that explored the consequences of variability at per capita and per population levels to the abundance of year-old algal recruits. We further simulated how this interaction affected fucoid germling abundance as the timing of their respective settlements varied relative to one another, as occurs regionally across the Gulf of Maine, USA. Juvenile S. balanoides have a weak estimated per capita effect on germlings. Germling populations are sensitive to variation in per capita effects of juvenile barnacles because of the typically large population sizes of the latter. However, high mortality of juvenile barnacles weakens the population interaction strength over time. Adult barnacles probably weakly facilitate fucoid germlings, but greater survival of adults sustains the strength of that interaction at the population level. Germling abundance is positively associated with densities of adult barnacles and negatively associated with that of juvenile barnacles. Metamorphosing cyprid larvae have the strongest per capita effect on germling abundance, but the interaction between the two stages is so short-lived that germling abundance is altered little. Variation in the timing of barnacle and A. nodosum settlement relative to one another had very little influence on the abundance of yearling germlings. Interactions between barnacles and germlings may influence the demographic structure of A. nodosum populations and the persistence of fucoid-dominated communities on sheltered rocky shores in New England.

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations.

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.

[Changes in parameters of Ascophyllum nodosum dark respiration induced by different salinities]. / izmenenie parametrov temnovogo dykhaniia Ascophyllum nodosum pri deistvii razlichnykh solenostei.

The changes in the rates of O2 consumption and CO2 release by Ascophyllum nodosum thalli in the dark were studied during a 12 day incubation at 34, 20, and 10/1000 salinity as well as at different pH. Depending on the initial pH of the medium, the algae demonstrated antipodal functional responses to the same salinities and, as a consequence, different capacity to overcome hypoosmotic stress. In addition, we observed a more pronounced effect of pH on the rate of O2 consumption than on the rate of CO2 release. Simultaneous desalination and acidification of the medium (pH 7-8) increased O2 consumption in the dark, which activated oxidative processes in the cells and increased their energy status. Forced pH stabilization (8.2) of the medium coupled with its desalination inhibited dark respiration of the algae, which, in our opinion, decreased their energy status. Specific manifestation of hypoosmotic stress and adaptation in A. nodosum are discussed considering the assumed differences in algal energy status.

Ascophyllum nodosum supplementation: a preharvest intervention for reducing Escherichia coli O157:H7 and Salmonella spp. in feedlot steers.

Ascophyllum nodosum (Tasco-14) decreased the prevalence of enterohemorrhagic Escherichia coli (EHEC) O157:H7 in animals fed prior to harvest. Tasco-14 was supplemented on a 2% dry matter basis 14 days prior to harvest to determine its effects on EHEC and Salmonella spp. prevalence. Two hundred mixed crossbred steers and heifers (Bos indicus x Bos taurus), in a large commercial finishing facility, served as experimental units. Treatment (TRT, n = 100) animals received a steam-rolled corn-based diet containing 2% Tasco-14 on a dry matter basis, and control (CON, n = 100) animals received only the steam-rolled corn-based diet. Hide swabs and fecal samples were obtained for EHEC and Salmonella spp. evaluations. Animals were sampled 1 day prior to (d - 1) the feeding of Tasco-14 and immediately following exsanguinations. The prevalence of EHEC O157 on hide swabs and in fecal samples (P = 0.0001 and P < 0.0001, respectively) and the prevalence of EHEC O157:H7 on hide swabs and in fecal samples (P < 0.0001 and P < 0.0001, respectively) was reduced by 33 and 36% from d - 1 levels on TRT hide swabs and by 9 and 11% in TRT fecal samples. The prevalence of EHEC O157 and EHEC O157:H7 was reduced by 33 and 36% from d - 1 levels on TRT hide swabs and by 9 and 11% in TRT fecal samples. The prevalence of Salmonella spp. on hide swabs did not change for TRT animals (P = 0.64). CON animals showed an increase in Salmonella spp. prevalence (P < 0.0001) from d - 1 feeding levels on hide swabs. The prevalence of Salmonella spp. increased in both TRT and CON fecal samples when compared to d - 1 levels (P = 0.002). However, TRT samples exhibited a lower (P < 0.05) postfeeding prevalence of Salmonella spp. in fecal samples than did CON samples. Results from this study indicate that 2% Tasco-14 supplementation in feedlot cattle diets reduces EHEC O157 and EHEC O157:H7 prevalence on hide swabs and in fecal samples and may suppress increases in Salmonella spp.

Removal of copper from aqueous solution by Ascophyllum nodosum immobilised in hydrophilic polyurethane foam.

The seaweed Ascophyllum nodosum was pre-treated by successive washes in distilled water and dilute acid, dried, and pulverised to produce particles of <150 microm. These were immobilised during the manufacturing process of Hypol 2002 polymer to form a biomass/polymer matrix that was stable and easy to handle. In making the composite a mixing speed of 360 rpm for 20-30 s with 2% (w/w) addition of surfactant to pre-polymer was found to be ideal. The average pore sizes for different water polymer mixes (expressed as volume ratios) were 1.66 mm +/- 0.98 (ratio 0.75:1), 1.58 mm +/- 0.76 (ratio 1:1), 1.64 mm +/- 0.6 (ratio 1.5:1) and 1.11 mm +/- 0.615 (ratio 2:1). The biomass/polymer was used alongside free native biomass in an initial adsorption experiment using a 0.0315 mmol dm(-3) Cu (II) solution and gave a copper uptake capacity (q)(max) of 0.037 mmol Cu g(-1) dry weight seaweed in both cases which represented approximately 85% of total initially available copper. In later adsorption isotherm experiments using Cu concentrations between 0.0315 and 0.944 mmol dm(-3) at pH 5.0 and immobilized biomass over five consecutive adsorption/desorption cycles the biomass/polymer showed an initial lowering of adsorption capacity but stabilised at 0.23 mmol g(-1) dry weight by the third re-use. The q(max) of the immobilised biomass decreased from 0.55 and 0.416 mmol of Cu g(-1) dry weight when pH was lowered from 4.0 to 3.0, and increased from 0.576 mmol g(-1) dry weight (biomass) at 283 K to 0.636 mmol g(-1)(biomass) at 303 K.