Hypoxia increases the behavioural activity of schooling herring: a response to physiological stress or respiratory distress?

Atlantic herring, Clupea harengus, increase their swimming speed during low O₂ (hypoxia) and it has been hypothesised that the behavioural response is modulated by the degree of “respiratory distress” (i.e. a rise in anaerobic metabolism and severe physiological stress). To test directly whether a deviation in physiological homeostasis is associated with any change in behavioural activity, we exposed C. harengus in a school to a progressive stepwise decline in water oxygen pressure and measured fish swimming speed and valid indicators of primary and secondary stress (i.e. blood cortisol, lactate, glucose and osmolality). Herring in hypoxia increased their swimming speed by 11–39% but only when was <8.5 kPa and in an unsteady (i.e. declining) state. In parallel with the shift in behaviour, plasma cortisol also exhibited an increase with plasma osmolality was subject to a transient rise at 8.5 kPa and plasma glucose was generally reduced at However, without any rise in anaerobically derived lactate levels, there was no evidence of respiratory distress at any set We show that a shift in physiological homeostasis is indeed linked with an increase in the swimming speed of herring but the physiological response reflects a hypoxia-induced shift in metabolic fuel-use rather than respiratory distress per se. The significance of this behavioural–physiological reaction is discussed in terms of behavioural-energetic trade-offs, schooling dynamics and the hypoxia tolerance of herring.     

Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.     

Extrinsic and intrinsic controls of zooplankton diversity in lakes

Pelagic crustacean zooplankton were collected from 336 Norwegian lakes covering a wide range of latitude, altitude, lake area, mean depth, production (as chlorophyll a), and fish community structure. Mean zooplankton species richness during the ice-free season was generally low at high latitudes and altitudes. Further, lower species richness was recorded in western lakes, possibly reflecting constraints on migration and dispersal. However, despite obvious spatial limitations, geographic boundaries were only weak predictors of mean zooplankton richness. Similarly, lake surface area did not contribute positively to mean richness such as seen in other ecosystem surveys. Rather, intrinsic factors such as primary production and fish community (planktivore) structure were identified by regression analysis as the major predictors of zooplankton diversity, while a positive correlation was observed between species richness and total zooplankton biomass. However, in spite of a large number of variables included in this study, the predictive power of multiple regression models was modest (<50% variance explained), pointing to a major role for within-lake properties, as yet unidentified intrinsic forces, stochasticity, or dispersal as constraints on zooplankton diversity in these lakes.     

Recruitment limitation constrains local species richness and productivity in dry grassland

Species coexistence and local-scale species richness are limited by the availability of seeds and microsites for germination and establishment. We conducted a seed addition experiment in seminatural grassland at three sites in southern Switzerland and repeated the experiment in two successive years to evaluate various circumstances under which seed limitation and establishment success affect community functioning. A collection of 144,000 seeds of 22 meadow species including grasses and forbs of local provenance was gathered, and seeds were individually sown in a density that resembled natural seed rain. The three communities were seed limited. Three years after sowing, single species varied in emergence (0-50%), survival (0-69%), and establishment rates (0-27%). One annual and 13 perennial species reached reproductive stage. Low establishment at one site and reduced growth at another site indicated stronger microsite limitation compared to the third site. Recruitment was influenced by differences in abiotic environmental conditions between sites (water availability, soil minerals) and by within-site differences in biotic interaction (competition). At the least water-limited site, sowing resulted in an increase in phytomass due to establishment of short-lived perennials in the second and third years after sowing. This increase persisted over the following two years due to establishment of longer-lived perennials. After sowing in a wetter year with higher phytomass, however, productivity did not increase, because higher intensity of competition in an early phase of establishment resulted in less vigorous plants later on. Due to the generally favorable weather conditions during this study, sowing year had a small effect on numbers of established individuals over all species. Recruitment limitation can thus constrain local-scale species richness and productivity, either by a lack of seeds or by reduced seedling growth, likely due to competition from the established vegetation.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Mechanisms generating modification of benthos following tidal flat invasion by a Spartina hybrid.

Many coastal habitats are being substantially altered by introduced plants. In San Francisco Bay, California, USA, a hybrid form of the eastern cordgrass Spartina alterniflora is rapidly invading open mudflats in southern and central sections of the Bay, altering habitat, reducing macrofaunal densities, and shifting species composition. The invasion has resulted in significant losses of surface-feeding amphipods, bivalves, and cirratulid polychaetes, while subsurface feeding groups such as tubificid oligochaetes and capitellid polychaetes have been unaffected. In the present paper, we document the causes and mechanisms underlying the changes observed. Through a series of in situ manipulative experiments we examined the influence of hybrid Spartina canopy on a range of physical, chemical, and biological properties. The hybrid Spartina canopy exerted a strong influence on the hydrodynamic regime, triggering a series of physical, chemical, and biological changes in the benthic system. Relative to tidal flats, water velocity was reduced in hybrid patches, promoting deposition of fine-grained, organic-rich particles. The resulting changes in the sediment environment included increased porewater sulfide concentrations and anoxia, which led to poor survivorship of surface feeders such as bivalves, amphipods, and polychaetes. These are key taxa that support higher trophic levels including migratory shorebirds that feed on tidal flats. Altered flow in the Spartina canopy further contributed to changes in barnacle recruitment and resuspension of adult benthic invertebrates. Increased crab-induced predation pressure associated with Spartina invasion also contributed to changes in benthic invertebrate communities. Our results suggest that multiple physical, chemical, biotic, and trophic impacts of the Spartina invasion have resulted in substantial changes in benthic communities that are likely to have important effects on the entire ecosystem.     

Carbon sequestration in California agriculture, 1980-2000.

To better understand agricultural carbon fluxes in California, USA, we estimated changes in soil carbon and woody material between 1980 and 2000 on 3.6 x 10(6) ha of farmland in California. Combining the CASA (Carnegie-Ames-Stanford Approach) model with data on harvest indices and yields, we calculated net primary production, woody production in orchard and vineyard crops, and soil carbon. Over the 21-yr period, two trends resulted in carbon sequestration. Yields increased an average of 20%, corresponding to greater plant biomass and more carbon returned to the soils. Also, orchards and vineyards increased in area from 0.7 x 10(6) ha to 1.0 x 10(6) ha, displacing field crops and sequestering woody carbon. Our model estimates that California's agriculture sequestered an average of 19 g C x m(-2) x yr(-1). Sequestration was lowest in non-rice annual cropland, which sequestered 9 g C x m(-2) x yr(-1) of soil carbon, and highest on land that switched from annual cropland to perennial cropland. Land that switched from annual crops to vineyards sequestered 68 g C x m(-2) x yr(-1), and land that switched from annual crops to orchards sequestered 85 g C x m(-2) x yr(-1). Rice fields, because of a reduction in field burning, sequestered 55 g C x m(-2) x yr(-1) in the 1990s. Over the 21 years, California's 3.6 x 10(6) ha of agricultural land sequestered 11.0 Tg C within soils and 3.5 Tg C in woody biomass, for a total of 14.5 Tg C statewide. This is equal to 0.7% of the state's total fossil fuel emissions over the same time period. If California's agriculture adopted conservation tillage, changed management of almond and walnut prunings, and used all of its orchard and vineyard waste wood in the biomass power plants in the state, California's agriculture could offset up to 1.6% of the fossil fuel emissions in the state.     

Modeling species-habitat relationships with spatially autocorrelated observation data.

Spatial autocorrelation in wildlife observation data arises when extrinsic environmental processes and patterns that influence the spatial distribution of wildlife are themselves spatially structured, or when species are subject to intrinsic population processes, causing contagion or dispersion effects. Territoriality, Allee effects, dispersal limitations, and social clustering are examples of intrinsic processes. Both forms of autocorrelation can violate the assumptions of generalized linear regression models, resulting in biased estimation of model coefficients and diminished predictive performance. Such consequences may be avoided for extrinsic autocorrelation when autocorrelated environmental variables are available for use as model covariates, whereas intrinsic spatial autocorrelation requires an alternative modeling approach. The autologistic model provides an approach suited to the binary observations often obtained in wildlife surveys, but its performance has not been tested across widely varying sampling intensities or strengths of intrinsic spatial structure. Here we use simulated data to test the autologistic model under a range of sampling conditions. The autologistic model obtains better fits and substantially better predictive performance than the standard logistic regression model over the full range of sampling designs and intensities tested. We provide a simple Bayesian implementation of the autologistic model, which until now has not been achieved with standard statistical software alone. A step-by-step procedure is given for characterizing and modeling spatial autocorrelation in binary observation data, along with computer code for fitting autologistic models in WinBUGS, a freeware Bayesian analysis package. This approach avoids normal approximations to the pseudo-likelihood, in contrast to previous Bayesian applications of the autologistic model. We provide a sample application of the autologistic model, fitted to survey data for a gliding marsupial in southeastern Australia.     

Feedbacks between bivalve density, flow, and suspended sediment concentration on patch stable states

We explore the role of biophysical feedbacks occurring at the patch scale (spatial scale of tens of meters) that influence bivalve physiological condition and affect patch stability by developing a numerical model for the pinnid bivalve, Atrina zelandica, in cohesive sediments. Simulated feedbacks involve bivalve density, flow conditions (assumed to be primarily influenced by local water depth and peak current speed), suspended sediment concentration (evaluated through a balance between background concentration, deposition, and erosion), and changes in the physiology of Atrina derived from empirical study. The model demonstrates that high bivalve density can lead to skimming flow and to a concomitant decrease in resuspension that will affect suspended sediment concentration over the patch directly feeding back on bivalve physiology. Consequently, for a given flow and background suspended sediment load, the stability of a patch directly depends on the size and density of bivalves in the patch. Although under a range of conditions patch stability is ensured independently of bivalve density, simulations clearly indicate that sudden changes in bivalve density or suspended sediment concentration can substantially affect patch structure and lead to different stable states. The model highlights the role of interactions between organisms, flow, and broader scale environmental conditions in providing a mechanistic explanation for the patchy occurrence of benthic suspension feeders.