Molecular chaperones, including the heat-shock proteins (Hsps), are a ubiquitous feature of cells in which these proteins cope with stress-induced denaturation of other proteins. Hsps have received the most attention in model organisms undergoing experimental stress in the laboratory, and the function of Hsps at the molecular and cellular level is becoming well understood in this context. A complementary focus is now emerging on the Hsps of both model and nonmodel organisms undergoing stress in nature, on the roles of Hsps in the stress physiology of whole multicellular eukaryotes and the tissues and organs they comprise, and on the ecological and evolutionary correlates of variation in Hsps and the genes that encode them. This focus discloses that (a) expression of Hsps can occur in nature, (b) all species have hsp genes but they vary in the patterns of their expression, (c) Hsp expression can be correlated with resistance to stress, and (d) species' thresholds for Hsp expression are correlated with levels of stress that they naturally undergo. These conclusions are now well established and may require little additional confirmation; many significant questions remain unanswered concerning both the mechanisms of Hsp-mediated stress tolerance at the organismal level and the evolutionary mechanisms that have diversified the hsp genes.

The effect of Ocean Acidification (OA) on marine biota is quasi-predictable at best. While perturbation studies, in the form of incubations under elevated pCO2, reveal sensitivities and responses of individual species, one missing link in the OA story results from a chronic lack of pH data specific to a given species' natural habitat. Here, we present a compilation of continuous, high-resolution time series of upper ocean pH, collected using autonomous sensors, over a variety of ecosystems ranging from polar to tropical, open-ocean to coastal, kelp forest to coral reef. These observations reveal a continuum of month-long pH variability with standard deviations from 0.004 to 0.277 and ranges spanning 0.024 to 1.430 pH units. The nature of the observed variability was also highly site-dependent, with characteristic diel, semi-diurnal, and stochastic patterns of varying amplitudes. These biome-specific pH signatures disclose current levels of exposure to both high and low dissolved CO2, often demonstrating that resident organisms are already experiencing pH regimes that are not predicted until 2100. Our data provide a first step toward crystallizing the biophysical link between environmental history of pH exposure and physiological resilience of marine organisms to fluctuations in seawater CO2. Knowledge of this spatial and temporal variation in seawater chemistry allows us to improve the design of OA experiments: we can test organisms with a priori expectations of their tolerance guardrails, based on their natural range of exposure. Such hypothesis-testing will provide a deeper understanding of the effects of OA. Both intuitively simple to understand and powerfully informative, these and similar comparative time series can help guide management efforts to identify areas of marine habitat that can serve as refugia to acidification as well as areas that are particularly vulnerable to future ocean change.

The interaction of climate and the timing of low tides along the West Coast of the United States creates a complex mosaic of thermal environments, in which northern sites can be more thermally stressful than southern sites. Thus, climate change may not lead to a poleward shift in the distribution of intertidal organisms, as has been proposed, but instead will likely cause localized extinctions at a series of "hot spots." Patterns of exposure to extreme climatic conditions are temporally variable, and tidal predictions suggest that in the next 3 to 5 years "hot spots" are likely to appear at several northern sites.

Comparative analysis of the sea urchin genome has broad implications for the primitive state of deuterostome host defense and the genetic underpinnings of immunity in vertebrates. The sea urchin has an unprecedented complexity of innate immune recognition receptors relative to other animal species yet characterized. These receptor genes include a vast repertoire of 222 Toll-like receptors, a superfamily of more than 200 NACHT domain-leucine-rich repeat proteins (similar to nucleotide-binding and oligomerization domain (NOD) and NALP proteins of vertebrates), and a large family of scavenger receptor cysteine-rich proteins. More typical numbers of genes encode other immune recognition factors. Homologs of important immune and hematopoietic regulators, many of which have previously been identified only from chordates, as well as genes that are critical in adaptive immunity of jawed vertebrates, also are present. The findings serve to underscore the dynamic utilization of receptors and the complexity of immune recognition that may be basal for deuterostomes and predicts features of the ancestral bilaterian form.

Abstract. We explicitly quantified spatial and temporal patterns in the body temperature of an ecologically important species of intertidal invertebrate, the mussel Mytilus californianus, along the majority of its latitudinal range from Washington to southern California, USA. Using long-term (five years), high-frequency temperature records recorded at multiple sites, we tested the hypothesis that local ''modifying factors'' such as the timing of low tide in summer can lead to large-scale geographic mosaics of body temperature. Our results show that patterns of body temperature during aerial exposure at low tide vary in physiologically meaningful and often counterintuitive ways over large sections of this species' geographic range. We evaluated the spatial correlations among sites to explore how body temperatures change along the latitudinal gradient, and these analyses show that ''hot spots'' and ''cold spots'' exist where temperatures are hotter or colder than expected based on latitude. We identified four major hot spots and four cold spots along the entire geographic gradient with at least one hot spot and one cold spot in each of the three regions examined (WashingtonOregon, Central California, and Southern California). Temporal autocorrelation analysis of year-to-year consistency and temporal predictability in the mussel body temperatures revealed that southern animals experience higher levels of predictability in thermal signals than northern animals. We also explored the role of wave splash at a subset of sites and found that, while average daily temperature extremes varied between sites with different levels of wave splash, yearly extreme temperatures were often similar, as were patterns of predictability. Our results suggest that regional patterns of tidal regime and local pattern of wave splash can overwhelm those of large-scale climate in driving patterns of body temperature, leading to complex thermal mosaics of temperature rather than simple latitudinal gradients. A narrow focus on population changes only at range margins may overlook climatically forced local extinctions and other population changes at sites well within a species range. Our results emphasize the importance of quantitatively examining biogeographic patterns in environmental variables at scales relevant to organisms, and in forecasting the impacts of changes in climate across species ranges.

Rising atmospheric carbon dioxide has resulted in scientific projections of changes in global temperatures, climate in general, and surface seawater chemistry. Although the consequences to ecosystems and communities of metazoans are only beginning to be revealed, a key to forecasting expected changes in animal communities is an understanding of species' vulnerability to a changing environment. For example, environmental stressors may affect a particular species by driving that organism outside a tolerance window, by altering the costs of metabolic processes under the new conditions, or by changing patterns of development and reproduction. Implicit in all these examples is the foundational understanding of physiological mechanisms and how a particular environmental driver (e.g., temperature and ocean acidification) will be transduced through the animal to alter tolerances and performance. In this review, we highlight examples of mechanisms, focusing on those underlying physiological plasticity, that operate in contemporary organisms as a means to consider physiological responses that are available to organisms in the future.

Thermal stress has been considered to be among the most important determinants of organismal distribution in the rocky intertidal zone. Yet our understanding of how body temperatures experienced under field conditions vary in space and time, and of how these temperatures translate into physiological performance, is still rudimentary. We continuously monitored temperatures at a site in central California for a period of two years, using loggers designed to mimic the thermal characteristics of mussels, Mytilus californianus. Model mussel temperatures were recorded on both a horizontal and a vertical, north-facing microsite, and in an adjacent tidepool. We periodically measured levels of heat shock proteins (Hsp70), a measure of thermal stress, from mussels at each microsite. Mussel temperatures were consistently higher on the horizontal surface than on the vertical surface, and differences in body temperature between these sites were reflected in the amount of Hsp70. Seasonal peaks in extreme high temperatures ("acute" high temperatures) did not always coincide with peaks in average daily maxima ("chronic" high temperatures), suggesting that the time history of body temperature may be an important factor in determining levels of thermal stress. Temporal patterns in body temperature during low tide were decoupled from patterns in water temperature, suggesting that water temperature is an ineffective metric of thermal stress for intertidal organisms. This study demonstrates that spatial and temporal variability in thermal stress can be highly complex, and "snapshot" sampling of temperature and biochemical indices may not always be a reliable method for defining thermal stress at a site.

Ocean acidification (OA), a consequence of anthropogenic carbon dioxide emissions, poses a serious threat to marine organisms in tropical, open-ocean, coastal, deep-sea, and high-latitude sea ecosystems. The diversity of taxonomic groups that precipitate calcium carbonate from seawater are at particularly high risk. Here we review the rapidly expanding literature concerning the biological and ecological impacts of OA on calcification, using a cross-scale, process-oriented approach. In comparison to calcification, we find that areas such as fertilization, early life-history stages, and interaction with synergistic stressors are understudied. Although understanding the long-term consequences of OA are critical, available studies are largely short-term experiments that do not allow for tests of long-term acclimatization or adaptation. Future research on the phenotypic plasticity of contemporary organisms and interpretations of performance in the context of current environmental heterogeneity of pCO2 will greatly aid in our understanding of how organisms will respond to OA in the future.