Accounting for direct and indirect cumulative effects of anthropogenic pressures on salmon- and herring-linked land and ocean ecosystems.

Salmon and herring support both land and ocean predators and are critical to ecosystem resilience. Their linkages across land and sea realms make them highly susceptible to human activities, which can have flow-on effects up the food web. We quantify and compare the potential cumulative effects of human-driven pressures on interdependent species in salmon- and herring-linked ecosystems of western Canada using a risk assessment methodology. Adding indirect risks resulted in 68% greater total risks for land species than for direct risk alone, versus 15% for marine species. Inclusion of climate change pressures resulted in the greatest change in risk for low trophic marine species and habitats (greater than 25% increase). Forestry-related pressures accounted for the highest risk to all species and projected management of these pressures resulted in a total reduction of risk across all ecosystem components that was more than 14% greater than management of fisheries pressures. Ignoring land food web linkages and pressures underestimated cumulative risk by more than 40% for salmon and herring. This simple framework can be used to evaluate potential risk of existing human uses and future change to inform immediate management of linked land-sea ecosystems and help species avoid the 'death by a thousand cuts'. This article is part of the theme issue 'Nurturing resilient marine ecosystems'.

Cascading social-ecological costs and benefits triggered by a recovering keystone predator.

Predator recovery often leads to ecosystem change that can trigger conflicts with more recently established human activities. In the eastern North Pacific, recovering sea otters are transforming coastal systems by reducing populations of benthic invertebrates and releasing kelp forests from grazing pressure. These changes threaten established shellfish fisheries and modify a variety of other ecosystem services. The diverse social and economic consequences of this trophic cascade are unknown, particularly across large regions. We developed and applied a trophic model to predict these impacts on four ecosystem services. Results suggest that sea otter presence yields 37% more total ecosystem biomass annually, increasing the value of finfish [+9.4 million Canadian dollars (CA$)], carbon sequestration (+2.2 million CA$), and ecotourism (+42.0 million CA$). To the extent that these benefits are realized, they will exceed the annual loss to invertebrate fisheries (-$7.3 million CA$). Recovery of keystone predators thus not only restores ecosystems but can also affect a range of social, economic, and ecological benefits for associated communities.

An approach to incorporating inferred connectivity of adult movement into marine protected area design with limited data.

Marine protected areas (MPAs) are important conservation tools that can support the resilience of marine ecosystems. Many countries, including Canada, have committed to protecting at least 10% of their marine areas under the Convention on Biological Diversity's Aichi Target 11, which includes connectivity as a key aspect. Connectivity, the movement of individuals among habitats, can enhance population stability and resilience within and among MPAs. However, little is known about regional spatial patterns of marine ecological connectivity, particularly adult movement. We developed a method to assess and design MPA networks that maximize inferred connectivity within habitat types for adult movement when ecological data are limited. We used the Northern Shelf Bioregion in British Columbia, Canada, to explore two different approaches: (1) evaluating sites important for inferred regional connectivity (termed hotspots) and (2) assessing MPA network configurations based on their overlap with connectivity hotspots and interconnectedness between MPAs. To assess inferred connectivity via adult movement, we used two different threshold distances (15 and 50 km) to capture moderate home ranges, which are most appropriate to consider in MPA design. We applied graph theory to assess inferred connectivity within 16 habitat and depth categories (proxies for distinct ecological communities), and used novel multiplex network methodologies to perform an aggregated assessment of inferred connectivity. We evaluated inferred regional connectivity hotspots based on betweenness and eigenvector centrality metrics, finding that the existing MPA network overlapped a moderate proportion of these regional hotspots and identified key areas to be considered as candidate MPAs. Network density among existing MPAs was low within the individual habitat networks, as well as the multiplex. This work informs an ongoing MPA planning process, and approaches for incorporating connectivity into MPA design when data are limited, with lessons for other contexts.

Supporting Risk Assessment: Accounting for Indirect Risk to Ecosystem Components.

The multi-scalar complexity of social-ecological systems makes it challenging to quantify impacts from human activities on ecosystems, inspiring risk-based approaches to assessments of potential effects of human activities on valued ecosystem components. Risk assessments do not commonly include the risk from indirect effects as mediated via habitat and prey. In this case study from British Columbia, Canada, we illustrate how such "indirect risks" can be incorporated into risk assessments for seventeen ecosystem components. We ask whether (i) the addition of indirect risk changes the at-risk ranking of the seventeen ecosystem components and if (ii) risk scores correlate with trophic prey and habitat linkages in the food web. Even with conservative assumptions about the transfer of impacts or risks from prey species and habitats, the addition of indirect risks in the cumulative risk score changes the ranking of priorities for management. In particular, resident orca, Steller sea lion, and Pacific herring all increase in relative risk, more closely aligning these species with their "at-risk status" designations. Risk assessments are not a replacement for impact assessments, but-by considering the potential for indirect risks as we demonstrate here-they offer a crucial complementary perspective for the management of ecosystems and the organisms within.

Managing small-scale commercial fisheries for adaptive capacity: insights from dynamic social-ecological drivers of change in Monterey Bay.

Globally, small-scale fisheries are influenced by dynamic climate, governance, and market drivers, which present social and ecological challenges and opportunities. It is difficult to manage fisheries adaptively for fluctuating drivers, except to allow participants to shift effort among multiple fisheries. Adapting to changing conditions allows small-scale fishery participants to survive economic and environmental disturbances and benefit from optimal conditions. This study explores the relative influence of large-scale drivers on shifts in effort and outcomes among three closely linked fisheries in Monterey Bay since the Magnuson-Stevens Fisheries Conservation and Management Act of 1976. In this region, Pacific sardine (Sardinops sagax), northern anchovy (Engraulis mordax), and market squid (Loligo opalescens) fisheries comprise a tightly linked system where shifting focus among fisheries is a key element to adaptive capacity and reduced social and ecological vulnerability. Using a cluster analysis of landings, we identify four modes from 1974 to 2012 that are dominated (i.e., a given species accounting for the plurality of landings) by squid, sardine, anchovy, or lack any dominance, and seven points of transition among these periods. This approach enables us to determine which drivers are associated with each mode and each transition. Overall, we show that market and climate drivers are predominantly attributed to dominance transitions. Model selection of external drivers indicates that governance phases, reflected as perceived abundance, dictate long-term outcomes. Our findings suggest that globally, small-scale fishery managers should consider enabling shifts in effort among fisheries and retaining existing flexibility, as adaptive capacity is a critical determinant for social and ecological resilience.

Sea otters homogenize mussel beds and reduce habitat provisioning in a rocky intertidal ecosystem.

Sea otters (Enhydra lutris) are keystone predators that consume a variety of benthic invertebrates, including the intertidal mussel, Mytilus californianus. By virtue of their competitive dominance, large size, and longevity, M. californianus are ecosystem engineers that form structurally complex beds that provide habitat for diverse invertebrate communities. We investigated whether otters affect mussel bed characteristics (i.e. mussel length distributions, mussel bed depth, and biomass) and associated community structure (i.e. biomass, alpha and beta diversity) by comparing four regions that varied in their histories of sea otter occupancy on the west coast of British Columbia and northern Washington. Mussel bed depth and average mussel lengths were 1.5 times lower in regions occupied by otters for >20 years than those occupied for <5 yrs. Diversity of mussel bed associated communities did not differ between regions; however, the total biomass of species associated with mussel beds was more than three-times higher where sea otters were absent. We examined alternative explanations for differences in mussel bed community structure, including among-region variation in oceanographic conditions and abundance of the predatory sea star Pisaster ochraceus. We cannot discount multiple drivers shaping mussel beds, but our findings indicate the sea otters are an important one. We conclude that, similar to their effects on subtidal benthic invertebrates, sea otters reduce the size distributions of intertidal mussels and, thereby, habitat available to support associated communities. Our study indicates that by reducing populations of habitat-providing intertidal mussels, sea otters may have substantial indirect effects on associated communities.

Using expert judgment to estimate marine ecosystem vulnerability in the California Current.

As resource management and conservation efforts move toward multi-sector, ecosystem-based approaches, we need methods for comparing the varying responses of ecosystems to the impacts of human activities in order to prioritize management efforts, allocate limited resources, and understand cumulative effects. Given the number and variety of human activities affecting ecosystems, relatively few empirical studies are adequately comprehensive to inform these decisions. Consequently, management often turns to expert judgment for information. Drawing on methods from decision science, we offer a method for eliciting expert judgment to (1) quantitatively estimate the relative vulnerability of ecosystems to stressors, (2) help prioritize the management of stressors across multiple ecosystems, (3) evaluate how experts give weight to different criteria to characterize vulnerability of ecosystems to anthropogenic stressors, and (4) identify key knowledge gaps. We applied this method to the California Current region in order to evaluate the relative vulnerability of 19 marine ecosystems to 53 stressors associated with human activities, based on surveys from 107 experts. When judging the relative vulnerability of ecosystems to stressors, we found that experts primarily considered two criteria: the ecosystem's resistance to the stressor and the number of species or trophic levels affected. Four intertidal ecosystems (mudflat, beach, salt marsh, and rocky intertidal) were judged most vulnerable to the suite of human activities evaluated here. The highest vulnerability rankings for coastal ecosystems were invasive species, ocean acidification, sea temperature change, sea level rise, and habitat alteration from coastal engineering, while offshore ecosystems were assessed to be most vulnerable to ocean acidification, demersal destructive fishing, and shipwrecks. These results provide a quantitative, transparent, and repeatable assessment of relative vulnerability across ecosystems to any ongoing or emerging human activity. Combining these results with data on the spatial distribution and intensity of human activities provides a systematic foundation for ecosystem-based management.

Impacts and interactions of multiple human perturbations in a California salt marsh.

Multiple disturbances to ecosystems can influence community structure by modifying resistance to and recovery from invasion by non-native species. Predicting how invasibility responds to multiple anthropogenic impacts is particularly challenging due to the variety of potential stressors and complex responses. Using manipulative field experiments, we examined the relative impact of perturbations that primarily change abiotic or biotic factors to promote invasion in coastal salt marsh plant communities. Specifically we test the hypotheses that nitrogen enrichment and human trampling facilitate invasion of upland weeds into salt marsh, and that the ability of salt marsh communities to resist and/or recover from invasion is modified by hydrological conditions. Nitrogen enrichment affected invasion of non-native upland plants at only one of six sites, and increased aboveground native marsh biomass at only two sites. Percent cover of native marsh plants declined with trampling at all sites, but recovered earlier at tidally flushed sites than at tidally restricted sites. Synergistic interactions between trampling and restricting tidal flow resulted in significantly higher cover of non-native upland plants in trampled plots at tidally restricted sites. Percent cover of non-native plants recovered to pre-trampling levels in fully tidal sites, but remained higher in tidally restricted sites after 22 months. Thus, perturbations that reduce biotic resistance interact with perturbations that alter abiotic conditions to promote invasion. This suggests that to effectively conserve or restore native biodiversity in altered systems, one must consider impacts of multiple human disturbances, and the interactions between them.

CREB binds to multiple loci on human chromosome 22.

The cyclic AMP-responsive element-binding protein (CREB) is an important transcription factor that can be activated by hormonal stimulation and regulates neuronal function and development. An unbiased, global analysis of where CREB binds has not been performed. We have mapped for the first time the binding distribution of CREB along an entire human chromosome. Chromatin immunoprecipitation of CREB-associated DNA and subsequent hybridization of the associated DNA to a genomic DNA microarray containing all of the nonrepetitive DNA of human chromosome 22 revealed 215 binding sites corresponding to 192 different loci and 100 annotated potential gene targets. We found binding near or within many genes involved in signal transduction and neuronal function. We also found that only a small fraction of CREB binding sites lay near well-defined 5' ends of genes; the majority of sites were found elsewhere, including introns and unannotated regions. Several of the latter lay near novel unannotated transcriptionally active regions. Few CREB targets were found near full-length cyclic AMP response element sites; the majority contained shorter versions or close matches to this sequence. Several of the CREB targets were altered in their expression by treatment with forskolin; interestingly, both induced and repressed genes were found. Our results provide novel molecular insights into how CREB mediates its functions in humans.

Distribution of NF-kappaB-binding sites across human chromosome 22.

We have mapped the chromosomal binding site distribution of a transcription factor in human cells. The NF-kappaB family of transcription factors plays an essential role in regulating the induction of genes involved in several physiological processes, including apoptosis, immunity, and inflammation. The binding sites of the NF-kappaB family member p65 were determined by using chromatin immunoprecipitation and a genomic microarray of human chromosome 22 DNA. Sites of binding were observed along the entire chromosome in both coding and noncoding regions, with an enrichment at the 5' end of genes. Strikingly, a significant proportion of binding was seen in intronic regions, demonstrating that transcription factor binding is not restricted to promoter regions. NF-kappaB binding was also found at genes whose expression was regulated by tumor necrosis factor alpha, a known inducer of NF-kappaB-dependent gene expression, as well as adjacent to genes whose expression is not affected by tumor necrosis factor alpha. Many of these latter genes are either known to be activated by NF-kappaB under other conditions or are consistent with NF-kappaB's role in the immune and apoptotic responses. Our results suggest that binding is not restricted to promoter regions and that NF-kappaB binding occurs at a significant number of genes whose expression is not altered, thereby suggesting that binding alone is not sufficient for gene activation.

The transcriptional activity of human Chromosome 22.

A DNA microarray representing nearly all of the unique sequences of human Chromosome 22 was constructed and used to measure global-transcriptional activity in placental poly(A)(+) RNA. We found that many of the known, related and predicted genes are expressed. More importantly, our study reveals twice as many transcribed bases as have been reported previously. Many of the newly discovered expressed fragments were verified by RNA blot analysis and a novel technique called differential hybridization mapping (DHM). Interestingly, a significant fraction of these novel fragments are expressed antisense to previously annotated introns. The coding potential of these novel expressed regions is supported by their sequence conservation in the mouse genome. This study has greatly increased our understanding of the biological information encoded on a human chromosome. To facilitate the dissemination of these results to the scientific community, we have developed a comprehensive Web resource to present the findings of this study and other features of human Chromosome 22 at http://array.mbb.yale.edu/chr22.