A primary aim of microbial ecology is to determine patterns and drivers of community distribution, interaction, and assembly amidst complexity and uncertainty. Microbial community composition has been shown to change across gradients of environment, geographic distance, salinity, temperature, oxygen, nutrients, pH, day length, and biotic factors 1-6 . These patterns have been identified mostly by focusing on one sample type and region at a time, with insights extra polated across environments and geography to produce generalized principles. To assess how microbes are distributed across environments globally-or whether microbial community dynamics follow funda mental ecological 'laws' at a planetary scale-requires either a massive monolithic cross environment survey or a practical methodology for coordinating many independent surveys. New studies of microbial environments are rapidly accumulating; however, our ability to extract meaningful information from across datasets is outstripped by the rate of data generation. Previous meta analyses have suggested robust gen eral trends in community composition, including the importance of salinity 1 and animal association 2 . These findings, although derived from relatively small and uncontrolled sample sets, support the util ity of meta analysis to reveal basic patterns of microbial diversity and suggest that a scalable and accessible analytical framework is needed.The Earth Microbiome Project (EMP, http://www.earthmicrobiome. org) was founded in 2010 to sample the Earth's microbial communities at an unprecedented scale in order to advance our understanding of the organizing biogeographic principles that govern microbial commu nity structure 7,8 . We recognized that open and collaborative science, including scientific crowdsourcing and standardized methods 8 , would help to reduce technical variation among individual studies, which can overwhelm biological variation and make general trends difficult to detect 9 . Comprising around 100 studies, over half of which have yielded peer reviewed publications (Supplementary Table 1), the EMP has now dwarfed by 100 fold the sampling and sequencing depth of earlier meta analysis efforts 1,2 ; concurrently, powerful analysis tools have been developed, opening a new and larger window into the distri bution of microbial diversity on Earth. In establishing a scalable frame work to catalogue microbiota globally, we provide both a resource for the exploration of myriad questions and a starting point for the guided acquisition of new data to answer them. As an example of using this Our growing awareness of the microbial world's importance and diversity contrasts starkly with our limited understanding of its fundamental structure. Despite recent advances in DNA sequencing, a lack of standardized protocols and common analytical frameworks impedes comparisons among studies, hindering the development of global inferences about microbial life on Earth. Here we present a meta-analysis of microbial community samples collected by hundreds of r...
Data on the partial pressure of carbon dioxide (CO(2)) in the surface waters from a large number of lakes (1835) with a worldwide distribution show that only a small proportion of the 4665 samples analyzed (less than 10 percent) were within +/-20 percent of equilibrium with the atmosphere and that most samples (87 percent) were supersaturated. The mean partial pressure of CO(2) averaged 1036 microatmospheres, about three times the value in the overlying atmosphere, indicating that lakes are sources rather than sinks of atmospheric CO(2). On a global scale, the potential efflux of CO(2) from lakes (about 0.14 x 10(15) grams of carbon per year) is about half as large as riverine transport of organic plus inorganic carbon to the ocean. Lakes are a small but potentially important conduit for carbon from terrestrial sources to the atmospheric sink.
Carbon in thawing permafrost soils may have global impacts on climate change; however, the factors that control its processing and fate are poorly understood. The dominant fate of dissolved organic carbon (DOC) released from soils to inland waters is either complete oxidation to CO2 or partial oxidation and river export to oceans. Although both processes are most often attributed to bacterial respiration, we found that photochemical oxidation exceeds rates of respiration and accounts for 70 to 95% of total DOC processed in the water column of arctic lakes and rivers. At the basin scale, photochemical processing of DOC is about one-third of the total CO2 released from surface waters and is thus an important component of the arctic carbon budget.
Arctic tundra has large amounts of stored carbon and is thought to be a sink for atmospheric carbon dioxide (CO(2)) (0.1 to 0.3 petagram of carbon per year) (1 petagram = 10(15) grams). But this estimate of carbon balance is only for terrestrial ecosystems. Measurements of the partial pressure of CO(2) in 29 aquatic ecosystems across arctic Alaska showed that in most cases (27 of 29) CO(2) was released to the atmosphere. This CO(2) probably originates in terrestrial environments; erosion of particulate carbon plus ground-water transport of dissolved carbon from tundra contribute to the CO(2) flux from surface waters to the atmosphere. If this mechanism is typical of that of other tundra areas, then current estimates of the arctic terrestrial sink for atmospheric CO(2) may be 20 percent too high.
Seasonal shifts in bacterioplankton community composition in Toolik Lake, a tundra lake on the North Slope of Alaska, were related to shifts in the source (terrestrial versus phytoplankton) and lability of dissolved organic matter (DOM). A shift in community composition, measured by denaturing gradient gel electrophoresis (DGGE) of 16S rRNA genes, occurred at 4°C in near-surface waters beneath seasonal ice and snow cover in spring. This shift was associated with an annual peak in bacterial productivity ([ 14 C]leucine incorporation) driven by the large influx of labile terrestrial DOM associated with snow meltwater. A second shift occurred after the flux of terrestrial DOM had ended in early summer as ice left the lake and as the phytoplankton community developed. Bacterioplankton communities were composed of persistent populations present throughout the year and transient populations that appeared and disappeared. Most of the transient populations could be divided into those that were advected into the lake with terrestrial DOM in spring and those that grew up from low concentrations during the development of the phytoplankton community in early summer. Sequencing of DNA in DGGE bands demonstrated that most bands represented single ribotypes and that matching bands from different samples represented identical ribotypes. Bacteria were identified as members of globally distributed freshwater phylogenetic clusters within the ␣-and -Proteobacteria, the Cytophaga-Flavobacteria-Bacteroides group, and the Actinobacteria.
Region 2 comprises arctic and subarctic North America and is underlain by continuous or discontinuous permafrost. Its freshwater systems are dominated by a low energy environment and cold region processes. Central northern areas are almost totally in¯uenced by arctic air masses while Paci®c air becomes more prominent in the west, Atlantic air in the east and southern air masses at the lower latitudes. Air mass changes will play an important role in precipitation changes associated with climate warming. The snow season in the region is prolonged resulting in long-term storage of water so that the spring¯ood is often the major hydrological event of the year, even though, annual rainfall usually exceeds annual snowfall. The unique character of ponds and lakes is a result of the long frozen period, which aects nutrient status and gas exchange during the cold season and during thaw. GCM models are in close agreement for this region and predict temperature increases as large as 48C in summer and 98C in winter for a 2 Â CO 2 scenario. Palaeoclimate indicators support the probability that substantial temperature increases have occurred previously during the Holocene. The historical record indicates a temperature increase of 418C in parts of the region during the last century. GCM predictions of precipitation change indicate an increase, but there is little agreement amongst the various models on regional disposition or magnitude. Precipitation change is as important as temperature change in determining the water balance. The water balance is critical to every aspect of hydrology and limnology in the far north. Permafrost close to the surface plays a major role in freshwater systems because it often maintains lakes and wetlands above an impermeable frost table, which limits the water storage capabilities of the subsurface. Thawing associated with climate change would, particularly in areas of massive ice, stimulate landscape changes, which can aect every aspect of the environment. The normal spring¯ooding of ice-jammed north-¯owing rivers, such as the Mackenzie, is a major event, which renews the water supply of lakes in delta regions and which determines the availability of habitat for aquatic organisms. Climate warming or river damming and diversion would probably lead to the complete drying of many delta lakes. Climate warming would also change the characteristics of ponds that presently freeze to the bottom and result in fundamental changes in their limnological characteristics. At present, the food chain is rather simple usually culminating in lake trout or arctic char. A lengthening of the growing season and warmer water temperature would aect the chemical, mineral and nutrient status of lakes and most likely have deleterious eects on the food chain. Peatlands are extensive in region 2. They would move northwards at their southern boundaries, and, with sustained drying, many would change form or become inactive. Extensive wetlands and peatlands are an important component of the global carbon budget, and warm...
SUMMARY1. Lake Victoria endured multiple stresses over the past century including population growth, increased cultivation of land, meteorological variability, resource extraction, intensive fishing, introduction of exotic species and more recently climate warming. These stressors became manifest through a fundamental and rapid change in the fish community and fishery in the early 1980s and visible eutrophication. However, the relation of these two phenomena and the possible interaction of the multiple stressors have been difficult to establish because of the temporally fragmented nature of the environmental data. 2. Comprehensive limnological observations from the 1960s were repeated in the 1990s and established the eutrophication of the lake, but these do not provide insight to the time course of when changes in trophic state occurred. Comprehensive fishery catch data from 1965 to the present provide a time course of the change in community composition and yield but cannot be correlated in time with discontinuous and sparse limnological data to determine possible cause-effect relationships. 3. Palaeolimnologic studies were conducted on three cores, two offshore and one nearshore, to establish a time course for the eutrophication of the lake that can be related to time-based data on the fishery. In the 1920s, the cores recorded an increase in nitrogen content of the sediments, but there was no significant response in the paleo-productivity indicators of biogenic Si deposition and change d 13 C of deposited organic matter.Phosphorus deposition began to increase in the 1940s in all three cores after which biogenic Si deposition increased steadily over time. Responses in d 13 C of organic matter begin in the 1960s at the coring sites. In the 1970s, the d 13 C of organic matter at the nearshore site increased nearly 3& in a 10-year period likely as a response to a dramatic increase in internal P loading caused by spreading anoxia. 4. Nile perch, the large predatory fish introduced in 1954, had become established through much of the lake at low abundances by the 1970s. In 1980, the catch of this fish began to increase, and by the end of the decade, the Lake Victoria fishery was the largest lake fishery in the world; and Nile perch dominated the catch. While catches of some other fishes also increased, the endemic haplochromines suffered a catastrophic decline in abundance and loss of biodiversity. 5. The detailed chronostratigraphies for these sediment cores established that the major changes in the trophic condition of the lake were accomplished prior to the change in the fish community and that the increased primary productivity of the lake likely contributed to the increased fish catches after 1980. The increased algal abundance also would have greatly reduced visibility and facilitated the emergence of Nile perch as the dominant top predator. 6. Thematic implications: multiple stresses were present in Lake Victoria over several decades, but transition to a new ecosystem state with a transformed food web and high...
Microbes are transported in hydrological networks through many environments, but the nature and dynamics of underlying microbial metacommunities and the impact of downslope inoculation on patterns of microbial diversity across landscapes are unknown. Pyrosequencing of small subunit ribosomal RNA gene hypervariable regions to characterize microbial communities along a hydrological continuum in arctic tundra showed a pattern of decreasing diversity downslope, with highest species richness in soil waters and headwater streams, and lowest richness in lake water. In a downstream lake, 58% and 43% of the bacterial and archaeal taxa, respectively, were also detected in diverse upslope communities, including most of the numerically dominant lake taxa. In contrast, only 18% of microbial eukaryotic taxa in the lake were detected upslope. We suggest that patterns of diversity in surface waters are structured by initial inoculation from microbial reservoirs in soils followed by a species-sorting process during downslope dispersal of both common and rare microbial taxa. Our results suggest that, unlike for metazoans, a substantial portion of bacterial and archaeal diversity in surface freshwaters may originate in complex soil environments.
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