We review the biogeography of microorganisms in light of the biogeography of macroorganisms A large body of research supports the idea that free-living microbial taxa exhibit biogeographic patterns Current evidence confirms that, as proposed by the Baas-Becking hypothesis, 'the environment selects' and is, in part, responsible for spatial variation in microbial diversity However, recent studies also dispute the idea that 'everything is everywhere' We also consider how the processes that generate and maintain biogeographic patterns in macroorganisms could operate in the microbial world

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Microbial biogeography : putting microorganisms on the map

Plants host distinct bacterial communities on and inside various plant organs, of which those associated with roots and the leaf surface are best characterized. The phylogenetic composition of these communities is defined by relatively few bacterial phyla, including Actinobacteria, Bacteroidetes, Firmicutes, and Proteobacteria. A synthesis of available data suggests a two-step selection process by which the bacterial microbiota of roots is differentiated from the surrounding soil biome. Rhizodeposition appears to fuel an initial substrate-driven community shift in the rhizosphere, which converges with host genotype–dependent finetuning of microbiota profiles in the selection of root endophyte assemblages. Substrate-driven selection also underlies the establishment of phyllosphere communities but takes place solely at the immediate leaf surface. Both the leaf and root microbiota contain bacteria that provide indirect pathogen protection, but root microbiota members appear to serve additional host functions through the acquisition of nutrients from soil for plant growth. Thus, the plant microbiota emerges as a fundamental trait that includes mutualism enabled through diverse biochemical mechanisms, as revealed by studies on plant growth–promoting and plant health–promoting bacteria.

Structure and functions of the bacterial microbiota of plants

The world's oceans contain a complex mixture of micro-organisms that are for the most part, uncharacterized both genetically and biochemically. We report here a metagenomic study of the marine planktonic microbiota in which surface (mostly marine) water samples were analyzed as part of the Sorcerer II Global Ocean Sampling expedition. These samples, collected across a several-thousand km transect from the North Atlantic through the Panama Canal and ending in the South Pacific yielded an extensive dataset consisting of 7.7 million sequencing reads (6.3 billion bp). Though a few major microbial clades dominate the planktonic marine niche, the dataset contains great diversity with 85% of the assembled sequence and 57% of the unassembled data being unique at a 98% sequence identity cutoff. Using the metadata associated with each sample and sequencing library, we developed new comparative genomic and assembly methods. One comparative genomic method, termed "fragment recruitment," addressed questions of genome structure, evolution, and taxonomic or phylogenetic diversity, as well as the biochemical diversity of genes and gene families. A second method, termed "extreme assembly," made possible the assembly and reconstruction of large segments of abundant but clearly nonclonal organisms. Within all abundant populations analyzed, we found extensive intra-ribotype diversity in several forms: (1) extensive sequence variation within orthologous regions throughout a given genome; despite coverage of individual ribotypes approaching 500-fold, most individual sequencing reads are unique; (2) numerous changes in gene content some with direct adaptive implications; and (3) hypervariable genomic islands that are too variable to assemble. The intra-ribotype diversity is organized into genetically isolated populations that have overlapping but independent distributions, implying distinct environmental preference. We present novel methods for measuring the genomic similarity between metagenomic samples and show how they may be grouped into several community types. Specific functional adaptations can be identified both within individual ribotypes and across the entire community, including proteorhodopsin spectral tuning and the presence or absence of the phosphate-binding gene PstS.

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The Sorcerer II Global Ocean Sampling Expedition: Northwest Atlantic through Eastern Tropical Pacific

The finding that total viral abundance is higher than total prokaryotic abundance and that a significant fraction of the prokaryotic community is infected with phages in aquatic systems has stimulated research on the ecology of prokaryotic viruses and their role in ecosystems. This review treats the ecology of prokaryotic viruses ('phages') in marine, freshwater and soil systems from a 'virus point of view'. The abundance of viruses varies strongly in different environments and is related to bacterial abundance or activity suggesting that the majority of the viruses found in the environment are typically phages. Data on phage diversity are sparse but indicate that phages are extremely diverse in natural systems. Lytic phages are predators of prokaryotes, whereas lysogenic and chronic infections represent a parasitic interaction. Some forms of lysogeny might be described best as mutualism. The little existing ecological data on phage populations indicate a large variety of environmental niches and survival strategies. The host cell is the main resource for phages and the resource quality, i.e., the metabolic state of the host cell, is a critical factor in all steps of the phage life cycle. Virus-induced mortality of prokaryotes varies strongly on a temporal and spatial scale and shows that phages can be important predators of bacterioplankton. This mortality and the release of cell lysis products into the environment can strongly influence microbial food web processes and biogeochemical cycles. Phages can also affect host diversity, e.g., by 'killing the winner' and keeping in check competitively dominant species or populations. Moreover, they mediate gene transfer between prokaryotes, but this remains largely unknown in the environment. Genomics or proteomics are providing us now with powerful tools in phage ecology, but final testing will have to be performed in the environment.

Ecology of prokaryotic viruses

The ocean's interior is Earth's largest biome. Recently, cultivation-independent ribosomal RNA gene surveys have indicated a potential importance for archaea in the subsurface ocean. But quantitative data on the abundance of specific microbial groups in the deep sea are lacking. Here we report a year-long study of the abundance of two specific archaeal groups (pelagic euryarchaeota and pelagic crenarchaeota) in one of the ocean's largest habitats. Monthly sampling was conducted throughout the water column (surface to 4,750 m) at the Hawai'i Ocean Time-series station. Below the euphotic zone (> 150 m), pelagic crenarchaeota comprised a large fraction of total marine picoplankton, equivalent in cell numbers to bacteria at depths greater than 1,000 m. The fraction of crenarchaeota increased with depth, reaching 39% of total DNA-containing picoplankton detected. The average sum of archaea plus bacteria detected by rRNA-targeted fluorescent probes ranged from 63 to 90% of total cell numbers at all depths throughout our survey. The high proportion of cells containing significant amounts of rRNA suggests that most pelagic deep-sea microorganisms are metabolically active. Furthermore, our results suggest that the global oceans harbour approximately 1.3 x 10(28) archaeal cells, and 3.1 x 10(28) bacterial cells. Our data suggest that pelagic crenarchaeota represent one of the ocean's single most abundant cell types.

Archaeal dominance in the mesopelagic zone of the Pacific Ocean

Total bacterial abundances estimated with different epifluorescence microscopy methods (4′,6-diamidino-2-phenylindole [DAPI], SYBR Green, and Live/Dead) and with flow cytometry (Syto13) showed good correspondence throughout two microcosm experiments with coastal Mediterranean water. In the Syto13-stained samples we could differentiate bacteria with apparent high DNA (HDNA) content and bacteria with apparent low DNA (LDNA) content. HDNA bacteria, “live” bacteria (determined as such with the Molecular Probes Live/Dead BacLight bacterial viability kit), and nucleoid-containing bacteria (NuCC) comprised similar fractions of the total bacterial community. Similarly, LDNA bacteria and “dead” bacteria (determined with the kit) comprised a similar fraction of the total bacterial community in one of the experiments. The rates of change of each type of bacteria during the microcosm experiments were also positively correlated between methods. In various experiments where predator pressure on bacteria had been reduced, we detected growth of the HDNA bacteria without concomitant growth of the LDNA bacteria, such that the percentage contribution of HDNA bacteria to total bacterial numbers (%HDNA) increased. This indicates that the HDNA bacteria are the dynamic members of the bacterial assemblage. Given how quickly and easily the numbers of HDNA and LDNA bacteria can be obtained, and given the similarity to the numbers of “live” cells and NuCC, the %HDNA is suggested as a reference value for the percentage of actively growing bacteria in marine planktonic environments.

https://aem.asm.org/content/aem/65/10/4475.full.pdf

Significance of size and nucleic acid content heterogeneity as measured by flow cytometry in natural planktonic bacteria.

Surface-water microbial populations were investigated in the northwest Mediterranean for possible indicators of phosphate deficiency and limitation. Low phosphorus availability was suggested by short turnover time (min. observed 0.68 h), high-alkaline phosphatase activity (V,,,,, = 28 nM hydrolyzed h-l), subsaturation of phosphate uptake (2.6-9% of V,,,), and high-pulse uptake capacity of added orthophosphate. Based on high pulse uptake capacity and subsaturated uptake in both the >l-pm and in the 0.2-l-pm size fractions, P deficiency is suggested for both phytoplankton and heterotrophic bacteria. P limitation of heterotrophic bacteria was also supported by fast positive responses after phosphate addition in both thymidine incorporation in whole-water samples and increased bacterial cell numbers in predator-free water. No effects were found after addition of carbon or nitrogen sources alone. Combined with other published evidence, we suggest that the growth rates of not only phytoplankton, but also of heterotrophic bacteria, are P limited in this environment in summer. The finding has important implications for the dynamics of accumulation of dissolved organic carbon in the photic zone and thus for the carbon cycle of oceans. The generally oligotrophic state of the Mediterranean Sea has traditionally been explained as a consequence of its antiestuarine circulation; the net evaporation from the Mediterranean produces an inflow of low-nutrient surface water through the Gibraltar and an eastward surface current in the Mediterranean (Redfield et al. 1963). If one combines an assumption of nitrogen-limited growth in the inflowing Atlantic water with the usual assumption of a faster recycling of phosphorus than of nitrogen, one might expect a strong nitrogen limitation in the upper photic zone during Mediterranean summer stratification. Contrary to such an expectation, accumulated experimental data seem to indicate that parts of the Mediterranean are characterized by a nitrate : phosphate ratio below Redfield (Krom et al. 1991). This seems to be particularly prominent in the eastern parts of the Mediterranean (Krom et al. 1991) and in the Adriatic Sea (Vukadin and Stojanski 1976). In an investigation of orthophosphate uptake in surface water from Villefranche Bay on the French Mediterranean coast, Dolan et al. (1995) found relatively short turnover times (down to 1.6 h). Estimates of bioavailable orthophosphate concentrations in this area (Thingstad et al. 1995) are also very low (0.8 nM), indicating, but not proving, that P may be the limiting nutrient also in this part of the western Mediterranean. The underlying biogeochemical mechanisms behind such an apparent shift toward P deficiency seem still to be unknown,

https://aslopubs.onlinelibrary.wiley.com/doi/epdf/10.4319/lo.1998.43.1.0088

P limitation of heterotrophic bacteria and phytoplankton in the northwest Mediterranean

Counts of heterotrophic bacteria in marine waters are usually in the order of 5 x 10(sup5) to 3 x 10(sup6) bacteria ml(sup-1). These numbers are derived from unspecific fluorescent staining techniques (J. E. Hobbie, R. J. Daley, and S. Jasper, Appl. Environ. Microbiol. 33:1225-1228, 1977; K. G. Porter and Y. S. Feig, Limnol. Oceanogr. 25:943-948, 1980) and are subsequently defined as total counts of bacteria. In samples from the Baltic Sea, the North Sea (Skagerrak), and the northeastern Mediterranean Sea, we found that only a minor fraction (2 to 32%) of total counts can be scored as bacteria with nucleoids. Lack of DNA no doubt means inactive cells; therefore, a much lower number of bacteria that grow at rates higher than those previously estimated must be responsible for the measured bacterial production in these seas. The remaining bacterium-sized and/or -shaped particles included in total counts may be cell residues of virus-lysed bacteria (ghosts) or remains of protozoan grazing.

Total counts of marine bacteria include a large fraction of non-nucleoid-containing bacteria (ghosts).

Consumption of dissolved organic carbon by marine bacteria and demand for inorganic nutrients.

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Consumption of dissolved organic carbon by marine bacteria and demand for inorganic nutrients

The density of specific aquatic bacteria was determined by use of whole-genome DNA hybridization towards community DNA. From a coastal marine environment (northern Baltic Sea), 48 specific bacteria were isolated on solid media over a 1-year period. Based on the presented hybridization protocol, the total density of the isolates ranged between 7 and 69% of the bacteria determined by acridine orange direct counts. When compared to the number of nucleoid-containing cells, the range increased to 29 to 111%. Thus, our results showed that bacteria able to form colonies on solid media accounted for a large fraction of the bacterioplankton. There were significant changes in the density of the different bacteria over the year, suggesting that bacterioplankton exhibit a seasonal succession analogous to phytoplankton. The bacteria studied were of diverse phylogenetic origin, being distributed among the alpha, beta, and gamma subdivisions of the class Proteobacteria and the cytophaga-flexibacter group. Partial 16S rRNA gene sequence analysis of 29 Baltic Sea isolates as well as of 30 Southern California Bight isolates showed that a majority of the isolates had low similarity (0.85 to 0.95) to reported sequence data. This indicated that the diversity of marine bacteria able to grow on solid media is largely unexplored.

Ulla Li Zweifel

Papers

Significance of size and nucleic acid content heterogeneity as measured by flow cytometry in natural planktonic bacteria.

P limitation of heterotrophic bacteria and phytoplankton in the northwest Mediterranean

Total counts of marine bacteria include a large fraction of non-nucleoid-containing bacteria (ghosts).

Consumption of dissolved organic carbon by marine bacteria and demand for inorganic nutrients

Dominant marine bacterioplankton species found among colony-forming bacteria