A Predictive Model of Bacterial Foraging by Means of Freely Released Extracellular Enzymes

Abstract Extracellular enzymes are important agents for microbial foraging and material cycling in diverse natural and man-made systems. Their abundance and effects are analyzed empirically on scales much larger than the forager. Here, we use a modelling approach to analyze the potential costs and benefits, to an individual immobile microbe, of freely releasing extracellular enzymes into a fluid-bathed, stable matrix of both inert and food-containing particles. The target environments are marine aggregates and sediments, but the results extend to biofilms, bioreactors, soils, stored foods, teeth, gut contents, and even soft tissues attacked by disease organisms. Model predictions, consistent with macroscopic observations of enzyme activity in laboratory and environmental samples, include: support of significant bacterial growth by cell-free enzymes; preponderance of particle-attached, as opposed to dissolved, cell-free enzymes; solubilization of particulate substrates in excess of resident microbe growth requirements; and constitutive, abundant enzyme release in some environments. Feeding with cell-free enzymes appears to be limited to substrates within a well-defined distance of the enzyme source. Fluxes of dissolved organic material out of pelagic oceanic aggregates and marine sediments, and difficulty detecting dissolved enzymes in such environments, may reflect characteristics of cell-free enzyme foraging and properties of the enzymes. Our calculations further suggest that cell-free enzymes may often be used by microorganisms as the fastest means to search for food. Received: 3 June 1997; Accepted: 25 November 1997     

Pelagic Bacteria–Particle Interactions and Community-Specific Growth Rates in Four Lakes along a Trophic Gradient

Abstract The relationships between bacterial concentration, bacterial production, and cell-specific activity of both free and attached bacteria and environmental factors such as suspended solids, nutrients, and temperature were examined in four lakes, two in New Zealand and two in Switzerland. Estimates of cell-specific production were obtained by microautoradiographic counts of [³H]thymidine-labeled cells. Bacteria attached to particles accounted for only 1.3 to 11.6% of the total bacterial abundance, but showed overall 20-fold higher specific growth rates and were relatively more active than their free counterparts. On average, 80 to 100% of epibacteria were attached to organic particles. The abundance and production of free and attached bacteria were positively correlated; however, relationships between these fractions and some environmental variables differed. Cell-specific activities of active bacteria were not equivalent to mean cellular activities of the entire bacterial community and differed in their relationship to trophic state. [³H]Thymidine-positive bacteria were more tightly linked to chlorophyll a than were total bacteria. Our findings indicate that production by attached bacteria, fueled by phytoplankton carbon, supplies ``new' free bacteria to the bacterial community. Our results support the idea that particulate organic matter acts as a source of dissolved nutrients to free bacteria. Bottom-up control of bacterial biomass, as shown by regressions of biomass vs production, appeared to be stronger in two ultraoligotrophic lakes than in two more eutrophic ones. Received: 17 April 1998; Accepted: 24 August 1998     

Consequences of protist-stimulated bacterial production for estimating protist growth efficiencies

The trophic link between bacteria and bacterivorous protists is a complex interaction that involves feedback of inorganic nutrients and growth substrates that are immeadiately available for prey growth. These interactions were examined in the laboratory and in incubations of concentrated natural assemblages of bacterioplankton. Growth dynamics of estuarine and marine bacterivorous protists were determined in laboratory culture using Vibrio natriegens as prey and were compared to growth of protists on bacterioplankton assemblages concentrated by tangential flow filtration from four northwest Florida Estuaries. Biomass transfers from bacteria to protists were monitored by tracing elemental carbon and nitrogen in particulate fractions of protist added and grazer free controls. Gross growth efficiencies of the protists on naturally occurring bacteria were within the range determined in lab estimates of growth efficiency on cultured bacteria (50%). However, bacterial response to protist excretion products was different in the lab and field incubations, and bacterial growth contributed to the biomass available to protists in the field incubations. As determined by radioisotope-labeled substrate incorporation, a time lag in bacterial reponse to protist excretion products was observed for laboratory batch cultures, allowing accurate estimation of growth efficiency. In incubations with concentrated natural bacterial assemblages, bacterial growth response coincided with protist growth and excretion. The additional bacterial production on protist excretion products reached a maximum of 2–3-fold higher than protist-free controls. In addition, ammonium concentrations increased with protist grazing and growth in lab cultures, but ammonium excreted by protists in concentrates did not accumulate. The C:N values for the bacterial concentrates suggests that these bacteria were nitrogen limited. It is speculated that dissolved organic carbon, concentrated by tangential flow filtration (> 100,000 MW membrane) with the bacterioplankton, was utilized by bacteria when nitrogen was supplied as ammonium and amino acids from protist excretion. Thus, estimates of protist growth efficiency on naturally occurring bacterioplankton, corrected for protist-stimulated bacterial production, were in the range of 13–21%.     

Coupling Between Rates of Bacterial Production and the Abundance of Metabolically Active Bacteria in Lakes, Enumerated Using CTC Reduction and Flow Cytometry

Abstract In natural bacterioplankton assemblages, only a fraction of the total cell count is active, and, therefore, rates of bacterial production should be more strongly correlated to the number of active cells than to the total number of bacteria. However, this hypothesis has seldom been tested. Herein we explore the relationship between rates of bacterial production (measured as leucine uptake) and the number of active bacteria in 14 lakes in southern Québec. Active bacteria are defined as those cells capable of reducing the tetrazolium salt CTC to its fluorescent formazan; these cells were enumerated using flow cytometry. Bacterial production varied two orders of magnitude in the lakes studied, as did the number of active bacteria, whereas the total number of bacteria varied by only sixfold. The number and proportion of active bacteria were similar among lake strata, but rates of bacterial production were highest in the epilimnion and lowest in the hypolimnion. As expected, bacterial production was better correlated to the number of active cells, and bacterial growth rates calculated for active cells ranged from 0.7 to 1.8 day⁻¹, on average threefold higher than those calculated on the basis of total bacterial abundance. Growth rates scaled to active cells were, on average, similar among lake strata and did not show any pattern along a gradient of increasing chlorophyll concentration, so there was no systematic change of bacterial growth rates with lake productivity. In contrast, growth rates scaled to the entire bacterial assemblage were positively correlated to chlorophyll, were tenfold more variable among lakes than growth rates of active cells, and showed larger differences among lake strata. Scaling bacterial production to either the total number or the number of active cells thus results in very different patterns in bacterial growth rates among aquatic systems. Received: 12 July 1996; Accepted: 24 September 1996     

Nutrient enrichment and nutrient regeneration stimulate bacterioplankton growth

Bacterial abundance results from predatory losses of individuals and replacement of losses through growth. Growth depends on sustained input of organic substrates and mineral nutrients. In this work we tested the hypothesis that bacterial growth in two oligotrophic Canadian shield lakes was limited by nitrogen (N) or phosphorus (P). We also determined whether consumer-regenerated resources contributed substantially to net bacterial growth. Two types of dilution assays were conducted to determine the response of bacteria to nutrient enrichment: diluted whole water (DWW, 1:9 whole/filtered with 0.2 m of filtered lake water) and diluted fractionated water (DFW, 1.0 m prefiltered then diluted as above). Replicate bottles in each dilution assay received either N (50 m), P (10 m), or both N and P enrichments. Controls received no nutrients. Resource-saturated growth rates and grazing rates were estimated from a standard dilution-growth approach. Bacterial growth was stimulated by addition of P alone and in combination with N. Consumers regenerated sufficient resources to support up to half the bacterial growth rate, but the benefit derived from consumers was minor when compared to mortality.     

Application of Traditional and Phylogenetically Based Comparative Methods to Test for a Trade-off in Bacterial Growth Rate at Low versus High Substrate Concentration

Abstract It is often hypothesized that those organisms that are superior competitors for sparse resources fare poorly in competition for abundant resources, and vice versa. If there is indeed such a systematic trade-off, then this has important implications for the choice of bacterial strains in bioremediation and other applications. We studied seven bacterial strains that can grow on either 2,4-dichlorophenoxyacetate (2,4-D) or succinate as a sole source of carbon. Growth rates were measured on each substrate at both low (5 μg/ml) and high (500 μg/ml) concentrations. We used two different methods to test the significance of correlations among growth rates, a traditional method that treats each strain as an independent observation and a newer method that takes into account phylogenetic relationships between strains, thereby avoiding spurious correlations caused by a lack of statistical independence of strains. In both 2,4-D and succinate, we observed significant positive correlations between growth rates measured at high and low substrate concentrations by the traditional comparative method. No significant correlations were detected after adjusting for the phylogenetic relationships among the strains. In neither case did we observe the negative correlation expected from a trade-off between growth rates at high and low substrate levels. Received: 5 April 1999; Accepted: 8 July 1999     

Influence of Three Contrasting Detrital Carbon Sources on Planktonic Bacterial Metabolism in a Mesotrophic Lake

Abstract Lakes receive organic carbon from a diversity of sources which vary in their contribution to planktonic microbial food webs. We conducted a mesocosm study to test the effects of three different detrital carbon sources (algae, aquatic macrophytes, terrestrial leaves) on several measures of microbial metabolism in a small meso-eutrophic lake (DOC ≈ 5 mg/L). Small DOC additions (ΔC < 1 mg/L) affected bacterial numbers, growth, and pathways of carbon acquisition. Macrophyte and leaf detritus significantly increased TDP and color, but bacterial densities initially (+12 h) were unaffected. After 168 h, densities in systems amended with terrestrial detritus were 60% less than in controls, while production rates in mesocosms with macrophyte detritus were 4-fold greater. Detritus treatments resulted in greater per-cell production rates either through stable cell numbers and greater growth rates (macrophyte-C) or lower densities with stable production rates (terrestrial-C). After only 12 h, rates of leucine aminopeptidase (LAPase) activity were 2.5× greater in macrophyte-C systems than in controls, but LAPase and β-N-acetylglucosamindase activities in systems amended with terrestrial-C were only 50% of rates in controls. After 168 h, β-xylosidase rates were significantly greater in communities with terrestrial and phytoplankton detritus. Microbial utilization of >20% of 102 carbon sources tested were affected by at least one detritus addition. Macrophyte-C had positive (6% of substrates) and negative (14%) effects on substrate use; terrestrial detritus had mainly positive effects. An ordination based on carbon-use profiles (+12 h) revealed a cluster of macrophyte-amended communities with greater use of psicose, lactulose, and succinamic acid; controls and algal-detritus systems were more effective in metabolizing two common sugars and cellobiose. After 168 h, communities receiving terrestrial detritus were most tightly clustered, exhibiting greater use of raffinose, pyroglutamic acid, and sebacic acid. Results suggest that pelagic bacterial communities respond to changes in organic carbon source rapidly and by different routes, including shifts in per-cell production rates and variations in degradation of a variety of compounds comprising the DOC pool. Received: 5 June 1998; Accepted: 24 August 1998     

The Formation of Large Bacterial Aggregates at Depth Within the Louisiana Hydrocarbon Seep Zone

Abstract Growth rate determinations on the individual components of size-fractionated water samples taken within the Louisiana hydrocarbon seep zone and westward of this region revealed that most of the biomass in the water was associated with particles larger than 64 μm (>60%). The aggregates in the seep zone were metabolically active and growing; whereas, 80 km to the west, the microbiota on the aggregates had negative growth rates, indicating a cessation of metabolic activity. This suggests that the raw materials for bacterial growth are the hydrocarbons, primarily methane, and that these particles may potentially provide a valuable food resource for regional benthic organisms. The large-scale geographic distribution of hydrocarbon seeps and the westward movement of water along the Louisiana slope suggest a nutritional resource capable of supporting a diverse marine heterotrophic community. Received: 15 August 1995; Revised: 14 May 1996     

Effects of Carbon Source, Carbon Concentration, and Chlorination on Growth Related Parameters of Heterotrophic Biofilm Bacteria

Abstract To investigate growth of heterotrophic biofilm bacteria, a model biofilm reactor was developed to simulate a drinking water distribution system. Controlled addition of three different carbon sources (amino acids, carbohydrates, and humics) at three different concentrations (500, 1,000, and 2,000 ppb carbon) in the presence and absence of chlorine were used in separate experiments. An additional experiment was run with a 1:1:2 mixture of the above carbon sources. Biofilm and effluent total and culturable cells in addition to total and dissolved organic carbon were measured in order to estimate specific growth rates (SGRs), observed yields, population densities, and bacterial carbon production rates. Bacterial carbon production rates (μg C/L day) were extremely high in the control biofilm communities (range = 295–1,738). Both growth rate and yield decreased with increasing carbon concentrations. Therefore, biofilm growth rates were zero-order with respect to the carbon concentrations used in these experiments. There was no correlation between growth rate and carbon concentration, but there was a significant negative correlation between growth rate and biofilm cell density (r=−0.637, p= 0.001 control and r=−0.57, p= 0.021 chlorinated biofilms). Growth efficiency was highest at the lowest carbon concentration (range = 12–4.5%, amino acids and humics respectively). Doubling times ranged from 2.3–15.4 days in the control biofilms and 1–12.3 days in the chlorinated biofilms. Growth rates were significantly higher in the presence of chlorine for the carbohydrates, humics, and mixed carbon sources (p= 0.004, < 0.0005, 0.013, respectively). The concept of r/K selection theory was used to explain the results with respect to specific growth rates and yields. Humic removal by the biofilm bacteria (78% and 56% for the control and chlorinated biofilms, respectively) was higher than previously reported literature values for planktonic bacteria. A number of control experiments indicated that filtration of drinking water was as effective as chlorination in controlling bacterial biofilm growth. Received: 26 March 1999; Accepted: 3 August 1999; Online Publication: 15 February 2000     

Effects of carbohydrates and leucine on growth of bacteria from Antarctic soils (Casey Station,Wilkes Land)

Growth rates of natural bacterial communities from Antarctic soils are analysed by an epifiuorescence microscopic method using data of microcolonies (colony number and colony areas). Incubations are performed on polycarbonate filters which are put on cellulose pads soaked with soil extracts, different concentrations of naturally occuring carbohydrates, polyols, and leucine. Concentrations of individual substrates were in the range of naturally occuring levels. The results showed that the growth of bacterial microcolonies could best be stimulated with glucose, sucrose, maltose, sorbitol, and mannitol. Leucine stimulated growth to a lower extent than glucose. Data on bacterial biomass production calculated from this approach are discussed in relation to those from tracer techniques carried out with ¹⁴C-labelled glucose from earlier experiments.     

Marine Heterotrophic Bacteria in Continuous Culture,the Bacterial Carbon Growth Efficiency,and Mineralization at Excess Substrate and Different Temperatures

To model the physiological potential of marine heterotrophic bacteria, their role in the food web, and in the biogeochemical carbon cycle, we need to know their growth efficiency response within a matrix of different temperatures and degrees of organic substrate limitation. In this work, we present one part of this matrix, the carbon growth efficiencies of marine bacteria under different temperatures and nonlimiting organic and inorganic substrate supply. We ran aerobic turbidostats with glucose enriched seawater, inoculated with natural populations of heterotrophic marine bacteria at 10, 14, 18, 22, and 26°C. The average cell-specific growth rates increased with temperature from 1.17 to 2.6 h−1. At steady-state total CO₂ production, biomass production [particulate organic carbon (POC) and nitrogen (PON)], and viruslike particle abundance was measured. CO₂ production and specific growth rate increased with increasing temperature. Bacterial carbon growth efficiency (BCGE), the particulate carbon produced per dissolved carbon utilized, varied between 0.12 and 0.70. Maximum BCGE values and decreased specific respiration rates occurred at higher temperatures (22 and 26°C) and growth rates. This trend was largely attributable to an increase in POC per cell abundance; when the BCGE was recalculated, parameterizing the biomass as the product of cell concentration and a constant cellular carbon content, the opposite trend was observed.     

Microbial attachment to particles in marine and freshwater ecosystems

Scanning electron microscopy observations ofin situ suspended marine and freshwater particles show diverse but similar modes of bacterial and fungal attachment. A survey of Sierra Nevada mountain lakes and pelagic and near-shore waters in the Pacific Ocean indicates that attachment is most noticeable in the near-surface waters where fresh dissolved and particulate input of carbon from phytoplankton and elevated temperatures favor microbial growth. The most common modes of attachment are: adhesive stalk formation, growth on adhesive webs, attachment by the use of pili-like appendages and slimy capsular secretions, and molecular or chemical sorption without the use of visualized structural appendages. Attached microbial growth is accelerated when particulate substrates are supplied, even when they are not rich in organic nutrients. This is the case in the Lake Tahoe basin, where microflora attached to eroded silts can significantly modify the organic carbon and nutrient content of such minerogenous particles.     

Moving Waves of Bacterial Populations and Total Organic Carbon along Roots of Wheat

Abstract To determine if spatial variation in soluble carbon sources along the root coincides with different trophic groups of bacteria, copiotrophic and oligotrophic bacteria were enumerated from bulk soil and rhizosphere samples at 2 cm intervals along wheat roots 2, 3, and 4 weeks after planting. There was a moderate rhizosphere effect in one experiment with soil rich in fresh plant debris, and a very pronounced rhizosphere effect in the second experiment with soil low in organic matter. We obtained wavelike patterns of both trophic groups of bacteria as well as water-soluble total organic carbon (TOC) along the whole root length (60 or 90 cm). TOC concentrations were maximal at the root tip and base and minimal in the middle part of the roots. Oscillations in populations of copiotrophic and oligotrophic bacteria had two maxima close to the root tip and at the root base, or three maxima close to the tip, in the middle section, and at the root base. The location and pattern of the waves in bacterial populations changed progressively from week to week and was not consistently correlated with TOC concentrations or the location of lateral root formation. Thus, the traditional view that patterns in bacterial numbers along the root directly reflect patterns in exudation and rhizodeposition from several fixed sources along the root may not be true. We attributed the observed wavelike patterns in bacterial populations to bacterial growth and death cycles (due to autolysis or grazing by predators). Considering the root tip as a moving nutrient source, temporal oscillations in bacterial populations at any location where the root tip passed would result in moving waves along the root. This change in concept about bacterial populations in the rhizosphere could have significant implications for plant growth promotion and bioremediation. Received: 11 May 1998; Accepted: 4 November 1998     

Response of Heterotrophic Planktonic Bacteria to the Zebra Mussel Invasion of the Tidal Freshwater Hudson River

Abstract Invasions of aquatic ecosystems by exotic bivalves are known to cause dramatic changes in phytoplankton and some other groups, but their effect on the microbial component is unknown. The invasion of the tidal freshwater Hudson River by the exotic zebra mussel (Dreissena polymorpha) has caused large changes in several components of the Hudson's food web. Planktonic bacteria in the tidal freshwater Hudson are a major part of the food web, and mediate important processes in the carbon budget. We used a long-term data set, spanning four years prior to the zebra mussel (ZM) invasion and four years post-invasion, to describe ZM effects on planktonic bacteria. Small and meso-scale experiments were conducted to specifically examine direct consumption of bacteria by ZM, as well as effects on protozoans. Bacterial abundances in the Hudson have increased roughly 2× since the ZM arrived, making it clear that direct consumption by Dreissena is a minor process. Experiments show that ZM do not remove bacteria from Hudson River water, but are very effective at clearing flagellated protozoans, the major predator of bacteria. The observed changes in bacterial abundance have not been accompanied by equally large changes in bacterial productivity, suggesting growth is primarily limited by carbon supply. Bacterial production has not declined despite a dramatic decline of phytoplankton, confirming previous suggestions that bacteria and phytoplankton are not strongly linked in the Hudson. As a result of the increase in bacterial abundance and removal of phytoplankton, the absolute and relative contributions of bacterial carbon to living particulate organic carbon (POC) standing stocks have increased dramatically. The maintenance of the bacterial component of the Hudson River's food web may be one mechanism whereby consumers are ``insulated' from effects of zebra mussel consumption of phytoplankton carbon. Received: 24 October 1997; Accepted: 9 February 1998     

Differing growth responses of major phylogenetic groups of marine bacteria to natural phytoplankton blooms in the western North Pacific Ocean

Growth and productivity of phytoplankton substantially change organic matter characteristics, which affect bacterial abundance, productivity, and community structure in aquatic ecosystems. We analyzed bacterial community structures and measured activities inside and outside phytoplankton blooms in the western North Pacific Ocean by using bromodeoxyuridine immunocytochemistry and fluorescence in situ hybridization (BIC-FISH). Roseobacter/Rhodobacter, SAR11, Betaproteobacteria, Alteromonas, SAR86, and Bacteroidetes responded differently to changes in organic matter supply. Roseobacter/Rhodobacter bacteria remained widespread, active, and proliferating despite large fluctuations in organic matter and chlorophyll a (Chl-a) concentrations. The relative contribution of Bacteroidetes to total bacterial production was consistently high. Furthermore, we documented the unexpectedly large contribution of Alteromonas to total bacterial production in the bloom. Bacterial abundance, productivity, and growth potential (the proportion of growing cells in a population) were significantly correlated with Chl-a and particulate organic carbon concentrations. Canonical correspondence analysis showed that organic matter supply was critical for determining bacterial community structures. The growth potential of each bacterial group as a function of Chl-a concentration showed a bell-shaped distribution, indicating an optimal organic matter concentration to promote growth. The growth of Alteromonas and Betaproteobacteria was especially strongly correlated with organic matter supply. These data elucidate the distinctive ecological role of major bacterial taxa in organic matter cycling during open ocean phytoplankton blooms.     

Evidence for an enhanced substrate requirement by marine mesophilic bacterial isolates at minimal growth temperatures

Bacterial isolates from the subtropical southeastern continental shelf were cultured in a matrix of temperature and substrate concentrations encompassing a range of temperature and substrate concentrations equal to and exceeding natural ones. At the annual minimum temperature, marine heterotrophic bacterial isolates required higher concentrations of dissolved substrates for active growth than are usually found in seawater. We show this to result from a nonlinear interaction of the combined effects of temperature and substrate concentration on bacterial growth and respiratory rate. As a result, bacterial and protozoan utilization of phytoplankton production during winter and early spring is low, permitting greater energy flow to zooplankton and benthic animals, while in late spring, summer, and fall, the microbial loop dominates energy flux and organic carbon utilization. Escherichia coli shows a similar nonlinear response to temperature at minimal substrate concentrations, albeit at a higher range of concentrations than were utilized by the marine isolates. Thus, bacteria from subtropical regions are shown to have a differential growth response near the minimum temperature for growth, depending on the concentration of available substrates. Offprint requests to: W.J. Wiebe.     

Factors controlling bacterial production in marine and freshwater sediments

We collected benthic bacterial production data measured by ³H thymidine incorporation (TTI) (25 studies), frequency of dividing cells (FDC) (3 studies), dark-C0₂ assimilation (1 study) and ³H-adenine uptake (2 studies) from the literature, which included 18 marine, 6 river, and 2 lake studies. In all of the studies that used the TTI method, ³H-DNA was isolated and incubations were carried out at in situ temperatures. Most of the researchers also determined ³H-DNA extraction efficiencies and isotope dilution, thus interpretable estimates of bacterial production were used in the analysis. In marine sediments, bacterial production rates were linked to bacterial biomass, bacterial abundance, sediment organic matter, temperature, and sediment chlorophyll a, with these variables explaining between 40% and 68% of the variation in production rates. Simple relationships between production and bacterial biomass or bacterial abundance, or between production and sediment organic matter, were improved by also including temperature in the analysis of marine sediments. Sediment organic matter explained an appreciable fraction (58%) of the observed production in freshwater sediments. Temperature was the most powerful predictor of the observed variability in specific growth rates (r ² = 0.48 and r ² = 0.58) in marine and freshwater sediments, respectively. Thus, bacterial production and specific growth rates are most closely linked to substrate supply and temperature in marine and freshwater sediments. Offprint requests to: B. C. Sander.     

Conductance measurements for data generation in predictive modeling

Summary The electrical resistance of a growth medium inoculated with bacteria may be automatically recorded throughout an incubation period without the necessity for sampling. The rate of change in conductance is dependent on the bacteria studied, the medium composition and the prevailing growth conditions.The effect of growth medium composition, growth conditions and inoculum level on the conductance response was studied forYersinia enterocolitica O:3. A large number of combinations of factors affecting the growth/activity of the bacteria could be studied simultaneously due to the large instrumental capacity of the Malthus 2000. A polynomial model based on conductance measurements was developed forY. enterocolitica describing the effect of temperature, pH andl-lactate level on conductance response curve parameters. The model was used for predicting growth rates. Growth rates calculated from bacterial counts ofY. enterocolitica growing in minced pork corresponded to growth rates predicted using the polynomial conductance models.     

Bacterial Growth Rate and Marine Virus–Host Dynamics

Abstract The dynamics of a marine virus–host system were investigated at different steady state growth rates in chemostat cultures and the data were analyzed using a simple model. The virus–host interactions showed strong dependence on host cell growth rate. The duration of the infection cycle and the virus burst size were found to depend on bacterial growth rate, and the rate of cell lysis and virus production were positively correlated with steady state growth rate in the cultures (r ² > 0.96, p < 0.05). At bacterial growth rates of 0.02 to 0.10 h⁻¹ in the chemostats the virus burst size increased from 12 ± 4 to 56 ± 4, and the latent period decreased from 2.0 to 1.7 h. Resistant clones of the host strain were present in the cultures from the beginning of the experiment and replaced the sensitive host cells following viral lysis in the cultures. Regrowth of resistant cells correlated significantly (r ²= 1.000, p < 0.02) with the lysis rate of sensitive cells, indicating that release of viral lysates stimulated growth of the non-infected, resistant cells. The constructed model was suitable for simulating the observed dynamics of the sensitive host cells, viruses and resistant clones in the cultures. The model was therefore used in an attempt to predict the dynamics of this virus–host interaction in a natural marine environment during a certain set of growth conditions. The simulation indicated that a steady state relationship between the specific viruses and sensitive and resistant bacterial clones may occur at densities that are reasonable to assume for natural environments. The study demonstrates that basic characterization and modeling of specific virus–host interactions may improve our understanding of the behavior of bacteria and viruses in natural systems. Received: 12 November 1999; Accepted: 2 May 2000; Online Publication: 11 August 2000     

Regulation of Bacterial Growth Rates by Dissolved Organic Carbon and Temperature in the Equatorial Pacific Ocean

Abstract The effect of dissolved organic matter (DOM) and temperature on bacterial production was examined in the equatorial Pacific Ocean. Addition of glucose, glucose plus ammonium, or free amino acids stimulated bacterial production ([³H]thymidine incorporation), whereas changes in bacterial abundance were either negligible or much less than changes in bacterial production. The average bacterial growth rate also greatly increased following DOM additions, whereas in contrast, addition of ammonium alone never affected production, bacterial abundance, or growth rates. Since the large glucose effect was not observed in previous studies of cold oceanic waters, several experiments were conducted to examine DOM-temperature interactions. These experiments suggest that bacteria respond more quickly and to a greater extent to DOM additions at higher temperatures, which may explain apparently conflicting results from previous studies. We also examined how temperate affects the kinetic parameters of sugar uptake. Maximum uptake rates (V_max) of glucose and mannose increased with temperature (Q₁₀= 2.4), although the half-saturation constant (K_m) was unaffected; K_m+ S was roughly equal to glucose concentrations (S) measured by a high pressure liquid chromographic technique. Bacterial production and growth rates appear to be limited by DOM in the equatorial Pacific, and thus bacterial production follows primary production over large spatial and temporal scales in this oceanic regime, as has been observed in other aquatic systems. Although temperature may not limit bacterial growth rates in the equatorial Pacific and similar warm waters, it could still affect how bacteria respond to changes in DOM supply and help set steady-state DOM concentrations. Received: 26 July 1995; Revised: 19 January 1996