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Microbial loop

The microbial loop is a key trophic pathway in aquatic ecosystems, especially marine environments, where dissolved organic matter (DOM) exuded by phytoplankton and other primary producers is assimilated by heterotrophic bacteria into bacterial biomass, which is subsequently grazed by protozoans such as heterotrophic nanoflagellates and ciliates, thereby recycling carbon and nutrients to higher trophic levels and integrating microbial processes with the classical plankton food chain.^[1] This loop highlights the dominant role of microbes in processing a substantial fraction of primary production, often 10–50% of photosynthetically fixed carbon, before it reaches larger grazers like zooplankton.^[1]^[2] The concept of the microbial loop was formally articulated in 1983 by Azam et al., who emphasized the ecological significance of water-column microbes in nutrient cycling and energy flow, building on earlier 20th-century observations of bacterial roles in organic matter decomposition dating back to the 1930s in Russian and Danish marine microbiology.^[1]^[3] Prior paradigms had largely overlooked microbes, focusing on direct grazing of phytoplankton by metazoans, but the microbial loop paradigm shift revealed how bacteria and their predators mediate rapid turnover of DOM, preventing its loss and sustaining productivity in nutrient-limited waters.^[4] Over the decades, the framework has evolved to incorporate viral lysis and other microbial interactions, yet the core loop remains central to understanding marine biogeochemistry.^[5] Key components include heterotrophic bacteria (typically 0.3–1 µm in size, with abundances up to 5–10 × 10⁶ cells ml⁻¹), which efficiently uptake low-molecular-weight DOM at concentrations as low as nanomolar levels; heterotrophic flagellates (3–10 µm, up to 3 × 10³ cells ml⁻¹), acting as primary bacterivores; and microzooplankton (10–80 µm), which consume these protists and link the loop to metazoan grazers.^[1] The process involves bacterial production of biomass from DOM, protozoan grazing that remineralizes nutrients like nitrogen and phosphorus, and efficient transfer efficiencies often exceeding 30% at each step, contrasting with lower efficiencies in the classical chain.^[6] In various marine systems, the loop can recycle up to 85% of net primary production—for example, in the English Channel—thereby retaining nutrients in the euphotic zone and fueling secondary production. In oligotrophic oceans, such recycling can reach ~86%, as observed in regions like the Sargasso Sea.^[6]^[7] Ecologically, the microbial loop is pivotal for global carbon cycling, as marine microbes fix approximately 50 gigatons of carbon annually—comparable to terrestrial net primary production—and the loop ensures much of this carbon is cycled internally rather than exported, influencing atmospheric CO₂ levels and oxygen production, which accounts for about 50% of the planet's total.^[2] It supports fisheries by channeling energy to harvestable species and modulates climate feedbacks through interactions with the microbial carbon pump, which sequesters refractory DOM for millennia.^[2] Disruptions, such as from ocean warming or acidification, could alter loop dynamics, potentially reducing carbon retention and biodiversity.^[4]

Overview

Definition and Components

The microbial loop refers to a trophic pathway in aquatic ecosystems, particularly marine environments, where dissolved organic matter (DOM) released primarily by phytoplankton is assimilated by heterotrophic bacteria, which in turn are grazed upon by protozoan consumers such as heterotrophic nanoflagellates and ciliates, thereby recycling organic carbon and nutrients while largely bypassing the classical herbivore-dominated food chain.^[1] This pathway highlights the central role of microbes in processing a significant portion of primary production that would otherwise remain inaccessible to higher trophic levels.^[1] The primary components of the microbial loop include DOM as the initial substrate, heterotrophic bacteria as decomposers, protozoan grazers, and viruses as lytic agents. DOM, comprising 5-50% of photosynthetically fixed carbon, serves as the entry point, originating from phytoplankton exudates, sloppy feeding, and cell lysis.^[1] Heterotrophic bacteria rapidly uptake and mineralize this low-molecular-weight DOM, converting it into bacterial biomass that can constitute a substantial fraction of total planktonic biomass.^[1] Heterotrophic nanoflagellates (typically 3-10 µm) and ciliates act as key grazers, efficiently filtering bacteria and controlling their populations through predation, with flagellates alone capable of processing a large volume of seawater daily.^[1] Viruses, predominantly bacteriophages, integrate into the loop by infecting and lysing bacterial cells, releasing intracellular contents—including DOM and nutrients—back into the water column, which sustains further bacterial growth and influences overall carbon flux.^[8] Conceptually, the loop can be outlined as a cycle: DOM → bacterial uptake and growth → protozoan grazing (releasing DOM and nutrients via excretion and sloppy feeding) and viral lysis (releasing DOM) → renewed DOM availability for bacteria.^[1]^[8] This contrasts with the classic food chain, where energy flows directly from primary producers to herbivores and then to carnivores; instead, the microbial loop mediates much of the carbon and nutrient turnover through rapid microbial interactions, often retaining 10-50% of fixed carbon within the microbial compartment before transfer to larger grazers.^[1] The microbial loop thus plays a pivotal role in global biogeochemical cycles by enhancing nutrient regeneration and organic matter recycling in the water column.^[1]

Ecological Role

The microbial loop plays a pivotal role in oceanic carbon cycling by channeling a substantial fraction of primary production through heterotrophic bacteria, which assimilate dissolved organic carbon (DOC) derived from phytoplankton exudates and mortality. In oligotrophic systems, where nutrient scarcity limits direct grazing on phytoplankton, bacterial production can account for 10–50% of net primary production, effectively recycling DOC and preventing its loss while regenerating bioavailable carbon for higher trophic levels.^[1] This process enhances overall carbon retention in the surface ocean, with estimates indicating that bacteria process approximately half of marine primary production globally, thereby influencing the efficiency of carbon transfer within planktonic food webs.^[9] Through bacterial uptake of DOC and subsequent grazing by protists, the microbial loop drives efficient nutrient recycling, particularly for nitrogen and phosphorus, which are remineralized into inorganic forms readily usable by phytoplankton. Bacteria rapidly incorporate dissolved organic nitrogen and phosphorus, converting them into biomass, while protistan grazers release ammonium and phosphate via sloppy feeding and excretion, closing the loop and sustaining primary productivity in nutrient-limited environments.^[1] This remineralization pathway ensures that up to 25% of respired carbon is accompanied by nutrient release, amplifying nutrient availability without relying on external inputs.^[1] The microbial loop supports microbial biodiversity by fostering dynamic interactions among bacteria, viruses, and protists, where grazing pressure prevents any single group from dominating and promotes diverse community structures that underpin aquatic food webs. This diversity at the base of the trophic pyramid facilitates energy and biomass transfer to higher levels, such as zooplankton and fish, enhancing overall ecosystem resilience.^[9] On a global scale, the loop contributes to ocean carbon sequestration by partitioning some processed carbon into recalcitrant DOC that evades rapid respiration, potentially accounting for 10–30% of total oceanic heterotrophic respiration and modulating atmospheric CO₂ drawdown through the biological pump.^[10]

Historical Development

Early Discoveries

In the 19th century, foundational observations on microbial roles in aquatic environments emerged from studies on decomposition processes. Louis Pasteur's experiments demonstrated that bacteria were responsible for putrefaction in water-based media, challenging prevailing notions of spontaneous generation and highlighting microbes as active agents in breaking down organic matter.^[11] These findings, detailed in Pasteur's 1861 memoir on organized corpuscles in the atmosphere and their relation to fermentation and putrefaction, established that airborne microbes could initiate decomposition in sterile liquids, laying early groundwork for understanding bacterial contributions to nutrient cycling in water.^[11] The early 20th century saw further advancements, particularly from the 1930s onward, when Russian marine bacteriologists such as Yurii Sorokin pioneered studies on bacterial decomposition of dissolved organic matter (DOM) in seawater, using direct microscopic counts to quantify microbial activity and demonstrate bacteria's role as intermediaries in nutrient cycling.^[3] Western researchers, including Selman Waksman and Claude ZoBell, employed culture-based methods like agar plate colony counts to assess bacterial abundances and their processing of organic substrates. By the 1960s, Mikhail Vinogradov's group in Russia developed the first numerical models incorporating microbial components into marine ecosystem dynamics, emphasizing bacteria's integration into food webs. These efforts, building on Danish and other European observations of marine microbiology, highlighted the significance of microbes in oceanic carbon and nutrient flows, setting the stage for later conceptual frameworks.^[3] During the 1970s, research in plankton ecology began to quantify bacterial uptake of dissolved organic matter (DOM) in seawater, revealing bacteria as primary consumers of this material. Farooq Azam and colleagues conducted studies showing that heterotrophic bacteria efficiently incorporated labile DOM, such as dissolved ATP, into their biomass, with uptake rates indicating that bacteria could process a significant portion of oceanic DOM pools.^[12] These investigations, based on seawater samples, demonstrated that bacterial metabolism linked phytoplankton-derived DOM to higher trophic levels, underscoring microbes' central role in marine carbon and nutrient dynamics. A pivotal 1977 study by Azam and R.E. Hodson, focusing on the Sargasso Sea, further illuminated interactions between bacteria and protozoa as key microheterotrophs. Analyzing size distributions and activities, the research found that bacteria dominated DOM consumption, while bacterivorous protozoa grazed on them, facilitating nutrient regeneration and hinting at a coupled microbial pathway in oligotrophic waters.^[13] This work provided empirical evidence from Sargasso Sea samples that protozoan predation on bacteria could recycle nutrients back to phytoplankton, prefiguring integrated microbial food web concepts. In parallel, limnological studies in the 1970s introduced precursor ideas to the "bacterial loop" in freshwater systems. Johannes Overbeck's research on lake ecosystems demonstrated that bacteria were crucial for the uptake and transformation of DOM, acting as intermediaries between algal exudates and higher trophic levels in planktonic communities.^[14] Overbeck showed that bacterial activity sustained a significant fraction of secondary production, emphasizing microbes' role in closing nutrient cycles in inland waters.^[15]

Key Milestones and Researchers

The concept of the microbial loop began to take shape in the 1970s with Lawrence R. Pomeroy's proposal of a "microbial garden," which emphasized the role of heterotrophic microorganisms in processing dissolved organic matter (DOM) and recycling nutrients in marine ecosystems, challenging the traditional view of a linear plankton food chain dominated by phytoplankton-zooplankton interactions.^[16] This idea evolved into a more formalized framework in the 1980s, when Farooq Azam, Tom Fenchel, and colleagues introduced the term "microbial loop" in a seminal 1983 paper, describing how bacteria assimilate DOM from primary production, are grazed by protists, and thereby channel energy back into higher trophic levels while also facilitating nutrient regeneration. Key researchers advanced this concept through focused studies on its components. Farooq Azam pioneered measurements of bacterial production and its linkage to DOM uptake, establishing bacteria as central processors in the loop.^[5] David L. Kirchman contributed extensively to understanding protist grazing on bacteria, quantifying how flagellates and ciliates control bacterial populations and influence carbon transfer efficiency within the loop. Curtis A. Suttle, starting in the 1990s, highlighted the role of viruses in microbial dynamics, demonstrating their impact on bacterial mortality and organic matter cycling.^[17] Major milestones in the 1990s included the integration of viral processes via the "viral shunt," proposed by Steven W. Wilhelm and Curtis A. Suttle, which showed that viral lysis of bacteria releases DOM and nutrients, bypassing grazing and promoting rapid recycling but reducing transfer to metazoans.^[17] Jed A. Fuhrman contributed foundational work on marine viral ecology in the 1990s, supporting these developments.^[18] In the 2000s, genomic approaches provided deeper insights into microbial diversity, as exemplified by J. Craig Venter and colleagues' 2004 metagenomic survey of the Sargasso Sea, which uncovered over 1,800 microbial species and revealed the genetic basis for diverse metabolic functions supporting loop processes in oligotrophic waters. Early adoption of the microbial loop faced skepticism regarding its efficiency, particularly debates over whether it primarily links carbon to higher trophic levels or acts as a sink via respiration, with evidence suggesting greater importance in oligotrophic systems compared to eutrophic ones where classical food chains prevail.

Core Processes

Bacterial Production and Grazing

Heterotrophic bacteria form the foundational step in the microbial loop by assimilating dissolved organic matter (DOM) into new biomass through the process of bacterial production (BP). This uptake converts low-molecular-weight DOM, primarily released from primary producers or other sources, into bacterial cellular material, thereby repackaging it for higher trophic levels. The efficiency of this conversion, known as bacterial growth efficiency (BGE), typically ranges from 20% to 50% in natural aquatic systems, reflecting the proportion of assimilated DOM incorporated into biomass versus that respired as CO₂. BGE = BP / (BP + BR), where BR denotes bacterial respiration, and BP itself can be expressed as BP = DOM uptake × growth yield, highlighting the direct linkage between substrate assimilation and biomass accrual.^[19]^[20] To quantify bacterial production rates, researchers commonly employ isotopic incorporation techniques. The thymidine incorporation method, introduced by Fuhrman and Azam, measures the rate of DNA synthesis by tracking the uptake of tritiated thymidine (³H-thymidine) into bacterial cells, providing an estimate of cell division and thus production. Complementarily, the leucine incorporation method assesses protein synthesis via the uptake of tritiated or ¹⁴C-labeled leucine, offering a robust alternative particularly suited for diverse bacterial communities. These methods have become staples for estimating BP in situ, with conversion factors calibrated to yield carbon production values.^[21]^[22] Bacterial production is tightly regulated by protistan grazing, primarily from heterotrophic nanoflagellates (2–5 μm) and ciliates (10–50 μm), which serve as the dominant bacterivores in the microbial loop. These predators exhibit size-selective grazing, favoring bacteria under 5 μm in diameter while largely avoiding larger cells or filaments that exceed this threshold, thereby shaping bacterial community structure and promoting morphological defenses in prey populations. Clearance rates—the volume of water cleared of bacteria per predator per hour—typically fall between 0.1 and 1 nl cell⁻¹ h⁻¹ for nanoflagellates, enabling them to consume substantial fractions of standing bacterial stocks daily. Ciliates contribute similarly but often at lower individual rates due to their larger size.^[23]^[24]^[25] Grazing facilitates trophic transfer within the microbial loop, with approximately 30–50% of bacterial production incorporated into protist biomass and passed to higher trophic levels, such as zooplankton, while the remainder is respired or excreted back into the DOM pool. This transfer efficiency underscores the loop's role in channeling energy from DOM to metazoans, though protist respiration and sloppy feeding can recycle up to 50% of grazed carbon as DOM, sustaining further bacterial growth. Such dynamics ensure that bacterivory not only controls bacterial abundances but also amplifies nutrient cycling efficiency in aquatic ecosystems.^[26]^[27]

Viral and Dissolution Pathways

In the microbial loop, the viral shunt represents a key non-predatory pathway for recycling organic matter, where bacteriophages infect and lyse bacterial cells, releasing cellular contents as dissolved organic matter (DOM) and colloidal particles that become available for uptake by other microbes. This process bypasses direct trophic transfer to grazers, instead channeling nutrients and carbon back into the lower levels of the food web, thereby influencing biogeochemical cycles in aquatic ecosystems. Lysis typically results in 25-50% of the lysed cell's carbon being released as labile DOM, with the remainder forming particulate or colloidal fractions that can aggregate or dissolve further.^[28] Viruses exert substantial control over bacterial populations, infecting 20-40% of marine bacteria daily and contributing to mortality rates that can reach up to 50% of daily bacterial production in some systems. Viral production, driven primarily by lytic cycles, accounts for approximately 10-25% of total bacterial production, with rates varying by environmental conditions such as nutrient availability and temperature. The impact of viral mortality on bacterial communities can be estimated using the equation for viral mortality:

\text{Viral mortality} = \text{VLP} \times \text{burst size} \times \text{infection rate}

where VLP denotes the abundance of virus-like particles, burst size is the average number of virions released per infected cell (typically 10-50), and infection rate reflects the fraction of susceptible hosts encountered per unit time. This dynamic ensures efficient recycling while preventing excessive accumulation of bacterial biomass.^[28]^[29]^[30] Complementing viral lysis, dissolution pathways involve the chemical and enzymatic breakdown of dead or moribund microbial cells, contributing to the DOM pool through passive leakage and autolysis. These processes release low-molecular-weight compounds that are rapidly assimilated by surviving bacteria, sustaining the microbial loop without the need for active infection. In oligotrophic environments, such dissolution enhances the bioavailability of refractory DOM, supporting basal production.^[31] From an evolutionary perspective, ongoing coevolution between phages and bacteria promotes microbial diversity by selectively targeting dominant strains, as encapsulated in the "killing-the-winner" hypothesis, which prevents any single bacterial genotype from monopolizing resources. This arms-race dynamic fosters genetic variation in bacterial defenses, such as CRISPR systems, and phage counter-adaptations, maintaining community stability and resilience within the microbial loop. Phage-mediated selection thus acts as a counterbalance to bacterial proliferation, ensuring diverse assemblages that underpin ecosystem function.^[32]

Influencing Factors

Environmental Controls

Temperature exerts a significant control on the microbial loop through its influence on bacterial production rates, with Q10 values typically ranging from 2 to 3 in marine systems, indicating that production approximately doubles or triples for every 10°C rise within suitable ranges.^[33] This temperature sensitivity arises from enzymatic processes in heterotrophic bacteria, where lower temperatures slow metabolism while extremes inhibit growth. In temperate marine environments, bacterial production peaks at 15–25°C, reflecting an optimal range for most marine bacterioplankton before heat stress reduces efficiency above 25°C.^[34] Nutrient availability modulates the microbial loop by constraining bacterial uptake of dissolved organic matter (DOM), with phosphorus (P) and nitrogen (N) often acting as key limiters in oligotrophic waters.^[35] When ambient concentrations fall below cellular demands, bacteria exhibit reduced growth and altered DOM assimilation, shifting the loop's carbon flux. Stoichiometric imbalances, particularly deviations from the Redfield ratio (C:N:P ≈ 106:16:1), further impact efficiency; for instance, P limitation can suppress bacterial production even with excess carbon, promoting co-limitation scenarios that bottleneck energy transfer to higher trophic levels.^[36] Light availability in surface waters drives photodegradation of DOM, breaking down complex molecules into more labile forms that enhance bioavailability for bacterial consumption and thereby accelerate the microbial loop.^[37] In contrast, low oxygen conditions, such as those in hypoxic zones, compel bacteria to shift from aerobic respiration to fermentation, yielding less ATP per glucose molecule and reducing overall loop efficiency while favoring acid production.^[38] pH influences protist grazing, a critical step in the loop, with optimal activity at 7–8, where ocean acidification (projected pH drop to ~7.8 by 2100) diminishes nanoflagellate growth and predation rates on bacteria.^[39] Salinity optima for marine protist grazing and bacterial processes align with oceanic norms of 30–35 practical salinity units (PSU), beyond which osmotic stress disrupts community dynamics and grazing efficiency.^[40]

Biological Regulations

The microbial loop is subject to various biotic regulations that influence its efficiency and stability through interspecies interactions. Beyond protistan grazing on bacteria, phytoplankton compete with heterotrophic bacteria for labile dissolved organic matter (DOM), particularly low-molecular-weight compounds released during phytoplankton blooms, which can limit bacterial access to this resource and alter carbon partitioning in the loop.^[41] Additionally, metazoan grazers, such as copepods and cladocerans, exert predation pressure on heterotrophic protists, reducing protist populations and thereby indirectly alleviating grazing on bacteria, which can enhance bacterial biomass accumulation within the loop.^[42] These interactions highlight how higher trophic levels modulate the flow of organic matter by targeting key grazers in the microbial compartment.^[4] Symbiotic relationships between bacteria and phytoplankton also play a critical role, where associated bacteria facilitate enhanced access to recalcitrant DOM fractions through extracellular enzyme production, allowing phytoplankton to recycle nutrients more effectively and supporting overall loop dynamics in nutrient-limited environments.^[43] These symbioses exemplify mutualistic feedbacks that stabilize carbon transfer from primary producers to decomposers.^[44] Top-down control from higher trophic levels, such as zooplankton, indirectly regulates the microbial loop by suppressing protist abundances through selective grazing on bacterivorous flagellates and ciliates, which in turn reduces predation pressure on bacteria and promotes microbial carbon cycling efficiency.^[4] This cascade effect is evident in mesocosm studies where increased zooplankton biomass shifts community structure, favoring smaller protists and altering the balance of bacterial production versus consumption in the loop.^[4] Higher microbial diversity contributes to the stability of the microbial loop by invoking the redundancy hypothesis, wherein functionally equivalent taxa compensate for losses in key populations, maintaining consistent rates of DOM processing and nutrient regeneration despite perturbations.^[45] This functional redundancy ensures resilient ecosystem processes, as diverse bacterial assemblages sustain decomposition efficiency even when dominant species are impacted by grazing or competition.^[45]

Ecosystem Applications

Marine Systems

In marine systems, the microbial loop plays a dominant role in carbon cycling, particularly in oligotrophic oceans where nutrient scarcity limits larger trophic pathways. In open ocean gyres such as the Sargasso Sea, bacteria and associated nanozooplankton account for approximately 70% of total heterotrophic carbon biomass in the photic zone, channeling a substantial portion of primary production—often up to 70-80%—through microbial processes rather than direct grazing by larger zooplankton.^[46]^[47] This dominance arises from the high efficiency of bacteria in assimilating dissolved organic matter (DOM) released by phytoplankton, which constitutes the primary energy source in these low-productivity environments, thereby retaining carbon within the surface layer and minimizing export to deeper waters.^[10] Vertically, the microbial loop exhibits distinct patterns across ocean depths, reflecting variations in organic matter availability. In surface waters, phytoplankton exudates and cell lysis provide abundant labile DOM, fueling elevated bacterial production and protozoan grazing that enhance loop activity and recycle nutrients efficiently.^[48] In contrast, deep-sea communities rely more heavily on refractory DOM and organic carbon from sinking particles, such as marine snow, which support slower but persistent microbial degradation and remineralization, contributing to the vertical flux of carbon.^[49] This stratification underscores the loop's role in modulating the biological carbon pump, with surface enhancement promoting retention and deep-sea processes facilitating gradual sequestration. Case studies illustrate the microbial loop's integration with broader marine productivity. In the Ross Sea, a high-latitude Antarctic polynya, the loop links phytoplankton blooms to bacterial and viral dynamics, where bacterial production is equivalent to about 25–30% of primary production, sustaining secondary production and influencing seasonal carbon export during productive summer periods.^[50] In coral reef ecosystems, microbial communities mediate nutrient cycling by processing DOM from algal symbionts and detritus, recycling nitrogen and phosphorus to support reef productivity and prevent nutrient limitation in oligotrophic tropical waters.^[51] Climate change poses significant threats to the microbial loop in marine systems, primarily through warming-induced shifts in respiration. Elevated temperatures accelerate bacterial respiration rates, increasing carbon mineralization and potentially reducing export efficiency by 10-20% in surface oceans, as more organic matter is respired as CO₂ rather than sinking as particles.^[52]^[53] This enhanced loop activity could diminish the ocean's capacity as a carbon sink, exacerbating atmospheric CO₂ accumulation.^[54]

Terrestrial Systems

In terrestrial ecosystems, the microbial loop functions within the heterogeneous matrix of soils, where bacteria and fungi decompose plant litter and other organic inputs into dissolved and particulate forms. These microbes incorporate carbon and nutrients into their biomass, which is then grazed by microfaunal predators such as nematodes and amoebae, accelerating nutrient mineralization—particularly nitrogen—and channeling resources back to plants. This process enhances overall soil nutrient cycling and supports primary production in nutrient-limited environments.^[55] A key hotspot for the microbial loop is the rhizosphere, the soil zone influenced by plant roots, where root exudates provide a rich source of dissolved organic matter (DOM), comprising up to 40% of a plant's photosynthetically fixed carbon. This DOM influx stimulates bacterial growth, resulting in rhizosphere bacterial biomass that can be 30-fold higher than in bulk soil, while fungi assimilate 10–20% of net fixed carbon to fuel decomposition and symbiotic interactions. Protozoan grazing in this zone further amplifies nutrient release, such as ammonia, promoting plant growth through increased root proliferation and nutrient uptake efficiency.^[55] The dynamics of the microbial loop adapt to varying terrestrial conditions, with higher activity in moist forest soils due to consistent water availability and abundant organic matter, fostering robust bacterial-fungal decomposition and faunal grazing. In arid and desert soils, loop processes are constrained by desiccation, leading to microbial and protozoan dormancy during dry periods; reactivation occurs with episodic wetting, as amoebae exploit thin water films for grazing, though overall turnover rates remain lower than in forests. Moisture thus serves as a primary environmental control on loop efficiency.^[55]^[56] Agricultural practices can impair the microbial loop, particularly through pesticide applications that reduce its efficiency and compromise soil fertility. Fungicides and herbicides diminish key microbial populations, including ammonia-oxidizing bacteria and archaea, while suppressing enzyme activities like urease and dehydrogenase essential for nutrient mineralization; this disrupts nitrogen cycling and organic matter decomposition, ultimately lowering nutrient bioavailability for crops.^[57]

Freshwater Systems

In freshwater lakes, the microbial loop processes a substantial fraction of bacterioplankton production, often exceeding 50% of primary production during key periods such as spring blooms, with allochthonous dissolved organic matter (DOM) from surrounding watersheds serving as the dominant carbon source for bacterial growth.^[58] This external DOM input, primarily from terrestrial runoff, can account for approximately 60% of bacterioplankton production in humic-influenced systems, decoupling bacterial activity from local phytoplankton-derived carbon and sustaining heterotrophic dominance in oligotrophic to mesotrophic lakes.^[59]^[60] Such dynamics highlight the loop's role in carbon cycling, where bacteria mineralize allochthonous DOM, releasing nutrients that fuel subsequent primary production while supporting protistan grazers. In riverine environments, the microbial loop operates under conditions of high flow and turbulence, which enhance encounter rates between bacterivores and bacteria, thereby increasing grazing efficiency by up to 19-fold compared to low-turbulence scenarios.^[61] This physical forcing promotes rapid turnover of bacterial biomass, integrating the loop into lotic food webs where dissolved organic carbon from upstream sources is efficiently recycled. Seasonal cyanobacterial blooms in eutrophic rivers further amplify loop activity, as excess nutrients drive phytoplankton pulses that supply labile carbon to bacteria, boosting heterotrophic production and grazer responses during summer low-flow periods.^[62] Case studies illustrate these processes in large freshwater systems. In the Laurentian Great Lakes, the microbial loop facilitates efficient carbon transfer from primary producers to higher trophic levels, with heterotrophic components comprising up to 75% of the organic carbon pool in Lake Ontario and supporting fisheries through nutrient regeneration and energy flow in Lakes Superior, Huron, and Erie.^[63] Similarly, in the hyporheic zone of rivers, the microbial loop drives the reintroduction of processed dissolved organic carbon back to surface waters, enhancing nutrient recycling via oxygen-dependent bacterial degradation and protistan grazing in this subsurface interface.^[64] Eutrophication from nutrient pollution intensifies microbial loop activity in freshwater systems by increasing organic matter availability, leading to higher bacterial production and nutrient recycling rates—such as bacteria remineralizing up to 95% of phosphorus for phytoplankton reuse—but often shifts pathways toward anaerobic metabolism in oxygen-depleted sediments due to bloom-induced hypoxia.^[65]^[66] This transition favors denitrifying and methanogenic microbes, altering carbon and nitrogen fluxes while potentially reducing overall loop efficiency in severe cases.^[67]

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... C:N:P ratios of the bioavailable nutrients. This means that BP in DOM-dominated systems can show C limitation (or co-limitation with N) even in cases where C:N: ...
[36]
Enhancement of Dissolved Organic Matter Bioavailability by ...
Rapid and sustained photochemical alterations of DOM bioavailability occurring in the euphotic zone of natural waters should accelerate the turnover of surface ...
[37]
Energy Conservation in Fermentations of Anaerobic Bacteria - PMC
Sep 13, 2021 · Anaerobic bacteria ferment carbohydrates and amino acids to obtain energy for growth. Due to the absence of oxygen and other inorganic electron acceptors.
[38]
Ocean acidification reduces growth and grazing impact of Antarctic ...
Aug 18, 2020 · We investigated the effect of increasing fCO2 on the growth of heterotrophic nanoflagellates (HNFs), nano- and picophytoplankton, and ...
[39]
(PDF) Response of a coastal tropical pelagic microbial community to ...
Jun 14, 2016 · Using this methodology, it became evident that temperature and salinity changes, individually and together, mediate direct and indirect effects ...
[40]
Chemotaxis toward phytoplankton drives organic matter partitioning ...
The microenvironment surrounding individual phytoplankton cells is often rich in dissolved organic matter (DOM), which can attract bacteria by chemotaxis. These ...
[41]
Grazing of protozoa and its effect on populations of aquatic bacteria
Several studies have revealed grazing by protists as the dominant factor controlling bacterial mortality. However, in some habitats and seasons, metazoan ...
[42]
Zooplankton grazing on bacteria and phytoplankton in a regulated ...
... bacterial standing crop can be suppressed by zooplankton grazing, i.e. that the microbial loop can experience top-down control, as occurs in the grazing food ...
[43]
Quorum sensing inhibits phage infection by regulating biofilm ... - NIH
Dec 31, 2024 · Our findings suggest that the inhibition of QS may enhance phage infectivity, potentially facilitating advanced phage therapy combined with QS interference.Missing: loop | Show results with:loop
[44]
A tripartite model system for Southern Ocean diatom-bacterial ...
Oct 3, 2022 · These findings revealed the natural coexistence of competing symbiotic strategies of diatom-associated bacteria in the SO, and the utility of this tripartite ...
[45]
Scaling down the microbial loop: data‐driven modelling of growth ...
The complex network of nutrients exchange among marine microorganisms is usually referred to as the microbial loop (Azam et al.,; Fenchel,). The foundation of ...Missing: components | Show results with:components
[46]
The influence of zooplankton enrichment on the microbial loop in a ...
Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by ...
[47]
Major contribution of both zooplankton and protists to the top-down ...
Here, we used a lake mesocosm experiment to address the top-down regulation of freshwater AAP by protists and zooplankton under 2 contrasting nutrient regimes.
[48]
Ecological and Evolutionary Forces Shaping Microbial Diversity in ...
Feb 24, 2006 · Functional redundancy in a microbiota confers stability (also known as the insurance hypothesis [Yachi and Loreau, 1999]) that will be ...
[49]
Microbial diversity, producer–decomposer interactions and ...
The model predicts that microbial diversity has a positive effect on nutrient recycling efficiency and ecosystem processes through either greater intensity ...<|control11|><|separator|>
[50]
Spatial and temporal changes in the partitioning of organic carbon in ...
... carbon in the surface waters of the Sargasso Sea near Bermuda cycles through bacteria and flagellates—the “microbial loop”. However, both seasonal (August ...
[51]
Large mesopelagic fishes biomass and trophic efficiency in the open ...
Feb 7, 2014 · Yet, ample evidence shows that microzooplankton, not mesozooplankton, are the major consumers of PP, consuming 70–80% of the PP on average. The ...
[52]
Metaproteomics Reveals Similar Vertical Distribution of Microbial ...
Mar 24, 2021 · More and more evidences suggest that the labile DOM is not the major carbon and energy resources for microbial community in the deep sea (Baltar ...
[53]
Functional vertical connectivity of microbial communities in the ocean
May 23, 2024 · This study highlights the functional vertical connectivity between surface and deep-sea microbial communities via sinking particles and reveals ...
[54]
Pelagic microbial heterotrophy in response to a highly productive ...
Apr 21, 2016 · In the Ross Sea Polynya, a high particulate concentration near the surface has been shown to yield large vertical fluxes of organic matter ( ...<|control11|><|separator|>
[55]
Microbial processes driving coral reef organic carbon flow
May 9, 2017 · Here we review microbial processes implicated in organic carbon flux in coral reefs displaying species phase shifts.Community Composition · Metabolism · Replacing Microbes In The...<|control11|><|separator|>
[56]
[PDF] Priorities for future research - Ocean Carbon & Biogeochemistry
Vertical migration of zooplankton spatially decouples consumption from fecal pellet production. Microbes attached to sinking particles excrete extracellular.
[57]
[PDF] Variable particle size distributions reduce the sensitivity of global ...
Jan 14, 2021 · Recent earth system models predict a 10 %–20 % decrease in particulate organic carbon export from the sur- face ocean by the end of the 21st ...
[58]
(PDF) Effect of anthropogenic warming on microbial respiration and ...
Our results showed that POC-normalised respiration increased with warming. We estimate that POC export (scaled to primary production) could decrease by 17 ± 7% ...
[59]
Protozoa and plant growth: the microbial loop in soil revisited
Apr 13, 2004 · Protozoa provide a model system that may considerably advance our understanding of the mechanisms underlying plant growth and community composition.Summary · The Rhizosphere – Interface of... · III. Victims and Benefactors...
[60]
A rather dry subject; investigating the study of arid-associated ...
Dec 1, 2020 · Here we aim to review the interactions between arid environments and the microbial communities which inhabit them, covering hot and cold deserts.
[61]
Impact of pesticides on soil health: identification of key soil microbial ...
We use a meta-analysis to assess the effects of pesticides on soil health parameters identifying key biological indicators for environmental risk assessment ...
[62]
Response of the microbial loop to the phytoplankton spring bloom in ...
During a spring bloom, over 50% of primary production went through the microbial loop. Bacteria and ciliates responded rapidly, with ciliates controlling ...Missing: rivers | Show results with:rivers
[63]
Allochthonous Carbon—a Major Driver of Bacterioplankton ... - NIH
The bacterioplankton production and the community composition are also influenced by the composition of the DOM, since different bacterial groups have varying ...
[64]
Allochthonous dissolved organic matter as an energy source for ...
Substantial evidence exists that allochthonous dissolved organic matter (DOM) can provide an important carbon source for pelagic bacteria.
[65]
Increased grazing rates of microplankton in response to small-scale ...
Dec 15, 2024 · Higher grazing in the turbulence treatment would then result in higher bacterial growth rates. A complicated picture of the physical effects ...Missing: loop | Show results with:loop
[66]
Food Web Responses to a Cyanobacterial Bloom in a Freshwater ...
May 5, 2021 · This study will help us to understand the role of microbial loop during algal blooms occurred in eutrophic lakes, and provide theoretical basis ...
[67]
Microbial foodweb comparison of the Laurentian Great Lakes during ...
Jul 1, 2013 · The role of the microbial loop in facilitating the transfer of organic carbon as well as recycling essential nutrients is an increasingly ...
[68]
(PDF) Inter-disciplinary perspectives on processes in the hyporheic zone
### Summary of the Role of the Microbial Loop in the Hyporheic Zone for Nutrient and Carbon Recycling
[69]
[PDF] Examination of the role of the microbial loop in regulating lake ... - BG
Jun 5, 2014 · This has been shown to play an im- portant role in shaping carbon fluxes in lakes and in enhanc- ing nutrient cycling at the base of food webs ( ...
[70]
Impacts of eutrophication on microbial community structure in ...
Aug 14, 2024 · Heavy eutrophication increased the relative abundance of phyllosphere microorganisms potentially involved in anaerobic metabolic processes, ...
[71]
The microbiology of Uganda's large freshwater lakes experiencing ...
Jun 11, 2025 · Thus, microbial OM degradation is expected to be the dominant attenuation pathway. Microbial OM degradation proceeds via either aerobic or ...