Adaptability and Viability of PNSB in Reef Aquaria with Respect to Salinity and Temperature
R. palustris strains can be found in wild and exotic places like a rusty bridge in Woods Hole, Massachusetts where TIE-1 was isolated from... Really, you can find these organisms everywhere.
—Dr. Arpita Bose, quoted in Ogliore (2019)
Within the marine aquarium hobby, certain assumptions persist not because aquarists lack curiosity or scientific understanding, but because broad textbook classifications are sometimes mistaken for absolute physiological limitations. One such assumption concerns purple nonsulfur bacteria (PNSB), particularly Rhodopseudomonas palustris. Because this species is frequently associated with freshwater environments, it is sometimes presumed to be inherently unsuitable for marine aquaria.
But bacteria do not necessarily observe the tidy ecological boundaries that armchair microbiologists on YouTube might assign to them. A species commonly encountered in freshwater may include strains adapted to very different environments. And an organism isolated from a temperate coastal wetland cannot be understood simply by looking at a map and noting the latitude.
Nature is considerably more interesting than that.
In a recent podcast discussion of bacterial supplementation in aquaria, a prominent hobby figure raised questions about the use of PNSB in reef tanks. Although the organism was not necessarily identified by strain designation, the geographic reference appears consistent with Rhodopseudomonas palustris TIE-1, one of three PNSB strains incorporated into our PNS ProBio preparation.
His comments vacillated between two major concerns: (1) Whether a strain originating from a salt marsh can tolerate full-strength seawater, and (2) whether it can tolerate the temperatures commonly maintained in tropical reef aquaria.
These are reasonable subjects for discussion. Indeed, skepticism toward bacterial additives is healthy, especially in an industry where commercial claims so flagrantly exceed the available experimental evidence. Too many of the microbial products currently available to the home aquarist–including some supposed PNSB based formulations–are, objectively, total crap.
However, skepticism must be applied consistently. Assertions that an organism cannot survive particular environmental conditions, and thus cannot provide meaningful biological benefits, require evidence just as much as affirmative product claims do.
In the case of R. palustris TIE-1, there is a substantial body of peer-reviewed research describing its environmental origin, physiology, metabolic versatility, and cellular adaptations. Much of this research complicates the assumptions put forth during the podcast.
When evaluating a specialized wild isolate such as R. palustris strain TIE-1, applying standard physiological limitations misinterprets the evolutionary history of the organism. To understand why these unique strains easily survive within a tropical marine aquarium, one must move past the generalized genus-level taxonomy and look closely at the specific physiological mechanisms, cellular structures, and metabolic adaptations that these unique intertidal isolates possess.
Part I: Deconstructing Claim #1–The Salinity Barrier and Osmotic Adaptability
The first misconception concerns the environmental origin of TIE-1. The core claim offered on the aforementioned podcast states that a Rhodopseudomonas palustris strain native to a temperate coastal zone like Massachusetts cannot survive in seawater because the salinity of a typical reef tank is too high. To evaluate this claim, one must look at the actual field hydrology of the source site from which these bacteria originated.
TIE-1 has a particularly informative isolation history.
In 2005, Jiao, Kappler, Croal, and Newman published a landmark study in Applied and Environmental Microbiology describing the isolation and characterization of this bacterium. Their work established that TIE-1 was recovered from an iron-rich microbial mat at School Street Marsh in Woods Hole, Massachusetts (specifically, just off the School Street bridge near 41.5264°N -70.6683°W).
This is an important distinction. School Street Marsh is world-renowned in the microbiological community. For decades, the Marine Biological Laboratory (MBL) in Woods Hole has used this exact marsh as a premier open-air classroom to isolate unusual bacteria due to its extreme environmental stress. Because the marsh experiences radical transitions between rainwater inundation, marine high tides, and total solar desiccation, the organisms residing in its muds are master metabolic shape-shifters. MBL field logs and scientific retrospectives highlight School Street Marsh as a fascinating ecological anomaly because it is covered alternately by saltwater and freshwater, creating a volatile wetland basin. The extreme environmental stress of switching from fresh rain to dried crystalline marine crusts is precisely why the site is prized for isolating rare, resilient bacteria.
As established by historical field data and academic surveys of Massachusetts intertidal systems like the School Street Marsh and Eel Pond networks in Woods Hole, these ecosystems are under continuous tidal influence. At high tide, full-strength seawater from Great Harbor pushes into the basin, exposing the resident microbial mats to salinities hovering between 3.1% and 3.4% (31 to 34 ppt). Conversely, during low-tide phases occurring during hot summer afternoons, shallow pools isolated by infrastructure or natural topography undergo intense solar evaporation. Because salts remain when water evaporates, localized surface salinities routinely spike well beyond the 35 ppt threshold of standard seawater, moving deep into hypersaline territory, and occasionally evaporating entirely into dry crystalline sea salt crusts.
Conversely, heavy New England rainfall flushes the exact same mudflats with pure freshwater, momentarily plunging the salinity down to less than 0.001%. Therefore, the resident R. palustris populations are not living in a static environment; they are continuously cycling through freshwater, brackish water, seawater, and hypersaline brine, oftentimes on a bi-daily basis. A bacterium incapable of surviving full marine salinity would be instantly eliminated from this habitat within a single tidal cycle.
The salinity variations on the west side of the marsh–specifically near the School Street bridge connection to Eel Pond–create an especially dynamic and volatile environmental gradient. This specific choke point is characterized by unique hydrological behaviors:
1. The "Culvert Effect" & Restricted Flushing
Because Eel Pond connects directly to Great Harbor via a drawbridge, its waters maintain a fairly steady marine salinity (~3.1% to 3.4%). However, as water moves toward School Street Marsh, the School Street bridge/culvert acts as a severe constriction point.
High Tide Inundation: During high tides and storm surges, full-strength seawater from Eel Pond is forced through the bridge opening, completely flooding the lower basin of the marsh.
Low Tide Impoundment: As the tide ebbs, the constricted bridge opening restricts the outflow. Instead of smoothly draining back into Eel Pond, pockets of seawater become trapped in the shallow depressions and mudflats immediately behind the School Street crossing.
2. Solar Radiation and Hypersaline Spikes
The restricted drainage behind the bridge sets the stage for extreme evaporation. During the summer months, the shallow, stagnant water trapped near the bridge is exposed to intense solar heating.
Water evaporates rapidly into the air, leaving behind all of its dissolved ions. The muddy, sun-exposed flats just inches away from the main channel experience the wildest swings—acting as a fresh pool during a rainstorm and transforming into a hypersaline salt pan under a hot afternoon sun.
Close to the bridge's muddy margins, salinities regularly spike past Eel Pond's marine baseline, concentrating heavily until the area undergoes complete desiccation, leaving visible crusts of dry, crystalline sea salts on top of the dark microbial mats.
3. The Freshwater Flash-Flushing
Conversely, because School Street Marsh is a small, coastal wetland basin, it receives freshwater from rainfall, land runoff, and a stream flowing through the marsh.
During heavy New England storms, freshwater can flow into the marsh basin and accumulate in low-lying areas near the School Street bridge restriction.
This influx of freshwater can cause a sudden, dramatic reduction in local salinity, potentially shifting conditions from predominantly marine or brackish water toward much fresher conditions, particularly in shallow pools where freshwater temporarily accumulates.
While there is no permanent, automated USGS gauge recording a long-term mathematical "average" specifically for the east side of the School Street crossing, published microbiological field data and hydrological surveys characterize the location as a brackish, tidally influenced environment, with reported salinity measurements ranging from 1.0% to 2.0% (10 to 20 ppt) from separate field studies. However, because this exact spot serves as a geographic choke point between the marine waters of Eel Pond and the interior basin of School Street Marsh, using a single average number can be highly misleading. The site behaves as an environmental pendulum swinging across three distinct phases:
Tidal and Weather Fluctuations: The marsh acts as a highly dynamic intertidal system influenced by freshwater inundation and rainwater dilution on one hand, and incoming marine waters approaching full seawater salinity on the other.
Evaporative Concentration: During periods of high sunlight, warm temperatures, and low tides, standing water and porewater undergo substantial evaporation. Salinity thus rises dramatically, potentially reaching hypersaline concentrations. Under sufficiently dry conditions, isolated pools may even undergo complete desiccation, potentially leaving behind salt crusts.
Microbial and Resilient Ecosystem: This extreme environment makes School Street Marsh an exceptional habitat for specialized, highly tolerant microbial communities, including unique cyanobacteria, amoebae, and specialized bacteria that thrive despite severe osmotic stress.
It is true that Rhodopseudomonas palustris is frequently described as a freshwater-associated bacterium. Many strains have been isolated from freshwater sediments, soils, and wetlands.
But species-level generalizations are not necessarily reliable descriptions of every strain's environmental tolerances.
Molecular and Cellular Mechanics of Salt Tolerance
The original isolation of TIE-1 from a brackish microbial mat makes it scientifically inappropriate to classify the strain as incapable of tolerating seawater solely because the species is commonly associated with freshwater habitats. The correct question is not whether R. palustris is sometimes described as a freshwater organism; it is whether this particular strain remains viable at marine salinity. That question requires strain-specific experimental evidence. How does an anoxygenic phototroph like R. palustris protect its inner cellular environment when the surrounding water reaches 35 ppt or higher?
Exposing a freshwater-adapted cell to elevated salinity, or vice-versa, creates a substantial osmotic gradient. Water moves out of the cell across its semipermeable membrane, potentially causing cellular dehydration, loss of turgor, impaired metabolic function, and, under sufficiently severe conditions, cell death. To counteract this, marine-adapted and intertidal strains of R. palustris utilize a sophisticated multi-tiered osmotic defense network:
1. Intracellular Osmolyte Accumulation (Compatible Solutes)
When salt levels rise in the external medium, salt-tolerant bacteria can respond by synthesizing or transporting low-molecular-weight organic compounds known as compatible solutes or osmolytes. These include molecules such as glycine betaine, proline, trehalose, and glutamate. Notably, trehalose accumulation under salt stress has been experimentally documented in another strain of R. palustris, while the TIE-1 genome contains genes annotated for glycine/betaine transport.
Because compatible solutes can accumulate without substantially interfering with internal enzymatic reactions or metabolic machinery, cells can increase their cytoplasmic concentrations to help counterbalance the higher salt concentration outside. This reduces osmotic water loss, helps maintain internal turgor pressure, and protects the metabolic machinery from the damaging effects of elevated salinity.
2. Hopanoid-Mediated Membrane Reinforcement
An important line of research concerns hopanoids, a class of bacterial membrane lipids sometimes compared functionally with sterols, including cholesterol, in eukaryotic membranes. Research specifically involving TIE-1 has shown that hopanoids contribute to membrane integrity and physiological resilience under certain environmental stresses.
In the study "Hopanoids Play a Role in Membrane Integrity and pH Homeostasis in Rhodopseudomonas palustris TIE-1," investigators examined the consequences of disrupting hopanoid biosynthesis. They found that TIE-1 cells unable to produce hopanoids exhibited substantial growth defects and cellular damage under acidic and alkaline conditions, even though they could still grow under normal laboratory conditions. A cornerstone discovery regarding this particular R. palustris strain, isolated from the Woods Hole marsh complex, involves the role of these unusual structural lipids.
Pioneering work in geobiology demonstrated that TIE-1 possesses the genetic machinery to synthesize squalene-derived hopanoids and produce chemically distinct forms of these molecules. Importantly, the researchers also observed changes in hopanoid methylation under certain stressful growth conditions, suggesting that the precise chemical composition of these lipids contributes to the bacterium's physiological resilience.
Hopanoids are thought to help organize and stabilize bacterial membranes, potentially influencing their permeability and resistance to environmental stress. By reinforcing the membrane's physical properties, these molecules may help cells maintain the controlled exchange of water, ions, and other substances essential for cellular function. Although the TIE-1 experiments directly demonstrated protection against pH-related stress rather than seawater-induced osmotic shock, they establish that this particular strain possesses a membrane-associated defense system that could be relevant to its ability to withstand changing environmental conditions (we also observe this phenomenon in the strain’s ability to survive freezing—a process heavily dependent on hopanoids).
3. Active Ion Antiports and Proton Pumps
In addition to physical membrane reinforcement, R. palustris strains possess membrane-bound transport proteins that help regulate their internal ion balance. Notably, the genome of TIE-1 contains genes annotated for Na⁺/H⁺ antiporters and related cation/proton transport systems. These molecular transporters can help remove excess sodium (Na⁺) from the cytoplasm by exchanging it for hydrogen ions (H⁺), using energy stored in the membrane's electrochemical gradient rather than directly consuming ATP. This process can help prevent excessive intracellular sodium accumulation while contributing to the maintenance of ion balance, pH regulation, and the electrochemical conditions necessary for cellular metabolism and ATP synthesis.
Part II: Deconstructing Claim #2–The Thermal Threshold and Tropical Resilience
The second concern raised is whether bacteria originating from a temperate climate zone like New England can survive and function at the steady tropical temperatures maintained in marine reef aquaria (25°C to 28°C). This line of reasoning raises a legitimate question about thermal compatibility, but geographic origin alone does not establish that an organism will suffer thermal damage or fail to survive when introduced into a warmer aquatic environment.
The central error is the assumption that an organism isolated from a temperate geographic region must itself be adapted exclusively to cold temperatures.
This is not how microbial temperature physiology works.
Microbiologists distinguish organisms according to their physiological temperature preferences and tolerances, not simply the climate of the place where they were collected. Many microorganisms recovered from temperate environments are mesophiles, meaning they grow at moderate temperatures that may overlap substantially with tropical aquatic conditions. TIE-1 is a useful example of this.
In the original characterization published by Jiao and colleagues, phototrophic cultures of TIE-1 were incubated at 30°C (86°F).
This is not a speculative estimate of environmental tolerance. It is a documented laboratory cultivation temperature used in the foundational study of the organism. By comparison, tropical reef aquaria are commonly maintained around 25–27°C (77–81°F), with some systems operating somewhat warmer.
This view relies on broad regional climate observations while overlooking the microclimate conditions of shallow estuarine ecosystems. A temperate salt marsh is not a steady, perpetually chilly body of water like the deep open ocean. During the summer season in Massachusetts, shallow mudflats, pannes, and restricted channels are directly exposed to long daylight hours and intense solar radiation. Because dark microbial mats and organic-rich muds absorb solar energy efficiently, they can act as natural heat sinks, warming the shallow water and sediments around them.
When the tide recedes, leaving just a few centimeters of standing water over these dark mats, the temperature of the microenvironment can rise rapidly; sun-exposed sediments and shallow water can become much warmer than nearby deeper water. Indeed, the microclimate of a shallow salt marsh can differ substantially from the regional climate. During July and August, shallow intertidal waters and sediments exposed to direct afternoon sunlight can reach temperatures exceeding 30°C (86°F), with particularly shallow, isolated pools potentially approaching 35°C (95°F) under sufficiently hot and sunny conditions. These summertime microclimates illustrate why microorganisms inhabiting temperate coastal marshes may encounter temperatures equal to, or significantly higher than, the controlled 25°C to 26°C environment maintained in a typical home reef aquarium.
Mesophilic Classification and Thermal Growth Curves
From a strict microbiological standpoint, Rhodopseudomonas palustris is universally classified as a mesophile. Rather than relying on generalized temperature boundaries for the species, the published cultivation history of TIE-1 provides direct evidence of its ability to grow at temperatures relevant to tropical reef aquaria.
Documented observation | Temperature °C | Temperature °F | Scientific significance |
Colony development during original isolation | 22°C | 71.6°F | Demonstrates growth under the reported isolation conditions |
Phototrophic laboratory cultivation | 30°C | 86°F | Demonstrates active cultivation above the typical temperature of many reef aquaria |
Aerobic laboratory cultivation | 30°C | 86°F | Shows that TIE-1 can also be cultivated aerobically at this temperature |
NCMA strain collection record | 30°C | 86°F | Independently documents the recommended cultivation temperature for authenticated TIE-1 |
Sources: Jiao et al. (2005), Applied and Environmental Microbiology; NCMA strain B111.
These observations do not establish the absolute minimum, optimum, or maximum growth temperatures of TIE-1. Nevertheless, they demonstrate that the strain is capable of growth under laboratory conditions at temperatures spanning 22°C to 30°C, including a temperature higher than that maintained in many tropical reef aquaria. Most importantly, cultivation at 30°C is a conventional baseline for reliable growth, not merely short-term survival. This directly undermines the assumption that TIE-1's geographic origin in Massachusetts makes it inherently incompatible with tropical aquarium temperatures.
When these documented cultivation conditions are compared to the operating parameters of a home reef aquarium, an important conclusion emerges: TIE-1 is easily cultivated at 30°C, a temperature higher than the 25°C to 27°C range maintained in many tropical reef aquaria. This provides direct evidence that the strain is not inherently restricted to the cooler temperatures associated with its Massachusetts origin. Furthermore, the relatively stable temperatures maintained in home reef aquaria eliminate the extreme seasonal temperature fluctuations characteristic of temperate coastal environments, potentially providing more consistent conditions for microbial activity throughout the year.
The significance is straightforward: TIE-1 is cultivated at a temperature higher than the operating temperature of many tropical reef aquaria. Consequently, the suggestion that TIE-1 is inherently incapable of tolerating tropical temperatures because it originated in Massachusetts is directly at odds with the published laboratory evidence. The original characterization of the strain documents successful cultivation at 30°C (86°F)—a temperature that exceeds the operating temperature of many tropical reef aquaria.
Part III: Conclusion
A personal anecdote may be relevant here—one that involves the late Jake Adams and the former Reef Builders Studio in Golden, Colorado.
Jake had taken an interest in our work with PNSB, was experimenting with one of our Rhodopseudomonas-based products (the original PNS ProBio), and received what would be his last package during one of my visits in September 2022. Sadly, Jake passed away the following month, leaving behind not only an enormous legacy in the reef aquarium community but also a remarkable (and huge) collection of living corals at the studio.
In the months that followed, his friend and assistant Evan faced the unenviable task of maintaining that collection under extraordinarily difficult circumstances. Understandably, in the following months, husbandry fell a bit behind schedule. And it showed, not just in some of the corals, but also in at least one microbial community.
Around March 2023, roughly six months after Jake's passing, I visited the studio for a research project of my own. I had previously collected samples of microbial communities associated with Porites corals in French Polynesia, and wanted to compare those wild coral-associated microbiomes with one from a captive Porites colony. Evan and I collected a water sample immediately adjacent to a Porites colony for this purpose and prepared the sample for Aquabiomics on site.
Unfortunately, when the report came back, the results were not particularly encouraging. The community appeared severely disrupted, with a single bacterial taxon dominating the sample. It was hardly the representative captive Porites microbiome I had hoped to obtain for comparison with my wild samples. But buried among the relatively small populations of other bacteria was something that caught my attention.
The analysis detected DNA assigned to Rhodopseudomonas palustris, representing approximately 0.15% of the sampled microbial community.
Now, 0.15% might not sound particularly impressive. But in this case, nearly everything other than the dominant bacterium was present at very low relative abundance. More importantly, to my knowledge, nobody had added any Rhodopseudomonas preparation to the system for several months—not since Jake's passing.
And there was another detail that made this finding especially intriguing. Jake was famous for his ultra-minimalistic approach to aquascaping, relying on ‘bare bottoms’ and relatively little live rock. These were not aquariums overflowing with the sorts of sheltered, oxygen-depleted substrates we might ordinarily associate with a facultatively anaerobic bacterium such as Rhodopseudomonas.
In other words, if one were deliberately trying to create an aquarium with relatively few obvious refuges for microorganisms that favor anaerobic or microaerobic conditions, Jake's systems would have been an interesting place to start. And yet, there it was. Not in a sample scraped from a deep sand bed. Not from the interior of a piece of live rock. Not even from a conspicuous microbial biofilm. It was detected in an open-water sample collected immediately adjacent to a living coral.
Of course, this particular DNA-based analysis cannot tell us whether those bacteria were descendants of the organisms I had supplied to Jake. Nor can we rule out the possibility that R. palustris had entered and established itself independently. The sample's unusually skewed microbial composition also limits what we can infer about the broader aquarium ecosystem.
Nevertheless, I find the observation fascinating. Months after the last known addition of my preparation, genetic material assigned to R. palustris was detectable in a mature marine aquarium—one with comparatively little of the habitat we might expect to favor such an organism.
Is it irrefutable proof of a successful colonization? Not really. Is it a noteworthy observation for a bacterium sometimes dismissed as fundamentally incompatible with very salty and very warm habitats? I think so.
And, quite apart from its scientific interest, I like to think Jake would have appreciated the irony.
Perhaps the most compelling evidence that TIE-1 can tolerate—and indeed grow at—full-strength seawater salinity comes not from its native salt marsh, its genetic adaptations, or even observations from established reef aquariums. It comes from our own cultivation practices.
At our facility, we have on occasion propagated authenticated Rhodopseudomonas palustris TIE-1 in marine culture media as a means of cleaning up mother culture lines when certain unwanted microorganisms appear. The principle is straightforward: Some contaminants that could grow readily in our standard freshwater medium cannot tolerate the salinity of full-strength seawater. By transferring the culture into marine medium, we can selectively suppress those organisms while retaining TIE-1.
The results are especially revealing. The first generation in marine medium tends to exhibit somewhat slower growth, but with successive propagation, the cultures eventually reach growth rates and densities comparable to those obtained in our freshwater medium. And these marine cultures are arguably a little extra salty: The seawater contributes its normal complement of dissolved salts, while other ingredients in the medium add additional sodium. And this, by the way, is at temperatures that hover around 88°F (31°C).
This doesn’t determine TIE-1's maximum salinity/temperature tolerance, nor does it establish that the organism will grow equally well under every conceivable condition encountered in a reef aquarium system. But it is direct, repeated, firsthand evidence of something far more specific than mere survival: TIE-1 can be cultivated through successive generations in a medium at least as saline as full-strength seawater in a tropical climate. Oh, and also, the other two purple nonsulfur bacteria species in PNS ProBio (our Rhodobacter sphaeroides in particular) are even more amenable to these conditions than our Rhodopseudomonas palustris.
And that is a rather inconvenient observation for any argument resting on the premise that this bacterium simply cannot grow in seawater at elevated temperatures.
So why, when we can make a sufficiently strong case with this first-hand evidence alone, would we go through the trouble to collect, interpret, and present all that Woods Hole stuff? The value of the site data isn't merely evidentiary; it introduces aquarists to the ecological complexity that shaped these organisms. We shouldn't sacrifice that joy of discovery in pursuit of brevity.
Nor should we trade that joy for the spoon-fed guidance of YouTube gurus–particularly when those ‘gurus’ themselves seemingly cannot be bothered to perform the most rudimentary investigation…to graze the surface of the science…to engage in basic scholarship.
Microbiology in general, and these bacteria in particular, are complicated enough. This author has easily put in 10,000 hours researching or working hands-on with these microbes (more than enough to earn a Bachelor of Science in PNSB-ology) and still finds the subject heady, if not downright freaking intimidating. Undoubtedly, there’s a lot more we could learn about PNSB aquarium applications, and should. Just as rebutting the same simplistic, ill-informed assumptions has become a tedious chore, I welcome any opportunity to discuss the ecology of phototrophic bacteria in a serious, competent, and collaborative setting.
If anyone associated with a certain podcast is listening—I’m always game, and you know where to find me.
And may we all rest assured that Rhodopseudomonas palustris TIE-1 can grow in seawater at 30°C.
References
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Bose, A., Fowler, R. E., Emerson, V. V., Dika, A. L., & Newman, D. K. (2020). Complete genome sequence of the photoferrotrophic bacterium Rhodopseudomonas palustris strain TIE-1. Microbiology Resource Announcements, 9(12), e00142-20.
Santiago-Ramirez, M. I., & Margulis, L. (2011). Growth of chromidia-forming vahlkampfiid amoebae from Laguna Figueroa, Baja California del Norte, Mexico and Eel Pond, Woods of Massachusetts Amherst, ScholarWorks Repository.
Welander, P. V., Hunter, R. C., Zhang, L., Sessions, A. L., Summons, R. E., & Newman, D. K. (2012). Hopanoids play a role in membrane integrity and pH tolerance in Rhodopseudomonas palustris TIE-1. Journal of Bacteriology, 194(20), 6140–6148.
Nixon, S. W. (1982). The ecology of New England high salt marshes: A community profile. U.S. Fish and Wildlife Service, Office of Biological Services. FWS/OBS-82/07.
Massachusetts Department of Environmental Protection. (2014). Cape Cod atlas of tidally restricted salt marshes. Commonwealth of Massachusetts Environmental Publications.
Expression of the Rhodopseudomonas palustris EcfG target gene blr2136 enhances salt tolerance in Arabidopsis thaliana. Bioscience, Biotechnology, and Biochemistry, Volume 86, Issue 3, March 2022, Pages 407–415. Hole, Massachusetts, U.S.A. under limited oxygen gas conditions (Master's thesis).
Harwood, C. S. Rhodopseudomonas palustris: An amazingly versatile bacterium. Harwood Lab - University of Washington.
Y. Oda, F.W. Larimer, P.S.G. Chain, S. Malfatti, M.V. Shin, L.M. Vergez, L. Hauser, M.L. Land, S. Braatsch, J.T. Beatty, D.A. Pelletier, A.L. Schaefer, & C.S. Harwood (2008). Multiple genome sequences reveal adaptations of a phototrophic bacterium to sediment microenvironments, Proc. Natl. Acad. Sci. U.S.A.105 (47) 18543-18548.
Ogliore, T. (2019, April 1). Study shows how electricity-eating microbes use electrons to fix carbon dioxide. WashU McKelvey School of Engineering. https://engineering.washu.edu/news/2019/Study-shows-how-electricity-eating-microbes-use-electrons-to-fix-carbon-dioxide.html
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