Phototrophic biofilms are microbial communities generally comprising both phototrophic microorganisms, which use light as their energy source, and chemoheterotrophs. [1] Thick laminated multilayered phototrophic biofilms are usually referred to as microbial mats or phototrophic mats (see also biofilm). [2] These organisms, which can be prokaryotic or eukaryotic organisms like bacteria, cyanobacteria, fungi, and microalgae, make up diverse microbial communities that are affixed in a mucous matrix, or film. These biofilms occur on contact surfaces in a range of terrestrial and aquatic environments. The formation of biofilms is a complex process and is dependent upon the availability of light as well as the relationships between the microorganisms. Biofilms serve a variety of roles in aquatic, terrestrial, and extreme environments; these roles include functions which are both beneficial and detrimental to the environment. In addition to these natural roles, phototrophic biofilms have also been adapted for applications such as crop production and protection, bioremediation, and wastewater treatment. [1] [2]
Biofilm formation is a complicated process which occurs in four general steps: attachment of cells, formation of the colony, maturation, and cell dispersal. These films can grow in sizes ranging from microns to centimeters in thickness. Most are green and/or brown, but can be more colorful. [1]
Biofilm development is dependent on the generation of extracellular polymeric substances (EPS) by microorganisms. The EPS, which is akin to a gel, is a matrix which provides structure for the biofilm and is essential for growth and functionality. It consists of organic compounds such as polysaccharides, proteins, and glycolipids and may also include inorganic substances like silt and silica. EPS join cells together in the biofilm and transmits light to organisms in the lower zone. Additionally, EPS serves as an adhesive for surface attachment and facilitates digestion of nutrients by extracellular enzymes. [1]
Microbial functions and interactions are also important for maintaining the well-being of the community. In general, phototrophic organisms in the biofilm provide a foundation for the growth of the community as a whole by mediating biofilm processes and conversions. The chemoheterotrophs use the photosynthetic waste products from the phototrophs as their carbon and nitrogen sources, and in turn perform nutrient regeneration for the community. [1] [2] Various groups of organisms are located in distinct layers based on availability of light, the presence of oxygen, and redox gradients produced by the species. [2] Light exposure early in biofilm development has an immense impact on growth and microbial diversity; greater light availability promotes more growth. Phototrophs such as cyanobacteria and green algae occupy the exposed layer of the biofilm while lower layers consist of anaerobic phototrophs and heterotrophs like bacteria, protozoa, and fungi. [1] Eukaryotic algae and cyanobacteria in the outer portion use light energy to reduce carbon dioxide, providing organic substrates and oxygen. This photosynthetic activity fuels processes and conversions in the total biofilm community, including the heterotrophic fraction. It also produces an oxygen gradient in the mat which inhibits most anaerobic phototrophs and chemotrophs from growing in the upper regions. [2]
Communication between the microorganisms is facilitated by quorum sensing or signal transduction pathways, which are accomplished through the secretion of molecules which diffuse through the biofilm. The identity of these substances varies depending on the type of microorganism from which it was secreted. [1]
While some of the organisms contributing to the formation of the biofilms can be identified, exact composition of the biofilms is difficult to determine because many of the organisms cannot be grown using pure culture methods. Though pure culture methods cannot be used to identify unculturable microorganisms and do not support the study of the complex interactions between photoautotrophs and heterotrophs, the use of metagenomics, proteomics, and transcriptomics has helped characterize these unculturable organisms and has provided some insight into molecular mechanisms, microbial organization, and interactions in biofilms. [1]
Phototrophic biofilms can be found on terrestrial and aquatic surfaces and can withstand environmental fluctuations and extreme environments. In aquatic systems, biofilms are prevalent on surfaces of rocks and plants, and in terrestrial environments they can be located in the soil, on rocks, and on buildings. [1] Phototrophic biofilms and microbial mats have been described in extreme environments like thermal springs, [3] hyper saline ponds, [4] desert soil crusts, and in lake ice covers in Antarctica. The 3.4-billion-year fossil record of benthic phototrophic communities, such as microbial mats and stromatolites, indicates that these associations represent the Earth's oldest known ecosystems. It is thought that these early ecosystems played a key role in the build-up of oxygen in the Earth’s atmosphere. [5]
A diverse array of roles is played by these microorganisms across the range of environments in which they can be found. In aquatic environments, these microbes are primary producers, a critical part of the food chain. They perform a key function in exchanging a substantial amount of nutrients and gases between the atmospheric and oceanic reservoirs. Biofilms in terrestrial systems can contribute to improving soil, reducing erosion, promoting growth of vegetation, and revitalizing desert-like land, but they can also accelerate the degradation of solid structures like buildings and monuments. [1]
There is a growing interest in the application of phototrophic biofilms, for instance in wastewater treatment in constructed wetlands, bioremediation, agriculture, and biohydrogen production. [2] A few are outlined below.
Agrochemicals such as pesticides, fertilizers, and food hormones are widely used to produce greater quality and quantity of food as well as provide crop protection. However, biofertilizers have been developed as a more environmentally cognizant method of assisting in plant development and protection by promoting the growth of microorganisms such as cyanobacteria. Cyanobacteria can augment plant growth by colonizing on plant roots to supply carbon and nitrogen, which they can provide to plants through the natural metabolic processes of carbon dioxide and nitrogen fixation. They can also produce substances which induce plant defense against harmful fungi, bacteria, and viruses. Other organisms can also produce secondary metabolites such as phytohormones which increase plants' resistance to pests and disease. [1] Promoting growth of phototrophic biofilms in agricultural settings improves the quality of the soil and water retention, reduces salinity, and protects against erosion. [2]
Organisms in mats such as cyanobacteria, sulfate reducers, and aerobic heterotrophs can aid in bioremediation of water systems through biodegradation of oils. [2] This is achieved by freeing oxygen, organic compounds, and nitrogen from hydrocarbon pollutants. Biofilm growth can also degrade other pollutants by oxidizing oils, pesticides, and herbicides and reducing heavy metals like copper, lead, and zinc. Aerobic processes to degrade pollutants can be achieved during the day and anaerobic processes are performed at night by biofilms. [1] Additionally, because biofilm response to pollutants during initial exposure suggested acute toxicity, biofilms can be used as sensors for pollution. [2]
Biofilms are used in wastewater treatment facilities and constructed wetlands for processes such as cleaning pesticide and fertilizer-laden water because it is simple to form flocs, or aggregates, using biofilms as compared to other floc materials. [1] [2] There are also many other benefits to using phototrophic biofilms in treating wastewater, particularly in nutrient removal. The organisms can sequester nutrients from the wastewater and use these along with carbon dioxide to build biomass. The biomass can capture nitrogen, which can be extracted and used in fertilizer production. [2] Due to their quick growth, phototrophic biofilms have greater nutrient uptake than other methods of nutrient removal utilizing algal biomass, and they are easier to harvest because they naturally grow on wastewater pond surfaces. [6]
Phototrophic activity of these films can precipitate dissolved phosphates due to an increase in pH; these phosphates are then removed by assimilation. Increase in pH of the wastewater also minimizes the presence of coliform bacteria. [2]
Heavy metal detoxification in wastewater treatment can also be achieved with these microbes primarily through passive mechanisms such as ion exchange, chelation, adsorption, and diffusion, which constitute biosorption. The active mode is known as bioaccumulation. Biosorption-mediated metal detoxification is influenced by factors including light intensity, pH, density of the biofilm, and organism tolerance of heavy metals. Though biosorption is an efficient process and inexpensive, methods to retrieve heavy metals from the biomass after biosorption still need further development. [2]
Using phototrophic biofilms for wastewater treatment is more energy efficient and economical and has the capability of producing byproducts which can be further processed into biofuels. [1] Specifically cyanobacteria are capable of producing biohydrogen, which is an alternative to fossil fuels and may become a viable source of renewable energy. [2]
Primary nutritional groups are groups of organisms, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the sources of carbon can be of organic or inorganic origin.
A heterotroph is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but not producers. Living organisms that are heterotrophic include all animals and fungi, some bacteria and protists, and many parasitic plants. The term heterotroph arose in microbiology in 1946 as part of a classification of microorganisms based on their type of nutrition. The term is now used in many fields, such as ecology in describing the food chain.
In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules and nutrients into organic matter using the oxidation of inorganic compounds or ferrous ions as a source of energy, rather than sunlight, as in photosynthesis. Chemoautotrophs, organisms that obtain carbon from carbon dioxide through chemosynthesis, are phylogenetically diverse. Groups that include conspicuous or biogeochemically-important taxa include the sulfur-oxidizing Gammaproteobacteria, the Campylobacterota, the Aquificota, the methanogenic archaea, and the neutrophilic iron-oxidizing bacteria.
Bioremediation is a process used to treat contaminated media, including water, soil and subsurface material, by altering environmental conditions to stimulate growth of microorganisms that degrade the target pollutants. Most bioremediation is inadvertent, involving native organisms. Research on bioremediation is heavily focused on stimulating the process by inoculation of a polluted site with organisms or supplying nutrients to promote the growth. In principle, bioremediation could be used to reduce the impact of byproducts created from anthropogenic activities, such as industrialization and agricultural processes. Bioremediation could prove less expensive and more sustainable than other remediation alternatives.
The purple sulfur bacteria (PSB) are part of a group of Pseudomonadota capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in stratified water environments including hot springs, stagnant water bodies, as well as microbial mats in intertidal zones. Unlike plants, algae, and cyanobacteria, purple sulfur bacteria do not use water as their reducing agent, and therefore do not produce oxygen. Instead, they can use sulfur in the form of sulfide, or thiosulfate (as well, some species can use H2, Fe2+, or NO2−) as the electron donor in their photosynthetic pathways. The sulfur is oxidized to produce granules of elemental sulfur. This, in turn, may be oxidized to form sulfuric acid.
Biofiltration is a pollution control technique using a bioreactor containing living material to capture and biologically degrade pollutants. Common uses include processing waste water, capturing harmful chemicals or silt from surface runoff, and microbiotic oxidation of contaminants in air. Industrial biofiltration can be classified as the process of utilizing biological oxidation to remove volatile organic compounds, odors, and hydrocarbons.
Phototrophs are organisms that carry out photon capture to produce complex organic compounds and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconception that phototrophs are obligatorily photosynthetic. Many, but not all, phototrophs often photosynthesize: they anabolically convert carbon dioxide into organic material to be utilized structurally, functionally, or as a source for later catabolic processes. All phototrophs either use electron transport chains or direct proton pumping to establish an electrochemical gradient which is utilized by ATP synthase, to provide the molecular energy currency for the cell. Phototrophs can be either autotrophs or heterotrophs. If their electron and hydrogen donors are inorganic compounds they can be also called lithotrophs, and so, some photoautotrophs are also called photolithoautotrophs. Examples of phototroph organisms are Rhodobacter capsulatus, Chromatium, and Chlorobium.
Algal mats are one of many types of microbial mat that forms on the surface of water or rocks. They are typically composed of blue-green cyanobacteria and sediments. Formation occurs when alternating layers of blue-green bacteria and sediments are deposited or grow in place, creating dark-laminated layers. Stromatolites are prime examples of algal mats. Algal mats played an important role in the Great Oxidation Event on Earth some 2.3 billion years ago. Algal mats can become a significant ecological problem, if the mats grow so expansive or thick as to disrupt the other underwater marine life by blocking the sunlight or producing toxic chemicals.
Lithotrophs are a diverse group of organisms using an inorganic substrate to obtain reducing equivalents for use in biosynthesis or energy conservation via aerobic or anaerobic respiration. While lithotrophs in the broader sense include photolithotrophs like plants, chemolithotrophs are exclusively microorganisms; no known macrofauna possesses the ability to use inorganic compounds as electron sources. Macrofauna and lithotrophs can form symbiotic relationships, in which case the lithotrophs are called "prokaryotic symbionts". An example of this is chemolithotrophic bacteria in giant tube worms or plastids, which are organelles within plant cells that may have evolved from photolithotrophic cyanobacteria-like organisms. Chemolithotrophs belong to the domains Bacteria and Archaea. The term "lithotroph" was created from the Greek terms 'lithos' (rock) and 'troph' (consumer), meaning "eaters of rock". Many but not all lithoautotrophs are extremophiles.
Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.
A photobioreactor is a bioreactor that utilizes a light source to cultivate phototrophic microorganisms. These organisms use photosynthesis to generate biomass from light and carbon dioxide and include plants, mosses, macroalgae, microalgae, cyanobacteria and purple bacteria. Within the artificial environment of a photobioreactor, specific conditions are carefully controlled for respective species. Thus, a photobioreactor allows much higher growth rates and purity levels than anywhere in nature or habitats similar to nature. Hypothetically, phototropic biomass could be derived from nutrient-rich wastewater and flue gas carbon dioxide in a photobioreactor.
Picoplankton is the fraction of plankton composed by cells between 0.2 and 2 μm that can be either prokaryotic and eukaryotic phototrophs and heterotrophs:
The microbial food web refers to the combined trophic interactions among microbes in aquatic environments. These microbes include viruses, bacteria, algae, heterotrophic protists.
Many microorganisms can naturally grow together on surfaces to form complex aggregations called biofilms. Much research has been done on methods to remove biofilms in clinical and food manufacturing processes, but biofilms are also used for constructive purposes in a variety of industries. One distinctive characteristic of biofilm formation is that microorganisms within biofilms are often much tougher and more resistant to environmental stress compared to individual microorganisms. The cells are stationary and are able to adapt to adverse environments. This phenomenon of enhanced resistance can be beneficial in industrial chemical production where microorganisms within biofilms may tolerate higher chemical concentration and act as robust biorefineries for various products. These microbes have also been used in bioremediation to remove contaminants from freshwater and wastewater. More novel uses of biofilms include generating electricity using microbial fuel cells. Challenges to scaling up this technology include cost, controlling the growth of biofilms, and membrane fouling.
A microbial mat is a multi-layered sheet of microorganisms, mainly bacteria and archaea, and also just bacterial. Microbial mats grow at interfaces between different types of material, mostly on submerged or moist surfaces, but a few survive in deserts. A few are found as endosymbionts of animals.
Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλανκτος, meaning "wanderer" or "drifter", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and freshwater.
Rhodopseudomonas palustris is a rod-shaped, Gram-negative purple nonsulfur bacterium, notable for its ability to switch between four different modes of metabolism.
An autotroph or primary producer is an organism that produces complex organic compounds using carbon from simple substances such as carbon dioxide, generally using energy from light (photosynthesis) or inorganic chemical reactions (chemosynthesis). They convert an abiotic source of energy into energy stored in organic compounds, which can be used by other organisms. Autotrophs do not need a living source of carbon or energy and are the producers in a food chain, such as plants on land or algae in water. Autotrophs can reduce carbon dioxide to make organic compounds for biosynthesis and as stored chemical fuel. Most autotrophs use water as the reducing agent, but some can use other hydrogen compounds such as hydrogen sulfide.
Oxygenic photogranules (OPGs) are a type of biological aggregate with an approximately spherical form, typically from a millimeter to a centimeter scale. OPGs are characterized by the cloth-like layer of phototrophic organisms, predominantly filamentous cyanobacteria of the order Oscillatoriales. Oxygen production by these phototrophs through photosynthesis is typically coupled to oxygen consumption of heterotrophic biomass, releasing CO2 that is presumably utilised in a syntrophic relationship by autotrophic phototrophs.
The viral shunt is a mechanism that prevents marine microbial particulate organic matter (POM) from migrating up trophic levels by recycling them into dissolved organic matter (DOM), which can be readily taken up by microorganisms. The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.