Nitrogen fixation is a chemical process by which molecular nitrogen (N
2), which has a strong triple covalent bond, is converted into ammonia (NH
3) or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. The nitrogen in air is molecular dinitrogen, a relatively nonreactive molecule that is metabolically useless to all but a few microorganisms. Biological nitrogen fixation or diazotrophy is an important microbe-mediated process that converts dinitrogen (N2) gas to ammonia (NH3) using the nitrogenase protein complex (Nif).
Photosynthesis is a biological process used by many cellular organisms to convert light energy into chemical energy, which is stored in organic compounds that can later be metabolized through cellular respiration to fuel the organism's activities. The term usually refers to oxygenic photosynthesis, where oxygen is produced as a byproduct, and some of the chemical energy produced is stored in carbohydrate molecules such as sugars, starch, glycogen and cellulose, which are synthesized from endergonic reaction of carbon dioxide with water. Most plants, algae and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the biological energy necessary for complex life on Earth.
Cyanobacteria, also called Cyanobacteriota or Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name cyanobacteria refers to their color, which similarly forms the basis of cyanobacteria's common name, blue-green algae, although they are not usually scientifically classified as algae. They appear to have originated in a freshwater or terrestrial environment. Sericytochromatia, the proposed name of the paraphyletic and most basal group, is the ancestor of both the non-photosynthetic group Melainabacteria and the photosynthetic cyanobacteria, also called Oxyphotobacteria.
Heterocysts or heterocytes are specialized nitrogen-fixing cells formed during nitrogen starvation by some filamentous cyanobacteria, such as Nostoc punctiforme, Cylindrospermum stagnale, and Anabaena sphaerica. They fix nitrogen from dinitrogen (N2) in the air using the enzyme nitrogenase, in order to provide the cells in the filament with nitrogen for biosynthesis.
Diazotrophs are bacteria and archaea that fix gaseous nitrogen in the atmosphere into a more usable form such as ammonia.
Phycocyanin is a pigment-protein complex from the light-harvesting phycobiliprotein family, along with allophycocyanin and phycoerythrin. It is an accessory pigment to chlorophyll. All phycobiliproteins are water-soluble, so they cannot exist within the membrane like carotenoids can. Instead, phycobiliproteins aggregate to form clusters that adhere to the membrane called phycobilisomes. Phycocyanin is a characteristic light blue color, absorbing orange and red light, particularly near 620 nm, and emits fluorescence at about 650 nm. Allophycocyanin absorbs and emits at longer wavelengths than phycocyanin C or phycocyanin R. Phycocyanins are found in cyanobacteria. Phycobiliproteins have fluorescent properties that are used in immunoassay kits. Phycocyanin is from the Greek phyco meaning “algae” and cyanin is from the English word “cyan", which conventionally means a shade of blue-green and is derived from the Greek “kyanos" which means a somewhat different color: "dark blue". The product phycocyanin, produced by Aphanizomenon flos-aquae and Spirulina, is for example used in the food and beverage industry as the natural coloring agent 'Lina Blue' or 'EXBERRY Shade Blue' and is found in sweets and ice cream. In addition, fluorescence detection of phycocyanin pigments in water samples is a useful method to monitor cyanobacteria biomass.
Anabaena is a genus of filamentous cyanobacteria that exist as plankton. They are known for nitrogen-fixing abilities, and they form symbiotic relationships with certain plants, such as the mosquito fern. They are one of four genera of cyanobacteria that produce neurotoxins, which are harmful to local wildlife, as well as farm animals and pets. Production of these neurotoxins is assumed to be an input into its symbiotic relationships, protecting the plant from grazing pressure.
Cyanophages are viruses that infect cyanobacteria, also known as Cyanophyta or blue-green algae. Cyanobacteria are a phylum of bacteria that obtain their energy through the process of photosynthesis. Although cyanobacteria metabolize photoautotrophically like eukaryotic plants, they have prokaryotic cell structure. Cyanophages can be found in both freshwater and marine environments. Marine and freshwater cyanophages have icosahedral heads, which contain double-stranded DNA, attached to a tail by connector proteins. The size of the head and tail vary among species of cyanophages. Cyanophages infect a wide range of cyanobacteria and are key regulators of the cyanobacterial populations in aquatic environments, and may aid in the prevention of cyanobacterial blooms in freshwater and marine ecosystems. These blooms can pose a danger to humans and other animals, particularly in eutrophic freshwater lakes. Infection by these viruses is highly prevalent in cells belonging to Synechococcus spp. in marine environments, where up to 5% of cells belonging to marine cyanobacterial cells have been reported to contain mature phage particles.
Anabaena circinalis is a species of Gram-negative, photosynthetic cyanobacteria common to freshwater environments throughout the world. Much of the scientific interest in A. circinalis owes to its production of several potentially harmful cyanotoxins, ranging in potency from irritating to lethal. Under favorable conditions for growth, A. circinalis forms large algae-like blooms, potentially harming the flora and fauna of an area.
An akinete is an enveloped, thick-walled, non-motile, dormant cell formed by filamentous, heterocyst-forming cyanobacteria under the order Nostocales and Stigonematales. Akinetes are resistant to cold and desiccation. They also accumulate and store various essential material, both of which allows the akinete to serve as a survival structure for up to many years. However, akinetes are not resistant to heat. Akinetes usually develop in strings with each cell differentiating after another and this occurs next to heterocysts if they are present. Development usually occurs during stationary phase and is triggered by unfavorable conditions such as insufficient light or nutrients, temperature, and saline levels in the environment. Once conditions become more favorable for growth, the akinete can then germinate back into a vegetative cell. Increased light intensity, nutrients availability, oxygen availability, and changes in salinity are important triggers for germination. In comparison to vegetative cells, akinetes are generally larger. This is associated with the accumulation of nucleic acids which is important for both dormancy and germination of the akinete. Despite being a resting cell, it is still capable of some metabolic activities such as photosynthesis, protein synthesis, and carbon fixation, albeit at significantly lower levels.
Cyanobionts are cyanobacteria that live in symbiosis with a wide range of organisms such as terrestrial or aquatic plants; as well as, algal and fungal species. They can reside within extracellular or intracellular structures of the host. In order for a cyanobacterium to successfully form a symbiotic relationship, it must be able to exchange signals with the host, overcome defense mounted by the host, be capable of hormogonia formation, chemotaxis, heterocyst formation, as well as possess adequate resilience to reside in host tissue which may present extreme conditions, such as low oxygen levels, and/or acidic mucilage. The most well-known plant-associated cyanobionts belong to the genus Nostoc. With the ability to differentiate into several cell types that have various functions, members of the genus Nostoc have the morphological plasticity, flexibility and adaptability to adjust to a wide range of environmental conditions, contributing to its high capacity to form symbiotic relationships with other organisms. Several cyanobionts involved with fungi and marine organisms also belong to the genera Richelia, Calothrix, Synechocystis, Aphanocapsa and Anabaena, as well as the species Oscillatoria spongeliae. Although there are many documented symbioses between cyanobacteria and marine organisms, little is known about the nature of many of these symbioses. The possibility of discovering more novel symbiotic relationships is apparent from preliminary microscopic observations.
Phototaxis is a kind of taxis, or locomotory movement, that occurs when a whole organism moves towards or away from a stimulus of light. This is advantageous for phototrophic organisms as they can orient themselves most efficiently to receive light for photosynthesis. Phototaxis is called positive if the movement is in the direction of increasing light intensity and negative if the direction is opposite.
Planktothrix is a diverse genus of filamentous cyanobacteria observed to amass in algal blooms in water ecosystems across the globe. Like all Oscillatoriales, Planktothrix species have no heterocysts and no akinetes. Planktothrix are unique because they have trichomes and contain gas vacuoles unlike typical planktonic organisms. Previously, some species of the taxon were grouped within the genus Oscillatoria, but recent work has defined Planktothrix as its own genus. A tremendous body of work on Planktothrix ecology and physiology has been done by Anthony E. Walsby, and the 55.6 kb microcystin synthetase gene which gives these organisms the ability to synthesize toxins has been sequenced. P. agardhii is an example of a type species of the genus. P. agardhii and P. rubescens are commonly observed in lakes of the Northern Hemisphere where they are known producers of potent hepatotoxins called microcystins.
CandidatusAtelocyanobacterium thalassa, also referred to as UCYN-A, is a diazotrophic species of cyanobacteria commonly found in measurable quantities throughout the world's oceans and some seas. Members of A. thalassa are spheroid in shape and are 1-2µm in diameter, and provide nitrogen to ocean regions by fixing non biologically available atmospheric nitrogen into biologically available ammonium that other marine microorganisms can use. Unlike many other cyanobacteria, the genome of A. thalassa does not contain genes for RuBisCO, photosystem II, or the TCA cycle. Consequently, A. thalassa lacks the ability to fix carbon via photosynthesis. Some genes specific to the cyanobacteria group are also absent from the A. thalassa genome despite being an evolutionary descendant of this group. With the inability to fix their own carbon, A. thalassa are obligate symbionts that have been found within photosynthetic picoeukaryote algae. Most notably, the UCYN-A2 sublineage has been observed as an endosymbiont in the alga Braarudosphaera bigelowii with a minimum of 1-2 endosymbionts per host. A. thalassa fixes nitrogen for the algae, while the algae provide carbon for A. thalassa through photosynthesis. There are many sublineages of A. thalassa that are distributed across a wide range of marine environments and host organisms. It appears that some sublineages of A. thalassa have a preference for oligotrophic ocean waters while other sublineages prefer coastal waters. Much is still unknown about all of A. thalassa's hosts and host preferences.
Raphidiopsis raciborskii is a freshwater cyanobacterium.)
Cyanothece is a genus of unicellular, diazotrophic, oxygenic photosynthesizing cyanobacteria.
Cyanophage N-1 is a myovirus bacteriophage that infects freshwater filamentous cyanobacteria of the Nostoc genus. The virus was first isolated by Kenneth Adolph and Robert Haselkorn in 1971 in the US, from the nitrogen-fixing cyanobacterium, Nostoc muscorum. N-1 is closely related to cyanophage A-1, but only distantly to other cyanophages of freshwater or marine origin.
Crocosphaera watsonii is an isolate of a species of unicellular diazotrophic marine cyanobacteria which represent less than 0.1% of the marine microbial population. They thrive in offshore, open-ocean oligotrophic regions where the waters are warmer than 24 degrees Celsius. Crocosphaera watsonii cell density can exceed 1,000 cells per milliliter within the euphotic zone; however, their growth may be limited by the concentration of phosphorus. Crocosphaera watsonii are able to contribute to the oceanic carbon and nitrogen budgets in tropical oceans due to their size, abundance, and rapid growth rate. Crocosphaera watsonii are unicellular nitrogen fixers that fix atmospheric nitrogen to ammonia during the night and contribute to new nitrogen in the oceans. They are a major source of nitrogen to open-ocean systems. Nitrogen fixation is important in the oceans as it not only allows phytoplankton to continue growing when nitrogen and ammonium are in very low supply but it also replenishes other forms of nitrogen, thus fertilizing the ocean and allowing more phytoplankton growth.
Richelia is a genus of nitrogen-fixing, filamentous, heterocystous and cyanobacteria. It contains the single species Richelia intracellularis. They exist as both free-living organisms as well as symbionts within potentially up to 13 diatoms distributed throughout the global ocean. As a symbiont, Richelia can associate epiphytically and as endosymbionts within the periplasmic space between the cell membrane and cell wall of diatoms.
Cyanobacterial morphology refers to the form or shape of cyanobacteria. Cyanobacteria are a large and diverse phylum of bacteria defined by their unique combination of pigments and their ability to perform oxygenic photosynthesis.
Page 12 Aurora by Kim Stanley Robinson.
