Some types of lichen are able to fix nitrogen from the atmosphere. This process relies on the presence of cyanobacteria as a partner species within the lichen. The ability to fix nitrogen enables lichen to live in nutrient-poor environments. Lichen can also extract nitrogen from the rocks on which they grow.
Nitrogen fixation, and hence the abundance of lichen and their host plants, may be decreased by application of nitrogen-based agricultural fertilizer and by atmospheric pollution.
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Biogeochemical cycles |
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The nitrogen cycle is one of the Earth's biogeochemical cycles. It involves the conversion of nitrogen into different chemical forms. The main processes of the nitrogen cycle are the fixation, ammonification, nitrification, and denitrification. As one of the macronutrients, nitrogen plays an important role in plant growth. The nitrogen cycle is affected by environmental factors. For example, in the subarctic heath, increase in temperature can cause nitrogen fixation to increase or decrease based on season, while overall climate warming indirectly caused the vegetation change which in turn affected the nitrogen fixation process. [1]
Lichens are symbiotic organisms that play an important role in the biogeochemical cycle on Earth. The characteristics of lichens, such as strong resistance to factors such as desiccation, ability to grow and break down rocks allow lichen to grow in different types of environment including highly nitrogen limited area such as subarctic heath. [1] [2] While it does not occur often, formation of akinetes (type of cell formed by cyanobacteria which are resistant to cold and desiccation) was observed in nitrogen fixing lichen. [2] Depending on its partner, lichens derive the carbon and nitrogen from algal and cyanobacteria photobionts (which fixes nitrogen from the air). [3] Lichen fungi can fix nitrogen during the day and night, as long the dark period is not too long. [2]
Both nitrogen-fixing lichens and non-nitrogen-fixing lichens take up nitrogen from the environment as a nutrient. [4] Both type of lichens secrete many different organic compounds to absorb minerals from the substrates.
Main difference between nitrogen fixing lichen and non-nitrogen fixing lichen is their photosynthetic partner: nitrogen fixing lichen partner with cyanobacteria which can fix nitrogen from the air, while green alga, partner of non nitrogen fixing lichen, does not perform the same process. [5] The nitrogen fixation is energetically costly due to chemical transformation and only about 10% of lichen are partnered with cyanobacteria. [5] [6] In agricultural regions, non nitrogen fixing lichen reflect uptake of ammonia emission indicating that it have lower nitrogen value. [7]
Some lichens such as Placopsis gelada contain both nitrogen fixing phototrophs and non nitrogen-fixing phototrophs in which Nostoc (cyanobacteria, the phototrophic nitrogen fixer) was dwelling within cephalodia (small gall like structure within lichen; contains cyanobacteria symbionts). [4] In such cases, heterocyst differentiation was greater in cephalodia when compared to having Nostoc as the primary symbionts in lichens, showing that, in the presence of non nitrogen-fixing phototroph, Nostoc specialize for nitrogen fixation. [4]
A lichen's response to nutrient enrichment depends on not only on species and environmental factors but also partially on thallus concentrations of nutrients such as nitrogen and phosphorus. [8]
Ammonium, nitrate and organic nitrogen can be assimilated by lichen along with phosphorus as an important stimulant for cyanolichens. The photobiont will become less dependent on fungal nutrient supply when nitrogen deposition increases as it will be able to access its own nitrogen and it will stimulate the photobiont, causing it to build up, resulting in increased photosynthesis which increases carbon input. [8] However, for lichens that cannot increase their photobiont growth, nitrogen deposition can be damaging due to higher nitrogen concentration than their biological requirements. [8]
Generally, when a lichenized algal cell is nitrogen limited, the addition of nitrogen caused the growth of algal cells. [8] Under nitrogen limiting condition, chlorophyll concentration was positively correlated with the growth of algal cells indicating that should the concentration of chlorophyll increase, the photobiont population will also increase. [8] As lichens absorb nitrogen through fixation, it will have a very strong negative reaction if the nitrogen availability changes, indicating its sensitivity to environmental changes. According to the experiment by Sparrius et al., when nitrogen fertilizer was added into the soil, lichen cover was reduced by ~50%, while the addition of phosphorus showed opposite result. [9] In the region such as boreal forest, where nitrogen and phosphorus are limiting nutrients and for symbiotic interaction to occur properly, their ratio must be balanced. [8] General pollution of climate that is indicated by the concentration of nitrogen oxides can also affect the growth of lichen. [10] When compared to bryophyte (non-vascular land plant), which is also sensitive to nitrogen fertilizer, lichen showed a much stronger response. [9]
There are many different species of lichens and each has its own way of allocating nitrogen. The non nitrogen fixing lichen invests a large amount of nitrogen into photosynthetic tissue, whereas nitrogen fixing lichen will invest into the fungal tissue. [3] Nitrogen-fixing lichen species can only attain a certain amount of nitrogen, as the addition of ammonium decreases its rate of nitrogen-fixation, which decreases the amount of nitrogen that is exported into the adjacent hyphae. [3] Nitrogen fixation is energy dependent and very costly for lichens. [11] In a region where nitrogen deposition is high, lichens have a lower uptake of nitrogen in comparison to the Antarctic green algal lichen, which takes up 90% of nitrogen deposition in both nitrate and ammonium form. [3] Some lichen species are able to refrain from assimilating excessive amount of nitrogen in order to maintain a balanced tissue concentration. [3] Majority of lichen species absorbs more NH4+ than NO3- and the impact of temperature on the rate of fixation is "consonant to the normal enzymatic kinetics of them". [3] [11]
Nitrogen fixing lichens actively fix atmospheric nitrogen using the nostoc, located in the cephalodia. Lichens are sensitive to nitrogen availability. [11] Upon nitrogen fixation, there will be an increase of algal cell growth, chlorophyll concentration, and photobiont population. While costly, in regions where nitrogen availability is low, fixation process is the main way for the lichen to absorb nitrogen which is macronutrient (essential nutrient).
Nitrogen, as a macronutrient and a biogeochemical cycle, also affects the ecology. Through the nitrogen cycle, it breaks down into the chemical form that allows plants to absorb as nutrients. There are certain regions in the world that most plants cannot live due to harsh environments as well as lack of nutrients such as nitrogen. That means that in some regions, the biogeochemical cycle (including nitrogen cycle and carbon cycle) is unlikely to run smoothly. Lichen is able to absorb nitrogen in multiple forms from soil, rock, and air, taking a part in carbon cycle at the same time. Even though only a small fraction of lichens have the ability to fix nitrogen, it helps the lichen to spread throughout the world and survive even in the harsh environment. [5] [6]
The industrial nitrogen fertilizer greatly affected the vegetation and agriculture throughout the world, resulting significantly increased the amount of food with better quality, but it has a negative impact on ecology in the long run. [12] Deposition of nitrogen causes soil acidification, and the nitrogen in the fertilizer are often leached through soil and water, running off the different area. [13] [14] Soil acidification increases toxicity of the soil which reduces plant biodiversity and based on the toxic level of soil acidification, heavy metal such as aluminum and iron can be related to soil water. [14]
Earth's mantle contains non-atmospheric nitrogen in the form of rocks and in the soil. [15] Weathering of the rocks and stone are normally caused by physical, chemical and biological processes. Plants cannot absorb nitrogen from rocks, but fungi can. Fungi within lichens can extract nutrients from mineral surfaces by secreting organic acids. The organic acids (e.g. phenolic acids) are important in solubilizing nutrients from inorganic substrates. [4] A study was conducted to test rock phosphate solubilization by lichen-forming fungi. Bacteria that were attached to biotic or abiotic surfaces stimulate exopolysaccharide synthesis. [4] While lichens have the ability to absorb nitrogen from rock, this only accounts for a small portion of the nitrogen cycle compared to the conversion of atmospheric nitrogen as it is more easily available.
Photobionts will become less dependent on fungal nutrient supply when nitrogen deposition increases, as it will be able to access its own nitrogen, and primary producers' nutrient limit will also be reduced. [8]
Nitrogen is one of the more limiting nutrients and the addition of nitrogen stimulates the photobiont, building up its cell, which subsequently increases its photosynthesis and its carbon input. Multiple nitrogen compounds can be assimilated by lichens, such as NH4+, NO3− and organic nitrogen compounds. [8] Nitrogen deposition reduces the nutrient limitation of primary production. Increase in nitrogen deposition will allow the photobiont to access its own nitrogen which makes it less fungal dependent but only up to certain point. [8]
Depending on the environmental nitrogen availability, the addition of nitrogen can either increase and decrease the growth of the lichen. If the lichen cannot increase its photobiont growth, high nitrogen uptake may result in a higher concentration than it physiologically requires which will negatively affect the lichen and its host plant as the other nutrients are too limiting.
Lichen's response to nutrient enrichment is both species-specific and dependent on environmental factors such as nutrient concentration, light availability and water supply. [8]
Lichen is nitrogen sensitive and change in nitrogen availability can affect its health greatly.
Two main nitrogen stress factors for lichens are nitrogen deficiency and high nitrogen deposition. [3] Both types of nitrogen stress result in the reduction of the rate of thallus expansion in lichen. Nitrogen stressed lichen did not show a significant change in chitin:chlorophyll ratios, but ergosterol concentration showed significant increase indicating a higher demand on the respiratory system.
According to an experiment, the ammonium toxicity due to nitrogen deposition reduced the vitality of lichen greatly at different regions such as inland dunes, boreal conditions, and subarctic heaths. [3] [9]
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).
A fertilizer or fertiliser is any material of natural or synthetic origin that is applied to soil or to plant tissues to supply plant nutrients. Fertilizers may be distinct from liming materials or other non-nutrient soil amendments. Many sources of fertilizer exist, both natural and industrially produced. For most modern agricultural practices, fertilization focuses on three main macro nutrients: nitrogen (N), phosphorus (P), and potassium (K) with occasional addition of supplements like rock flour for micronutrients. Farmers apply these fertilizers in a variety of ways: through dry or pelletized or liquid application processes, using large agricultural equipment or hand-tool methods.
An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term algae encompasses many types of aquatic photosynthetic organisms, both macroscopic multicellular organisms like seaweed and microscopic unicellular organisms like cyanobacteria. Algal bloom commonly refers to the rapid growth of microscopic unicellular algae, not macroscopic algae. An example of a macroscopic algal bloom is a kelp forest.
Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytoplankton productivity". Water bodies with very low nutrient levels are termed oligotrophic and those with moderate nutrient levels are termed mesotrophic. Advanced eutrophication may also be referred to as dystrophic and hypertrophic conditions. Eutrophication can affect freshwater or salt water systems. In freshwater ecosystems it is almost always caused by excess phosphorus. In coastal waters on the other hand, the main contributing nutrient is more likely to be nitrogen, or nitrogen and phosphorus together. This depends on the location and other factors.
The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biological and physical processes. Important processes in the nitrogen cycle include fixation, ammonification, nitrification, and denitrification. The majority of Earth's atmosphere (78%) is atmospheric nitrogen, making it the largest source of nitrogen. However, atmospheric nitrogen has limited availability for biological use, leading to a scarcity of usable nitrogen in many types of ecosystems.
Plant nutrition is the study of the chemical elements and compounds necessary for plant growth and reproduction, plant metabolism and their external supply. In its absence the plant is unable to complete a normal life cycle, or that the element is part of some essential plant constituent or metabolite. This is in accordance with Justus von Liebig’s law of the minimum. The total essential plant nutrients include seventeen different elements: carbon, oxygen and hydrogen which are absorbed from the air, whereas other nutrients including nitrogen are typically obtained from the soil.
Diazotrophs are bacteria and archaea that fix gaseous nitrogen in the atmosphere into a more usable form such as ammonia.
Biological soil crusts are communities of living organisms on the soil surface in arid and semi-arid ecosystems. They are found throughout the world with varying species composition and cover depending on topography, soil characteristics, climate, plant community, microhabitats, and disturbance regimes. Biological soil crusts perform important ecological roles including carbon fixation, nitrogen fixation and soil stabilization; they alter soil albedo and water relations and affect germination and nutrient levels in vascular plants. They can be damaged by fire, recreational activity, grazing and other disturbances and can require long time periods to recover composition and function. Biological soil crusts are also known as biocrusts or as cryptogamic, microbiotic, microphytic, or cryptobiotic soils.
The phosphorus cycle is the biogeochemical cycle that describes the movement of phosphorus through the lithosphere, hydrosphere, and biosphere. Unlike many other biogeochemical cycles, the atmosphere does not play a significant role in the movement of phosphorus, because phosphorus and phosphorus-based compounds are usually solids at the typical ranges of temperature and pressure found on Earth. The production of phosphine gas occurs in only specialized, local conditions. Therefore, the phosphorus cycle should be viewed from whole Earth system and then specifically focused on the cycle in terrestrial and aquatic systems.
Human impact on the nitrogen cycle is diverse. Agricultural and industrial nitrogen (N) inputs to the environment currently exceed inputs from natural N fixation. As a consequence of anthropogenic inputs, the global nitrogen cycle (Fig. 1) has been significantly altered over the past century. Global atmospheric nitrous oxide (N2O) mole fractions have increased from a pre-industrial value of ~270 nmol/mol to ~319 nmol/mol in 2005. Human activities account for over one-third of N2O emissions, most of which are due to the agricultural sector. This article is intended to give a brief review of the history of anthropogenic N inputs, and reported impacts of nitrogen inputs on selected terrestrial and aquatic ecosystems.
In agriculture, leaching is the loss of water-soluble plant nutrients from the soil, due to rain and irrigation. Soil structure, crop planting, type and application rates of fertilizers, and other factors are taken into account to avoid excessive nutrient loss. Leaching may also refer to the practice of applying a small amount of excess irrigation where the water has a high salt content to avoid salts from building up in the soil. Where this is practiced, drainage must also usually be employed, to carry away the excess water.
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.
Lobaria pulmonaria is a large epiphytic lichen consisting of an ascomycete fungus and a green algal partner living together in a symbiotic relationship with a cyanobacterium—a symbiosis involving members of three kingdoms of organisms. Commonly known by various names like tree lungwort, lung lichen, lung moss, lungwort lichen, oak lungs or oak lungwort, it is sensitive to air pollution and is also harmed by habitat loss and changes in forestry practices. Its population has declined across Europe and L. pulmonaria is considered endangered in many lowland areas. The species has a history of use in herbal medicines, and recent research has corroborated some medicinal properties of lichen extracts.
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.
Agricultural pollution refers to biotic and abiotic byproducts of farming practices that result in contamination or degradation of the environment and surrounding ecosystems, and/or cause injury to humans and their economic interests. The pollution may come from a variety of sources, ranging from point source water pollution to more diffuse, landscape-level causes, also known as non-point source pollution and air pollution. Once in the environment these pollutants can have both direct effects in surrounding ecosystems, i.e. killing local wildlife or contaminating drinking water, and downstream effects such as dead zones caused by agricultural runoff is concentrated in large water bodies.
A biofertilizer is a substance which contains living micro-organisms which, when applied to seeds, plant surfaces, or soil, colonize the rhizosphere or the interior of the plant and promotes growth by increasing the supply or availability of primary nutrients to the host plant. Biofertilizers add nutrients through the natural processes of nitrogen fixation, solubilizing phosphorus, and stimulating plant growth through the synthesis of growth-promoting substances. The micro-organisms in biofertilizers restore the soil's natural nutrient cycle and build soil organic matter. Through the use of biofertilizers, healthy plants can be grown, while enhancing the sustainability and the health of the soil. Biofertilizers can be expected to reduce the use of synthetic fertilizers and pesticides, but they are not yet able to replace their use. Since they play several roles, a preferred scientific term for such beneficial bacteria is "plant-growth promoting rhizobacteria" (PGPR).
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.
Orchid mycorrhizae are endomycorrhizal fungi which develop symbiotic relationships with the roots and seeds of plants of the family Orchidaceae. Nearly all orchids are myco-heterotrophic at some point in their life cycle. Orchid mycorrhizae are critically important during orchid germination, as an orchid seed has virtually no energy reserve and obtains its carbon from the fungal symbiont.
Mycorrhiza helper bacteria (MHB) are a group of organisms that form symbiotic associations with both ectomycorrhiza and arbuscular mycorrhiza. MHBs are diverse and belong to a wide variety of bacterial phyla including both Gram-negative and Gram-positive bacteria. Some of the most common MHBs observed in studies belong to the phylas Pseudomonas and Streptomyces. MHBs have been seen to have extremely specific interactions with their fungal hosts at times, but this specificity is lost with plants. MHBs enhance mycorrhizal function, growth, nutrient uptake to the fungus and plant, improve soil conductance, aid against certain pathogens, and help promote defense mechanisms. These bacteria are naturally present in the soil, and form these complex interactions with fungi as plant root development starts to take shape. The mechanisms through which these interactions take shape are not well-understood and needs further study.
Seventeen elements or nutrients are essential for plant growth and reproduction. They are carbon (C), hydrogen (H), oxygen (O), nitrogen (N), phosphorus (P), potassium (K), sulfur (S), calcium (Ca), magnesium (Mg), iron (Fe), boron (B), manganese (Mn), copper (Cu), zinc (Zn), molybdenum (Mo), nickel (Ni) and chlorine (Cl). Nutrients required for plants to complete their life cycle are considered essential nutrients. Nutrients that enhance the growth of plants but are not necessary to complete the plant's life cycle are considered non-essential. With the exception of carbon, hydrogen and oxygen, which are supplied by carbon dioxide and water, and nitrogen, provided through nitrogen fixation, the nutrients derive originally from the mineral component of the soil. The Law of the Minimum expresses that when the available form of a nutrient is not in enough proportion in the soil solution, then other nutrients cannot be taken up at an optimum rate by a plant. A particular nutrient ratio of the soil solution is thus mandatory for optimizing plant growth, a value which might differ from nutrient ratios calculated from plant composition.