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A biogeochemical cycle, or more generally a cycle of matter, [1] is the movement and transformation of chemical elements and compounds between living organisms, the atmosphere, and the Earth's crust. Major biogeochemical cycles include the carbon cycle, the nitrogen cycle and the water cycle. Every cycle involves the transformation and cycling of a chemical element or molecule by living things as well as by means of different geological forms and reservoirs, such as the atmosphere, soil, and oceans. It can be thought of as the pathway by which a chemical substance cycles (is turned over or moves through) the biotic compartment and the abiotic compartments of Earth. The biotic compartment is the biosphere and the abiotic compartments are the atmosphere, lithosphere and hydrosphere.
For example, in the carbon cycle, atmospheric carbon dioxide is absorbed by plants through photosynthesis, which converts it into organic compounds that are used by organisms for energy and growth. Carbon is then released back into the atmosphere through respiration and decomposition. Additionally, carbon is stored in fossil fuels and is released into the atmosphere through human activities such as burning fossil fuels. In the nitrogen cycle, atmospheric nitrogen gas is converted by plants into usable forms such as ammonia and nitrates through the process of nitrogen fixation. These compounds can be used by other organisms, and nitrogen is returned to the atmosphere through denitrification and other processes. In the water cycle, the universal solvent water evaporates from land and oceans to form clouds in the atmosphere, and then precipitates back to different parts of the planet. Precipitation can seep into the ground and become part of groundwater systems used by plants and other organisms, or can runoff the surface to form lakes and rivers. Subterranean water can then seep into the ocean along with river discharges, rich with dissolved and particulate organic matter and other nutrients.
There are biogeochemical cycles for many other elements, such as for oxygen, hydrogen, phosphorus, calcium, iron, sulfur, mercury and selenium. There are also cycles for molecules, such as water and silica. In addition there are macroscopic cycles such as the rock cycle, and human-induced cycles for synthetic compounds such as for polychlorinated biphenyls (PCBs). In some cycles there are geological reservoirs where substances can remain or be sequestered for long periods of time.
Biogeochemical cycles involve the interaction of biological, geological, and chemical processes. Biological processes include the influence of microorganisms, which are critical drivers of biogeochemical cycling. Microorganisms have the ability to carry out wide ranges of metabolic processes essential for the cycling of nutrients and chemicals throughout global ecosystems. Without microorganisms many of these processes would not occur, with significant impact on the functioning of land and ocean ecosystems and the planet's biogeochemical cycles as a whole. Changes to cycles can impact human health. The cycles are interconnected and play important roles regulating climate, supporting the growth of plants, phytoplankton and other organisms, and maintaining the health of ecosystems generally. Human activities such as burning fossil fuels and using large amounts of fertilizer can disrupt cycles, contributing to climate change, pollution, and other environmental problems.
Energy flows directionally through ecosystems, entering as sunlight (or inorganic molecules for chemoautotrophs) and leaving as heat during the many transfers between trophic levels. However, the matter that makes up living organisms is conserved and recycled. The six most common elements associated with organic molecules — carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur — take a variety of chemical forms and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth's surface. Geologic processes, such as weathering, erosion, water drainage, and the subduction of the continental plates, all play a role in this recycling of materials. Because geology and chemistry have major roles in the study of this process, the recycling of inorganic matter between living organisms and their environment is called a biogeochemical cycle. [3]
The six aforementioned elements are used by organisms in a variety of ways. Hydrogen and oxygen are found in water and organic molecules, both of which are essential to life. Carbon is found in all organic molecules, whereas nitrogen is an important component of nucleic acids and proteins. Phosphorus is used to make nucleic acids and the phospholipids that comprise biological membranes. Sulfur is critical to the three-dimensional shape of proteins. The cycling of these elements is interconnected. For example, the movement of water is critical for leaching sulfur and phosphorus into rivers which can then flow into oceans. Minerals cycle through the biosphere between the biotic and abiotic components and from one organism to another. [4]
Ecological systems (ecosystems) have many biogeochemical cycles operating as a part of the system, for example, the water cycle, the carbon cycle, the nitrogen cycle, etc. All chemical elements occurring in organisms are part of biogeochemical cycles. In addition to being a part of living organisms, these chemical elements also cycle through abiotic factors of ecosystems such as water (hydrosphere), land (lithosphere), and/or the air (atmosphere). [5]
The living factors of the planet can be referred to collectively as the biosphere. All the nutrients — such as carbon, nitrogen, oxygen, phosphorus, and sulfur — used in ecosystems by living organisms are a part of a closed system; therefore, these chemicals are recycled instead of being lost and replenished constantly such as in an open system. [5]
The major parts of the biosphere are connected by the flow of chemical elements and compounds in biogeochemical cycles. In many of these cycles, the biota plays an important role. Matter from the Earth's interior is released by volcanoes. The atmosphere exchanges some compounds and elements rapidly with the biota and oceans. Exchanges of materials between rocks, soils, and the oceans are generally slower by comparison. [2]
The flow of energy in an ecosystem is an open system; the Sun constantly gives the planet energy in the form of light while it is eventually used and lost in the form of heat throughout the trophic levels of a food web. Carbon is used to make carbohydrates, fats, and proteins, the major sources of food energy. These compounds are oxidized to release carbon dioxide, which can be captured by plants to make organic compounds. The chemical reaction is powered by the light energy of sunshine.
Sunlight is required to combine carbon with hydrogen and oxygen into an energy source, but ecosystems in the deep sea, where no sunlight can penetrate, obtain energy from sulfur. Hydrogen sulfide near hydrothermal vents can be utilized by organisms such as the giant tube worm. In the sulfur cycle, sulfur can be forever recycled as a source of energy. Energy can be released through the oxidation and reduction of sulfur compounds (e.g., oxidizing elemental sulfur to sulfite and then to sulfate).
Although the Earth constantly receives energy from the Sun, its chemical composition is essentially fixed, as the additional matter is only occasionally added by meteorites. Because this chemical composition is not replenished like energy, all processes that depend on these chemicals must be recycled. These cycles include both the living biosphere and the nonliving lithosphere, atmosphere, and hydrosphere.
Biogeochemical cycles can be contrasted with geochemical cycles. The latter deals only with crustal and subcrustal reservoirs even though some process from both overlap.
The global ocean covers more than 70% of the Earth's surface and is remarkably heterogeneous. Marine productive areas, and coastal ecosystems comprise a minor fraction of the ocean in terms of surface area, yet have an enormous impact on global biogeochemical cycles carried out by microbial communities, which represent 90% of the ocean's biomass. [8] Work in recent years has largely focused on cycling of carbon and macronutrients such as nitrogen, phosphorus, and silicate: other important elements such as sulfur or trace elements have been less studied, reflecting associated technical and logistical issues. [9] Increasingly, these marine areas, and the taxa that form their ecosystems, are subject to significant anthropogenic pressure, impacting marine life and recycling of energy and nutrients. [10] [11] [12] A key example is that of cultural eutrophication, where agricultural runoff leads to nitrogen and phosphorus enrichment of coastal ecosystems, greatly increasing productivity resulting in algal blooms, deoxygenation of the water column and seabed, and increased greenhouse gas emissions, [13] with direct local and global impacts on nitrogen and carbon cycles. However, the runoff of organic matter from the mainland to coastal ecosystems is just one of a series of pressing threats stressing microbial communities due to global change. Climate change has also resulted in changes in the cryosphere, as glaciers and permafrost melt, resulting in intensified marine stratification, while shifts of the redox-state in different biomes are rapidly reshaping microbial assemblages at an unprecedented rate. [14] [15] [16] [17] [9]
Global change is, therefore, affecting key processes including primary productivity, CO2 and N2 fixation, organic matter respiration/remineralization, and the sinking and burial deposition of fixed CO2. [17] In addition to this, oceans are experiencing an acidification process, with a change of ~0.1 pH units between the pre-industrial period and today, affecting carbonate/bicarbonate buffer chemistry. In turn, acidification has been reported to impact planktonic communities, principally through effects on calcifying taxa. [18] There is also evidence for shifts in the production of key intermediary volatile products, some of which have marked greenhouse effects (e.g., N2O and CH4, reviewed by Breitburg in 2018, [15] due to the increase in global temperature, ocean stratification and deoxygenation, driving as much as 25 to 50% of nitrogen loss from the ocean to the atmosphere in the so-called oxygen minimum zones [19] or anoxic marine zones, [20] driven by microbial processes. Other products, that are typically toxic for the marine nekton, including reduced sulfur species such as H2S, have a negative impact for marine resources like fisheries and coastal aquaculture. While global change has accelerated, there has been a parallel increase in awareness of the complexity of marine ecosystems, and especially the fundamental role of microbes as drivers of ecosystem functioning. [16] [9]
Microorganisms drive much of the biogeochemical cycling in the earth system. [21] [22]
The chemicals are sometimes held for long periods of time in one place. This place is called a reservoir, which, for example, includes such things as coal deposits that are storing carbon for a long period of time. [23] When chemicals are held for only short periods of time, they are being held in exchange pools. Examples of exchange pools include plants and animals. [23]
Plants and animals utilize carbon to produce carbohydrates, fats, and proteins, which can then be used to build their internal structures or to obtain energy. Plants and animals temporarily use carbon in their systems and then release it back into the air or surrounding medium. Generally, reservoirs are abiotic factors whereas exchange pools are biotic factors. Carbon is held for a relatively short time in plants and animals in comparison to coal deposits. The amount of time that a chemical is held in one place is called its residence time or turnover time (also called the renewal time or exit age). [23]
Box models are widely used to model biogeochemical systems. [24] [25] Box models are simplified versions of complex systems, reducing them to boxes (or storage reservoirs) for chemical materials, linked by material fluxes (flows). Simple box models have a small number of boxes with properties, such as volume, that do not change with time. The boxes are assumed to behave as if they were mixed homogeneously. [25] These models are often used to derive analytical formulas describing the dynamics and steady-state abundance of the chemical species involved.
The diagram at the right shows a basic one-box model. The reservoir contains the amount of material M under consideration, as defined by chemical, physical or biological properties. The source Q is the flux of material into the reservoir, and the sink S is the flux of material out of the reservoir. The budget is the check and balance of the sources and sinks affecting material turnover in a reservoir. The reservoir is in a steady state if Q = S, that is, if the sources balance the sinks and there is no change over time. [25]
The residence or turnover time is the average time material spends resident in the reservoir. If the reservoir is in a steady state, this is the same as the time it takes to fill or drain the reservoir. Thus, if τ is the turnover time, then τ = M/S. [25] The equation describing the rate of change of content in a reservoir is
When two or more reservoirs are connected, the material can be regarded as cycling between the reservoirs, and there can be predictable patterns to the cyclic flow. [25] More complex multibox models are usually solved using numerical techniques.
Global biogeochemical box models usually measure:
- reservoir masses in petagrams (Pg)
- flow fluxes in petagrams per year (Pg yr−1)
The diagram on the left shows a simplified budget of ocean carbon flows. It is composed of three simple interconnected box models, one for the euphotic zone, one for the ocean interior or dark ocean, and one for ocean sediments. In the euphotic zone, net phytoplankton production is about 50 Pg C each year. About 10 Pg is exported to the ocean interior while the other 40 Pg is respired. Organic carbon degradation occurs as particles (marine snow) settle through the ocean interior. Only 2 Pg eventually arrives at the seafloor, while the other 8 Pg is respired in the dark ocean. In sediments, the time scale available for degradation increases by orders of magnitude with the result that 90% of the organic carbon delivered is degraded and only 0.2 Pg C yr−1 is eventually buried and transferred from the biosphere to the geosphere. [26]
The diagram on the right shows a more complex model with many interacting boxes. Reservoir masses here represents carbon stocks, measured in Pg C. Carbon exchange fluxes, measured in Pg C yr−1, occur between the atmosphere and its two major sinks, the land and the ocean. The black numbers and arrows indicate the reservoir mass and exchange fluxes estimated for the year 1750, just before the Industrial Revolution. The red arrows (and associated numbers) indicate the annual flux changes due to anthropogenic activities, averaged over the 2000–2009 time period. They represent how the carbon cycle has changed since 1750. Red numbers in the reservoirs represent the cumulative changes in anthropogenic carbon since the start of the Industrial Period, 1750–2011. [28] [29] [27]
There are fast and slow biogeochemical cycles. Fast cycle operate in the biosphere and slow cycles operate in rocks. Fast or biological cycles can complete within years, moving substances from atmosphere to biosphere, then back to the atmosphere. Slow or geological cycles can take millions of years to complete, moving substances through the Earth's crust between rocks, soil, ocean and atmosphere. [31]
As an example, the fast carbon cycle is illustrated in the diagram below on the left. This cycle involves relatively short-term biogeochemical processes between the environment and living organisms in the biosphere. It includes movements of carbon between the atmosphere and terrestrial and marine ecosystems, as well as soils and seafloor sediments. The fast cycle includes annual cycles involving photosynthesis and decadal cycles involving vegetative growth and decomposition. The reactions of the fast carbon cycle to human activities will determine many of the more immediate impacts of climate change. [32] [33] [34] [35]
The slow cycle is illustrated in the diagram above on the right. It involves medium to long-term geochemical processes belonging to the rock cycle. The exchange between the ocean and atmosphere can take centuries, and the weathering of rocks can take millions of years. Carbon in the ocean precipitates to the ocean floor where it can form sedimentary rock and be subducted into the Earth's mantle. Mountain building processes result in the return of this geologic carbon to the Earth's surface. There the rocks are weathered and carbon is returned to the atmosphere by degassing and to the ocean by rivers. Other geologic carbon returns to the ocean through the hydrothermal emission of calcium ions. In a given year between 10 and 100 million tonnes of carbon moves around this slow cycle. This includes volcanoes returning geologic carbon directly to the atmosphere in the form of carbon dioxide. However, this is less than one percent of the carbon dioxide put into the atmosphere by burning fossil fuels. [31] [32]
The terrestrial subsurface is the largest reservoir of carbon on earth, containing 14–135 Pg of carbon [36] and 2–19% of all biomass. [37] Microorganisms drive organic and inorganic compound transformations in this environment and thereby control biogeochemical cycles. Current knowledge of the microbial ecology of the subsurface is primarily based on 16S ribosomal RNA (rRNA) gene sequences. Recent estimates show that <8% of 16S rRNA sequences in public databases derive from subsurface organisms [38] and only a small fraction of those are represented by genomes or isolates. Thus, there is remarkably little reliable information about microbial metabolism in the subsurface. Further, little is known about how organisms in subsurface ecosystems are metabolically interconnected. Some cultivation-based studies of syntrophic consortia [39] [40] [41] and small-scale metagenomic analyses of natural communities [42] [43] [44] suggest that organisms are linked via metabolic handoffs: the transfer of redox reaction products of one organism to another. However, no complex environments have been dissected completely enough to resolve the metabolic interaction networks that underpin them. This restricts the ability of biogeochemical models to capture key aspects of the carbon and other nutrient cycles. [45] New approaches such as genome-resolved metagenomics, an approach that can yield a comprehensive set of draft and even complete genomes for organisms without the requirement for laboratory isolation [42] [46] [47] have the potential to provide this critical level of understanding of biogeochemical processes. [48]
Some of the more well-known biogeochemical cycles are shown below:
Many biogeochemical cycles are currently being studied for the first time. Climate change and human impacts are drastically changing the speed, intensity, and balance of these relatively unknown cycles, which include:
Biogeochemical cycles always involve active equilibrium states: a balance in the cycling of the element between compartments. However, overall balance may involve compartments distributed on a global scale.
As biogeochemical cycles describe the movements of substances on the entire globe, the study of these is inherently multidisciplinary. The carbon cycle may be related to research in ecology and atmospheric sciences. [53] Biochemical dynamics would also be related to the fields of geology and pedology. [54]
The carbon cycle is that part of the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of Earth. Other major biogeochemical cycles include the nitrogen cycle and the water cycle. Carbon is the main component of biological compounds as well as a major component of many minerals such as limestone. A series of processes known as the carbon cycle are essential to Earth's ability to support life. It describes the movement of carbon as it is recycled and reused throughout the biosphere, as well as long-term processes of carbon sequestration (storage) to and release from carbon sinks.
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.
The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. In other words, it is a biologically mediated process which results in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).
Biogeochemistry is the scientific discipline that involves the study of the chemical, physical, geological, and biological processes and reactions that govern the composition of the natural environment. In particular, biogeochemistry is the study of biogeochemical cycles, the cycles of chemical elements such as carbon and nitrogen, and their interactions with and incorporation into living things transported through earth scale biological systems in space and time. The field focuses on chemical cycles which are either driven by or influence biological activity. Particular emphasis is placed on the study of carbon, nitrogen, oxygen, sulfur, iron, and phosphorus cycles. Biogeochemistry is a systems science closely related to systems ecology.
The pedosphere is the outermost layer of the Earth that is composed of soil and subject to soil formation processes. It exists at the interface of the lithosphere, atmosphere, hydrosphere and biosphere. The pedosphere is the skin of the Earth and only develops when there is a dynamic interaction between the atmosphere, biosphere, lithosphere and the hydrosphere. The pedosphere is the foundation of terrestrial life on Earth.
The iron cycle (Fe) is the biogeochemical cycle of iron through the atmosphere, hydrosphere, biosphere and lithosphere. While Fe is highly abundant in the Earth's crust, it is less common in oxygenated surface waters. Iron is a key micronutrient in primary productivity, and a limiting nutrient in the Southern ocean, eastern equatorial Pacific, and the subarctic Pacific referred to as High-Nutrient, Low-Chlorophyll (HNLC) regions of the ocean.
The sulfur cycle is a biogeochemical cycle in which the sulfur moves between rocks, waterways and living systems. It is important in geology as it affects many minerals and in life because sulfur is an essential element (CHNOPS), being a constituent of many proteins and cofactors, and sulfur compounds can be used as oxidants or reductants in microbial respiration. The global sulfur cycle involves the transformations of sulfur species through different oxidation states, which play an important role in both geological and biological processes. Steps of the sulfur cycle are:
In oceanic biogeochemistry, the f-ratio is the fraction of total primary production fuelled by nitrate. The ratio was originally defined by Richard Eppley and Bruce Peterson in one of the first papers estimating global oceanic production. This fraction was originally believed significant because it appeared to directly relate to the sinking (export) flux of organic marine snow from the surface ocean by the biological pump. However, this interpretation relied on the assumption of a strong depth-partitioning of a parallel process, nitrification, that more recent measurements has questioned.
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.
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.
Marine chemistry, also known as ocean chemistry or chemical oceanography, is influenced by plate tectonics and seafloor spreading, turbidity currents, sediments, pH levels, atmospheric constituents, metamorphic activity, and ecology. The field of chemical oceanography studies the chemistry of marine environments including the influences of different variables. Marine life has adapted to the chemistries unique to Earth's oceans, and marine ecosystems are sensitive to changes in ocean chemistry.
Rare biosphere refers to a large number of rare species of microbial life, i.e. bacteria, archaea and fungi, that can be found in very low concentrations in an environment.
The carbon cycle is an essential part of life on Earth. About half the dry weight of most living organisms is carbon. It plays an important role in the structure, biochemistry, and nutrition of all living cells. Living biomass holds about 550 gigatons of carbon, most of which is made of terrestrial plants (wood), while some 1,200 gigatons of carbon are stored in the terrestrial biosphere as dead biomass.
The oceanic carbon cycle is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many interacting forces across multiple time and space scales that circulates carbon around the planet, ensuring that carbon is available globally. The Oceanic carbon cycle is a central process to the global carbon cycle and contains both inorganic carbon and organic carbon. Part of the marine carbon cycle transforms carbon between non-living and living matter.
Chemical cycling describes systems of repeated circulation of chemicals between other compounds, states and materials, and back to their original state, that occurs in space, and on many objects in space including the Earth. Active chemical cycling is known to occur in stars, many planets and natural satellites.
Mary Ann Moran is a distinguished research professor of marine sciences at the University of Georgia in Athens. She studies the role of bacteria in Earth's marine nutrient cycles, and is a leader in the fields of marine sciences and biogeochemistry. Her work is focused on how microbes interact with dissolved organic matter and the impact of microbial diversity on the global carbon and sulfur cycles. By defining the roles of diverse bacteria in the carbon and sulfur cycles, she connects the biogeochemical and organismal approaches in marine science.
Marine biogeochemical cycles are biogeochemical cycles that occur within marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. These biogeochemical cycles are the pathways chemical substances and elements move through within the marine environment. In addition, substances and elements can be imported into or exported from the marine environment. These imports and exports can occur as exchanges with the atmosphere above, the ocean floor below, or as runoff from the land.
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.
The hydrothermal vent microbial community includes all unicellular organisms that live and reproduce in a chemically distinct area around hydrothermal vents. These include organisms in the microbial mat, free floating cells, or bacteria in an endosymbiotic relationship with animals. Chemolithoautotrophic bacteria derive nutrients and energy from the geological activity at Hydrothermal vents to fix carbon into organic forms. Viruses are also a part of the hydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.
The gold cycle is the biogeochemical cycling of gold through the lithosphere, hydrosphere, atmosphere, and biosphere. Gold is a noble transition metal that is highly mobile in the environment and subject to biogeochemical cycling, driven largely by microorganisms. Gold undergoes processes of solubilization, stabilization, bioreduction, biomineralization, aggregation, and ligand utilization throughout its cycle. These processes are influenced by various microbial populations and cycling of other elements such as carbon, nitrogen, and sulfur. Gold exists in several forms in the Earth's surface environment including Au(I/III)-complexes, nanoparticles, and placer gold particles. The gold biogeochemical cycle is highly complex and strongly intertwined with cycling of other metals including silver, copper, iron, manganese, arsenic, and mercury. Gold is important in the biotech field for applications such as mineral exploration, processing and remediation, development of biosensors and drug delivery systems, industrial catalysts, and for recovery of gold from electronic waste.
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