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Soil respiration refers to the production of
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
when soil organisms respire. This includes respiration of
plant root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...
s, the
rhizosphere The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microo ...
, microbes and
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is ''funga''. Flora, fauna, funga and other forms of life are collectively referred to as ''Biota (ecology ...
. Soil respiration is a key
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
process that releases
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon mak ...
from the soil in the form of CO2. CO2 is acquired by plants from the atmosphere and converted into organic compounds in the process of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. Plants use these organic compounds to build structural components or respire them to release energy. When
plant respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
occurs below-ground in the roots, it adds to soil respiration. Over time, plant structural components are consumed by
heterotroph 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 ...
s. This heterotrophic consumption releases CO2 and when this CO2 is released by below-ground organisms, it is considered soil respiration. The amount of soil respiration that occurs in an ecosystem is controlled by several factors. The temperature, moisture, nutrient content and level of
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
in the soil can produce extremely disparate rates of respiration. These rates of respiration can be measured in a variety of methods. Other methods can be used to separate the source components, in this case the type of photosynthetic pathway ( C3/ C4), of the respired plant structures. Soil respiration rates can be largely affected by human activity. This is because humans have the ability to and have been changing the various controlling factors of soil respiration for numerous years. Global
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
is composed of numerous changing factors including rising atmospheric CO2, increasing temperature and shifting
precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hail. ...
patterns. All of these factors can affect the rate of global soil respiration. Increased
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
by humans also has the potential to affect rates over the entire
planet A planet is a large, rounded astronomical body that is neither a star nor its remnant. The best available theory of planet formation is the nebular hypothesis, which posits that an interstellar cloud collapses out of a nebula to create a you ...
. Soil respiration and its rate across ecosystems is extremely important to understand. This is because soil respiration plays a large role in global carbon cycling as well as other
nutrient cycle A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cyc ...
s. The respiration of plant structures releases not only CO2 but also other nutrients in those structures, such as nitrogen. Soil respiration is also associated with positive feedback with global climate change. Positive feedback is when a change in a system produces response in the same direction of the change. Therefore, soil respiration rates can be affected by climate change and then respond by enhancing climate change.


Sources of carbon dioxide in soil

All cellular respiration releases energy, water and CO2 from organic compounds. Any respiration that occurs below-ground is considered soil respiration. Respiration by plant roots, bacteria,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
and soil animals all release CO2 in soils, as described below.


Tricarboxylic acid (TCA) cycle

The
tricarboxylic acid A tricarboxylic acid is an organic carboxylic acid whose chemical structure contains three carboxyl functional groups (-COOH). The best-known example of a tricarboxylic acid is citric acid. Uses Citric acid cycle Citric acid, a type of tricar ...
(TCA) cycle – or
citric acid cycle The citric acid cycle (CAC)—also known as the Krebs cycle or the TCA cycle (tricarboxylic acid cycle)—is a series of chemical reactions to release stored energy through the oxidation of acetyl-CoA derived from carbohydrates, fats, and protein ...
– is an important step in cellular respiration. In the TCA cycle, a six carbon sugar is
oxidized Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a ...
.Berg J, Tymoczko J, Stryer L. (2002). Biochemistry. WH Freeman and Company. This oxidation produces the CO2 and H2O from the sugar. Plants, fungi, animals and bacteria all use this cycle to convert organic compounds to energy. This is how the majority of soil respiration occurs at its most basic level. Since the process relies on oxygen to occur, this is referred to as aerobic respiration.


Fermentation

Fermentation is another process in which cells gain energy from organic compounds. In this
metabolic pathway In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical reac ...
, energy is derived from the carbon compound without the use of oxygen. The products of this reaction are carbon dioxide and usually either
ethyl alcohol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a hyd ...
or
lactic acid Lactic acid is an organic acid. It has a molecular formula . It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as well as nat ...
.Klein D, Prescott L, Harley J. (2005). Microbiology. McGraw-Hill. Due to the lack of oxygen, this pathway is described as
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing r ...
. This is an important source of CO2 in soil respiration in waterlogged ecosystems where oxygen is scarce, as in
peat bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
s and
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The p ...
s. However, most CO2 released from the soil occurs via respiration and one of the most important aspects of below-ground respiration occurs in the plant roots.


Root respiration

Plants respire some of the carbon compounds which were generated by photosynthesis. When this respiration occurs in roots, it adds to soil respiration. Root respiration accounts for approximately half of all soil respiration. However, these values can range from 10 to 90% depending on the dominate plant types in an
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
and conditions under which the plants are subjected. Thus, the amount of CO2 produced through root respiration is determined by the root biomass and specific root respiration rates.Shibistova O, Lloyd J, Evgrafova S, Savushkina N, Zrazhevskaya G, Arneth A, Knohl A, Kolle O. (2002) Seasonal and spatial variability in soil CO2 efflux rates for a central Siberian Pinus sylvestris forest. Tellus 54B, 552–567. Directly next to the root is the area known as the rhizosphere, which also plays an important role in soil respiration.


Rhizosphere respiration

The
rhizosphere The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microo ...
is a zone immediately next to the root surface with its neighboring soil. In this zone there is a close interaction between the plant and microorganisms. Roots continuously release substances, or exudates, into the soil. These exudates include sugars,
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...
s,
vitamin A vitamin is an organic molecule (or a set of molecules closely related chemically, i.e. vitamers) that is an essential micronutrient that an organism needs in small quantities for the proper functioning of its metabolism. Essential nutrie ...
s, long chain
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or m ...
s,
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s and
lysate Lysis ( ) is the breaking down of the membrane of a cell, often by viral, enzymic, or osmotic (that is, "lytic" ) mechanisms that compromise its integrity. A fluid containing the contents of lysed cells is called a ''lysate''. In molecular bio ...
s which are released when roots cells break. The amount of carbon lost as exudates varies considerably between plant species. It has been demonstrated that up to 20% of carbon acquired by photosynthesis is released into the soil as root exudates.Hutsch B, Augustin J, Merbach W. (2002) Plant rhizodeposition – an important source for carbon turnover in soils. Journal of Plant Nutrition and Soil Science. 165, 4, 397–407. These exudates are decomposed primarily by bacteria. These bacteria will respire the carbon compounds through the TCA cycle; however, fermentation is also present. This is due to the lack of oxygen due to greater oxygen consumption by the root as compared to the bulk soil, soil at a greater distance from the root.Vance E, Chapin III F. (2001) Substrate limitations to microbial activity in
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruc ...
forest floors. Soil Biology and Biochemistry. 33, 2, 173–188.
Another important organism in the rhizosphere are root-infecting fungi or
mycorrhizae   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
. These fungi increase the surface area of the plant root and allow the root to encounter and acquire a greater amount of soil nutrients necessary for plant growth. In return for this benefit, the plant will transfer sugars to the fungi. The fungi will respire these sugars for energy thereby increasing soil respiration.Harrison M. (2005) Peace Talks and Trade Deals. Keys to Long-Term Harmony in Legume-Microbe Symbioses. Plant Physiology. 137, 4, 1205–1210. Fungi, along with bacteria and soil animals, also play a large role in the decomposition of
litter Litter consists of waste products that have been discarded incorrectly, without consent, at an unsuitable location. Litter can also be used as a verb; to litter means to drop and leave objects, often man-made, such as aluminum cans, paper cups ...
and soil organic matter.


Soil animals

Soil animals graze on populations of bacteria and fungi as well as ingest and break up litter to increase soil respiration.
Microfauna Microfauna (Ancient Greek ''mikros'' "small" + New Latin ''fauna'' "animal") refers to microscopic animals and organisms that exhibit animal-like qualities. Microfauna are represented in the animal kingdom (e.g., nematodes, small arthropods) and t ...
are made up of the smallest soil animals. These include nematodes and mites. This group specializes on soil bacteria and fungi. By ingesting these organisms, carbon that was initially in plant organic compounds and was incorporated into bacterial and fungal structures will now be respired by the soil animal. Mesofauna are soil animals from in length and will ingest soil litter. The
fecal Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
material will hold a greater amount of moisture and have a greater surface area. This will allow for new attack by microorganisms and a greater amount of soil respiration.
Macrofauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zoo ...
are organisms from , such as
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
s and
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blatto ...
s. Most macrofauna fragment litter, thereby exposing a greater amount of area to microbial attack. Other macrofauna burrow or ingest litter, reducing soil bulk density, breaking up soil aggregates and increasing soil aeration and the infiltration of water.Chapin III F, Matson P, Mooney H. (2002) Principles of terrestrial ecosystem ecology. Springer-Verlag, New York.


Regulation of soil respiration

Regulation of CO2 production in soil is due to various
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
, or non-living, factors. Temperature, soil moisture and
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
all contribute to the rate of respiration in soil.


Temperature

Temperature affects almost all aspects of respiration processes. Temperature will increase respiration exponentially to a maximum, at which point respiration will decrease to zero when enzymatic activity is interrupted. Root respiration increases exponentially with temperature in its low range when the respiration rate is limited mostly by the TCA cycle. At higher temperatures the transport of sugars and the products of metabolism become the limiting factor. At temperatures over , root respiration begins to shut down completely.Atkin O, Edwards E, Loveys B. (2000) Response of root respiration to changes in temperature and its relevance to global warming. New Phytologist. 147, 141–154. Microorganisms are divided into three temperature groups; cryophiles, mesophiles and
thermophile A thermophile is an organism—a type of extremophile—that thrives at relatively high temperatures, between . Many thermophiles are archaea, though they can be bacteria or fungi. Thermophilic eubacteria are suggested to have been among the earl ...
s. Cryophiles function optimally at temperatures below , mesophiles function best at temperatures between 20 and and thermophiles function optimally at over . In natural soils many different cohorts, or groups of microorganisms exist. These cohorts will all function best at different conditions, so respiration may occur over a very broad range.Mikan C, Schimel J, Doyle A. (2002) Temperature controls of microbial respiration in Arctic tundra soils above and below freezing. Soil Biology and Biochemistry. 34, 1785–1795. Temperature increases lead to greater rates of soil respiration until high values retard microbial function, this is the same pattern that is seen with soil moisture levels.


Soil moisture

Soil moisture is another important factor influencing soil respiration. Soil respiration is low in dry conditions and increases to a maximum at intermediate moisture levels until it begins to decrease when moisture content excludes oxygen. This allows anaerobic conditions to prevail and depress aerobic microbial activity. Studies have shown that soil moisture only limits respiration at the lowest and highest conditions with a large plateau existing at intermediate soil moisture levels for most ecosystems.Xu L, Baldocchi D, Tang J. (2004) How soil moisture, rain pulses, and growth alter the response of ecosystem respiration and temperature. Global Biogeochemical Cycles. 18. Many microorganisms possess strategies for growth and
survival Survival, or the act of surviving, is the propensity of something to continue existing, particularly when this is done despite conditions that might kill or destroy it. The concept can be applied to humans and other living things (or, hypotheti ...
under low soil moisture conditions. Under high soil moisture conditions, many bacteria take in too much water causing their cell membrane to lyse, or break. This can decrease the rate of soil respiration temporarily, but the lysis of bacteria causes for a spike in resources for many other bacteria. This rapid increase in available labile substrates causes short-term enhanced soil respiration. Root respiration will increase with increasing soil moisture, especially in dry ecosystems; however, individual species' root respiration response to soil moisture will vary widely from species to species depending on life history traits. Upper levels of soil moisture will depress root respiration by restricting access to atmospheric oxygen. With the exception of wetland plants, which have developed specific mechanisms for root aeration, most plants are not adapted to wetland soil environments with low oxygen.Lambers H, Chapin III F, Pons T. (1998) Plant physiological ecology. Springer-Verlag, New York. The respiration dampening effect of elevated soil moisture is amplified when soil respiration also lowers soil
redox Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a ...
through bioelectrogenesis. Soil-based microbial fuel cells are becoming popular educational tools for science classrooms.


Nitrogen

Nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
directly affects soil respiration in several ways. Nitrogen must be taken in by roots to promote plant growth and life. Most available nitrogen is in the form of NO3, which costs 0.4 units of CO2 to enter the root because energy must be used to move it up a
concentration gradient Molecular diffusion, often simply called diffusion, is the thermal motion of all (liquid or gas) particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size (mass) o ...
. Once inside the root the NO3 must be reduced to NH3. This step requires more energy, which equals 2 units of CO2 per molecule reduced. In plants with bacterial symbionts, which fix atmospheric nitrogen, the energetic cost to the plant to acquire one molecule of NH3 from atmospheric N2 is 2.36 CO2. It is essential that plants uptake nitrogen from the soil or rely on symbionts to fix it from the atmosphere to assure growth, reproduction and long-term survival. Another way nitrogen affects soil respiration is through litter
decomposition Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is e ...
. High nitrogen litter is considered high quality and is more readily decomposed by microorganisms than low quality litter. Degradation of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
, a tough plant structural compound, is also a nitrogen limited process and will increase with the addition of nitrogen to litter.Sinsabaugh R, Carreiro M, Repert D. (2002) Allocation of extracellular enzymatic activity in relation to litter composition, N deposition, and mass loss. Biogeochemistry. 60, 1–24.


Methods of measurement

Different methods exist for the measurement of soil respiration rate and the determination of sources. Methods can be divided into field- and laboratory-based methods. The most common field methods include the use of long-term stand alone soil flux systems for measurement at one location at different times; survey soil respiration systems for measurement of different locations and at different times. The use of
stable isotope ratio The term stable isotope has a meaning similar to stable nuclide, but is preferably used when speaking of nuclides of a specific element. Hence, the plural form stable isotopes usually refers to isotopes of the same element. The relative abunda ...
s can be used both in laboratory of field measurements. Soil respiration can be measured alone or with added nutrients and (carbon) substrates that supply food sources to the microorganisms. Soil respiration without any additions of nutrients and substrates is called the basal soil respiration (BR). With the addition of nutrients (often nitrogen and phosphorus) and substrates (e.g. sugars), it is called the substrate-induced soil respiration (SIR). In both BR and SIR measurements, the moisture content can be adjusted with water.


Field methods


Long-term stand-alone soil flux systems for measurement at one location over time

These systems measure at one location over long periods of time. Since they only measure at one location, it is common to use multiple stations to reduce measuring error caused by soil variability over small distances. Soil variability may be tested with survey soil respiration instruments. The long-term instruments are designed to expose the measuring site to ambient conditions as much as is possible between measurements.


Types of long-term stand-alone instruments


= Closed, non-steady state systems

= Closed systems take short-term measurements (typically over few minutes only) in a chamber sealed over the soil. The rate of soil CO2 efflux is calculated on the basis of CO2 increased inside the chamber. As it is within the nature of closed chambers that CO2 continues to accumulate, measurement periods are reduced to a minimum to achieve a detectable, linear concentration increase, avoiding an excessive build-up of CO2 inside the chamber over time. Both individual assay information and diurnal CO2 respiration measuring information is accessible. It is also common for such systems to also measure soil temperature, soil moisture and PAR (
photosynthetically active radiation Photosynthetically active radiation (PAR) designates the spectral range (wave band) of solar radiation from 400 to 700 nanometers that photosynthetic organisms are able to use in the process of photosynthesis. This spectral region corresponds more o ...
). These variables are normally recorded in the measuring file along with CO2 values. For determination of soil respiration and the slope of CO2 increase, researchers have used linear regression analysis, the Pedersen (2001) algorithm, and exponential regression. There are more published references for linear regression analysis; however, the Pedersen algorithm and exponential regression analysis methods also have their following. Some systems offer a choice of mathematical methods.Wayson C, Randolph J, Hanson P, Grimmond P, Schmid H. (2006) Comparison of soil respiration methods in a mid-latitude deciduous forest. Biogeochemistry. 80, 173–189. When using linear regression, multiple data points are graphed and the points can be fitted with a linear regression equation, which will provide a slope. This slope can provide the rate of soil respiration with the equation F=bV/A, where ''F'' is the rate of soil respiration, ''b'' is the slope, ''V'' is the volume of the chamber and ''A'' is the surface area of the soil covered by the chamber.Field C, Ball J, Berry J. (1989) Photosynthesis, Principles and field techniques. Plant physiological ecology, field methods and instrumentation. Chapman and Hall, New York. It is important that the measurement is not allowed to run over a longer period of time as the increase in CO2 concentration in the chamber will also increase the concentration of CO2 in the porous top layer of the soil profile. This increase in concentration will cause an underestimation of soil respiration rate due to the additional CO2 being stored within the soil.Conen F, and Smith K. (2000) An explanation of linear increases in gas concentration under closed chambers used to measure gas exchange between soil and the atmosphere. European Journal of Soil Science. 51, 111–117.


= Open, steady-state systems

= Open mode systems are designed to find soil flux rates when measuring chamber equilibrium has been reached. Air flows through the chamber before the chamber is closed and sealed. This purges any non-ambient CO2 levels from the chamber before measurement. After the chamber is closed, fresh air is pumped into the chamber at a controlled and programmable flow rate. This mixes with the CO2 from the soil, and after a time, equilibrium is reached. The researcher specifies the equilibrium point as the difference in CO2 measurements between successive readings, in an elapsed time. During the assay, the rate of change slowly reduces until it meets the customer's rate of change criteria, or the maximum selected time for the assay. Soil flux or rate of change is then determined once equilibrium conditions are reached within the chamber. Chamber flow rates and times are programmable, accurately measured, and used in calculations. These systems have vents that are designed to prevent a possible unacceptable buildup of partial CO2 pressure discussed under closed mode systems. Since the air movement inside the chamber might cause increased chamber pressure, or external winds may produce reduced chamber pressure, a vent is provided that is designed to be as wind proof as possible. Open systems are also not as sensitive to soil structure variation, or to boundary layer resistance issues at the soil surface. Air flow in the chamber at the soil surface is designed to minimize boundary layer resistance phenomena.


= Hybrid Mode Systems

= A hybrid system also exists. It has a vent that is designed to be as wind proof as possible, and prevent possible unacceptable partial CO2 pressure buildup, but is designed to operate like a closed mode design system in other regards.


Survey soil respiration systems – for testing the variation of CO2 respiration at different locations and at different times

These are either open or closed mode instruments that are portable or semi-portable. They measure CO2 soil respiration variability at different locations and at different times. With this type of instrument, soil collars that can be connected to the survey measuring instrument are inserted into the ground and the soil is allowed to stabilize for a period of time. The insertion of the soil collar temporarily disturbs the soil, creating measuring artifacts. For this reason, it is common to have several soil collars inserted at different locations. Soil collars are inserted far enough to limit lateral diffusion of CO2. After soil stabilization, the researcher then moves from one collar to another according to experimental design to measure soil respiration. Survey soil respiration systems can also be used to determine the number of long-term stand-alone temporal instruments that are required to achieve an acceptable level of error. Different locations may require different numbers of long-term stand-alone units due to greater or lesser soil respiration variability.


Isotope methods

Plants acquire CO2 and produce organic compounds with the use of one of three photosynthetic pathways. The two most prevalent pathways are the C3 and C4 processes. C3 plants are best adapted to cool and wet conditions while C4 plants do well in hot and dry ecosystems. Due to the different photosynthetic enzymes between the two pathways, different
carbon isotope Carbon (6C) has 15 known isotopes, from to , of which and are stable. The longest-lived radioisotope is , with a half-life of years. This is also the only carbon radioisotope found in nature—trace quantities are formed cosmogenically by t ...
s are acquired preferentially.
Isotope Isotopes are two or more types of atoms that have the same atomic number (number of protons in their nuclei) and position in the periodic table (and hence belong to the same chemical element), and that differ in nucleon numbers (mass numb ...
s are the same element that differ in the number of neutrons, thereby making one isotope heavier than the other. The two stable carbon isotopes are 12C and 13C. The C3 pathway will discriminate against the heavier isotope more than the C4 pathway. This will make the plant structures produced from C4 plants more enriched in the heavier isotope and therefore root exudates and litter from these plants will also be more enriched. When the carbon in these structures is respired, the CO2 will show a similar ratio of the two isotopes. Researchers will grow a C4 plant on soil that was previously occupied by a C3 plant or vice versa. By taking soil respiration measurements and analyzing the isotopic ratios of the CO2 it can be determined whether the soil respiration is mostly old versus recently formed carbon. For example, maize, a C4 plant, was grown on soil where
spring wheat Winter wheat (usually '' Triticum aestivum'') are strains of wheat that are planted in the autumn to germinate and develop into young plants that remain in the vegetative phase during the winter and resume growth in early spring. Classification ...
, a C3 plant, was previously grown. The results showed respiration of C3 SOM in the first 40 days, with a gradual linear increase in heavy isotope enrichment until day 70. The days after 70 showed a slowing enrichment to a peak at day 100.Rochette P, Flanagan L, Gregorich E. (1999) Separating soil respiration into plant and soil components using analysis of natural abundance of carbon-13. Soil Science Society of America Journal. 63, 1207–1213. By analyzing stable carbon isotope data it is possible to determine the source components of respired SOM that was produced by different photosynthetic pathways.


Substrate-induced respiration in the field using stable isotopes

One problem in the measurement of soil respiration in the field is that respiration of microorganisms can not be distinguished from respiration from plant roots and soil animals. This can be overcome using stable isotope techniques. Cane sugar is a C4 – sugar which can act as an isotopic tracer. Cane sugar has a slightly higher abundance of 13C (δ13C ≈ −10‰) than the endogenous (natural) carbon in a C3 ecosystem (δ13C=−25 to −28‰). Cane sugar can be sprayed on the soil in a solution and will infiltrate the upper soil, Only microorganisms will respire the added sugar because roots exclusively respire carbon products that are assimilated by the plant via photosynthesis. By analyses of the δ13C of the CO2 evolving from the soil with or without adding cane sugar, the fraction of C3 (root and microbial) and C4 (microbial respiration) can be calculated. Field respiration using stable isotopes can be used as a tool to measure microbial respiration in-situ without disturbing the microbial communities by mixing soil nutrients, oxygen, and soil contaminants that may be present.


Responses to human disturbance

Throughout the past 160 years, humans have changed
land use Land use involves the management and modification of natural environment or wilderness into built environment such as settlements and semi-natural habitats such as arable fields, pastures, and managed woods. Land use by humans has a long ...
and industrial practices, which have altered the climate and global biogeochemical cycles. These changes have affected the rate of soil respiration around the planet. In addition, increasingly frequent extreme climatic events such as heat waves (involving high temperature disturbances and associated intense droughts), followed by intense rainfall, impact on microbial communities and soil physico-chemistry and may induce changes in soil respiration.


Elevated carbon dioxide

Since the
Industrial Revolution The Industrial Revolution was the transition to new manufacturing processes in Great Britain, continental Europe, and the United States, that occurred during the period from around 1760 to about 1820–1840. This transition included going f ...
, humans have emitted vast amounts of CO2 into the atmosphere. These emissions have increased greatly over time and have increased global atmospheric CO2 levels to their highest in over 750,000 years. Soil respiration increases when ecosystems are exposed to elevated levels of CO2. Numerous free air CO2 enrichment (FACE) studies have been conducted to test soil respiration under predicted future elevated CO2 conditions. Recent FACE studies have shown large increases in soil respiration due to increased root biomass and microbial activity.Lipson D, Wilson R, Oechel W. (2005) Effects of Elevated Atmospheric CO2 on Soil Microbial Biomass, Activity, and Diversity in a Chaparral Ecosystem. Applied and Environmental Microbiology. 71, 12, 8573–8580 Soil respiration has been found to increase up to 40.6% in a sweetgum forest in
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
and poplar forests in
Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...
under elevated CO2 conditions.King J, Hanson P, Bernhardt E, Deangelis P, Norby R, Pregitzer K. (2004) A multiyear synthesis of soil respiration responses to elevated atmospheric CO2 from four forest FACE experiments. Global Change Biology. 10, 1027–1042. It is extremely likely that CO2 levels will exceed those used in these FACE experiments by the middle of this century due to increased human use of fossil fuels and land use practices.


Climate warming

Due to the increase in temperature of the soil, CO2 levels in our atmosphere increase, and as such the mean average temperature of the Earth is rising. This is due to human activities such as forest clearing, soil denuding, and developments that destroy
autotrophic An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
processes. With the loss of photosynthetic plants covering and cooling the surface of the soil, the infrared energy penetrates the soil heating it up and causing a rise in heterotrophic bacteria. Heterotrophs in the soil quickly degrade the organic matter and soil structure crumbles, thus it dissolves into streams and rivers into the sea. Much of the organic matter swept away in floods caused by forest clearing goes into
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
, wetlands and eventually into the open ocean. Increased
turbidity Turbidity is the cloudiness or haziness of a fluid caused by large numbers of individual particles that are generally invisible to the naked eye, similar to smoke in air. The measurement of turbidity is a key test of water quality. Fluids ...
of surface waters causes biological oxygen demand and more autotrophic organisms die. Carbon dioxide levels rise with increased respiration of soil bacteria after temperatures rise due to loss of soil cover. As mentioned earlier, temperature greatly affects the rate of soil respiration. This may have the most drastic influence in the
Arctic The Arctic ( or ) is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenland), Finland, Iceland, N ...
. Large stores of carbon are locked in the frozen permafrost. With an increase in temperature, this permafrost is melting and aerobic conditions are beginning to prevail, thereby greatly increasing the rate of respiration in that ecosystem. Oechel W, Vourlitis G, Hastings S. (1995) Change in Arctic CO2 flux over two decades, Effects of climate change at Barrow, Alaska. Ecological Applications. 5, 3, 846–855.


Changes in precipitation

Due to the shifting patterns of temperature and changing oceanic conditions, precipitation patterns are expected to change in location, frequency and intensity. Larger and more frequent storms are expected when oceans can transfer more energy to the forming storm systems. This may have the greatest impact on
xeric Deserts and xeric shrublands are a biome defined by the World Wide Fund for Nature. Deserts and xeric (ancient Greek xērós, “dry") shrublands form the largest terrestrial biome, covering 19% of Earth's land surface area. Ecoregions in this h ...
, or arid, ecosystems. It has been shown that soil respiration in arid ecosystems shows dynamic changes within a raining cycle. The rate of respiration in dry soil usually bursts to a very high level after rainfall and then gradually decreases as the soil dries. With an increase in rainfall frequency and intensity over area without previous extensive rainfall, a dramatic increase in soil respiration can be inferred.


Nitrogen fertilization

Since the onset of the
Green Revolution The Green Revolution, also known as the Third Agricultural Revolution, was a period of technology transfer initiatives that saw greatly increased crop yields and agricultural production. These changes in agriculture began in developed countrie ...
in the middle of the last century, vast amounts of nitrogen fertilizers have been produced and introduced to almost all agricultural systems. This has led to increases in plant available nitrogen in ecosystems around the world due to agricultural runoff and wind-driven
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
. As discussed earlier, nitrogen can have a significant positive effect on the level and rate of soil respiration. Increases in soil nitrogen have been found to increase plant dark respiration, stimulate specific rates of root respiration and increase total root biomass.Lutze J, Gifford R, Adams H. (2000) Litter quality and decomposition in Danthonia richardsonii swards in response to CO2 and nitrogen supply over four years of growth. Global Change Biology. 6, 13–24. This is because high nitrogen rates are associated with high plant growth rates. High plant growth rates will lead to the increased respiration and biomass found in the study. With this increase in productivity, an increase in soil activities and therefore respiration can be assured.


Importance

Soil respiration plays a significant role in the global carbon and nutrient cycles as well as being a driver for changes in climate. These roles are important to our understanding of the natural world and human preservation.


Global carbon cycling

Soil respiration plays a critical role in the regulation of carbon cycling at the ecosystem level and at global scales. Each year approximately 120
petagram To help compare different orders of magnitude, the following lists describe various mass levels between 10−59  kg and 1052 kg. The least massive thing listed here is a graviton, and the most massive thing is the observable universe. ...
s (Pg) of carbon are taken up by land plants and a similar amount is released to the atmosphere through ecosystem respiration. The global soils contain up to 3150 Pg of carbon, of which 450 Pg exist in wetlands and 400 Pg in permanently frozen soils. The soils contain more than four times the carbon as the atmosphere.Sabine C, Hemann M, Artaxo P, Bakker D, Chen C, Field C, Gruber N, Le Quere C, Prinn R, Richey J, Romero-Lankao P, Sathaye J, Valentini R. (2003) Current status and past trends of the carbon cycle. Toward CO2 Stabilization: Issues, strategies, and consequences. Island Press. Washington DC. Researchers have estimated that soil respiration accounts for 77 Pg of carbon released to the atmosphere each year.Raich J, and Potter C. (1995) Global patterns of carbon dioxide emissions from soils. Global Biogeochemical Cycles. 9, 23–36. This level of release is greater than the carbon release due to anthropogenic sources (56 Pg per year) such as fossil fuel burning. Thus, a small change in soil respiration can seriously alter the balance of atmosphere CO2 concentration versus soil carbon stores. Much like soil respiration can play a significant role in the global carbon cycle, it can also regulate global
nutrient cycling A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cycli ...
.


Nutrient cycling

A major component of soil respiration is from the decomposition of litter which releases CO2 to the environment while simultaneously immobilizing or mineralizing nutrients. During decomposition, nutrients such as nitrogen are immobilized by microbes for their own growth. As these microbes are ingested or die, nitrogen is added to the soil. Nitrogen is also mineralized from the degradation of proteins and nucleic acids in litter. This mineralized nitrogen is also added to the soil. Due to these processes, the rate of nitrogen added to the soil is coupled with rates of microbial respiration. Studies have shown that rates of soil respiration were associated with rates of microbial turnover and nitrogen mineralization. Alterations of the global cycles can further act to change the climate of the planet.


Climate change

As stated earlier, the CO2 released by soil respiration is a greenhouse gas that will continue to trap energy and increase the global mean temperature if concentrations continue to rise. As global temperature rises, so will the rate of soil respiration across the globe thereby leading to a higher concentration of CO2 in the atmosphere, again leading to higher global temperatures. This is an example of a positive feedback loop. It is estimated that a rise in temperature by 2° Celsius will lead to an additional release of 10 Pg carbon per year to the atmosphere from soil respiration.Friedlingstein P, Dufresne J, Cox P. (2003) How positive is the feedback between climate change and the global carbon cycle? Tellus. 55B, 692–700. This is a larger amount than current anthropogenic carbon emissions. There also exists a possibility that this increase in temperature will release carbon stored in permanently frozen soils, which are now melting. Climate models have suggested that this positive feedback between soil respiration and temperature will lead to a decrease in soil stored carbon by the middle of the 21st century.Cox P, Betts R, Jones C, Spall S, Totterdell I. (2000) Acceleration of global warming due to carbon-cycle feedbacks in a coupled climate model. Nature. 408, 184–187.


Summary

Soil respiration is a key ecosystem process that releases carbon from the soil in the form of carbon dioxide. Carbon is stored in the soil as organic matter and is respired by plants, bacteria, fungi and animals. When this respiration occurs below ground, it is considered soil respiration. Temperature, soil moisture and nitrogen all regulate the rate of this conversion from carbon in soil organic compounds to CO2. Many methods are used to measure soil respiration; however, the closed dynamic chamber and use of stable isotope ratios are two of the most prevalent techniques. Humans have altered atmospheric CO2 levels, precipitation patterns and fertilization rates, all of which have had a significant role on soil respiration rates. The changes in these rates can alter the global carbon and nutrient cycles as well as play a significant role in climate change.


References

* * Su B. (2005
Interactions between ecosystem carbon, nitrogen and water cycles under global change: Results from field and mesocosm experiments.
University of Oklahoma, Norman, OK. * Flanagan L, and Veum A. (1974) Relationships between respiration, weight loss, temperature and moisture in organic residues in tundra. Soil Organisms and decomposition in Tundra. 249–277.


External links



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