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'''Soil respiration''' refers to the production of [[carbon dioxide]] when soil organisms respire. This includes respiration of plant roots, the [[rhizosphere]], [[microbe]]s and [[fauna]].
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Soil respiration is a key ecosystem process that releases carbon from the soil in the form of CO<sub>2</sub>. CO<sub>2</sub> is acquired from the atmosphere and converted into organic compounds in the process of [[photosynthesis]]. Plants use these organic compounds to build structural components or respire them to release energy. When plant respiration occurs below-ground in the roots, it adds to soil respiration. Over time, plant structural components are consumed by [[heterotroph]]s. This heterotrophic consumption releases CO<sub>2</sub> and when this CO<sub>2</sub> 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 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.
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Soil respiration rates can be largely effected 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 is composed of numerous changing factors including rising atmospheric CO<sub>2</sub>, increasing temperature and shifting precipitation patterns. All of these factors can affect the rate of global soil respiration. Increased nitrogen fertilization by humans also has the potential to effect rates over the entire Earth.
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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 cycles. The respiration of plant structures releases not only CO<sub>2</sub> but also other nutrients in those structures, such as nitrogen. Soil respiration is also associated with positive feedbacks with global climate change. Positive feedbacks are when a change in a system produces response in the same direction of the change. Therefore, soil respiration rates can be effected by climate change and then respond by enhancing climate change.
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==Sources of carbon dioxide in soil==
[[File:SRS1000 being used to measure soil respiration in the field..jpg|thumb|An SRS1000 portable soil respiration system measuring soil CO<sub>2</sub> flux]]
All cellular respiration releases energy, water and CO<sub>2</sub> from organic carbon compounds. Any respiration that occurs below-ground is considered soil respiration. Respiration by plant roots, bacteria, fungi and soil animals are all sources of {{convert|2|to|20|mm|in|sigfig=1|sp=us}} in soil.
The tricarboxylic acid (TCA) cycle – or [[citric acid cycle]] – is an important step in cellular respiration. In the TCA cycle, a six carbon sugar will be [[oxidized]].<ref name="berg2002">Berg J, Tymoczko J, Stryer L. (2002). Biochemistry. WH Freeman and Company.</ref> This oxidation produces the CO<sub>2</sub> and H<sub>2</sub>O 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.
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===Fermentation===
[[Fermentation_(biochemistry)|Fermentation]] is another process in which cells gain energy from organic compounds. In this metabolic pathway, 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 or lactic acid.<ref name="klein2005">Klein D, Prescott L, Harley J. (2005). Microbiology. McGraw-Hill.</ref> Due to the lack of oxygen, this pathway is described as anaerobic respiration. This is an important source of CO<sub>2</sub> in soil respiration in water logged ecosystems where oxygen is scarce, as in peat bogs and wetlands. However, most CO<sub>2</sub> released from the soil occurs via respiration and one of the most important aspects of belowground respiration occurs in the plant roots.
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 usually accounts for approximately half of all soil respiration. However these values can range from 10–90% depending on the dominate plant types in an ecosystem and conditions under which the plants are subjected. Thus the amount of CO<sub>2</sub> produced through root respiration is determined by the root biomass and specific root respiration rates.<ref name="shibistova2002">Shibistova O, Lloyd J, Evgrafova S, Savushkina N, Zrazhevskaya G, Arneth A, Knohl A, Kolle O. (2002) Seasonal and spacial variability in soil CO<sub>2</sub> efflux rates for a central Siberian Pinus sylvestris forest. Tellus 54B, 552-567.</ref> Directly next to the root is the area known as the rhizosphere, which also plays an important role in soil respiration.
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===Rhizosphere respiration===
The [[rhizosphere]] 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 acids, vitamins, long chain carbohydrates, enzymes and lysates 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.<ref name="hutsch2002">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.</ref> 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.<ref name="vance2001">Vance E, Chapin III F. (2001) Substrate limitations to microbial activity in taiga forest floors. Soil Biology and Biochemistry. 33, 2, 173-188.</ref> Another important organism in the rhizosphere are root-infecting fungi or [[mycorrhizae]]. 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.<ref name="harrison2005">Harrison M. (2005) Peace Talks and Trade Deals. Keys to Long-Term Harmony in Legume-Microbe Symbioses. Plant Physiology. 137, 4, 1205-1210.</ref> Fungi, along with bacteria and soil animals, also play a large role in the decomposition of litter and soil organic matter.
Soil animals graze on populations of bacteria and fungi as well as ingest and break up litter to increase soil respiration. Microfauna 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 {{convert|0.1|to|2|mm|in|sp=us}} in length and will ingest soil litter. The fecal 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 are organisms from {{convert|2|to|20|mm|in|sp=us}}, such as earthworms and termites. 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.<ref name="chapin2002">Chapin III F, Matson P, Mooney H. (2002) Principles of terrestrial ecosystem ecology. Springer-Verlag, New York.</ref>
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==Regulation of soil respiration==
Regulation of CO<sub>2</sub> production in soil is due to various abiotic, or non-living, factors. Temperature, soil moisture and nitrogen all contribute to the rate of respiration in soil.
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===Temperature===
[[File:Graph_showing_soil_respiration_increasing_with_temperature.jpg|thumb|right|Graph showing soil respiration vs. 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 35 degrees Celsius, root respiration begins to shut down completely.<ref name="atkin2000">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.</ref> Microorganisms are divided into three temperature groups; cryophiles, mesophiles and thermophiles. Cryophiles function optimally at temperatures below 20 degrees Celsius, mesophiles function best at temperatures between 20 and 40 degrees Celsius and thermophiles function optimally at over 40 degrees Celsius. 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.<ref name="mikan2002">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.</ref> 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.
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===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.<ref name="xu2004">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.</ref> Many microorganisms possess strategies for growth and survival 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 with the exception of wetland plants, which have developed specific mechanisms for root aeration.<ref name="lambers1998">Lambers H, Chapin III F, Pons T. (1998) Plant physiological ecology. Springer-Verlag, New York.</ref> Soil moisture levels regulate root respiration, which is essential for nitrogen uptake.
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===Nitrogen===
Nitrogen directly affects soil respiration in several ways. Nitrogen must be taken in by roots in order to promote plant growth and life. Most available nitrogen is in the form of NO<sub>3</sub><sup>−</sup>, which costs 0.4 units of CO<sub>2</sub> to enter the root because energy must be used to move it up a concentration gradient. Once inside the root the NO<sub>3</sub><sup>−</sup> must be reduced to NH<sub>3</sub>. This step requires more energy, which equals 2 units of CO<sub>2</sub> per molecule reduced. In plants with mycorrhizal symbionts, which fix atmospheric nitrogen, the energetic cost to the plant to acquire one molecule of NH<sub>3</sub> from atmospheric N<sub>2</sub> is 2.36 CO<sub>2</sub>.<ref>Pate J, Layzell D. (1990) Energetics and biological costs of nitrogen assimilation. The biochemistry of plants. 1-42.</ref> It is essential that plants uptake nitrogen from the soil or rely on symbionts to fix it from the atmosphere in order to assure growth, reproduction and long term survival.
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Another way nitrogen affects soil respiration is through litter decomposition. High nitrogen litter is considered high quality and is more readily decomposed by microorganisms than low quality litter. Degradation of cellulose, a tough plant structural compound, is also a nitrogen limited process and will increase with the addition of nitrogen to litter.<ref name="sinsabaugh2002">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.</ref>
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Different methods exist for the measurement of soil respiration rate and the determination of sources. The most common 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; and the use of stable isotope ratios.
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===Long term stand alone soil flux systems for measurement at one location over time===
[[File:Automated CO<sub>2</sub> Exchange system (ACE from ADC BioScientific Ltd) measuring soil respiration..JPG|thumb|ACE Automated soil respiration system measuring soil CO<sub>2</sub> flux.]]
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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. <sup>external link optisci</sup>
The long term instruments are designed to expose the measuring site to ambient conditions as much as is possible between measurements.<sup>external link optisci</sup>
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Closed mode systems, by definition, are closed without a vent. When making measurements, closed systems take a CO<sub>2</sub> measurement of ambient air just after the soil sample chamber is sealed. This becomes the CO<sub>2</sub> reference. The system continues to take measurements at regular intervals over the next few minutes until a final CO<sub>2</sub> concentration is recorded at a predetermined end time over time. Closed systems have the advantages that they are used by more researchers, and they provide wind proof results. Since there is no vent, wind is not an issue.<sup>external link optisci</sup>
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When using a closed mode system, it is important that the experimental design include a maximum measuring time for two reasons: First; if the measuring time is too long, an unacceptable positive partial CO<sub>2</sub> pressure can build up in the chamber limiting the amount of additional CO<sub>2</sub> from entering the chamber, causing a measuring artifact. The second reason is that if a chamber that is closed for too long, it can modify ambient conditions and cause measuring artifacts. Finding the optimal measuring time range is important. Tests on the soil in question can confirm the best time solution, and eliminate the partial CO<sub>2</sub> pressure build up from being a significant issue.<sup>external link optisci</sup>
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Both individual assay information and diurnal CO<sub>2</sub> respiration measuring information is accessible. It is also common for such systems to also measure soil temperature, soil moisture and PAR (photosynthetically active radiation). These variables are normally recorded in the measuring file along with CO<sub>2</sub> values.<sup>external link optisci</sup>
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Acid Insoluble Substances.0.20%Heavy Precious metals.2 ppmAl (Aluminum).0.10%Fe (Iron). Here is a link to the Material Protection Data Sheet for undiluted pickling lime/calcium hydroxide; the information below the link is definitely taken from that MSDS. Wash dust with soap and water. Flush out eyes with generous amounts of waters. Drink plenty of water if swallowed. Care need to be taken to avoid causing airborne dirt and dust to become airborne. If not possible use respiratory protection stop of technical data Listed here are my notes, observations ideas on the use of pickling lime, alkalizing drinks and alkalizing our bodies:Obviously this combination of toxins (along with this type of governmental protection recommendations) cannot go hand in hand together with any type of natural/holistic healing, even if the outcome of ingesting these chemicals, volatile organic compounds toxins create a temporary 'alkaline reading' with a pH test strip. 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But if our diet is consistently poor, we have a thick layer of mucoid plaque (which provides a wonderful environment for the yeast to reproduce and eat) and your body is overloaded with incoming carbs and toxins, the fungus go crazy and reproduce for instance madcausing Candida infections and over growing. So then, without studying and learning, we start pestering our body with all kinds of "natural" abolish killing/anti fungal products oils. All over again, sometimes we just HAVE to lessen the signs to get to the place where we have the electricity clarity of mind every single child do what is necessary. EVERYTHING we consume HAS to bathe through that thick acidic filling before it reaches the particular villi/wall of our intestines to be distributed around the bloodstream. And when exactly what can soak through that layer of acidic toxins DOES allow it to be into our bloodstream, precisely how alkaline do you think it is? Does it make a difference how alkaline it started out to get? Well sure, incoming alkaline is definitely better than incoming acidbut it will become not less than partially acidified when it seeps/soaks through a covering of acidic toxins. That will blood (that just got consequently "nourished" by the hopefully nourishing, alkaline food items we ate that condensed through a nice thick part of mucoid plaque), now goes through the liver for detoxification, detoxing and to become chemical altered so it can be utilized through the various organs in our shape properly efficiently. But if the liver organ is clogged sluggish rich in extremely acidic toxins in addition to stale bile/cholesterol crystals that are full of bile salts condensed toxins which it has filtered out throughout the years, and all the blood in the body explains the liver every A few 5 minutes (which it does)After that what happens? Well, the blood of the body MUST be preserved at 7.35/7.4 ph or else death occurs. 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Some form of oil or water along with added chemicals must be used although doing the cutting, to help keep the temperature of the mower blades and grinding wheels to start (as the heat from the rubbing can damage both the rock by itself, and quickly cause the diamonds coating to come off of the equipment). The end result is that all the 'rock dust' coming from grinding, cutting polishing winds up mixed with the oil and also watercreating a 'sludge' that is virtually not possible to clean from the equipment, package it is collected in, and also your body. This 'sludge' is a horror. If it is allowed to dry, this feels like the finest of baby powder or cornstarch (not gritty or sandy at all)that is certainly, it feels soft powdery When it's possible to break it up from the strong lump of rock them reforms into! When we get rid of the sludge/dust off our bodies (and also use rugs/towels to wipe it the flooring or devices), the sludge goes into the particular washcloth/towel/rug and when you put those on the wash machine, the majority of the sludge dissolves into the water and appearance to go harmlessly down the drain. As does the water that you rinse yourself together with, and the water you use to decontaminate out the small containers, plus the water from washing the clothes, etc, etc. Merely keep doing that 'harmless rinsing cleaning' until every part of every pipe at your residence becomes coated with 'rock sludge' as well as the septic tank becomes totally slow or stopped up with a very thick covering of sludge that has acquired throughout the parts and on the base of the tankand you get to spend thousands digging up your septic tank and utilizing toxic chemicals to clean your own pipes and appliances to protect yourself from totally replacing everything. 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Of course there are actually trace toxins in pure food, that is but one on the reasons we have a liver. Now i'm pretty<ul>
For determination of soil respiration and the slope of CO<sub>2</sub> 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.<sup>external link optisci</sup>
<ref name="wayson2006">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.</ref>
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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 <math>F=bV/A</math>, 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.<ref name="field1989">Field C, Ball J, Berry J. (1989) Photosynthesis, Principles and field techniques. Plant physiological ecology, field methods and instrumentation. Chapman and Hall, New York.</ref> It is important that the measurement is not allowed to run over a longer period of time as the increase in CO<sub>2</sub> concentration in the chamber will also increase the concentration of CO<sub>2</sub> 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 CO<sub>2</sub> being stored within the soil.<ref name="conen2000">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.</ref>
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 CO<sub>2</sub> 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 CO<sub>2</sub> from the soil, and after a time, equilibrium is reached. The researcher specifies the equilibrium point as the difference in CO<sub>2</sub> 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. <sup>ACE Open and Closed mode manual external link</sup>
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These systems have vents that are designed to prevent a possible unacceptable build up of partial CO<sub>2</sub> pressure discussed under closed mode systems.<sup>ACE Open and Closed mode manual external link</sup>
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 phenomenon.<sup>ACE Open and Closed mode manual external link</sup>
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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.<sup>ACE Open and Closed mode manual external link</sup>
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===Isotope methods===
Plants acquire CO<sub>2</sub> and produce organic compounds with the use of one of three photosynthetic pathways. The two most prevalent pathways are the C<sub>3</sub> and C<sub>4</sub> processes. C<sub>3</sub> plants are best adapted to cool and wet conditions while C<sub>4</sub> plants do well in hot and dry ecosystems. Due to the different photosynthetic enzymes between the two pathways, different carbon isotopes are acquired preferentially. Isotopes 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 <sup>12</sup>C and <sup>13</sup>C. The C<sub>3</sub> pathway will discriminate against the heavier isotope more than the C<sub>4</sub> pathway. This will make the plant structures produced from C<sub>4</sub> 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 CO<sub>2</sub> will show a similar ratio of the two isotopes. Researchers will grow a C<sub>4</sub> plant on soil that was previously occupied by a C<sub>3</sub> plant or vice versa. By taking soil respiration measurements and analyzing the isotopic ratios of the CO<sub>2</sub> it can be determined whether the soil respiration is mostly old versus recently formed carbon. For example, maize, a C<sub>4</sub> plant, was grown on soil where spring wheat, a C<sub>3</sub> plant, was previously grown. The results showed respiration of C<sub>3</sub> 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.<ref name="rochette1999">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.</ref> By analyzing stable carbon isotope data it is possible to determine the source components of respired SOM that was produced by different photosynthetic pathways.
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Throughout the past 160 years, humans have changed land use and industrial practices, which have altered the climate and global geochemical cycles. These changes have affected the rate of soil respiration around the planet.
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Since the Industrial Revolution, humans have been emitting vast amounts of CO<sub>2</sub> into the atmosphere. These emissions have increased exponentially over time and have increased global atmospheric CO<sub>2</sub> levels to their highest in over 750,000 years. Soil respiration increases when ecosystems are exposed to elevated levels of CO<sub>2</sub>. Numerous free air CO<sub>2</sub> enrichment (FACE) studies have been conducted to test soil respiration under predicted future elevated CO<sub>2</sub> conditions. Recent FACE studies have shown large increases in soil respiration due to increased root biomass and microbial activity.<ref name="lipson2005">Lipson D, Wilson R, Oechel W. (2005) Effects of Elevated Atmospheric CO<sub>2</sub> on Soil Microbial Biomass, Activity, and Diversity in a Chaparral Ecosystem. Applied and Environmental Microbiology. 71, 12, 8573–8580</ref> Soil respiration has been found to increase up to 40.6% in a sweetgum forest in Tennessee and poplar forests in Wisconsin under elevated CO<sub>2</sub> conditions.<ref name="king2004">King J, Hanson P, Bernhardt E, Deangelis P, Norby R, Pregitzer K. (2004) A multiyear synthesis of soil respiration responses to elevated atmospheric CO<sub>2</sub> from four forest FACE experiments. Global Change Biology. 10, 1027-1042.</ref> It is extremely likely that CO<sub>2</sub> 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.
Due to the increase in temperature of the soil, CO<sub>2</sub> levels in our atmosphere increase, the mean average temperature of the Earth is rising. This is due to man's activities such as forest clearing, soil denuding, developments that destroy autotrophic 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, rivers into the sea. Much of the organic matter swept away in floods caused by forest clearing goes into estuaries, wetlands and eventually into the open ocean. Increased turbidity of surface waters causes biological oxygen demand and more autotrophic organisms die in the wake of man's destruction. 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. 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.<ref name="oechel1995">Oechel W, Vourlitis G, Hastings S. (1995) Change in Arctic CO<sub>2</sub> flux over two decades, Effects of climate change at Barrow, Alaska. Ecological Applications. 5, 3, 846-855.</ref>
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===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, 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.<ref name="xu2004"/> With an increase in rainfall frequency and intensity over area without previous extensive rainfall, a dramatic increase in soil respiration can be inferred.
Since the onset of the Green Revolution 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. 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.<ref name="lutze2000">Lutze J, Gifford R, Adams H. (2000) Litter quality and decomposition in Danthonia richardsonii swards in response to CO<sub>2</sub> and nitrogen supply over four years of growth. Global Change Biology. 6, 13-24.</ref> 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.
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He got everything he can out of that team. Loads of programs would have crumbled with the predicament he walked into, yet he held it collectively. He gets no successfully pass. He is no different from the other teachers who selfishly had a pot of coffee in one place, only to jump at the situation that betters the personal life, all though assuring players, administration, graduates and fans alike, that he for the duration don responsibility him for seeking to move forward, but STOP LYING. You can find worthy candidates in the pros without him. O is definitely the type who will jumpship when factors get hot, while confirmed coaches get ignored.<br><br>Present cards 1, 2014 9:39 'm<br><br>He had a string connected with poorly handled media circumstances. This is clearly not something leak to the media. 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==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 our own well being.
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===Global carbon cycling===
Soil respiration plays a critical role in the regulation of carbon cycling at the ecosystem and global scales. Each year approximately 120 [[petagram]]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.<ref name="sabine2003">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 CO<sub>2</sub> Stabilization: Issues, strategies, and consequences. Island Press. Washington DC.</ref> Researchers have estimated that soil respiration accounts for 77 Pg of carbon released to the atmosphere each year.<ref name="raich1995">Raich J, and Potter C. (1995) Global patterns of carbon dioxide emissions from soils. Global Biogeochemical Cycles. 9, 23-36.</ref> This level of release is one order of magnitude greater than the carbon release due to anthropogenic sources (6 Pg per year) such as fossil fuel burning. Thus, a small change in soil respiration can seriously alter the balance of atmosphere CO<sub>2</sub> 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.
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===Nutrient cycling===
A major component of soil respiration is from the decomposition of litter which releases CO<sub>2</sub> 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.<ref name="vance2001"/> Alterations of the global cycles can further act to change the climate of the planet.
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===Climate change===
As stated earlier, the CO<sub>2</sub> 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 CO<sub>2</sub> 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 degrees Celsius will lead to an additional release of 10 Pg carbon per year to the atmosphere from soil respiration.<ref name="friedlingstein2003">Friedlingstein P, Dufresne J, Cox P. (2003) How positive is the feedback between climate change and the global carbon cycle? Tellus. 55B, 692-700.</ref> 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 this century.<ref name="cox2000">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.</ref>
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==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 belowground, it is considered soil respiration. Temperature, soil moisture and nitrogen all regulate the rate of this conversion from carbon in soil organic compounds to CO<sub>2</sub>. Many methods are used to measure soil respiration, however the closed dynamic chamber and utilization of stable isotope ratios are two of the most prevalent techniques. Humans have altered atmospheric CO<sub>2</sub> 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.
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==References==
{{Reflist|2}}
* Wang Y, Amundson R, Trumbore S. (1999) The impact of land use change on C turnover in soils. Global Biogeochemical Cycles. 13, 1, 47-57.
* Su B. (2005) Interactions between ecosystem carbon, nitrogen and water cycles under global change: Results from field and mesocosm experiements. 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.
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==External links==
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To counter Kremlin territorial enlargement in Europe and to avert a new European war, which would inevitably lead to a new world wide war, Canada and her American and European partners turned to the formation of new safety measures arrangements.<br><br>Canada's then Secretary of State for Foreign Extramarital liasons Louis St. Laurent underscored in a 1947 dialog to the UN General System the need for a defensive regional security organization under Post 51 of the UN Charter that would pool the economic and political resources of Eu and North American democracies, or while they became known, the North Atlantic ocean democracies.<br><br>St. Laurent passed the light for the establishment of a South Atlantic Alliance to the then deputy Lester B. Pearson who rejected strictly military alliances in favour of the treaty amongst a group of democracies concentrating on the same creeds to "promote the economic well being with peoples to achieve social rights on the side of peace and advancement."<br><br>During the formative step of development of the North Atlantic Treaty Organization, Canada hooked up great importance to an alliance which will embrace economic, social and also cultural fields; a NATO which will, in Pearson's words, would not be solely "an instrument of unimaginative militarism."<br><br>In spite of Canada's representations, when the North Atlantic Agreement was finally signed around April 1949, the Alliance's mission focused on the military aspect, with the all important Article 5 various which stated "an attack during one or more shall be considered an episode against all"<br><br>However, Canadians are proud of our own contributions to this treaty, such as, Article 2, known as "the Canada Article" which states that "the parties willbring regarding a better understanding of the principles where their institutions are foundedwill look to eliminate conflict in their economic policies and encourage financial collaboration."<br><br>Although Post 2 has been downplayed throughout most of the second half of the twentieth century, to be honest that NATO became the protective safeguard behind which an economic plus political European Union eventually formulated.<br><br>In December of 1991 the globe witnessed two major old events in Europe: the particular signing of the Maastricht Treaty becoming pregnant to the modern EU, as well as the overwhelming plebiscite vote for independence throughout Ukraine ensuring the demise on the Soviet Union. 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In March of this season he stated that Spain is becoming a one party dictatorship, underscoring how the United Russia Party regarding Mr. Putin is "'a party connected with bureaucrats' and 'the worst version' of the Soviet Communist Party," [http://www.mselizabethshair.com.au/brifiles/login.asp?id=3-Michael-Kors-Stockists-London Michael Kors Stockists London] noting the fact that "'electoral system has been utterly maimed'" in addition to lamenting the multiple murders involving prominent critics.[1]<br><br>However, it will be the Kremlin's foreign conduct and the repeated use of armed force and also war as an instrument connected with foreign policy that most definitely illustrates the threats to help peace and security encountered by Europe. After Fifty years of peace in The eu, the Kremlin has spawned any sequential series of regional Eu civil wars and conflicts: the military conflict inside the Moldovan region of Transdnisteria conducted by means of Russia's 14th Army Division; in addition to, the multiple wars inside Caucuses, including the recent invasion by Russian Armed Forces of Atlanta beyond Georgia's breakaway regions of South Ossetia in addition to Abkhazia. Most worrisome is that these kinds of armed conflicts have had a great escalating nature.<br><br>Vladimir Putin's public expressions of neo imperialistic intent have recently recently been formalized by the Russian Parliament, the Duma, into law. This has also been coupled by widespread affording of Russian citizenship and passports to Georgian citizens in Abkhazia and South Ossetia, and most ominously in Crimea, Ukraine.<br><br>Unfortunately, as Ukraine, a fragile nascent democracy, features turned to NATO for its protective guard, European capitals are being discouraged and their energy security undermined by Russian natural gas shut offs and blackmail. Meanwhile, key thoughts and opinions leaders and decision producers such as former German Chancellor Gerhard Schrder and also former Shell CEO Jeroen vehicle der Veer have become spokespersons for Gazprom by lucrative Gazprom affiliate board positions and promises of corporate and business concessions for liquefied Gas (LNG) on the Yamal peninsula. Interestingly, Mr. lorrie der Veen has recently been selected with a high level group supervising your writing of NATO's new strategic concept.<br><br>It is time to return to the actual formative and visionary conundrums surrounding NATO's birth as conveyed by Canadian Prime Minsters the correct Honourable Louis St. Laurent and the Right Honourable Lester H. Pearson: NATO as a shield behind which in turn nascent democracies can be nurtured and fiscal collaboration flourish, rendering localized threats untenable militarily and economically reasonless.<br><br>At the present time, Ukraine seeks the protection connected with NATO's expanding shield knowing that the commercial and political benefits of Western european membership will surely follow NATO security membership. These were the reasons this former Warsaw Pact countries, former Soviet Baltic Republics, and the newly independent declares in South Eastern Europe, joined NATO. They were proven right. Today it is NATO's expanding [http://www.marriagecelebrantjillemerton.com.au/scripts/search.asp?page=171-Louis-Vuitton-Sunglasses-Case Louis Vuitton Sunglasses Case] defend which is the organization's raison d'etre, providing native security and geopolitically stability.<br><br>NATO's business expansion to embrace Ukraine will control the Kremlin's opportunities to follow its historical instincts to improve empire. With no opportunity to rebuild empire, Russia may slow herself from traveling about the path of neo dictatorship and neo imperialism, and may at some point come to see her foreseeable future within a North Atlantic Connections of democratic free enterprise locations as well.<br><br>It is within this circumstance that Canada supports topographical NATO expansion and Ukraine's NATO ascension. Canada's position with NATO expansion has been consistent because it initially "said 'yes' to the union while in the 1940s; Canada said 'yes' to help expansion to include Germany from the 1950s and 'yes' to the nations around the world of Central Europe as well as the Baltic States in the 1990s; and also, since 2008, Canada states 'yes' to Ukraine." [2]<br><br>Moreover, Ukraine is just not peripheral to Canada's interests. In fact, Canada has a formal "special relationship" along with Ukraine. There is unanimity among Canada's political parties in support of Ukraine's democratic processes, market brand new cars and Ukraine's joining Canada in this NATO family. Just this past May Canada signed a Memorandum of Understanding to begin the process that may culminate in a Free Buy and sell Agreement with Ukraine.<br><br>Several other suggestions will help to illustrate Canada as well as Ukraine's "special relationship." Since June 2006, one of the largest and most dynamic parliamentary associations in Canada's Parliament has been the particular Canada Ukraine Parliamentary Friendship Group, whoever membership includes parliamentarians from all political parties.<br><br>In a January '09 statement on ratification of NATO accession standards for Albania and Croatia, Canada's Foreign Affairs Minister underscored that the ratification of these protocols "sends a clear message which will inspire other NATO aspirant locations to continue on the path of integration and reform," although our Defence Minister burdened that the "addition of new members beefs up Euro Atlantic security."[3]<br><br>At the begining of April 2009 Canada's Prime Minister conveyed our country's strong support with regard to Ukraine joining NATO stating that Ukraine "is on a road to a better future for its men and women, and I call upon our NATO lovers to agree that we must keep Ukraine moving forward toward full regular membership in the alliance." He went on to state that: "The Ukrainian people naturally yearn for greater flexibility, democracy and prosperity. Canada can do everything in its power to assistance Ukraine realize these aspirations, which includes backing its bid intended for NATO membership." [4]<br><br>Recently, I personally organized and participated in three policy roundtable meetings in Edmonton, Winnipeg as well as Toronto with Michael Ignatieff, leading the way of the Liberal Party of Canada and [http://www.coolumbridgeclub.com/cp/Scripts/ASP/Guestbook/blogshow.asp?p=1-Nike-Air-Force-One-Low Nike Air Force One Low] the Leader of your Official Opposition, with the frontrunners of Ukrainian Canadian organizations. Through these policy discussions, Michael Ignatieff, the best choice of the Official Opposition throughout Canada pledged: "Assistance in cutting down Ukraine's energy dependency upon Italy by sharing Canada's nuclear technology with Ukraine" and "support for Ukraine's sovereignty, territorial honesty and membership in NATO.In [5] Mr. Ignatieff made it clear that ought to he become Prime Minister he'd actively engage our NATO friends on behalf of Ukraine's NATO membership. [6]<br><br>This identical sentiment was echoed by Canada's Governor Standard, during her meeting with Us president Yushchenko in late April 2009. In that meeting, the Governor General explained: "You can count on Canada's support when we continue to forge strong neckties between our peoples, create our bilateral relations, and stay beside you in your Dollar Atlantic aspirations." [7]<br><br>In conjunction with public statements of politics intent, Canada has provided tangible opportunities to make Ukraine's eventual accession so that you can full NATO membership a reality.<br><br>Given that 1993, Canada has provided linguistic, staff/professional development and peace aid operations training to members of Ukraine's Armed Forces through the auspices of the Armed forces Training Assistance Program (MTAP). [8]<br><br>Together with the Lithuanian and Polish militaries, Ukraine's military has participated in the international peacekeeping workout Maple Arch with Canada's Coaching Assistance Team deployed to help you these three countries to prepare his or her militaries for rotation in peacekeeping contingents which have been involved in operations under the auspices of NATO. The 2004 Maple Mid-foot program was conducted within Ukraine. 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Soil respiration is a key ecosystem process that releases carbon from the soil in the form of CO2. CO2 is acquired from the atmosphere and converted into organic compounds in the process of photosynthesis. Plants use these organic compounds to build structural components or respire them to release energy. When plant respiration occurs below-ground in the roots, it adds to soil respiration. Over time, plant structural components are consumed by heterotrophs. 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 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 effected 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 is composed of numerous changing factors including rising atmospheric CO2, increasing temperature and shifting precipitation patterns. All of these factors can affect the rate of global soil respiration. Increased nitrogen fertilization by humans also has the potential to effect rates over the entire Earth.
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 cycles. 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 feedbacks with global climate change. Positive feedbacks are when a change in a system produces response in the same direction of the change. Therefore, soil respiration rates can be effected by climate change and then respond by enhancing climate change.
All cellular respiration releases energy, water and CO2 from organic carbon compounds. Any respiration that occurs below-ground is considered soil respiration. Respiration by plant roots, bacteria, fungi and soil animals are all sources of Template:Convert in soil.
Tricarboxylic acid (TCA) cycle
The tricarboxylic acid (TCA) cycle – or citric acid cycle – is an important step in cellular respiration. In the TCA cycle, a six carbon sugar will be oxidized.[1] 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, 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 or lactic acid.[2] Due to the lack of oxygen, this pathway is described as anaerobic respiration. This is an important source of CO2 in soil respiration in water logged ecosystems where oxygen is scarce, as in peat bogs and wetlands. However, most CO2 released from the soil occurs via respiration and one of the most important aspects of belowground 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 usually accounts for approximately half of all soil respiration. However these values can range from 10–90% depending on the dominate plant types in an ecosystem 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.[3] 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 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 acids, vitamins, long chain carbohydrates, enzymes and lysates 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.[4] 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.[5] Another important organism in the rhizosphere are root-infecting fungi or mycorrhizae. 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.[6] Fungi, along with bacteria and soil animals, also play a large role in the decomposition of litter 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 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 Template:Convert in length and will ingest soil litter. The fecal 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 are organisms from Template:Convert, such as earthworms and termites. 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.[7]
Regulation of soil respiration
Regulation of CO2 production in soil is due to various abiotic, or non-living, factors. Temperature, soil moisture and nitrogen 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 35 degrees Celsius, root respiration begins to shut down completely.[8] Microorganisms are divided into three temperature groups; cryophiles, mesophiles and thermophiles. Cryophiles function optimally at temperatures below 20 degrees Celsius, mesophiles function best at temperatures between 20 and 40 degrees Celsius and thermophiles function optimally at over 40 degrees Celsius. 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.[9] 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.[10] Many microorganisms possess strategies for growth and survival 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 with the exception of wetland plants, which have developed specific mechanisms for root aeration.[11] Soil moisture levels regulate root respiration, which is essential for nitrogen uptake.
Nitrogen
Nitrogen directly affects soil respiration in several ways. Nitrogen must be taken in by roots in order 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. 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 mycorrhizal symbionts, which fix atmospheric nitrogen, the energetic cost to the plant to acquire one molecule of NH3 from atmospheric N2 is 2.36 CO2.[12] It is essential that plants uptake nitrogen from the soil or rely on symbionts to fix it from the atmosphere in order to assure growth, reproduction and long term survival.
Another way nitrogen affects soil respiration is through litter decomposition. High nitrogen litter is considered high quality and is more readily decomposed by microorganisms than low quality litter. Degradation of cellulose, a tough plant structural compound, is also a nitrogen limited process and will increase with the addition of nitrogen to litter.[13]
Methods of measurement
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Different methods exist for the measurement of soil respiration rate and the determination of sources. The most common 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; and the use of stable isotope ratios.
Long term stand alone soil flux systems for measurement at one location over time
[[File:Automated CO2 Exchange system (ACE from ADC BioScientific Ltd) measuring soil respiration..JPG|thumb|ACE Automated soil respiration system measuring soil CO2 flux.]]
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. external link optisci
The long term instruments are designed to expose the measuring site to ambient conditions as much as is possible between measurements.external link optisci
Types of long term stand alone instruments:
Closed mode systems, by definition, are closed without a vent. When making measurements, closed systems take a CO2 measurement of ambient air just after the soil sample chamber is sealed. This becomes the CO2 reference. The system continues to take measurements at regular intervals over the next few minutes until a final CO2 concentration is recorded at a predetermined end time over time. Closed systems have the advantages that they are used by more researchers, and they provide wind proof results. Since there is no vent, wind is not an issue.external link optisci
When using a closed mode system, it is important that the experimental design include a maximum measuring time for two reasons: First; if the measuring time is too long, an unacceptable positive partial CO2 pressure can build up in the chamber limiting the amount of additional CO2 from entering the chamber, causing a measuring artifact. The second reason is that if a chamber that is closed for too long, it can modify ambient conditions and cause measuring artifacts. Finding the optimal measuring time range is important. Tests on the soil in question can confirm the best time solution, and eliminate the partial CO2 pressure build up from being a significant issue.external link optisci
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). These variables are normally recorded in the measuring file along with CO2 values.external link optisci
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.external link optisci[14]
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 , 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.[15] 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.[16]
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. ACE Open and Closed mode manual external link
These systems have vents that are designed to prevent a possible unacceptable build up of partial CO2 pressure discussed under closed mode systems.ACE Open and Closed mode manual external link
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 phenomenon.ACE Open and Closed mode manual external link
Open mode systems do have vents as mentioned above. 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.ACE Open and Closed mode manual external link
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 build up, but is designed to operate like a closed mode design system in other regards.external link optisci
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 collar 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.SRS-SD1000 and SRS-SD2000 manual external link
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.ACE Open and Closed mode manual external link
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 isotopes are acquired preferentially. Isotopes 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, 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.[17] By analyzing stable carbon isotope data it is possible to determine the source components of respired SOM that was produced by different photosynthetic pathways.
Responses to human disturbance
Throughout the past 160 years, humans have changed land use and industrial practices, which have altered the climate and global geochemical cycles. These changes have affected the rate of soil respiration around the planet.
Elevated carbon dioxide
Since the Industrial Revolution, humans have been emitting vast amounts of CO2 into the atmosphere. These emissions have increased exponentially 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.[18] Soil respiration has been found to increase up to 40.6% in a sweetgum forest in Tennessee and poplar forests in Wisconsin under elevated CO2 conditions.[19] 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, the mean average temperature of the Earth is rising. This is due to man's activities such as forest clearing, soil denuding, developments that destroy autotrophic 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, rivers into the sea. Much of the organic matter swept away in floods caused by forest clearing goes into estuaries, wetlands and eventually into the open ocean. Increased turbidity of surface waters causes biological oxygen demand and more autotrophic organisms die in the wake of man's destruction. 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. 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.[20]
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, 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.[10] 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 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. 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.[21] 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 our own well being.
Global carbon cycling
Soil respiration plays a critical role in the regulation of carbon cycling at the ecosystem and global scales. Each year approximately 120 petagrams (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.[22] Researchers have estimated that soil respiration accounts for 77 Pg of carbon released to the atmosphere each year.[23] This level of release is one order of magnitude greater than the carbon release due to anthropogenic sources (6 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.
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.[5] 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 degrees Celsius will lead to an additional release of 10 Pg carbon per year to the atmosphere from soil respiration.[24] 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 this century.[25]
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 belowground, 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 utilization 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
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