Respiratory Exchange Ratio
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The respiratory quotient (RQ or respiratory coefficient) is a
dimensionless number Dimensionless quantities, or quantities of dimension one, are quantities implicitly defined in a manner that prevents their aggregation into unit of measurement, units of measurement. ISBN 978-92-822-2272-0. Typically expressed as ratios that a ...
used in calculations of
basal metabolic rate Basal metabolic rate (BMR) is the rate of energy expenditure per unit time by endothermic animals at rest.. In other words it is the energy required by body organs to perform normal It is reported in energy units per unit time ranging from watt ( ...
(BMR) when estimated from carbon dioxide production. It is calculated from the ratio of carbon dioxide produced by the body to oxygen consumed by the body, when the body is in a steady state. Such measurements, like measurements of oxygen uptake, are forms of indirect
calorimetry In chemistry and thermodynamics, calorimetry () is the science or act of measuring changes in '' state variables'' of a body for the purpose of deriving the heat transfer associated with changes of its state due, for example, to chemical reac ...
. It is measured using a respirometer. The respiratory quotient value indicates which macronutrients are being metabolized, as different energy pathways are used for fats, carbohydrates, and proteins. If metabolism consists solely of lipids, the respiratory quotient is approximately 0.7, for proteins it is approximately 0.8, and for carbohydrates it is 1.0. Most of the time, however, energy consumption is composed of both fats and carbohydrates. The approximate respiratory quotient of a mixed diet is 0.8. Some of the other factors that may affect the respiratory quotient are energy balance, circulating insulin, and insulin sensitivity. It can be used in the
alveolar gas equation The alveolar gas equation is the method for calculating partial pressure of alveolar oxygen (). The equation is used in assessing if the lungs are properly transferring oxygen into the blood. The alveolar air equation is not widely used in clini ...
.


Respiratory exchange ratio

The respiratory exchange ratio (RER) is the
ratio In mathematics, a ratio () shows how many times one number contains another. For example, if there are eight oranges and six lemons in a bowl of fruit, then the ratio of oranges to lemons is eight to six (that is, 8:6, which is equivalent to the ...
between the
metabolic Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the ...
production of
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
(CO2) and the uptake of
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
(O2). The ratio is determined by comparing exhaled gases to room air. Measuring this ratio is equal to RQ only at rest or during mild to moderate
aerobic exercise Aerobic exercise, also known as cardio, is physical exercise of low to high intensity that depends primarily on the aerobic energy-generating process. "Aerobic" is defined as "relating to, involving, or requiring oxygen", and refers to the use of ...
without the accumulation of lactate. The loss of accuracy during more intense
anaerobic exercise Anaerobic exercise is a type of exercise that breaks down glucose in the body without using oxygen; ''anaerobic'' means "without oxygen". This type of exercise leads to a buildup of lactic acid. In practical terms, this means that anaerobic exe ...
is among others due to factors including the
bicarbonate buffer system The bicarbonate buffer system is an acid-base homeostatic mechanism involving the balance of carbonic acid (H2CO3), bicarbonate ion (HCO), and carbon dioxide (CO2) in order to maintain pH in the blood and duodenum, among other tissues, to suppo ...
. The body tries to compensate for the accumulation of lactate and minimize the acidification of the blood by expelling more CO2 through the
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...
. The RER can exceed 1.0 during intense exercise. A value above 1.0 cannot be attributed to the substrate metabolism, but rather to the aforementioned factors regarding bicarbonate buffering. Calculation of RER is commonly done in conjunction with exercise tests such as the VO2 max test. This can be used as an indicator that the participants are nearing exhaustion and the limits of their cardio-respiratory system. An RER greater than or equal to 1.0 is often used as a secondary endpoint criterion of a VO2 max test.


Calculation

The respiratory quotient (RQ) is the ratio: RQ = CO2 eliminated / O2 consumed where the term "eliminated" refers to carbon dioxide (CO2) removed from the body in a steady state. In this calculation, the CO2 and O2 must be given in the same units, and in quantities proportional to the number of molecules. Acceptable inputs would be either moles, or else volumes of gas at standard temperature and pressure. Many metabolized substances are compounds containing only the elements
carbon Carbon () is a chemical element; it has chemical symbol, symbol C and atomic number 6. It is nonmetallic and tetravalence, tetravalent—meaning that its atoms are able to form up to four covalent bonds due to its valence shell exhibiting 4 ...
,
hydrogen Hydrogen is a chemical element; it has chemical symbol, symbol H and atomic number 1. It is the lightest and abundance of the chemical elements, most abundant chemical element in the universe, constituting about 75% of all baryon, normal matter ...
, and
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
. Examples include
fatty acids In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
,
glycerol Glycerol () is a simple triol compound. It is a colorless, odorless, sweet-tasting, viscous liquid. The glycerol backbone is found in lipids known as glycerides. It is also widely used as a sweetener in the food industry and as a humectant in pha ...
,
carbohydrates A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ma ...
,
deamination Deamination is the removal of an amino group from a molecule. Enzymes that catalysis, catalyse this reaction are called deaminases. In the human body, deamination takes place primarily in the liver; however, it can also occur in the kidney. In s ...
products, and
ethanol Ethanol (also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound with the chemical formula . It is an Alcohol (chemistry), alcohol, with its formula also written as , or EtOH, where Et is the ps ...
. For complete oxidation of such compounds, the chemical equation is CxHyOz + (x + y/4 - z/2) O2 → x CO2 + (y/2) H2O and thus metabolism of this compound gives an RQ of x/(x + y/4 - z/2). For glucose, with the molecular formula, C6H12O6, the complete oxidation equation is C6H12O6 + 6 O2 → 6 CO2 + 6 H2O. Thus, the RQ= 6 CO2/ 6 O2=1. For oxidation of a fatty acid molecule, namely
palmitic acid Palmitic acid (hexadecanoic acid in IUPAC nomenclature) is a fatty acid with a 16-carbon chain. It is the most common saturated fatty acid found in animals, plants and microorganisms.Gunstone, F. D., John L. Harwood, and Albert J. Dijkstra. The ...
: : 23\ \mathrm O_2 + \mathrm C_\mathrm H_\mathrm O_2 \to 16\ \mathrm_2 + 16\ \mathrm H_2\mathrm O + 129\ \mathrm : \mathrm = \frac = \frac \approx 0.7 A RQ near 0.7 indicates that
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...
is the predominant fuel source, a value of 1.0 is indicative of
carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...
being the predominant fuel source, and a value between 0.7 and 1.0 suggests a mix of both fat and carbohydrate. In general a mixed diet corresponds with an RER of approximately 0.8. For fats, the RQ depends on the specific fatty acids present. Amongst the commonly stored fatty acids in vertebrates, RQ varies from 0.692 (
stearic acid Stearic acid ( , ) is a saturated fatty acid with an 18-carbon chain. The IUPAC name is octadecanoic acid. It is a soft waxy solid with the formula . The triglyceride derived from three molecules of stearic acid is called stearin. Stearic acid ...
) to as high as 0.759 (
docosahexaenoic acid Docosahexaenoic acid (DHA) is an omega−3 fatty acid that is an important component of the human brain, cerebral cortex, skin, and retina. It is given the fatty acid notation 22:6(''n''−3). It can be synthesized from alpha-linolenic acid or ...
). Historically, it was assumed that 'average fat' had an RQ of about 0.71, and this holds true for most mammals including humans. However, a recent survey showed that aquatic animals, especially fish, have fat that should yield higher RQs on oxidation, reaching as high as 0.73 due to high amounts of docosahexaenoic acid. The range of respiratory coefficients for organisms in metabolic balance usually ranges from 1.0 (representing the value expected for pure carbohydrate oxidation) to ~0.7 (the value expected for pure fat oxidation). In general, molecules that are more oxidized (e.g., glucose) require less oxygen to be fully metabolized and, therefore, have higher respiratory quotients. Conversely, molecules that are less oxidized (e.g., fatty acids) require more oxygen for their complete metabolism and have lower respiratory quotients. See BMR for a discussion of how these numbers are derived. A mixed diet of fat and carbohydrate results in an average value between these numbers. RQ value corresponds to a caloric value for each liter (L) of CO2 produced. If O2 consumption numbers are available, they are usually used directly, since they are more direct and reliable estimates of energy production. RQ as measured includes a contribution from the energy produced from protein. However, due to the complexity of the various ways in which different amino acids can be metabolized, no single RQ can be assigned to the oxidation of protein in the diet. Insulin, which increases lipid storage and decreases fat oxidation, is positively associated with increases in the respiratory quotient. A positive energy balance will also lead to an increased respiratory quotient.


Applications

Practical applications of the respiratory quotient can be found in severe cases of
chronic obstructive pulmonary disease Chronic obstructive pulmonary disease (COPD) is a type of progressive lung disease characterized by chronic respiratory symptoms and airflow limitation. GOLD defines COPD as a heterogeneous lung condition characterized by chronic respiratory s ...
, in which patients spend a significant amount of energy on respiratory effort. By increasing the proportion of fats in the diet, the respiratory quotient is driven down, causing a relative decrease in the amount of CO2 produced. This reduces the respiratory burden to eliminate CO2, thereby reducing the amount of energy spent on respirations. Respiratory Quotient can be used as an indicator of over or underfeeding. Underfeeding, which forces the body to utilize fat stores, will lower the respiratory quotient, while overfeeding, which causes
lipogenesis In biochemistry, lipogenesis is the conversion of fatty acids and glycerol into Adipose tissue, fats, or a metabolic process through which acetyl-CoA is converted to triglyceride for storage in adipose, fat. Lipogenesis encompasses both fatty aci ...
, will increase it. Underfeeding is marked by a respiratory quotient below 0.85, while a respiratory quotient greater than 1.0 indicates overfeeding. This is particularly important in patients with compromised respiratory systems, as an increased respiratory quotient significantly corresponds to increased respiratory rate and decreased
tidal volume Tidal is the adjectival form of tide. Tidal may also refer to: * ''Tidal'' (album), a 1996 album by Fiona Apple * Tidal (king), a king involved in the Battle of the Vale of Siddim * TidalCycles TidalCycles (also known as Tidal) is a live coding ...
, placing compromised patients at a significant risk. Because of its role in metabolism, respiratory quotient can be used in analysis of liver function and diagnosis of liver disease. In patients with
liver cirrhosis Cirrhosis, also known as liver cirrhosis or hepatic cirrhosis, chronic liver failure or chronic hepatic failure and end-stage liver disease, is a chronic condition of the liver in which the normal functioning tissue, or parenchyma, is replaced ...
, non-protein respiratory quotient (npRQ) values act as good indicators in the prediction of overall survival rate. Patients having a npRQ < 0.85 show considerably lower survival rates as compared to patients with a npRQ > 0.85. A decrease in npRQ corresponds to a decrease in glycogen storage by the liver. Similar research indicates that non-alcoholic fatty liver diseases are also accompanied by a low respiratory quotient value, and the non protein respiratory quotient value was a good indication of disease severity. Recently the respiratory quotient is also used from aquatic scientists to illuminate its environmental applications. Experimental studies with natural
bacterioplankton Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word (), meaning "wandering" or "drifting", and , a Latin term coined in the 19th century by Christian Got ...
using different single substrates suggested that RQ is linked to the elemental composition of the respired compounds. By this way, it is demonstrated that bacterioplankton RQ is not only a practical aspect of Bacterioplankton Respiration determination, but also a major ecosystem state variable that provides unique information about
aquatic ecosystem An aquatic ecosystem is an ecosystem found in and around a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms—aquatic life—that are dependent on each other and on their environ ...
functioning. Based on the stoichiometry of the different metabolized substrates, the scientists can predict that
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It can ...
(O2) and
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
(CO2) in aquatic ecosystems should covary inversely due to the processing of
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
and
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
. Using this quotient we could shed light on the metabolic behavior and the simultaneous roles of chemical and physical forcing that shape the
biogeochemistry Biogeochemistry is the Branches of science, scientific discipline that involves the study of the chemistry, chemical, physics, physical, geology, geological, and biology, biological processes and reactions that govern the composition of the natu ...
of aquatic ecosystems. Moving from a molecular and cellular level to an ecosystem level, various processes account for the exchange of O2 and CO2 between the biosphere and atmosphere. Field measurements of the concurrent consumption of oxygen (-ΔO2) and production of carbon dioxide (ΔCO2) can be used to derive an apparent respiratory quotient (ARQ). This value reflects a cumulative effect of not only the
aerobic respiration Cellular respiration is the process of oxidizing biological fuels using an inorganic electron acceptor, such as oxygen, to drive production of adenosine triphosphate (ATP), which stores chemical energy in a biologically accessible form. Cellu ...
of all organisms (microorganisms and higher consumers) in the sample, but also all the other
biogeochemical Biogeochemistry is the scientific discipline that involves the study of the chemical, physical, geological, and biological processes and reactions that govern the composition of the natural environment (including the biosphere, the cryosphere, ...
processes which consume O2 without a corresponding CO2 production and vice versa influencing the observed RQ.


Respiratory quotients of some substances


See also

* *


References


External links

{{authority control Biochemistry methods Energy conversion Metabolism Respiratory physiology Underwater diving physiology