Biological thermodynamics is the quantitative study of the energy transductions that occur in and between living organisms, structures, and cells and of the nature and function of the chemical processes underlying these transductions. Biological thermodynamics may address the question of whether the benefit associated with any particular phenotypic trait is worth the energy investment it requires.
German-British medical doctor and biochemist Hans Krebs' 1957 book Energy Transformations in Living Matter (written with Hans Kornberg) was the first major publication on the thermodynamics of biochemical reactions. In addition, the appendix contained the first-ever published thermodynamic tables, written by Kenneth Burton, to contain equilibrium constants and Gibbs free energy of formations for chemical species, able to calculate biochemical reactions that had not yet occurred.
Non-equilibrium thermodynamics has been applied for explaining how biological organisms can develop from disorder. Ilya Prigogine developed methods for the thermodynamic treatment of such systems. He called these systems dissipative systems, because they are formed and maintained by the dissipative processes that exchange energy between the system and its environment, and because they disappear if that exchange ceases. It may be said that they live in symbiosis with their environment. Energy transformations in biology are dependent primarily on photosynthesis. The total energy captured by photosynthesis in green plants from the solar radiation is about 2 x 1023 joules of energy per year. Annual energy captured by photosynthesis in green plants is about 4% of the total sunlight energy that reaches Earth. The energy transformations in biological communities surrounding hydrothermal vents are exceptions; they oxidize sulfur, obtaining their energy via chemosynthesis rather than photosynthesis.
The focus of thermodynamics in biology
The field of biological thermodynamics is focused on principles of chemical thermodynamics in biology and biochemistry. Principles covered include the first law of thermodynamics, the second law of thermodynamics, Gibbs free energy, statistical thermodynamics, reaction kinetics, and on hypotheses of the origin of life. Presently, biological thermodynamics concerns itself with the study of internal biochemical dynamics as: ATP hydrolysis, protein stability, DNA binding, membrane diffusion, enzyme kinetics, and other such essential energy controlled pathways. In terms of thermodynamics, the amount of energy capable of doing work during a chemical reaction is measured quantitatively by the change in the Gibbs free energy. The physical biologist Alfred Lotka attempted to unify the change in the Gibbs free energy with evolutionary theory.
Energy Transformation in Biological Systems
The sun is the primary source of energy for living organisms.Some living organisms like plants need sunlight directly while other organisms like humans can acquire energy from the sun indirectly. There is however evidence that some bacteria can thrive in harsh environments like Antarctica as evidence by the blue-green algae beneath thick layers of ice in the lakes. No matter what the type of living species, all living organisms must capture, transduce, store, and use energy to live.
The energy can be represented by the following mathematical model:
Where h is Planck's constant (6.63x10−34Js) and c is the speed of light (2.998x108 m/s). Plants trap this energy from the sunlight and undergo photosynthesis, effectively converting solar energy into chemical energy. To transfer the energy once again, animals will feed on plants and use the energy of digested plant materials to create biological macromolecules.
Energy and Dieting
Most diets are based on calorie counting. This is the conservation of energy principle in that a person who consumes more calories than his or her body burns will gain weight whereas a person who consumes less calories than his or her body burns will lose weight. However, there is more to dieting than just keeping track of calories. There are several theories on dieting. One theory suggests that some people have food efficient bodies, meaning these people need fewer calories than other people do for the same activity. Sometimes, the body interprets dieting as starvation and starts using the energy reserves of the body more cautiously. This is why switching to half the number of calories needed can lower the basal metabolic rate by 10 and 20 percent. Extended periods of low-calorie dieting without adequate exercise may result in the loss of considerable muscle tissue together with fat. With less muscle to burn the calories, the metabolic rate declines even after the diet has been stopped. This will most likely result in the person regaining the weight. The basal metabolic rate remains about the same in people who exercise while dieting. Another theory suggests that people with too many fat cells developed during childhood are more likely to gain weight. Research shows that the fat content of the bodies is controlled by the setting of a fat control mechanism, similar to the temperature of a house is controlled by the thermostat setting. Another study suggests that weight gain is due to genes. Considering that 80 percent of the children of overweight parents are also overweight, heredity may play an important role in the way a body stores fat. According to research from the University of Washington and the Rockefeller University, the body tries to keep the body fat at a particular level called the set point, which differs from person to person. This is done by speeding up the metabolism and burning extra calories much faster when a person tends to gain weight and by slowing down the metabolism and burning calories at a slower rate when a person tend to lose weight. In those with high metabolic rates, the body dissipates the extra calories as body heat instead of storing them as fat, and there is no violation of the conservation of energy principle. The scientific criterion used to determine physical fitness is somewhat useful when trying to gauge if the proper weight is in the proper range. The range of healthy weight for adults is usually expressed in terms of the body mass index (BMI) as the following:
BMI= W(kg)/H2(m2) BMI< 19 underweight 19 ≤ BMI ≤ 25 healthy weight BMI > 25 overweight
Where W is the weight (which in scientific terms is actually mass) of the person in kg and H is the height in m.
First Law of Thermodynamics
The First Law of Thermodynamics is a statement of the conservation of energy; though it can be changed from one form to another, energy can be neither created nor destroyed. From the first law, a principle called Hess's Law arises. Hess’s Law states that the heat absorbed or evolved in a given reaction must always be constant and independent of the manner in which the reaction takes place. Although some intermediate reactions may be endothermic and others may be exothermic, the total heat exchange is equal to the heat exchange had the process occurred directly. This principle is the basis for the calorimeter, a device used to determine the amount of heat in a chemical reaction. Since all incoming energy enters the body as food and is ultimately oxidized, the total heat production may be estimated by measuring the heat produced by the oxidation of food in a calorimeter. This heat is expressed in kilocalories, which are the common unit of food energy found on nutrition labels.
Second Law of Thermodynamics
The Second Law of Thermodynamics is concerned primarily with whether or not a given process is possible. The Second Law states that no natural process can occur unless it is accompanied by an increase in the entropy of the universe. Stated differently, an isolated system will always tend to disorder. Living organisms are often mistakenly believed to defy the Second Law because they are able to increase their level of organization. To correct this misinterpretation, one must refer simply to the definition of systems and boundaries. A living organism is an open system, able to exchange both matter and energy with its environment. Take, for example, the assembly of a virus molecule from its subunits, which involves an increase of order. If the virus is considered an isolated system, this process would be in defiance of the Second Law. However, a virus molecule interacts directly with its environment. The assembly of a virus molecule results in an increase of entropy in the system as a whole due to the liberation of water of solvation from the components and the resulting increase in rotational and translational entropy of the solvent.
Gibbs Free Energy
In biological systems, in general energy and entropy change together. Therefore, it is necessary to be able to define a state function that accounts for these changes simultaneously. This state function is the Gibbs Free Energy, G.
- G = H − TS
- H is the enthalpy (SI unit: joule)
- T is the temperature (SI unit: kelvin)
- S is the entropy (SI unit: joule per kelvin)
The change in Gibbs Free Energy can be used to determine whether a given chemical reaction can occur spontaneously. If ∆G is negative, the reaction can occur spontaneously. Likewise, if ∆G is positive, the reaction is nonspontaneous. Chemical reactions can be “coupled” together if they share intermediates. In this case, the overall Gibbs Free Energy change is simply the sum of the ∆G values for each reaction. Therefore, an unfavorable reaction (positive ∆G1) can be driven by a second, highly favorable reaction (negative ∆G2 where the magnitude of ∆G2 > magnitude of ∆G1). For example, the reaction of glucose with fructose to form sucrose has a ∆G value of +5.5 kcal/mole. Therefore, this reaction will not occur spontaneously. The breakdown of ATP to form ADP and inorganic phosphate has a ∆G value of -7.3 kcal/mole. These two reactions can be coupled together, so that glucose binds with ATP to form glucose-1-phosphate and ADP. The glucose-1-phosphate is then able to bond with fructose yielding sucrose and inorganic phosphate. The ∆G value of the coupled reaction is -1.8 kcal/mole, indicating that the reaction will occur spontaneously. This principle of coupling reactions to alter the change in Gibbs Free Energy is the basic principle behind all enzymatic action in biological organisms.
- Alberty R (2004). "A short history of the thermodynamics of enzyme-catalyzed reactions". J Biol Chem 279 (27): 27831–6. doi:10.1074/jbc.X400003200. PMID 15073189.
- Akihiko Ito and Takehisa Oikawa. "Global Mapping of Terrestrial Primary Productivity and Light-Use Efficiency with a Process-Based Model". In M. Shiyomi et al. Global Environmental Change in the Ocean and on Land. pp. 343–358.
- M.J. Farabee. "Reactions and Enzymes". On-Line Biology Book. Estrella Mountain Community College.
- Haynie, Donald T. (2001). Biological Thermodynamics. Cambridge University Press. pp. 1–16.
- Cengel, Yunus A. (2011). Thermodynamics An Engineering Approach. New York: McGraw Hill Companies, Inc. pp. 191–194. ISBN 978-0-07-107254-0.
- Haynie, Donald T. (2001). Biological Thermodynamics. Cambridge UP. ISBN 9780521795494.
- Stacy, Ralph W., David T. Williams, Ralph E. Worden, and Rex O. McMorris. Essentials of Biological and Medical Physics. New York: McGraw-Hill Book, 1955. Print.
- Haynie, Donald T. Biological Thermodynamics. Cambridge: Cambridge UP, 2001. Print.
- Edsall, John T., and H. Gutfreund. Biothermodynamics: The Study of Biochemical Processes at Equilibrium. Chichester, West Sussex: Wiley, 1983. Print.
- Bergethon, P. R. The Physical Basis of Biochemistry: The Foundations of Molecular Biophysics. New York: Springer, 1998. Print.
- Alberts, Bruce. Essential Cell Biology. New York: Garland Science, 2009. Print.
- Haynie, D. (2001). Biological Thermodynamics (textbook). Cambridge: Cambridge University Press.
- Lehninger, A., Nelson, D., & Cox, M. (1993). Principles of Biochemistry, 2nd Ed (textbook). New York: Worth Publishers.
- Alberty, Robert, A. (2006). Biochemical Thermodynamics: Applications of Mathematica (Methods of Biochemical Analysis), Wiley-Interscience.