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Homeostasis (from Greek: á½…î¼î¿î¹î¿Ï‚, homoios, "similar"; and ἵστî·î¼î¹, histä“mi, "standing still"; defined by Claude Bernard and later by Walter Bradford Cannon in 1929[1] and 1932[2][3]) is the property of a system, either open or closed, that regulates its internal environment and tends to maintain a stable, constant condition. Typically used to refer to a living organism, the concept came from that of milieu interieur that was created by Claude Bernard and published in 1865. Multiple dynamic equilibrium adjustment and regulation mechanisms make homeostasis possible.


[edit] Biological

With regards to any given life system parameter, an organism may be a conformer or a regulator. On one hand, regulators try to maintain the parameter at a constant level over possibly wide ambient environmental variations. On the other hand, conformers allow the environment to determine the parameter. For instance, endothermic animals maintain a constant body temperature, while exothermic (both ectotherm and poikilotherm) animals exhibit wide body temperature variation. Examples of endothermic animals include mammals and birds, examples of exothermic animals include reptiles and some sea animals. Humans are nature's ultimate example of "regulators" because they control their parameters in a variety of climates and conditions.

Behavioral adaptations allow endothermic animals to exert some control over a given parameter. For instance, reptiles often rest on sun-heated rocks in the morning to raise their body temperature. Regulators are also responsive to external circumstances, however: if the same sun-baked boulder happens to host a ground squirrel, the animal's metabolism will adjust to the lesser need for internal heat production.

Thermal image of a cold-blooded tarantula (cold-blooded or exothermic) on a warm-blooded human hand (endothermic).

An advantage of homeostatic regulation is that it allows an organism to function effectively in a broad range of environmental conditions. For example, ectotherms tend to become sluggish at low temperatures, whereas a co-located endotherm may be fully active. That thermal stability comes at a price since an automatic regulation system requires additional energy. One reason snakes may eat only once a week is that they use much less energy to maintain homeostasis.

Most homeostatic regulation is controlled by the release of hormones into the bloodstream. However, other regulatory processes rely on simple diffusion to maintain a balance.

Homeostatic regulation extends far beyond the control of temperature. All animals also regulate their blood glucose, as well as the concentration of their blood. Mammals regulate their blood glucose with insulin and glucagon. The human body maintains glucose levels constant most of the day, even after a 24-hour fast. Even during long periods of fasting, glucose levels are reduced only very slightly.[4] Insulin, secreted by the beta cells of the pancreas, transports glucose to the body's cells and causes the liver to store some of the glucose received by it, thus lowering blood glucose levels. Insulin helps to prevent hyperglycemia. When insulin is deficient or cells become resistant to it, diabetes occurs. Glucagon, secreted by the alpha cells of the pancreas, helps the body utilise stored glycogen or convert non-carbohydrate carbon sources to glucose via gluconeogenesis, thus preventing hypoglycemia. The kidneys are used to remove excess water and ions from the blood. These are then expelled as urine. The kidneys perform a vital role in homeostatic regulation in mammals, removing excess water, salt, and urea from the blood. These are the body's main waste products.

Another homeostatic regulation occurs in the gut. Homeostasis of the gut is not fully understood but it is believed that Toll-like receptor (TLR) expression profiles contribute to it. Intestinal epithelial cells exhibit important factors that contribute to homeostasis: 1) They have different cellular distribution of TLR’s compared to the normal gut mucosa. An example of this is how TLR5 (activated by flagellin) can redistribute to the basolateral membrane, which is the perfect place where flagellin can be detected.[5] 2) The enterocytes express high levels of TLR inhibitor Toll-interacting protein (TOLLIP). TOLLIP is a human gene that is a part of the innate immune system and is highest in a healthy gut; it correlates to luminal bacterial load.[5] 3) Surface enterocytes also express high levels of Interleukin-1 receptor (IL-1R) -containing inhibitory molecule. IL-1R are also referred to as single immunoglobulin IL-1R (SIGIRR). Animals deficient in this are more susceptible to induced colitis, implying that SIGIRR might possibly play a role in tuning mucosal tolerance towards commensal flora.[5] Nucleotide-binding oligomerisation domain containing 2 (NOD2) is suggested to have an effect on suppressing inflammatory cascades based on recent evidence.[5] It is believed to modulate signals transmitted through TLRs, TLR3, 4, and 9 specifically. Mutation of it has resulted in Crohn's disease. Excessive T-helper 1 responses to resident flora in the gut are controlled by inhibiting the controlling influence of regulatory T cells and tolerance-inducing dendritic cells.

Sleep timing depends upon a balance between homeostatic sleep propensity, the need for sleep as a function of the amount of time elapsed since the last adequate sleep episode, and circadian rhythms that determine the ideal timing of a correctly structured and restorative sleep episode.[6]

[edit] Control mechanisms

All homeostatic control mechanisms have at least three interdependent components for the variable being regulated: The receptor is the sensing component that monitors and responds to changes in the environment. When the receptor senses a stimulus, it sends information to a control center, the component that sets the range at which a variable is maintained. The control center determines an appropriate response to the stimulus. In most homeostatic mechanisms the control center is the brain. The control center then sends signals to an effector, which can be muscles, organs or other structures that receive signals from the control center. After receiving the signal, a change occurs to correct the deviation by either enhancing it with positive feedback or depressing it with negative feedback [7]

[edit] Positive feedback

Positive feedback is a mechanism by which an output is enhanced, such as protein levels. However, in order to avoid any fluctuation in the protein level, the mechanism is inhibited stochastically (I), therefore when the concentration of the activated protein (A) is past the threshold ([I]), the loop mechanism is activated and the concentration of A increases exponentially if d[A]=k [A]

Positive feedback mechanisms are designed to accelerate or enhance the output created by a stimulus that has already been activated.

Unlike negative feedback mechanisms that initiate to maintain or regulate physiological functions within a set and narrow range, the positive feedback mechanisms are designed to push levels out of normal ranges. To achieve this purpose, a series of events initiates a cascading process that builds to increase the effect of the stimulus. This process can be beneficial but is rarely used by the body due to risks of the acceleration's becoming uncontrollable.

One positive feedback example event in the body is blood platelet accumulation, which, in turn, causes blood clotting in response to a break or tear in the lining of blood vessels. Another example is the release of oxytocin to intensify the contractions that take place during childbirth.[7]

[edit] Negative feedback

Negative feedback mechanisms consist of reducing the output or activity of any organ or system back to its normal range of functioning. A good example of this is regulating blood pressure. Blood vessels can sense resistance of blood flow against the walls when blood pressure increases. The blood vessels act as the receptors and they relay this message to the brain. The brain then sends a message to the heart and blood vessels, both of which are the effectors. The heart rate would decrease as the blood vessels increase in diameter (or vasodilation). This change would cause the blood pressure to fall back to its normal range. The opposite would happen when blood pressure decreases, and would cause vasoconstriction.

Another important example is seen when the body is deprived of food. The body would then reset the metabolic set point to a lower than normal value. This would allow the body to continue to function, at a slower rate, even though the body is starving. Therefore, people who deprive themselves of food while trying to lose weight would find it easy to shed weight initially and much harder to lose more after. This is due to the body readjusting itself to a lower metabolic set point to allow the body to survive with its low supply of energy. Exercise can change this effect by increasing the metabolic demand.

Another good example of negative feedback mechanism is temperature control. The hypothalamus, which monitors the body temperature, is capable of determining even the slightest of variation of normal body temperature (37 degrees Celsius). Response to such variation could be stimulation of glands that produces sweat to reduce the temperature or signaling various muscles to shiver to increase body temperature.

Both feedbacks are equally important for the healthy functioning of one's body. Complications can arise if any of the two feedbacks are affected or altered in any way.

[edit] Homeostatic imbalance

Much disease results from disturbance of homeostasis, a condition known as homeostatic imbalance. As it ages, every organism will lose efficiency in its control systems. The inefficiencies gradually result in an unstable internal environment that increases the risk for illness. In addition, homeostatic imbalance is also responsible for the physical changes associated with aging. Even more serious than illness and other characteristics of aging is death. Heart failure has been seen where nominal negative feedback mechanisms become overwhelmed, and destructive positive feedback mechanisms then take over.[7]

Diseases that result from a homeostatic imbalance include diabetes, dehydration, hypoglycemia, hyperglycemia, gout, and any disease caused by a toxin present in the bloodstream. All of these conditions result from the presence of an increased amount of a particular substance. In ideal circumstances, homeostatic control mechanisms should prevent this imbalance from occurring, but, in some people, the mechanisms do not work efficiently enough or the quantity of the substance exceeds the levels at which it can be managed. In these cases, medical intervention is necessary to restore the balance, or permanent damage to the organs may result.

[edit] Varieties

The Dynamic Energy Budget theory for metabolic organization delineates structure and (one or more) reserves in an organism. Its formulation is based on three forms of homeostasis:

  • Strong homeostasis, wherein structure and reserve do not change in composition. Because the amount of reserve and structure can vary, this allows a particular change in the composition of the whole body (as explained by the Dynamic Energy Budget theory).
  • Weak homeostasis, wherein the ratio of the amounts of reserve and structure becomes constant as long as food availability is constant, even when the organism grows. This means that the whole body composition is constant during growth in constant environments.
  • Structural homeostasis, wherein the sub-individual structures grow in harmony with the whole individual; the relative proportions of the individuals remain constant.

[edit] Ecological

Historically, ecological succession was seen as having a stable end-stage called the climax (see Frederic Clements), sometimes referred to as the 'potential biodiversity' of a site, shaped primarily by the local climate. This idea has been largely abandoned by modern ecologists in favor of nonequilibrium ideas of how ecosystems function, as most natural ecosystems experience disturbance at a rate that makes a "climax" community unattainable.

Only on small, isolated habitats known as ecological islands can the phenomenon be observed. One such case study is the island of Krakatoa after its major eruption in 1883: the established stable homeostasis of the previous forest climax ecosystem was destroyed, and all life was eliminated from the island. In the years after the eruption, Krakatoa went through a sequence of ecological changes in which successive groups of new plant or animal species followed one another, leading to increasing biodiversity and eventually culminating in a re-established climax community. This ecological succession on Krakatoa occurred in a number of stages; a sere is defined as "a stage in a sequence of events by which succession occurs". The complete chain of seres leading to a climax is called a prisere. In the case of Krakatoa, the island reached its climax community, with eight hundred different recorded species, in 1983, one hundred years after the eruption that cleared all life off the island. Evidence confirms that this number has been homeostatic for some time, with the introduction of new species rapidly leading to elimination of old ones. The evidence of Krakatoa, and other disturbed island ecosystems, has confirmed many principles of Island Biogeography, mimicking general principles of ecological succession albeit in a virtually closed system comprised almost exclusively of endemic species.

[edit] Biosphere

In the Gaia hypothesis, James Lovelock stated that the entire mass of living matter on Earth (or any planet with life) functions as a vast homeostatic superorganism that actively modifies its planetary environment to produce the environmental conditions necessary for its own survival. In this view, the entire planet maintains homeostasis. Whether this sort of system is present on Earth is still open to debate. However, some relatively simple homeostatic mechanisms are generally accepted. For example, when atmospheric carbon dioxide levels rise, certain plants are able to grow better and thus act to remove more carbon dioxide from the atmosphere. When sunlight is plentiful and atmospheric temperature climbs, the phytoplankton of the ocean surface waters thrive and produce more dimethyl sulfide, DMS. The DMS molecules act as cloud condensation nuclei, which produce more clouds, and thus increase the atmospheric albedo, and this feeds back to lower the temperature of the atmosphere. As scientists discover more about Gaia, vast numbers of positive and negative feedback loops are being discovered, that, together, maintain a metastable condition, sometimes within very broad range of environmental conditions. Environmental pressure, such as competition or change in temperature, can lead to adaptation/extinction of species over time.

[edit] Reactive

Example of use: "Reactive homeostasis is an immediate homeostasic response to a challenge such as predation."

However, any homeostasis is impossible without reaction - because homeostasis is and must be a "feedback" phenomenon.

The phrase "reactive homeostasis" is simply short for "reactive compensation reestablishing homeostasis", that is to say, "reestablishing a point of homeostasis." - it should not be confused with a separate kind of homeostasis or a distinct phenomenon from homeostasis; it is simply the compensation (or compensatory) phase of homeostasis.

[edit] Other fields

The term has come to be used in other fields, as well.

[edit] Risk

An actuary may refer to risk homeostasis, where (for example) people that have anti-lock brakes have no better safety record than those without anti-lock brakes, because the former unconsciously compensate for the safer vehicle via less-safe driving habits. Previous to the innovation of anti-lock brakes, certain maneuvers involved minor skids, evoking fear and avoidance: now the anti-lock system moves the boundary for such feedback, and behavior patterns expand into the no-longer punitive area. It has also been suggested[citation needed] that ecological crises are an instance of risk homeostasis in which a particular behavior continues until proven dangerous or dramatic consequences actually occur.

[edit] Stress

Sociologists and psychologists may refer to stress homeostasis, the tendency of a population or an individual to stay at a certain level of stress, often generating artificial stresses if the "natural" level of stress is not enough.[citation needed]

Jean-François Lyotard, a postmodern theorist, has applied this term to societal 'power centers' that he describes as being 'governed by a principle of homeostasis,' for example, the scientific hierarchy, which will sometimes ignore a radical new discovery for years because it destabilises previously-accepted norms. (See The Postmodern Condition: A Report on Knowledge by Jean-François Lyotard)

[edit] See also

[edit] References

  1. ^ Cannon WB. Organization For Physiological Homeostasis. Physiol Rev. 1929; 9: 399-431.
  2. ^ Cannon WB. The Wisdom of the Body. 1932. W.W. Norton & Company, Inc., New York.
  3. ^ Karl Ludwig von Bertalanffy: ... aber vom Menschen wissen wir nichts, (English title: Robots, Men and Minds), translated by Dr. Hans-Joachim Flechtner. page 115. Econ Verlag GmbH (1970), Düsseldorf, Wien. 1st edition.
  4. ^ Bhagavan, N. V. (2002). Medical biochemistry (4th ed.). Academic Press. pp. 499. ISBN 9780120954407. http://books.google.com/?id=vT9YttFTPi0C&pg=PA499&dq=%22in+very+prolonged+fasts+the+plasma+glucose+level+decreases+very+slightly%22#v=onepage&q=%22in%20very%20prolonged%20fasts%20the%20plasma%20glucose%20level%20decreases%20very%20slightly%22. 
  5. ^ a b c d Ann M O'Hara, Fergus Shanahan The gut flora as a forgotten organ. EMBO reports 7, 688 - 693 (01 Jul 2006)
  6. ^ Wyatt, James K.; Ritz-De Cecco, Angela; Czeisler, Charles A.; Dijk, Derk-Jan (1 October 1999). "Circadian temperature and melatonin rhythms, sleep, and neurobehavioral function in humans living on a 20-h day". Am J Physiol 277 (4): R1152–R1163. Fulltext. PMID 10516257. http://ajpregu.physiology.org/cgi/content/full/277/4/R1152. Retrieved 2007-11-25. "... significant homeostatic and circadian modulation of sleep structure, with the highest sleep efficiency occurring in sleep episodes bracketing the melatonin maximum and core body temperature minimum". 
  7. ^ a b c Marieb, Elaine N. & Hoehn, Katja (2007). Human Anatomy & Physiology (Seventh ed.). San Francisco, CA: Pearson Benjamin Cummings.

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