Homeostasis (from Greek: ὅμος, "hómos", "equal"; and ιστημι, "istēmi", "to stand" lit. "to stand equally"; coined by
Walter Bradford Cannon) is the property of either an open system or a closed system, especially a living organism, that regulates its internal environment so as to maintain a stable, constant condition. Multiple dynamic equilibriumadjustment and regulation mechanisms make homeostasis possible. The concept was created by Claude Bernard, often considered as the father of physiology, and published in 1865.
With regard to any given life system
parameter, an organism may be a "conformer" or a "regulator". 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 ectothermic animals exhibit wide body temperature variation. Examples of endothermic animals include mammalsand birds, examples of ectothermic animals include reptilesand some sea animals.
This is not to say that conformers don't have
behavioural adaptations allowing them 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. Vice versa, regulators are usually responsive to external circumstances: if the same sun-baked boulder happens to host a ground squirrel, its metabolism will adjust to the lesser need for internal heat production.
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 insulinand glucagon. These hormones are released by the pancreas. If the pancreas is for any reason unable to produce enough of these two hormones diabetesresults. The kidneysare 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 ureafrom the blood. These are the body's main waste products.
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 which determine the ideal timing of a correctly structured and restorative sleep episode. [cite journal |last=Wyatt |first=James K. |coauthors=Ritz-De Cecco, Angela; Czeisler, Charles A.; Dijk, Derk-Jan |year=1999 |month=October |title= Circadian temperature and melatonin rhythms, sleep, and neurobehavioral function in humans living on a 20-h day |journal=Am J Physiol |volume=277 |issue=4 |pages=R1152–R1163. Fulltext |id= |url= http://ajpregu.physiology.org/cgi/content/full/277/4/R1152
accessdate=2007-11-25 |quote=... 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 |pmid= 10516257 ]
All homeostatic control mechanisms have at least three interdependent components for the variable being regulated: The 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. The result of that response feeds to the effector, either enhancing it with positive feedback or depressing it with negative feedback Marieb, Elaine N. & Hoehn, Katja (2007). "Human Anatomy & Physiology" (Seventh ed.). San Francisco, CA: Pearson Benjamin Cummings.]
Positive Feedback Mechanisms
Positive feedbackmechanisms 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 becoming uncontrollable.
One positive feedback example event in the body is blood
plateletaccumulation, which, in turn, causes blood clottingin response to a break or tear in the lining of blood vessels. Another example is the release of oxytocinto intensify the contractions that take place during childbirth.
Positive feedback can also be harmful. One particular example is when a
fevercauses a positive feedback within homeostasis that pushes the temperature continually higher. Body temperature can reach extremes of 45°C (113°F), at which cellular proteins denature, causing the active site in proteins to change, thus causing metabolism to stop, resulting in death.
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.
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 imbalance, or permanent damage to the organs may result.
Varieties of homeostasis
Dynamic Energy Budgettheory for metabolic organisation 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. Since 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 Budgettheory).
*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.
Ecological homeostasis is found in a
climax communityof maximum permitted biodiversity, given the prevailing ecological conditions.
An example of a disturbed
ecosystems or sub-climax biological communities was the island of Krakatoaafter its major eruption in 1883: the established stable homeostasis of the previous forestclimax ecosystem was destroyed and all life 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 or virgin ecosystems, shows that the initial colonization by "pioneer" or "
R strategy" species occurs through positive feedback reproduction strategies, wherein species are weeds, producing huge numbers of possible offspring, but investing little in the success of any one. Rapid boom and bustplague or pest cycles are observed with such species. As an ecosystem starts to approach climax, these species get replaced by more sophisticated climax species, which, through negative feedback, adapt themselves to specific environmental conditions. These species, closely controlled by " carrying capacity", follow "K strategies", wherein species produce fewer numbers of potential offspring, but invest more heavily in securing the reproductive success of each one to the micro-environmental conditions of its specific ecological niche.
It begins with a pioneer community and ends with a climax community. This climax community occurs when the ultimate vegetation has achieved equilibrium with the local environment. Such ecosystems form nested communities or "heterarchies", in which homeostasis at one level contributes to homeostatic processes at another holonic level. For example, the loss of leaves on a mature rainforest tree creates space for new growth, and contributes to the
plant litterand soil humusbuild-up upon which such growth depends. Of equal importance, a mature rainforest tree reduces the sunlight falling on the forest floor and helps prevent invasion by other species. But trees too fall to the forest floor, and a healthy forest glade is dependent upon a constant rate of forest regrowth, produced by the fall of logs, and the recycling of forest nutrients through the respiration of termites and other insect, fungal, and bacterial decomposers. In a similar manner, such forest glades contribute ecological services, such as the regulation of microclimates or of the hydrological cyclefor an ecosystem, and a number of different ecosystems act together to maintain homeostasis, perhaps of a number of river catchmentswithin a bioregion. A diversity of bioregions, in like manner, makes up a stable homeostatic biological region or biome.
Gaia hypothesis, James Lovelockstated that the entire mass of living matter on Earth (or any planet with life) functions as a vast homeostatic superorganismthat 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 phytoplanktonof 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 albedoand 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.
Example of use: "Reactive homeostasis is an immediate response to a homeostatic 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.
The term has come to be used in other fields, as well.
actuarymay 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 that ecological crises are an instance of risk homeostasis in which behavior known to be dangerous continues until dramatic consequences actually occur.
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. Fact|date=February 2007
Jean Francois 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 destabilizes previously-accepted norms. (See "The Postmodern Condition: A Report on Knowledge" by J.F. Lyotard)
Andrew Potterhas used the term " waste homeostasis" in reference to the lack of net gain from energy-saving technologies. [Harvard reference | Surname=Potter| Given=Andrew| Authorlink=Andrew Potter | Title=Planet-friendly design? Bah, humbug. | Journal=MacLean's | Volume=120 | Issue=5 | Year=2007 | Page=14 | URL=http://www.macleans.ca]
A 2007 study purported to find (and show clinically) "conversational homeostasis" in which overly-familiar people (such as spouses) condense their speech so much that they are actually worse at communicating novel information than strangers are, while not being conscious of this problem. [Harvard reference | Surname=Keysar| Given=Boaz| Authorlink= | Title=The Effect of Information Overlap on Communication Effectiveness | Journal=Cognitive Science | Year=2007 | URL=http://www.eurekalert.org/pub_releases/2007-02/uoc-wec022007.php]
Some herbal medicines, known as
adaptogens, have been defined to function as non-toxic metabolic regulators that can enhance metabolic homeostasis during stress.Winston, David & Maimes, Steven. “ADAPTOGENS: Herbs for Strength, Stamina, and Stress Relief,” Healing Arts Press, 2007.]
Le Chatelier's principle
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