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Thermoregulation
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Thermoregulation in animals
Ectotherm
Endotherm
Mesotherm
Poikilotherm
Homeothermy
Heterothermy
Stenotherm
Eurytherm
Thermolabile
Thermostability
Gigantothermy
Kleptothermy
Bradymetabolism
Tachymetabolism
Thermogenesis
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t
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Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature as its own body temperature, thus avoiding the need for internal thermoregulation. The internal thermoregulation process is one aspect of homeostasis: a state of dynamic stability in an organism's internal conditions, maintained far from thermal equilibrium with its environment (the study of such processes in zoology has been called physiological ecology). If the body is unable to maintain a normal temperature and it increases significantly above normal, a condition known as hyperthermia occurs. For humans, this occurs when the body is exposed to constant temperatures of approximately 55 °C (131 °F), and with prolonged exposure (longer than a few hours) at this temperature and up to around 75 °C (167 °F) death is almost inevitable.[citation needed] Humans may also experience lethal hyperthermia when the wet bulb temperature is sustained above 35 °C (95 °F) for six hours.[1] The opposite condition, when body temperature decreases below normal levels, is known as hypothermia. It results when the homeostatic control mechanisms of heat within the body malfunction, causing the body to lose heat faster than producing it. Normal body temperature is around 37 °C (99 °F), and hypothermia sets in when the core body temperature gets lower than 35 °C (95 °F).[2] Usually caused by prolonged exposure to cold temperatures, hypothermia is usually treated by methods that attempt to raise the body temperature back to a normal range.[3]
It was not until the introduction of thermometers that any exact data on the temperature of animals could be obtained. It was then found that local differences were present, since heat production and heat loss vary considerably in different parts of the body, although the circulation of the blood tends to bring about a mean temperature of the internal parts. Hence it is important to identify the parts of the body that most closely reflect the temperature of the internal organs. Also, for such results to be comparable, the measurements must be conducted under comparable conditions. The rectum has traditionally been considered to reflect most accurately the temperature of internal parts, or in some cases of sex or species, the vagina, uterus or bladder.[4]
Occasionally the temperature of the urine as it leaves the urethra may be of use in measuring body temperature. More often the temperature is taken in the mouth, axilla, ear or groin.[5]
Some animals undergo one of various forms of dormancy where the thermoregulation process temporarily allows the body temperature to drop, thereby conserving energy. Examples include hibernating bears and torpor in bats.
Contents
[hide]
1 Classification of animals by thermal characteristics
1.1 Endothermy vs. ectothermy
1.2 Ectotherms
1.3 Ectothermic cooling
1.4 Ectothermic heating (or minimizing heat loss)
1.5 Endothermy
1.6 Homeothermy compared with poikilothermy
2 Vertebrates
2.1 Brain control
2.2 In birds and mammals
2.3 In humans
3 In plants
4 Behavioral temperature regulation
4.1 Hibernation, estivation and daily torpor
5 Variation in animals
5.1 Normal human temperature
5.2 Variations due to circadian rhythms
5.3 Variations due to women's menstrual cycles
5.4 Variations due to fever
5.5 Variations due to biofeedback
6 Low body temperature increases lifespan
7 Limits compatible with life
7.1 Arthropoda
8 See also
9 References
10 Further reading
11 External links
Classification of animals by thermal characteristics[edit]
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Endothermy vs. ectothermy[edit]
Thermoregulation in organisms runs along a spectrum from endothermy to ectothermy. Endotherms create most of their heat via metabolic processes, and are colloquially referred to as warm-blooded. When the surrounding temperatures are cold, endotherms increase metabolic heat production to keep their body temperature constant, thus making the internal body temperature of an endotherm more or less independent of the temperature of the environment.[6] One metabolic activity, in terms of generating heat, that endotherms are able to do is that they possess a larger number of mitochondria per cell than ectotherms, enabling them to generate more heat by increasing the rate at which they metabolize fats and sugars.[7] Ectotherms use external sources of temperature to regulate their body temperatures. They are colloquially referred to as cold-blooded despite the fact that body temperatures often stay within the same temperature ranges as warm-blooded animals. Ectotherms are the opposite of endotherms when it comes to regulating internal temperatures. In ectotherms, the internal physiological sources of heat are of negligible importance; the biggest factor that enables them to maintain adequate body temperatures is due to environmental influences. Living in areas that maintain a constant temperature throughout the year, like the tropics or the ocean, have enabled ectotherms to develop a wide range of behavioral mechanisms that enable them to respond to external temperatures, such as sun-bathing to increase body temperature, or seeking the cover of shade to lower body temperature.[7][6]
Ectotherms[edit]
Main article: Ectotherm
Seeking shade is one method of cooling. Here sooty tern chicks are using a black-footed albatross chick for shade.
Ectothermic cooling[edit]
Vaporization:
Evaporation of sweat and other bodily fluids.
Convection:
Increasing blood flow to body surfaces to maximize heat transfer across the advective gradient.
Conduction:
Losing heat by being in contact with a colder surface. For instance:
Lying on cool ground.
Staying wet in a river, lake or sea.
Covering in cool mud.
Radiation:
releasing heat by radiating it away from the body.
Ectothermic heating (or minimizing heat loss)[edit]
Convection:
Climbing to higher ground up trees, ridges, rocks.
Entering a warm water or air current.
Building an insulated nest or burrow.
The red line represents the air temperature.
The purple line represents the body temperature of the lizard.
The green line represents the base temperature of the burrow.
Lizards are ectotherms and use behavioral adaptations to control their temperature. They regulate their behavior based on the temperature outside, if it is warm they will go outside up to a point and return to their burrow as necessary.
Conduction:
Lying on a hot surface.
Radiation:
Lying in the sun (heating this way is affected by the body's angle in relation to the sun).
Folding skin to reduce exposure.
Concealing wing surfaces.
Exposing wing surfaces.
Insulation:
Changing shape to alter surface/volume ratio.
Inflating the body.
Thermographic image of a snake around an arm
To cope with low temperatures, some fish have developed the ability to remain functional even when the water temperature is below freezing; some use natural antifreeze or antifreeze proteins to resist ice crystal formation in their tissues. Amphibians and reptiles cope with heat loss by evaporative cooling and behavioral adaptations. An example of behavioral adaptation is that of a lizard lying in the sun on a hot rock in order to heat through conduction.
Endothermy[edit]
Main article: Endotherm
An endotherm is an animal that regulates its own body temperature, typically by keeping it at a constant level. To regulate body temperature, an organism may need to prevent heat gains in arid environments. Evaporation of water, either across respiratory surfaces or across the skin in those animals possessing sweat glands, helps in cooling body temperature to within the organism's tolerance range. Animals with a body covered by fur have limited ability to sweat, relying heavily on panting to increase evaporation of water across the moist surfaces of the lungs and the tongue and mouth. Mammals like cats, dogs and pigs, rely on panting or other means for thermal regulation and have sweat glands only in foot pads and snout. The sweat produced on pads of paws and on palms and soles mostly serves to increase friction and enhance grip. Birds also counteract overheating by gular fluttering, or rapid vibrations of the gular (throat) skin. Down feathers trap warm air acting as excellent insulators just as hair in mammals acts as a good insulator. Mammalian skin is much thicker than that of birds and often has a continuous layer of insulating fat beneath the dermis. In marine mammals, such as whales, or animals that live in very cold regions, such as the polar bears, this is called blubber. Dense coats found in desert endotherms also aid in preventing heat gain such as in the case of the camels.
A cold weather strategy is to temporarily decrease metabolic rate, decreasing the temperature difference between the animal and the air and thereby minimizing heat loss. Furthermore, having a lower metabolic rate is less energetically expensive. Many animals survive cold frosty nights through torpor, a short-term temporary drop in body temperature. Organisms when presented with the problem of regulating body temperature have not only behavioural, physiological, and structural adaptations but also a feedback system to trigger these adaptations to regulate temperature accordingly. The main features of this system are stimulus, receptor, modulator, effector and then the feedback of the newly adjusted temperature to the stimulus. This cyclical process aids in homeostasis.
Homeothermy compared with poikilothermy[edit]
Homeothermy and poikilothermy refer to how stable an organism's deep-body temperature is. Most endothermic organisms are homeothermic, like mammals. However, animals with facultative endothermy are often poikilothermic, meaning their temperature can vary considerably. Most fish are ectotherms, as most of their heat comes from the surrounding water. However, almost all fish are poikilothermic.
Vertebrates[edit]
By numerous observations upon humans and other animals, John Hunter showed that the essential difference between the so-called warm-blooded and cold-blooded animals lies in observed constancy of the temperature of the former, and the observed variability of the temperature of the latter. Almost all birds and mammals have a high temperature almost constant and independent of that of the surrounding air (homeothermy). Almost all other animals display a variation of body temperature, dependent on their surroundings (poikilothermy).[5]
Brain control[edit]
Thermoregulation in both ectotherms and endotherms is controlled mainly by the preoptic area of the anterior hypothalamus.[8] Such homeostatic control is separate from the sensation of temperature.[8]
In birds and mammals[edit]
Kangaroo licking its arms to cool down on a very hot day
In cold environments, birds and mammals employ the following adaptations and strategies to minimize heat loss:
Using small smooth muscles (arrector pili in mammals), which are attached to feather or hair shafts; this distorts the surface of the skin making feather/hair shaft stand erect (called goose bumps or pimples) which slows the movement of air across the skin and minimizes heat loss.
Increasing body size to more easily maintain core body temperature (warm-blooded animals in cold climates tend to be larger than similar species in warmer climates (see Bergmann's Rule))
Having the ability to store energy as fat for metabolism
Have shortened extremities
Have countercurrent blood flow in extremities – this is where the warm arterial blood travelling to the limb passes the cooler venous blood from the limb and heat is exchanged warming the venous blood and cooling the arterial (e.g., Arctic wolf[9] or penguins[10][11])
In warm environments, birds and mammals employ the following adaptations and strategies to maximize heat loss:
Behavioural adaptations like living in burrows during the day and being nocturnal
Evaporative cooling by perspiration and panting
Storing fat reserves in one place (e.g., camel's hump) to avoid its insulating effect
Elongated, often vascularized extremities to conduct body heat to the air
In humans[edit]
Main article: Thermoregulation in humans
Simplified control circuit of human thermoregulation.[12]
As in other mammals, thermoregulation is an important aspect of human homeostasis. Most body heat is generated in the deep organs, especially the liver, brain, and heart, and in contraction of skeletal muscles.[13] Humans have been able to adapt to a great diversity of climates, including hot humid and hot arid. High temperatures pose serious stresses for the human body, placing it in great danger of injury or even death. For example, one of the most common reactions to hot temperatures is heat exhaustion, which is an illness that could happen if one is exposed to high temperatures, resulting in some symptoms such as dizziness, fainting, or a rapid heartbeat.[14][15] For humans, adaptation to varying climatic conditions includes both physiological mechanisms resulting from evolution and behavioural mechanisms resulting from conscious cultural adaptations.[16][17] The physiological control of the body’s core temperature takes place primarily through the hypothalamus, which assumes the role as the body’s “thermostat.”[18] This organ possesses control mechanisms as well as key temperature sensors, which are connected to nerve cells called thermoreceptors.[19] Thermoreceptors come in two subcategories; ones that respond to cold temperatures and ones that respond to warm temperatures. Scattered throughout the body in both peripheral and central nervous systems, these nerve cells are sensitive to changes in temperature and are able to provide useful information to the hypothalamus through the process of negative feedback, thus maintaining a constant core temperature.[20][21]
There are three avenues of heat loss: convection, conduction, and radiation. If skin temperature is greater than that of the surroundings, the body can lose heat by radiation and conduction. But, if the temperature of the surroundings is greater than that of the skin, the body actually gains heat by radiation and conduction. In such conditions, the only means by which the body can rid itself of heat is by evaporation. So, when the surrounding temperature is higher than the skin temperature, anything that prevents adequate evaporation will cause the internal body temperature to rise.[22] During intense physical activity (e.g. sports), evaporation becomes the main avenue of heat loss.[23] Humidity affects thermoregulation by limiting sweat evaporation and thus heat loss.[24]
Play media
A dog panting after exercise
In plants[edit]
Thermogenesis occurs in the flowers of many plants in the Araceae family as well as in cycad cones.[25] In addition, the sacred lotus (Nelumbo nucifera) is able to thermoregulate itself,[26] remaining on average 20 °C (36 °F) above air temperature while flowering. Heat is produced by breaking down the starch that was stored in their roots,[27] which requires the consumption of oxygen at a rate approaching that of a flying hummingbird.[28]
One possible explanation for plant thermoregulation is to provide protection against cold temperature. For example, the skunk cabbage is not frost-resistant, yet it begins to grow and flower when there is still snow on the ground.[25] Another theory is that thermogenicity helps attract pollinators, which is borne out by observations that heat production is accompanied by the arrival of beetles or flies.[29]
Behavioral temperature regulation[edit]
Animals other than humans regulate and maintain their body temperature with physiological adjustments and behavior. Desert lizards are ectotherms and so unable to metabolically control their temperature but can do this by altering their location. They may do this, in the morning only by raising their head from its burrow and then exposing their entire body. By basking in the sun, the lizard absorbs solar heat. It may also absorb heat by conduction from heated rocks that have stored radiant solar energy. To lower their temperature, lizards exhibit varied behaviors. Sand seas, or ergs, produce up to 57.7 °C (135.9 °F), and the sand lizard will hold its feet up in the air to cool down, seek cooler objects with which to contact, find shade or return to their burrow. They also go to their burrows to avoid cooling when the sun goes down or the temperature falls. Aquatic animals can also regulate their temperature behaviorally by changing their position in the thermal gradient.[30]
During cold weather many animals increase their thermal inertia by huddling.
Animals also engage in kleptothermy in which they share or even steal each other's body warmth. In endotherms such as bats[31] and birds (such as the mousebird[32] and emperor penguin[33]) it allows the sharing of body heat (particularly amongst juveniles). This allows the individuals to increase their thermal inertia (as with gigantothermy) and so reduce heat loss.[34] Some ectotherms share burrows of ectotherms. Other animals exploit termite mounds.[35][36]
Some animals living in cold environments maintain their body temperature by preventing heat loss. Their fur grows more densely to increase the amount of insulation. Some animals are regionally heterothermic and are able to allow their less insulated extremities to cool to temperatures much lower than their core temperature—nearly to 0 °C (32 °F). This minimizes heat loss through less insulated body parts, like the legs, feet (or hooves), and nose.
An ostrich can keep its body temperature relatively constant, even though the environment can be very hot during the day and cold at night.
Hibernation, estivation and daily torpor[edit]
To cope with limited food resources and low temperatures, some mammals hibernate during cold periods. To remain in "stasis" for long periods, these animals build up brown fat reserves and slow all body functions. True hibernators (e.g., groundhogs) keep their body temperatures low throughout hibernation whereas the core temperature of false hibernators (e.g., bears) varies; occasionally the animal may emerge from its den for brief periods. Some bats are true hibernators and rely upon a rapid, non-shivering thermogenesis of their brown fat deposit to bring them out of hibernation.
Estivation is similar to hibernation, however, it usually occurs in hot periods to allow animals to avoid high temperatures and desiccation. Both terrestrial and aquatic invertebrate and vertebrates enter into estivation. Examples include lady beetles (Coccinellidae),[37] North American desert tortoises, crocodiles, salamanders, cane toads,[38] and the water-holding frog.[39]
Daily torpor occurs in small endotherms like bats and hummingbirds, which temporarily reduces their high metabolic rates to conserve energy.[40]
Variation in animals[edit]
Chart showing diurnal variation in body temperature.
Normal human temperature[edit]
Main article: Normal human body temperature
Previously, average oral temperature for healthy adults had been considered 37.0 °C (98.6 °F), while normal ranges are 36.1 °C (97.0 °F) to 37.8 °C (100.0 °F). In Poland and Russia, the temperature had been measured axillary. 36.6 °C was considered "ideal" temperature in these countries, while normal ranges are 36 °C to 36.9 °C.
Recent studies suggest that the average temperature for healthy adults is 36.8 °C (98.2 °F) (same result in three different studies). Variations (one standard deviation) from three other studies are:
36.4–37.1 °C (97.5–98.8 °F)
36.3–37.1 °C (97.3–98.8 °F) for males, 36.5–37.3 °C (97.7–99.1 °F) for females
36.6–37.3 °C (97.9–99.1 °F)[41]
Measured temperature varies according to thermometer placement, with rectal temperature being 0.3–0.6 °C (0.5–1 °F) higher than oral temperature, while axillary temperature is 0.3–0.6 °C (0.5–1 °F) lower than oral temperature.[42] The average difference between oral and axillary temperatures of Indian children aged 6–12 was found to be only 0.1 °C (standard deviation 0.2 °C),[43] and the mean difference in Maltese children aged 4–14 between oral and axillary temperature was 0.56 °C, while the mean difference between rectal and axillary temperature for children under 4 years old was 0.38 °C.[44]
Variations due to circadian rhythms[edit]
In humans, a diurnal variation has been observed dependent on the periods of rest and activity, lowest at 11 p.m. to 3 a.m. and peaking at 10 a.m. to 6 p.m. Monkeys also have a well-marked and regular diurnal variation of body temperature that follows periods of rest and activity, and is not dependent on the incidence of day and night; nocturnal monkeys reach their highest body temperature at night and lowest during the day. Sutherland Simpson and J.J. Galbraith observed that all nocturnal animals and birds – whose periods of rest and activity are naturally reversed through habit and not from outside interference – experience their highest temperature during the natural period of activity (night) and lowest during the period of rest (day).[5] Those diurnal temperatures can be reversed by reversing their daily routine.[45]
In essence, the temperature curve of diurnal birds is similar to that of man and other homoeothermal animals, except that the maximum occurs earlier in the afternoon and the minimum earlier in the morning. Also, the curves obtained from rabbits, guinea pigs, and dogs were quite similar to those from man.[5] These observations indicate that body temperature is partially regulated by circadian rhythms.
Variations due to women's menstrual cycles[edit]
During the follicular phase (which lasts from the first day of menstruation until the day of ovulation), the average basal body temperature in women ranges from 36.45 to 36.7 °C (97.6 to 98.1 °F). Within 24 hours of ovulation, women experience an elevation of 0.15–0.45 °C (0.2–0.9 °F) due to the increased metabolic rate caused by sharply elevated levels of progesterone. The basal body temperature ranges between 36.7–37.3 °C (98.1–99.2 °F) throughout the luteal phase, and drops down to pre-ovulatory levels within a few days of menstruation.[46] Women can chart this phenomenon to determine whether and when they are ovulating, so as to aid conception or contraception.
Variations due to fever[edit]
Fever is a regulated elevation of the set point of core temperature in the hypothalamus, caused by circulating pyrogens produced by the immune system. To the subject, a rise in core temperature due to fever may result in feeling cold in an environment where people without fever do not.
Variations due to biofeedback[edit]
Some monks are known to practice Tummo, biofeedback meditation techniques, that allow them to raise their body temperatures substantially.[47]
Low body temperature increases lifespan[edit]
It has been theorised that low body temperature may increase lifespan. In 2006, it was reported that transgenic mice with a body temperature 0.3–0.5 C lower than normal mice lived longer than normal mice. This mechanism is due to overexpressing the uncoupling protein 2 in hypocretin neurons (Hcrt-UCP2), which elevated hypothalamic temperature, thus forcing the hypothalamus to lower body temperature. Lifespan was increased by 12% and 20% for males and females, respectively. The mice were fed ad libitum.[48][49][50] The effects of such a genetic change in body temperature on longevity is more difficult to study in humans; in 2011, the UCP2 genetic alleles in humans were associated with obesity.[51]
Limits compatible with life[edit]
There are limits both of heat and cold that an endothermic animal can bear and other far wider limits that an ectothermic animal may endure and yet live. The effect of too extreme a cold is to decrease metabolism, and hence to lessen the production of heat. Both catabolic and anabolic pathways share in this metabolic depression, and, though less energy is used up, still less energy is generated. The effects of this diminished metabolism become telling on the central nervous system first, especially the brain and those parts concerning consciousness;[52] both heart rate and respiration rate decrease; judgment becomes impaired as drowsiness supervenes, becoming steadily deeper until the individual loses consciousness; without medical intervention, death by hypothermia quickly follows. Occasionally, however, convulsions may set in towards the end, and death is caused by asphyxia.[53][52]
In experiments on cats performed by Sutherland Simpson and Percy T. Herring, the animals were unable to survive when rectal temperature fell below 16 °C.[52] At this low temperature, respiration became increasingly feeble; heart-impulse usually continued after respiration had ceased, the beats becoming very irregular, appearing to cease, then beginning again. Death appeared to be mainly due to asphyxia, and the only certain sign that it had taken place was the loss of knee-jerks.[53]
However, too high a temperature speeds up the metabolism of different tissues to such a rate that their metabolic capital is soon exhausted. Blood that is too warm produces dyspnea by exhausting the metabolic capital of the respiratory centre;[citation needed] heart rate is increased; the beats then become arrhythmic and eventually cease. The central nervous system is also profoundly affected by hyperthermia and delirium, and convulsions may set in. Consciousness may also be lost, propelling the person into a comatose condition. These changes can sometimes also be observed in patients suffering from an acute fever.[citation needed] Mammalian muscle becomes rigid with heat rigor at about 50 °C, with the sudden rigidity of the whole body rendering life impossible.[53]
H.M. Vernon performed work on the death temperature and paralysis temperature (temperature of heat rigor) of various animals. He found that species of the same class showed very similar temperature values, those from the Amphibia examined being 38.5 °C, fish 39 °C, reptiles 45 °C, and various molluscs 46 °C.[citation needed] Also, in the case of pelagic animals, he showed a relation between death temperature and the quantity of solid constituents of the body. In higher animals, however, his experiments tend to show that there is greater variation in both the chemical and physical characteristics of the protoplasm and, hence, greater variation in the extreme temperature compatible with life.[53]
Arthropoda[edit]
The maximum temperatures tolerated by certain thermophilic arthropods exceeds the lethal temperatures for most vertebrates.[54]
The most heat-resistant insects are three genera of desert ants recorded from three different parts of the world. The ants have developed a lifestyle of scavenging for short durations during the hottest hours of the day, in excess of 50 °C (122 °F), for the carcasses of insects and other forms of life which have succumbed to heat stress.[55]
In April 2014, the South Californian mite Paratarsotomus macropalpis has been recorded as the world's fastest land animal relative to body length, at a speed of 322 body lengths per second. Besides the unusually great speed of the mites, the researchers were surprised to find the mites running at such speeds on concrete at temperatures up to 60 °C (140 °F), which is significant because this temperature is well above the lethal limit for the majority of animal species. In addition, the mites are able to stop and change direction very quickly.[54]
See also[edit]
Innate heat
Normal human body temperature
Thermal neutral zone
Insect thermoregulation
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Jump up ^ Bob Moore (29 September 2009). "Estivation: The Survial Siesta". Audubon Guides. Retrieved 24 October 2013.
Jump up ^ F.H. Pough, R.M. Andrews, J.E. Cadle, M.L. Crump, A.H. Savitzky and K.D. Wells (2001). Herpetology, second edition. Upper Saddle River, New Jersey: Prentice Hall. CS1 maint: Multiple names: authors list (link)
Jump up ^ Starr, Cecie (2005). Biology: Concepts and Applications. Thomson Brooks/Cole. p. 639. ISBN 0-534-46226-X.
Jump up ^ Wong, Lena; Forsberg, C; Wahren, LK (2005). "Temperature of a Healthy Human (Body Temperature)". Scandinavian Journal of Caring Sciences, The Physics Factbook. 16 (2): 122–128. doi:10.1046/j.1471-6712.2002.00069.x. PMID 12000664. Retrieved 24 October 2013.
Jump up ^ Rectal, ear, oral, and axillary temperature comparison. Yahoo Health.
Jump up ^ Deepti Chaturvedi; K.Y. Vilhekar; Pushpa Chaturvedi; M.S. Bharambe (17 June 2004). "Comparison of Axillary Temperature with Rectal or Oral Temperature and Determination of Optimum Placement Time in Children" (PDF). Indian Pediatrics. 41 (6): 600–603. PMID 15235167.
Jump up ^ Quintana, E.C. (June 2004). How reliable is axillary temperature measurement?. Annuals of Emergency Medicine. 43. pp. 797–798. doi:10.1016/j.annemergmed.2004.03.010.
Jump up ^ Simpson, Sutherland; Galbraith, J.J. (1905). An investigation into the diurnal variation of the body temperature of nocturnal and other birds, and a few mammals (PDF). The Journal of Physiology Online.
Jump up ^ Swedan, Nadya Gabriele (2001). Women's Sports Medicine and Rehabilitation. Lippincott Williams & Wilkins. p. 149. ISBN 0-8342-1731-7.
Jump up ^ Cromie, William J. (2002). Meditation changes temperatures: Mind controls body in extreme experiments. Harvard Gazette.
Jump up ^ Transgenic Mice with a Reduced Core Body Temperature Have an Increased Life Span, by Bruno Conti et al. Science, 3, November 2006
Jump up ^ Reduced Body Temperature Extends Lifespan, Study Finds
Jump up ^ Bee cool, live long
Jump up ^ "OMIM entry on human UnCoupling Protein 2 (UCP2)".
^ Jump up to: a b c Simpson S, Herring PT (1905-05-09). "The effect of cold narcosis on reflex action in warm-blooded animals". J. Physiol. 32 (5 Suppl 8): 305–11. PMC 1465681  . PMID 16992777.
^ Jump up to: a b c d Chisholm 1911, p. 50.
^ Jump up to: a b Federation of American Societies for Experimental Biology (FASEB) (27 April 2014). "Mite sets new record as world's fastest land animal". Featured Research. ScienceDaily. Retrieved 28 April 2014.
Jump up ^ Sherwood, Van (1 May 1996). "Chapter 21: Most heat tolerant". Book of Insect Records. University of Florida. Retrieved 30 April 2014.
Further reading[edit]
Blatteis, Clark M, ed. (2001) [First published 1998]. Physiology and Pathophysiology of Temperature Regulation. Singapore & River Edge, NJ: World Scientific Publishing Co. ISBN 981-02-3172-5. Retrieved 8 September 2010.
Charkoudian, Nisha (May 2003). Skin Blood Flow in Adult Human Thermoregulation: How It Works, When It Does Not, and Why. Mayo Clinic Proceedings. 78. pp. 603–612. doi:10.4065/78.5.603. PMID 12744548. full pdf
This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed. (1911). "Animal Heat". Encyclopædia Britannica. 2 (11th ed.). Cambridge University Press. pp. 48–50. This cites work of Simpson & Galbraith
Green, Charles Wilson (1917). Kirke's Handbook of Physiology. North American Revision. New York: William Wood & Co. Retrieved 8 September 2010. Other Internet Archive listings
Hall, John E. (2010). Guyton and Hall Textbook of Medical Physiology with Student Consult Online Access (12th ed.). Philadelphia: Elsevier Saunders. ISBN 978-1-4160-4574-8. see Table of Contents link (Previously Guyton's Textbook of Medical Physiology. Earlier editions back to at least 5th edition 1976, contain useful information on the subject of thermoregulation, the concepts of which have changed little in that time).
Hardy, James D; Gagge, A. Pharo; Stolwijk, Jan A, eds. (1970). Physiological and Behavioral Temperature Regulation. Springfield, Illinois: Charles C Thomas.
Havenith, George; Coenen, John M.L; Kistemaker, Lyda; Kenney, W. Larry (1998). Relevance of individual characteristics for human heat stress response is dependent on exercise intensity and climate type. European Journal of Applied Physiology. 77. pp. 231–241. doi:10.1007/s004210050327.
Kakuta, Naoto; Yokoyama, Shintaro; Nakamura, Mitsuyoshi; Mabuchi, Kunihiko (March 2001). Estimation of Radiative Heat Transfer Using a Geometric Human Model. IEEE Transactions on Biomedical Engineering. 48. pp. 324–331. doi:10.1109/10.914795. PMID 11327500. link to abstract
Marino, Frank E (2008). Thermoregulation and Human Performance: Physiological and Biological Aspects. Medicine and Sport Science. Vol.53. Basel, Switzerland: Karger. ISBN 978-3-8055-8648-1. Retrieved 9 September 2010.
Mitchell, John W (1 June 1976). Heat transfer from spheres and other animal forms. Biophysical Journal. 16. pp. 561–569. doi:10.1016/S0006-3495(76)85711-6. PMC 1334880  . PMID 1276385.
Milton, Anthony Stewart (1994). Temperature Regulation: Recent Physiological and Pharmacological Advances. Switzerland: Birkhäuser Verlag. ISBN 0-8176-2992-0. Retrieved 9 September 2010.
Selkirk, Glen A & McLellan, Tom M (November 2001). Influence of aerobic fitness and body fatness on tolerance to uncompensable heat stress. Journal of Applied Physiology. 91. pp. 2055–2063. PMID 11641344. Retrieved 9 September 2010.
Simpson, S. & Galbraith, J.J (1905). Observations on the normal temperatures of the monkey and its diurnal variation, and on the effects of changes in the daily routine on this variation. Transactions of the Royal Society of Edinburgh. 45. pp. 65–104. doi:10.1017/S0080456800011649.
Helmut Tributsch (Author), Miriam Varon (Translator) (May 1985). How Life Learned to Live: Adaptation in Nature. Mit Pr. ISBN 978-0262700283.
Weldon Owen Pty Ltd. (1993). Encyclopedia of animals - Mammals, Birds, Reptiles, Amphibians. Reader's Digest Association, Inc. Pages 567-568. ISBN 1-875137-49-1.
External links[edit]
Australian Government Bureau of Meteorology. Thermal Comfort Observations. Retrieved 28 January 2013.
Royal Institution Christmas Lectures 1998
Wong, Lena (1997). "Temperature of a Healthy Human (Body Temperature)". The Physics Factbook. Retrieved 24 October 2013.
Thermoregulation at the US National Library of Medicine Medical Subject Headings (MeSH)
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text:
Geography of Somalia
From Wikipedia, the free encyclopedia
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Geography of Somalia

Region
Horn of Africa
Coordinates
5°35′39″N 47°13′09″E / 5.5941182°N 47.2192383°E / 5.5941182; 47.2192383
Area
Ranked 42nd
• Total
637,657 km2 (246,201 sq mi)
Coastline
3,025 km (1,880 mi)
Borders
Total land borders:[1]
Djibouti:
60 km (37 mi)
2,340 km (1,450 mi)
Ethiopia:
1,600 km (990 mi)
Kenya:
682 km (424 mi)
Highest point
Shimbiris
2,460 m (8,070.9 ft)
Lowest point
Indian Ocean
0 m (0.0 ft)
Longest river
Shebelle River
1,130 km (702.1 mi)
Somalia map of Köppen climate classification.
Somalia is a country located in the Horn of Africa. It is bordered by Ethiopia to the west, Djibouti to the northwest, the Gulf of Aden to the north, the Indian Ocean and Guardafui Channel to the east, and Kenya to the southwest. With a land area of 637,657 square kilometers, Somalia's terrain consists mainly of plateaus, plains and highlands.[2] Its coastline is more than 3,333 kilometers in length, the longest of mainland Africa and the Middle East.[3] It has been described as being roughly shaped "like a tilted number seven".[4]
In the far north, the rugged east-west ranges of the Ogo Mountains lie at varying distances from the Gulf of Aden coast. Hot conditions prevail year-round, along with periodic monsoon winds and irregular rainfall.[5] Geology suggests the presence of valuable mineral deposits.
Contents
1 Climate
2 Terrain, vegetation and drainage
3 Data
4 Extreme points
5 See also
6 References
6.1 Notes
6.2 General references
Climate[edit]
Due to Somalia's proximity to the equator, there is not much seasonal variation in its climate. However, there are some very unpredictable rainfalls that occur sometimes. Hot conditions prevail year-round along with monsoon (periodic seasonal reversing wind accompanied by corresponding changes in precipitation) winds and irregular rainfall. Mean daily maximum temperatures range from 30 to 40 °C (86 to 104 °F), except at higher elevations and along the eastern seaboard, where the effects of a cold offshore current can be felt. In Mogadishu, for instance, average afternoon highs range from 28 °C (82 °F) to 32 °C (90 °F) in April. Some of the highest mean annual temperatures in the world have been recorded in the country; Berbera on the northwestern coast has an afternoon high that averages more than 38 °C (100 °F) from June through September. Nationally, mean daily minimums usually vary from about 15 to 30 °C (59 to 86 °F).[6] The greatest range in climate occurs in northern Somalia, where temperatures sometimes surpass 45 °C (113 °F) in July on the littoral plains and drop below the freezing point during December in the highlands.[5][6] In this region, relative humidity ranges from about 40 percent in the mid-afternoon to 85 percent at night, changing somewhat according to the season.[6]
Unlike the climates of most other countries at this latitude, conditions in Somalia range from arid in the northeastern and central regions to semiarid in the northwest and south. In the northeast, annual rainfall is less than 4 inches (100 mm); in the central plateaus, it is about 8 to 12 inches (200 to 300 mm). The northwestern and southwestern parts of the nation, however, receive considerably more rain, with an average of 20 to 24 inches (510 to 610 mm) falling per year. Although the coastal regions are hot and humid throughout the year, the hinterland is typically dry and hot.[6]
Most of the country receives less than 500 millimeters (19.7 in) of rain annually, and a large area encompassing the northeast and much of northern Somalia receives as little as 50 to 150 millimeters (1.97 to 5.91 in). Certain higher areas in the north, however, record more than 500 millimeters (19.7 in) a year, as do some coastal sites. The southwest receives 330 to 500 millimeters (13.0 to 19.7 in). Generally, rainfall takes the form of showers or localized torrential rains and is extremely variable. Mean daily maximum temperatures throughout the country range from 30 to 40 °C (86 to 104 °F), except at higher elevations and along the Indian Ocean coast. Mean daily minimum temperatures vary from 20 °C (68 °F) to more than 30 °C (86 °F). Northern Somalia experiences the greatest temperature extremes, with readings ranging from below freezing in the highlands in December to more than 45 °C (113 °F) in July in the coastal plain skirting the Gulf of Aden. The north's relative humidity ranges from about 40 percent in midafternoon to 85 percent at night, varying somewhat with the season. During the colder months, December to February, visibility at higher elevations is often restricted by fog. Temperatures in the south are less extreme, ranging from about 20 to 40 °C (68 to 104 °F). The hottest months are February through April. Coastal readings are usually five to ten degrees cooler than those inland. The coastal zone's relative humidity usually remains about 70 percent even during the dry seasons.
There are four main seasons around which pastoral and agricultural life revolve, and these are dictated by shifts in the wind patterns. From December to March is the Jilal, the harshest dry season of the year. The main rainy season, referred to as the Gu, lasts from April to June. This period is characterized by the southwest monsoons, which rejuvenate the pasture land, especially the central plateau, and briefly transform the desert into lush vegetation. From July to September is the second dry season, the Xagaa (pronounced "Hagaa"). The Dayr, which is the shortest rainy season, lasts from October to December.[6] The tangambili periods that intervene between the two monsoons (October–November and March–May) are hot and humid.[6]
hideClimate data for Somalia
Month
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Year
Average high °C (°F)
28
(82) 28
(82) 34
(93) 38
(100) 40
(104) 40
(104) 40
(104) 30
(86) 30
(86) 30
(86) 30
(86) 30
(86) 30
(86)
Average low °C (°F)
15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59) 15
(59)
Average precipitation mm (inches)
40
(1.57) 40
(1.57) 40
(1.57) 40
(1.57) 40
(1.57) 40
(1.57) 50
(1.97) 50
(1.97) 40
(1.57) 40
(1.57) 40
(1.57) 40
(1.57) 500
(19.69)
Source: Country Studies - Somalia[5]
Terrain, vegetation and drainage[edit]
Satellite image of Somalia
Topography of Somalia
Physio-graphically, Somalia is a land of limited contrast. In the north, a maritime semidesert plain parallels the Gulf of Aden coast, varying in width from roughly twelve kilometers in the west to as little as two kilometers in the east. Scrub-covered, this plain, known as the Guban (scrub land), is crossed by broad, shallow watercourses that are beds of dry sand except in the rainy seasons. When the rains arrive, the vegetation, which is a combination of low bushes and grass clumps, is quickly renewed, and for a time the guban provides some grazing for nomad livestock. This coastal strip is part of the Ethiopian xeric grasslands and shrublands ecoregion.
Inland from the gulf coast, the plain rises to the precipitous northward-facing cliffs of the dissected highlands. These form the rugged Karkaar mountain ranges that extend from the northwestern border with Ethiopia eastward to the tip of the Horn of Africa, where they end in sheer cliffs at Caseyr. The general elevation along the crest of these mountains averages about 1,800 meters above sea level south of the port town of Berbera, and eastward from that area it continues at 1,800 to 2,100 meters almost to Caseyr. The country's highest point, Shimber Berris, which rises to 2,460 meters, is located near the town of Erigavo.
Southward the mountains descend, often in scarped ledges, to an elevated plateau devoid of perennial rivers. This region of broken mountain terrain, shallow plateau valleys, and usually dry watercourses is known to the Somalis as the Ogo.
In the Ogo's especially arid eastern part, the plateau—broken by several isolated mountain ranges—gradually slopes toward the Indian Ocean and in central Somalia constitutes the Mudug Plain. A major feature of this eastern section is the long and broad Nugaal Valley, with its extensive network of intermittent seasonal watercourses. The Nugaal river enters the Indian Ocean at Eyl. The eastern area's population consists mainly of pastoral nomads eking a living in a zone of low and erratic rainfall.
The western part of the Ogo plateau region is crossed by numerous shallow valleys and dry watercourses. Annual rainfall is greater than in the east, and there are flat areas of arable land that provide a home for dryland cultivators. Most important, the western area has permanent wells to which the predominantly nomadic population returns during the dry seasons. The western plateau slopes gently southward and merges imperceptibly into an area known as the Haud, a broad, undulating terrain that constitutes some of the best grazing lands for Somali nomads, despite the lack of appreciable rainfall more than half the year. Enhancing the value of the Haud are the natural depressions that during periods of rain become temporary lakes and ponds.
The Haud zone continues for more than sixty kilometers into Ethiopia, and the vast Somali Plateau, which lies between the northern Somali mountains and the highlands of southeast Ethiopia, extends south and eastward through Ethiopia into central and southwest Somalia. The portion of the Haud lying within Ethiopia was the subject of an agreement made during the colonial era. In 1948, under pressure from their World War II allies and to the dismay of the Somalis,[7] the British "returned" the Haud (an important Somali grazing area that was presumably 'protected' by British treaties with the Somalis in 1884 and 1886) and the Ogaden to Ethiopia, based on an 1897 treaty in which the British ceded Somali territory to the Ethiopian Emperor Menelik in exchange for his help against plundering by Somali clans.[8] Britain included the proviso that the Somali nomads would retain their autonomy, but Ethiopia immediately claimed sovereignty over them.[9] This prompted an unsuccessful bid by Britain in 1956 to buy back the Somali lands it had turned over.[9] The stretch of land has since been a considerable source of regional strife.
Southwestern Somalia is dominated by the country's only two permanent rivers, the Jubba and the Shabeelle. With their sources in the Ethiopian highlands, these rivers flow in a generally southerly direction, cutting wide valleys in the Somali Plateau as it descends toward the sea; the plateau's elevation falls off rapidly in this area. The adjacent coastal zone, which includes the lower reaches of the rivers and extends from the Mudug Plain to the Kenyan border, averages 180 meters above sea level.
The Jubba River enters the Indian Ocean at Kismaayo. Although the Shabeelle River at one time apparently also reached the sea near Merca, its course is thought to have changed in prehistoric times. The Shabeelle now turns southwestward near Balcad (about thirty kilometers north of Mogadishu) and parallels the coast for more than eighty-five kilometers. The river is perennial only to a point southwest of Mogadishu; thereafter it consists of swampy areas and dry reaches and is finally lost in the sand east of Jilib, not far from the Jubba River. During the flood seasons, the Shabeelle River may fill its bed to a point near Jilib and occasionally may even break through to the Jubba River farther south. Favorable rainfall and soil conditions make the entire riverine region a fertile agricultural area and the center of the country's largest sedentary population.
In most of northern, northeastern, and north-central Somalia, where rainfall is low, the vegetation consists of scattered low trees, including various acacias, and widely scattered patches of grass. This vegetation gives way to a combination of low bushes and grass clumps in the highly arid areas of the northeast and along the Gulf of Aden.
As elevations and rainfall increase in the maritime ranges of the north, the vegetation becomes denser. Aloes are common, and on the higher plateau areas of the Ogo are woodlands. At a few places above 1,500 meters, the remnants of juniper forests (protected by the state) and areas of Euphorbia candelabrum (a chandelier-type spiny plant) occur. In the more arid highlands of the northeast, Boswellia and Commiphora trees are sources, respectively, of the frankincense and myrrh for which Somalia has been known since ancient times.
Hargeisa and much of northwestern Somalia is desert or hilly terrain. Here, the thelarchic-shaped Naasa Hablood hills are shown.
A broad plateau encompassing the northern city of Hargeysa, which receives comparatively heavy rainfall, is covered naturally by woodland (much of which has been degraded by overgrazing) and in places by extensive grasslands. Parts of this area have been under cultivation since the 1930s, producing sorghum and maize; in the 1990s it constituted the only significant region of sedentary cultivation outside southwestern Somalia.
The Haud south of Hargeysa is covered mostly by a semiarid woodland of scattered trees, mainly acacias, underlain by grasses that include species especially favored by livestock as forage. There vegetation forms spatially periodic patterns reminiscent of a tiger skin when viewed from above and therefore knowns as "tiger bush".[10] As the Haud merges into the Mudug Plain in central Somalia, the aridity increases and the vegetation takes on a subdesert character. Farther southward the terrain gradually changes to semiarid woodlands and grasslands as the annual precipitation increases.
The region encompassing the Shabeelle and Jubba rivers is relatively well watered and constitutes the country's most arable zone. The lowland between the rivers supports rich pasturage. It features arid to subarid savanna, open woodland, and thickets that include frequently abundant underlying grasses. There are areas of grassland, and in the far southwest, near the Kenyan border, some dry evergreen forests are found.
Along the Indian Ocean from Hobyo southwestward to near Mogadishu lies a stretch of dry coastal sand dunes, the Hobyo grasslands and shrublands ecoregion. This area is covered with scattered scrub and grass clumps where rainfall is sufficient although unpredictable. Much of this coast is thinly populated but overgrazing, particularly south of Mogadishu, has resulted in the destruction of the protective vegetation cover and the gradual movement of the once-stationary dunes inland. The original flora of this coast contains a number of endemic species and beginning in the early 1970s, efforts were made to stabilize these dunes by replanting. Endemic mammals are the silver dik-dik (Madoqua piacentinii), one of the world's smallest antelopes, and the Somali golden mole (Calcochloris tytonis). Other endemic species include two reptiles, a skink Haackgreerius miopus and a lizard Latastia cherchii and two birds, Ash's lark (Mirafra ashi) and the Obbia lark (Spizocorys obbiensis). The long-term political strife in Somalia has meant the coast is poorly studied and the state of the habitat unknown.[11]
Other vegetation includes plants and grasses found in the swamps into which the Shabeelle River empties most of the year and in other large swamps in the course of the lower Jubba River. Swamps of East African mangroves are found at points along the coast, particularly from Kismaayo to near the Kenyan border. Uncontrolled exploitation appears to have caused some damage to forests in that area. Other mangroves are located near Mogadishu and at a number of places along the northeastern and northern coasts.
Data[edit]
The Socotra Passage, between the tip of Somalia and Socotra island.
Location: Horn of Africa
Geographic coordinates: 10°00′N 49°00′E / 10.000°N 49.000°E / 10.000; 49.000
Map references: Africa
Area:
total: 637,657 km2 (246,600 sq mi)
land: 627,337 km2 (242,216 sq mi)
water: 10,320 km2 (3,984 sq mi)
Area - comparative: slightly smaller than Texas
Land boundaries:
total: 2,366 km (1,470 mi)
border countries: Djibouti 60 km (36 mi), Ethiopia 1,626 km (1,010 mi), and Kenya for 682 km (423 mi)
Coastline: 3,300 km (1,879 mi)
Maritime claims:
territorial sea: 200 nmi (370.4 km; 230.2 mi)
Climate: principally desert; December to February - northeast monsoon, moderate temperatures in north and very hot in south; May to October - southwest monsoon, torrid in the north and hot in the south, irregular rainfall, hot and humid periods (tangambili) between monsoons
Terrain: mostly flat to undulating plateau rising to hills in north
Elevation extremes:
lowest point: Indian Ocean 0 m (0 ft)
highest point: Shimbiris 2,460 m (8,070.9 ft)
Natural resources: uranium and largely unexploited reserves of iron ore, tin, gypsum, bauxite, copper, salt, natural gas
Land use:
arable land: 1.64%
permanent crops: 0.04%
other: 98.32% (2005)
Irrigated land: 2,000 km2 (772 sq mi) (2003)
Natural hazards: recurring droughts; frequent dust storms over eastern plains in summer; floods during rainy season
Man-made hazards: desertification; deforestation; overgrazing; soil erosion; waste dumping; overfishing
International environment agreements:
party to: Biodiversity, Desertification, Endangered Species, Law of the Sea, Ozone Layer Protection
signed, but not ratified: Marine Dumping, Nuclear Test Ban
Geography - note: Strategic location on Horn of Africa along southern approaches to Bab el Mandeb and route through Red Sea and Suez Canal. The Provisional Administrative Line forms the de facto southern border between Somalia and Ethiopia.
Extreme points[edit]
This is a list of the extreme points of Somalia, the points that are farther north, south, east or west than any other location.
Northernmost point - Alula Lagoon, Bari region
Easternmost point - Ras Hafun, Bari region*
Southernmost point - the point at which the border with Kenya enters the Indian Ocean, Jubbada Hoose region
Westernmost point - the western section of the border with Kenya**
* This is also the eastern-most point of the African continent
** Somalia does not have a western-most point, the western section of the border being formed by the 41° longitude
See also[edit]
Somalia
List of cities in Somalia
References[edit]
Notes[edit]
Jump up ^ "Geography and Map of Somalia". Retrieved 26 January 2014.
Jump up ^ "Somalia". World Factbook. Central Intelligence Agency. 2009-05-14. Retrieved 2009-05-31.
Jump up ^ "Coastline". World Factbook. Central Intelligence Agency. Retrieved 2013-08-03.
Jump up ^ Ganzglass, Martin R. "The Somali Refugees-Africa's Open Wound Refuses to Heal." Hum. Rts. 8 (1979): 28.
^ Jump up to: a b c "Somalia – Climate". countrystudies.us. 14 May 2009.
^ Jump up to: a b c d e f Hadden, Robert Lee. 2007. "The Geology of Somalia: A Selected Bibliography of Somalian Geology, Geography and Earth Science." Engineer Research and Development Laboratories, Topographic Engineering Center
Jump up ^ Federal Research Division, Somalia: A Country Study, (Kessinger Publishing, LLC: 2004), p.38
Jump up ^ David D. Laitin, Politics, Language, and Thought: The Somali Experience, (University Of Chicago Press: 1977), p.73
^ Jump up to: a b Zolberg, Aristide R., et al., Escape from Violence: Conflict and the Refugee Crisis in the Developing World, (Oxford University Press: 1992), p.106
Jump up ^ MacFadyen W.A. (1950). Vegetation patterns in the semi-desert plains of British Somaliland. Geographical Journal, 116, 199-210.
Jump up ^ "Hobyo grasslands and shrublands". Terrestrial Ecoregions. World Wildlife Fund.
General references[edit]
Hadden, Robert Lee. 2007. "The Geology of Somalia: A Selected Bibliography of Somalian Geology, Geography and Earth Science." Engineer Research and Development Laboratories, Topographic Engineering Center (now known as the Army Geospatial Center, US Army Corps of Engineers), in Alexandria, VA. Includes data and citations on the geology, geography, water and transportation in Somalia.
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Umalohokan
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This article is an orphan, as no other articles link to it. Please introduce links to this page from related articles; try the Find link tool for suggestions. (February 2009)
Umalohokan refers to the town criers of ancient barangays in the Philippines. They were responsible for going around and making people aware of new laws and policies enacted by the Datu or chieftain.[1]
Some historians, however, have a different interpretation. In cases of large scale disputes between barangays in the Visayas, the respective Datus of the barangays may elect a head Datu, called an Umalohokan, to serve as judge. The Spanish colonizers noted that elections were held in the Philippines before they arrived, specifically for this post. When the dispute was settled, the term of the Umalohokan was over.[2]
References[edit]
Jump up ^ Agoncillo, Teodoro A.; Oscar M. Alfonso; Milagros Guerrero (1970). History of the Filipino people. Malaya books. p. 47. Retrieved 2008-08-12.
Jump up ^ Laput, Ernesto J. "Buhay sa Nayon". Pinas: Munting Kasaysayan Ng Pira-pirasong Bayan. Retrieved 2007-08-06.
Scott, William Henry (1994). Barangay: Sixteenth Century Philippine Culture and Society. Quezon City: Ateneo de Manila University Press. ISBN 971-550-135-4.

This Philippines-related article is a stub. You can help Wikipedia by expanding it.
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