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Saturday, February 7, 2015

Tornado


From Wikipedia, the free encyclopedia


A tornado near Anadarko, Oklahoma. The funnel is the thin tube reaching from the cloud to the ground. The lower part of this tornado is surrounded by a translucent dust cloud, kicked up by the tornado's strong winds at the surface. The wind of the tornado has a much wider radius than the funnel itself.

All tornadoes in the US, 1950-2013, plotted by midpoint, highest F-scale on top, Alaska and Hawaii negligible, source NOAA Storm Prediction Center.

A tornado is a violently rotating column of air that is in contact with both the surface of the earth and a cumulonimbus cloud or, in rare cases, the base of a cumulus cloud. They are often referred to as twisters or cyclones,[1] although the word cyclone is used in meteorology, in a wider sense, to name any closed low pressure circulation. Tornadoes come in many shapes and sizes, but they are typically in the form of a visible condensation funnel, whose narrow end touches the earth and is often encircled by a cloud of debris and dust. Most tornadoes have wind speeds less than 110 miles per hour (177 km/h), are about 250 feet (76 m) across, and travel a few miles (several kilometers) before dissipating. The most extreme tornadoes can attain wind speeds of more than 300 miles per hour (483 km/h), stretch more than two miles (3.2 km) across, and stay on the ground for dozens of miles (more than 100 km).[2][3][4]

Various types of tornadoes include the landspout, multiple vortex tornado, and waterspout. Waterspouts are characterized by a spiraling funnel-shaped wind current, connecting to a large cumulus or cumulonimbus cloud. They are generally classified as non-supercellular tornadoes that develop over bodies of water, but there is disagreement over whether to classify them as true tornadoes. These spiraling columns of air frequently develop in tropical areas close to the equator, and are less common at high latitudes.[5] Other tornado-like phenomena that exist in nature include the gustnado, dust devil, fire whirls, and steam devil; downbursts are frequently confused with tornadoes, though their action is dissimilar.

Tornadoes have been observed on every continent except Antarctica. However, the vast majority of tornadoes occur in the Tornado Alley region of the United States, although they can occur nearly anywhere in North America.[6] They also occasionally occur in south-central and eastern Asia, northern and east-central South America, Southern Africa, northwestern and southeast Europe, western and southeastern Australia, and New Zealand.[7] Tornadoes can be detected before or as they occur through the use of Pulse-Doppler radar by recognizing patterns in velocity and reflectivity data, such as hook echoes or debris balls, as well as through the efforts of storm spotters.

There are several scales for rating the strength of tornadoes. The Fujita scale rates tornadoes by damage caused and has been replaced in some countries by the updated Enhanced Fujita Scale. An F0 or EF0 tornado, the weakest category, damages trees, but not substantial structures. An F5 or EF5 tornado, the strongest category, rips buildings off their foundations and can deform large skyscrapers. The similar TORRO scale ranges from a T0 for extremely weak tornadoes to T11 for the most powerful known tornadoes.[8] Doppler radar data, photogrammetry, and ground swirl patterns (cycloidal marks) may also be analyzed to determine intensity and assign a rating.[9][10]

Etymology

The word tornado is an altered form of the Spanish word tronada, which means "thunderstorm". This in turn was taken from the Latin tonare, meaning "to thunder". It most likely reached its present form through a combination of the Spanish tronada and tornar ("to turn"); however, this may be a folk etymology.[11][12] A tornado is also commonly referred to as a "twister", and is also sometimes referred to by the old-fashioned colloquial term cyclone.[13][14] The term "cyclone" is used as a synonym for "tornado" in the often-aired 1939 film The Wizard of Oz. The term "twister" is also used in that film, along with being the title of the 1996 tornado-related film Twister.

Definitions


A tornado near Seymour, Texas 1987

A tornado is "a violently rotating column of air, in contact with the ground, either pendant from a cumuliform cloud or underneath a cumuliform cloud, and often (but not always) visible as a funnel cloud".[15] For a vortex to be classified as a tornado, it must be in contact with both the ground and the cloud base. Scientists have not yet created a complete definition of the word; for example, there is disagreement as to whether separate touchdowns of the same funnel constitute separate tornadoes.[4] Tornado refers to the vortex of wind, not the condensation cloud.[16][17]

Funnel cloud


This tornado has no funnel cloud; however, the rotating dust cloud indicates that strong winds are occurring at the surface, and thus it is a true tornado.

A tornado is not necessarily visible; however, the intense low pressure caused by the high wind speeds (as described by Bernoulli's principle) and rapid rotation (due to cyclostrophic balance) usually causes water vapor in the air to condense into cloud droplets due to adiabatic cooling. This results in the formation of a visible funnel cloud or condensation funnel.[18]

There is some disagreement over the definition of funnel cloud and condensation funnel. According to the Glossary of Meteorology, a funnel cloud is any rotating cloud pendant from a cumulus or cumulonimbus, and thus most tornadoes are included under this definition.[19] Among many meteorologists, the funnel cloud term is strictly defined as a rotating cloud which is not associated with strong winds at the surface, and condensation funnel is a broad term for any rotating cloud below a cumuliform cloud.[4]

Tornadoes often begin as funnel clouds with no associated strong winds at the surface, and not all funnel clouds evolve into tornadoes. Most tornadoes produce strong winds at the surface while the visible funnel is still above the ground, so it is difficult to discern the difference between a funnel cloud and a tornado from a distance.[4]

Outbreaks and families

Occasionally, a single storm will produce more than one tornado, either simultaneously or in succession. Multiple tornadoes produced by the same storm cell are referred to as a "tornado family".[20] Several tornadoes are sometimes spawned from the same large-scale storm system. If there is no break in activity, this is considered a tornado outbreak (although the term "tornado outbreak" has various definitions). A period of several successive days with tornado outbreaks in the same general area (spawned by multiple weather systems) is a tornado outbreak sequence, occasionally called an extended tornado outbreak.[15][21][22]

Characteristics

Size and shape


A wedge tornado, nearly a mile wide, which hit Binger, Oklahoma in 1981

Most tornadoes take on the appearance of a narrow funnel, a few hundred yards (meters) across, with a small cloud of debris near the ground. Tornadoes may be obscured completely by rain or dust. These tornadoes are especially dangerous, as even experienced meteorologists might not see them.[23] Tornadoes can appear in many shapes and sizes.

Small, relatively weak landspouts may be visible only as a small swirl of dust on the ground. Although the condensation funnel may not extend all the way to the ground, if associated surface winds are greater than 40 mph (64 km/h), the circulation is considered a tornado.[16] A tornado with a nearly cylindrical profile and relative low height is sometimes referred to as a "stovepipe" tornado. Large single-vortex tornadoes can look like large wedges stuck into the ground, and so are known as "wedge tornadoes" or "wedges". The "stovepipe" classification is also used for this type of tornado, if it otherwise fits that profile. A wedge can be so wide that it appears to be a block of dark clouds, wider than the distance from the cloud base to the ground. Even experienced storm observers may not be able to tell the difference between a low-hanging cloud and a wedge tornado from a distance. Many, but not all major tornadoes are wedges.[24]

A rope tornado in its dissipating stage, Tecumseh, Oklahoma.

Tornadoes in the dissipating stage can resemble narrow tubes or ropes, and often curl or twist into complex shapes. These tornadoes are said to be "roping out", or becoming a "rope tornado". When they rope out, the length of their funnel increases, which forces the winds within the funnel to weaken due to conservation of angular momentum.[25] Multiple-vortex tornadoes can appear as a family of swirls circling a common center, or they may be completely obscured by condensation, dust, and debris, appearing to be a single funnel.[26]

In the United States, tornadoes are around 500 feet (150 m) across on average and travel on the ground for 5 miles (8.0 km).[23] However, there is a wide range of tornado sizes. Weak tornadoes, or strong yet dissipating tornadoes, can be exceedingly narrow, sometimes only a few feet or couple meters across. One tornado was reported to have a damage path only 7 feet (2 m) long.[23] On the other end of the spectrum, wedge tornadoes can have a damage path a mile (1.6 km) wide or more. A tornado that affected Hallam, Nebraska on May 22, 2004, was up to 2.5 miles (4.0 km) wide at the ground, and a tornado in El Reno, Oklahoma on May 31, 2013 was approximately 2.6 miles (4.2 km) wide, the widest on record.[3][27]

In terms of path length, the Tri-State Tornado, which affected parts of Missouri, Illinois, and Indiana on March 18, 1925, was on the ground continuously for 219 miles (352 km). Many tornadoes which appear to have path lengths of 100 miles (160 km) or longer are composed of a family of tornadoes which have formed in quick succession; however, there is no substantial evidence that this occurred in the case of the Tri-State Tornado.[21] In fact, modern reanalysis of the path suggests that the tornado may have begun 15 miles (24 km) further west than previously thought.[28]

Appearance

Tornadoes can have a wide range of colors, depending on the environment in which they form. Those that form in dry environments can be nearly invisible, marked only by swirling debris at the base of the funnel. Condensation funnels that pick up little or no debris can be gray to white. While traveling over a body of water (as a waterspout), tornadoes can turn very white or even blue. Slow-moving funnels, which ingest a considerable amount of debris and dirt, are usually darker, taking on the color of debris. Tornadoes in the Great Plains can turn red because of the reddish tint of the soil, and tornadoes in mountainous areas can travel over snow-covered ground, turning white.[23]

Photographs of the Waurika, Oklahoma tornado of May 30, 1976, taken at nearly the same time by two photographers. In the top picture, the tornado is lit with the sunlight focused from behind the camera, thus the funnel appears bluish. In the lower image, where the camera is facing the opposite direction, the sun is behind the tornado, giving it a dark appearance.[29]

Lighting conditions are a major factor in the appearance of a tornado. A tornado which is "back-lit" (viewed with the sun behind it) appears very dark. The same tornado, viewed with the sun at the observer's back, may appear gray or brilliant white. Tornadoes which occur near the time of sunset can be many different colors, appearing in hues of yellow, orange, and pink.[13][30]

Dust kicked up by the winds of the parent thunderstorm, heavy rain and hail, and the darkness of night are all factors which can reduce the visibility of tornadoes. Tornadoes occurring in these conditions are especially dangerous, since only weather radar observations, or possibly the sound of an approaching tornado, serve as any warning to those in the storm's path. Most significant tornadoes form under the storm's updraft base, which is rain-free,[31] making them visible.[32] Also, most tornadoes occur in the late afternoon, when the bright sun can penetrate even the thickest clouds.[21] Night-time tornadoes are often illuminated by frequent lightning.

There is mounting evidence, including Doppler On Wheels mobile radar images and eyewitness accounts, that most tornadoes have a clear, calm center with extremely low pressure, akin to the eye of tropical cyclones. Lightning is said to be the source of illumination for those who claim to have seen the interior of a tornado.[33][34][35]

Rotation

Tornadoes normally rotate cyclonically (when viewed from above, this is counterclockwise in the northern hemisphere and clockwise in the southern). While large-scale storms always rotate cyclonically due to the Coriolis effect, thunderstorms and tornadoes are so small that the direct influence of the Coriolis effect is unimportant, as indicated by their large Rossby numbers. Supercells and tornadoes rotate cyclonically in numerical simulations even when the Coriolis effect is neglected.[36][37] Low-level mesocyclones and tornadoes owe their rotation to complex processes within the supercell and ambient environment.[38]

Approximately 1 percent of tornadoes rotate in an anticyclonic direction in the northern hemisphere. Typically, systems as weak as landspouts and gustnadoes can rotate anticyclonically, and usually only those which form on the anticyclonic shear side of the descending rear flank downdraft (RFD) in a cyclonic supercell.[39] On rare occasions, anticyclonic tornadoes form in association with the mesoanticyclone of an anticyclonic supercell, in the same manner as the typical cyclonic tornado, or as a companion tornado either as a satellite tornado or associated with anticyclonic eddies within a supercell.[40]

Sound and seismology

Tornadoes emit widely on the acoustics spectrum and the sounds are caused by multiple mechanisms. Various sounds of tornadoes have been reported, mostly related to familiar sounds for the witness and generally some variation of a whooshing roar. Popularly reported sounds include a freight train, rushing rapids or waterfall, a nearby jet engine, or combinations of these. Many tornadoes are not audible from much distance; the nature and propagation distance of the audible sound depends on atmospheric conditions and topography.

The winds of the tornado vortex and of constituent turbulent eddies, as well as airflow interaction with the surface and debris, contribute to the sounds. Funnel clouds also produce sounds. Funnel clouds and small tornadoes are reported as whistling, whining, humming, or the buzzing of innumerable bees or electricity, or more or less harmonic, whereas many tornadoes are reported as a continuous, deep rumbling, or an irregular sound of "noise".[41]

Since many tornadoes are audible only when very near, sound is not reliable warning of a tornado. Tornadoes are also not the only source of such sounds in severe thunderstorms; any strong, damaging wind, a severe hail volley, or continuous thunder in a thunderstorm may produce a roaring sound.[42]

An illustration of generation of infrasound in tornadoes by the Earth System Research Laboratory's Infrasound Program

Tornadoes also produce identifiable inaudible infrasonic signatures.[43]

Unlike audible signatures, tornadic signatures have been isolated; due to the long distance propagation of low-frequency sound, efforts are ongoing to develop tornado prediction and detection devices with additional value in understanding tornado morphology, dynamics, and creation.[44] Tornadoes also produce a detectable seismic signature, and research continues on isolating it and understanding the process.[45]

Electromagnetic, lightning, and other effects

Tornadoes emit on the electromagnetic spectrum, with sferics and E-field effects detected.[44][46][47] There are observed correlations between tornadoes and patterns of lightning. Tornadic storms do not contain more lightning than other storms and some tornadic cells never produce lightning. More often than not, overall cloud-to-ground (CG) lightning activity decreases as a tornado reaches the surface and returns to the baseline level when the tornado lifts. In many cases, intense tornadoes and thunderstorms exhibit an increased and anomalous dominance of positive polarity CG discharges.[48] Electromagnetics and lightning have little or nothing to do directly with what drives tornadoes (tornadoes are basically a thermodynamic phenomenon), although there are likely connections with the storm and environment affecting both phenomena.

Luminosity has been reported in the past and is probably due to misidentification of external light sources such as lightning, city lights, and power flashes from broken lines, as internal sources are now uncommonly reported and are not known to ever have been recorded. In addition to winds, tornadoes also exhibit changes in atmospheric variables such as temperature, moisture, and pressure. For example, on June 24, 2003 near Manchester, South Dakota, a probe measured a 100 mbar (hPa) (2.95 inHg) pressure decrease. The pressure dropped gradually as the vortex approached then dropped extremely rapidly to 850 mbar (hPa) (25.10 inHg) in the core of the violent tornado before rising rapidly as the vortex moved away, resulting in a V-shape pressure trace. Temperature tends to decrease and moisture content to increase in the immediate vicinity of a tornado.[49]

Life cycle


A sequence of images showing the birth of a tornado. First, the rotating cloud base lowers. This lowering becomes a funnel, which continues descending while winds build near the surface, kicking up dust and other debris. Finally, the visible funnel extends to the ground, and the tornado begins causing major damage. This tornado, near Dimmitt, Texas, was one of the best-observed violent tornadoes in history.

Supercell relationship

Tornadoes often develop from a class of thunderstorms known as supercells. Supercells contain mesocyclones, an area of organized rotation a few miles up in the atmosphere, usually 1–6 miles (2–10 km) across. Most intense tornadoes (EF3 to EF5 on the Enhanced Fujita Scale) develop from supercells. In addition to tornadoes, very heavy rain, frequent lightning, strong wind gusts, and hail are common in such storms.

Most tornadoes from supercells follow a recognizable life cycle. That begins when increasing rainfall drags with it an area of quickly descending air known as the rear flank downdraft (RFD). This downdraft accelerates as it approaches the ground, and drags the supercell's rotating mesocyclone towards the ground with it.[16]

Formation

As the mesocyclone lowers below the cloud base, it begins to take in cool, moist air from the downdraft region of the storm. This convergence of warm air in the updraft, and this cool air, causes a rotating wall cloud to form. The RFD also focuses the mesocyclone's base, causing it to siphon air from a smaller and smaller area on the ground. As the updraft intensifies, it creates an area of low pressure at the surface. This pulls the focused mesocyclone down, in the form of a visible condensation funnel. As the funnel descends, the RFD also reaches the ground, creating a gust front that can cause severe damage a good distance from the tornado. Usually, the funnel cloud begins causing damage on the ground (becoming a tornado) within a few minutes of the RFD reaching the ground.[16]

Maturity

Initially, the tornado has a good source of warm, moist inflow to power it, so it grows until it reaches the "mature stage". This can last anywhere from a few minutes to more than an hour, and during that time a tornado often causes the most damage, and in rare cases can be more than one mile (1.6 km) across. Meanwhile, the RFD, now an area of cool surface winds, begins to wrap around the tornado, cutting off the inflow of warm air which feeds the tornado.[16]

Dissipation

As the RFD completely wraps around and chokes off the tornado's air supply, the vortex begins to weaken, and become thin and rope-like. This is the "dissipating stage", often lasting no more than a few minutes, after which the tornado fizzles. During this stage the shape of the tornado becomes highly influenced by the winds of the parent storm, and can be blown into fantastic patterns.[21][29][30] Even though the tornado is dissipating, it is still capable of causing damage. The storm is contracting into a rope-like tube and, like the ice skater who pulls her arms in to spin faster, winds can increase at this point.[25]

As the tornado enters the dissipating stage, its associated mesocyclone often weakens as well, as the rear flank downdraft cuts off the inflow powering it. Sometimes, in intense supercells, tornadoes can develop cyclically. As the first mesocyclone and associated tornado dissipate, the storm's inflow may be concentrated into a new area closer to the center of the storm. If a new mesocyclone develops, the cycle may start again, producing one or more new tornadoes. Occasionally, the old (occluded) mesocyclone and the new mesocyclone produce a tornado at the same time.

Although this is a widely accepted theory for how most tornadoes form, live, and die, it does not explain the formation of smaller tornadoes, such as landspouts, long-lived tornadoes, or tornadoes with multiple vortices. These each have different mechanisms which influence their development—however, most tornadoes follow a pattern similar to this one.[50]

Types

Multiple vortex


A multiple-vortex tornado outside Dallas, Texas on April 2, 1957.

A multiple-vortex tornado is a type of tornado in which two or more columns of spinning air rotate around a common center. A multi-vortex structure can occur in almost any circulation, but is very often observed in intense tornadoes. These vortices often create small areas of heavier damage along the main tornado path.[4][16] This is a distinct phenomenon from a satellite tornado, which is a smaller tornado which forms very near a large, strong tornado contained within the same mesocyclone. The satellite tornado may appear to "orbit" the larger tornado (hence the name), giving the appearance of one, large multi-vortex tornado. However, a satellite tornado is a distinct circulation, and is much smaller than the main funnel.[4]

Waterspout


A waterspout near the Florida Keys in 1969.

A waterspout is defined by the National Weather Service as a tornado over water. However, researchers typically distinguish "fair weather" waterspouts from tornadic waterspouts. Fair weather waterspouts are less severe but far more common, and are similar to dust devils and landspouts. They form at the bases of cumulus congestus clouds over tropical and subtropical waters. They have relatively weak winds, smooth laminar walls, and typically travel very slowly. They occur most commonly in the Florida Keys and in the northern Adriatic Sea.[51][52][53] In contrast, tornadic waterspouts are stronger tornadoes over water. They form over water similarly to mesocyclonic tornadoes, or are stronger tornadoes which cross over water. Since they form from severe thunderstorms and can be far more intense, faster, and longer-lived than fair weather waterspouts, they are more dangerous.[54] In official tornado statistics, waterspouts are generally not counted unless they affect land, though some European weather agencies count waterspouts and tornadoes together.[4][55]

Landspout

A landspout, or dust-tube tornado, is a tornado not associated with a mesocyclone. The name stems from their characterization as a "fair weather waterspout on land". Waterspouts and landspouts share many defining characteristics, including relative weakness, short lifespan, and a small, smooth condensation funnel which often does not reach the surface. Landspouts also create a distinctively laminar cloud of dust when they make contact with the ground, due to their differing mechanics from true mesoform tornadoes. Though usually weaker than classic tornadoes, they can produce strong winds which could cause serious damage.[4][16]

Similar circulations

Gustnado


A dust devil in Arizona

A gustnado, or gust front tornado, is a small, vertical swirl associated with a gust front or downburst. Because they are not connected with a cloud base, there is some debate as to whether or not gustnadoes are tornadoes. They are formed when fast moving cold, dry outflow air from a thunderstorm is blown through a mass of stationary, warm, moist air near the outflow boundary, resulting in a "rolling" effect (often exemplified through a roll cloud). If low level wind shear is strong enough, the rotation can be turned vertically or diagonally and make contact with the ground. The result is a gustnado.[4][56] They usually cause small areas of heavier rotational wind damage among areas of straight-line wind damage.

Dust devil

A dust devil resembles a tornado in that it is a vertical swirling column of air. However, they form under clear skies and are no stronger than the weakest tornadoes. They form when a strong convective updraft is formed near the ground on a hot day. If there is enough low level wind shear, the column of hot, rising air can develop a small cyclonic motion that can be seen near the ground. They are not considered tornadoes because they form during fair weather and are not associated with any clouds. However, they can, on occasion, result in major damage in arid areas.[23][57]

Fire whirls

Small-scale, tornado-like circulations can occur near any intense surface heat source. Those that occur near intense wildfires are called fire whirls. They are not considered tornadoes, except in the rare case where they connect to a pyrocumulus or other cumuliform cloud above. Fire whirls usually are not as strong as tornadoes associated with thunderstorms. They can, however, produce significant damage.[21]

Steam devils

A steam devil is a rotating updraft that involves steam or smoke. Steam devils are very rare. They most often form from smoke issuing from a power plant's smokestack. Hot springs and deserts may also be suitable locations for a steam devil to form. The phenomenon can occur over water, when cold arctic air passes over relatively warm water.[23]

Intensity and damage

Tornado rating classifications[21][58]
F0
EF0
F1
EF1
F2
EF2
F3
EF3
F4
EF4
F5
EF5
Weak Strong Violent
Significant
Intense
The Fujita scale and the Enhanced Fujita Scale rate tornadoes by damage caused. The Enhanced Fujita (EF) Scale was an update to the older Fujita scale, by expert elicitation, using engineered wind estimates and better damage descriptions. The EF Scale was designed so that a tornado rated on the Fujita scale would receive the same numerical rating, and was implemented starting in the United States in 2007. An EF0 tornado will probably damage trees but not substantial structures, whereas an EF5 tornado can rip buildings off their foundations leaving them bare and even deform large skyscrapers. The similar TORRO scale ranges from a T0 for extremely weak tornadoes to T11 for the most powerful known tornadoes. Doppler weather radar data, photogrammetry, and ground swirl patterns (cycloidal marks) may also be analyzed to determine intensity and award a rating.[4][59][60]

A house displaying EF1 damage. The roof and garage door have been damaged, but walls and supporting structures are still intact.

Tornadoes vary in intensity regardless of shape, size, and location, though strong tornadoes are typically larger than weak tornadoes. The association with track length and duration also varies, although longer track tornadoes tend to be stronger.[61] In the case of violent tornadoes, only a small portion of the path is of violent intensity, most of the higher intensity from subvortices.[21]

In the United States, 80% of tornadoes are EF0 and EF1 (T0 through T3) tornadoes. The rate of occurrence drops off quickly with increasing strength—less than 1% are violent tornadoes (EF4, T8 or stronger).[62] Outside Tornado Alley, and North America in general, violent tornadoes are extremely rare. This is apparently mostly due to the lesser number of tornadoes overall, as research shows that tornado intensity distributions are fairly similar worldwide. A few significant tornadoes occur annually in Europe, Asia, southern Africa, and southeastern South America, respectively.[63]

Climatology


Areas worldwide where tornadoes are most likely, indicated by orange shading

The United States has the most tornadoes of any country, nearly four times more than estimated in all of Europe, excluding waterspouts.[64] This is mostly due to the unique geography of the continent. North America is a large continent that extends from the tropics north into arctic areas, and has no major east-west mountain range to block air flow between these two areas. In the middle latitudes, where most tornadoes of the world occur, the Rocky Mountains block moisture and buckle the atmospheric flow, forcing drier air at mid-levels of the troposphere due to downsloped winds, and causing the formation of a low pressure area downwind to the east of the mountains. Increased westerly flow off the Rockies force the formation of a dry line when the flow aloft is strong,[65] while the Gulf of Mexico fuels abundant low-level moisture in the southerly flow to its east. This unique topography allows for frequent collisions of warm and cold air, the conditions that breed strong, long-lived storms throughout the year. A large portion of these tornadoes form in an area of the central United States known as Tornado Alley.[6] This area extends into Canada, particularly Ontario and the Prairie Provinces, although southeast Quebec, the interior of British Columbia, and western New Brunswick are also tornado-prone.[66] Tornadoes also occur across northeastern Mexico.[4]

The United States averages about 1,200 tornadoes per year. The Netherlands has the highest average number of recorded tornadoes per area of any country (more than 20, or 0.0013 per sq mi (0.00048 per km2), annually), followed by the UK (around 33, or 0.00035 per sq mi (0.00013 per km2), per year),[67][68] but most are small and cause minor damage. In absolute number of events, ignoring area, the UK experiences more tornadoes than any other European country, excluding waterspouts.[64]

Intense tornado activity in the United States. The darker-colored areas denote the area commonly referred to as Tornado Alley.

Tornadoes kill an average of 179 people per year in Bangladesh, the most in the world.[69] This is due to high population density, poor quality of construction and lack of tornado safety knowledge, as well as other factors.[69][70] Other areas of the world that have frequent tornadoes include South Africa, the La Plata Basin area, portions of Europe, Australia and New Zealand, and far eastern Asia.[7][71]

Tornadoes are most common in spring and least common in winter, but tornadoes can occur any time of year that favorable conditions occur.[21] Spring and fall experience peaks of activity as those are the seasons when stronger winds, wind shear, and atmospheric instability are present.[72] Tornadoes are focused in the right front quadrant of landfalling tropical cyclones, which tend to occur in the late summer and autumn. Tornadoes can also be spawned as a result of eyewall mesovortices, which persist until landfall.[73]

Tornado occurrence is highly dependent on the time of day, because of solar heating.[74] Worldwide, most tornadoes occur in the late afternoon, between 3 pm and 7 pm local time, with a peak near 5 pm.[75][76][77][78][79] Destructive tornadoes can occur at any time of day. The Gainesville Tornado of 1936, one of the deadliest tornadoes in history, occurred at 8:30 am local time.[21]

Associations with climate and climate change


U. S. Annual January - December Tornado Count 1976-2011 from NOAA National Climatic Data Center

Associations with various climate and environmental trends exist. For example, an increase in the sea surface temperature of a source region (e.g. Gulf of Mexico and Mediterranean Sea) increases atmospheric moisture content. Increased moisture can fuel an increase in severe weather and tornado activity, particularly in the cool season.[80]

Some evidence does suggest that the Southern Oscillation is weakly correlated with changes in tornado activity, which vary by season and region, as well as whether the ENSO phase is that of El Niño or La Niña.[81]

Climatic shifts may affect tornadoes via teleconnections in shifting the jet stream and the larger weather patterns. The climate-tornado link is confounded by the forces affecting larger patterns and by the local, nuanced nature of tornadoes. Although it is reasonable to suspect that global warming may affect trends in tornado activity,[82] any such effect is not yet identifiable due to the complexity, local nature of the storms, and database quality issues. Any effect would vary by region.[83]

Detection


Path of a tornado across Wisconsin on August 21, 1857

Rigorous attempts to warn of tornadoes began in the United States in the mid-20th century. Before the 1950s, the only method of detecting a tornado was by someone seeing it on the ground. Often, news of a tornado would reach a local weather office after the storm. However, with the advent of weather radar, areas near a local office could get advance warning of severe weather. The first public tornado warnings were issued in 1950 and the first tornado watches and convective outlooks in 1952. In 1953 it was confirmed that hook echoes are associated with tornadoes.[84] By recognizing these radar signatures, meteorologists could detect thunderstorms probably producing tornadoes from dozens of miles away.[85]

Radar

Today, most developed countries have a network of weather radars, which remains the main method of detecting signatures probably associated with tornadoes. In the United States and a few other countries, Doppler weather radar stations are used. These devices measure the velocity and radial direction (towards or away from the radar) of the winds in a storm, and so can spot evidence of rotation in storms from more than a hundred miles (160 km) away. When storms are distant from a radar, only areas high within the storm are observed and the important areas below are not sampled.[86] Data resolution also decreases with distance from the radar. Some meteorological situations leading to tornadogenesis are not readily detectable by radar and on occasion tornado development may occur more quickly than radar can complete a scan and send the batch of data. Also, most populated areas on Earth are now visible from the Geostationary Operational Environmental Satellites (GOES), which aid in the nowcasting of tornadic storms.[87]

A Doppler on Wheels radar loop of a hook echo and associated mesocyclone in Goshen County, Wyoming on June 5, 2009. Strong mesocyclones show up as adjacent areas of yellow and blue (on other radars, bright red and bright green), and usually indicate an imminent or occurring tornado.

Storm spotting

In the mid-1970s, the U.S. National Weather Service (NWS) increased its efforts to train storm spotters to spot key features of storms which indicate severe hail, damaging winds, and tornadoes, as well as damage itself and flash flooding. The program was called Skywarn, and the spotters were local sheriff's deputies, state troopers, firefighters, ambulance drivers, amateur radio operators, civil defense (now emergency management) spotters, storm chasers, and ordinary citizens. When severe weather is anticipated, local weather service offices request that these spotters look out for severe weather, and report any tornadoes immediately, so that the office can warn of the hazard.

Usually spotters are trained by the NWS on behalf of their respective organizations, and report to them. The organizations activate public warning systems such as sirens and the Emergency Alert System (EAS), and forward the report to the NWS.[88] There are more than 230,000 trained Skywarn weather spotters across the United States.[89]

In Canada, a similar network of volunteer weather watchers, called Canwarn, helps spot severe weather, with more than 1,000 volunteers.[87] In Europe, several nations are organizing spotter networks under the auspices of Skywarn Europe[90] and the Tornado and Storm Research Organisation (TORRO) has maintained a network of spotters in the United Kingdom since 1974.[91]

Storm spotters are needed because radar systems such as NEXRAD do not detect a tornado; merely signatures which hint at the presence of tornadoes.[92] Radar may give a warning before there is any visual evidence of a tornado or imminent tornado, but ground truth from an observer can either verify the threat or determine that a tornado is not imminent.[93] The spotter's ability to see what radar cannot is especially important as distance from the radar site increases, because the radar beam becomes progressively higher in altitude further away from the radar, chiefly due to curvature of Earth, and the beam also spreads out.[86]

Visual evidence


A rotating wall cloud with rear flank downdraft clear slot evident to its left rear

Storm spotters are trained to discern whether a storm seen from a distance is a supercell. They typically look to its rear, the main region of updraft and inflow. Under the updraft is a rain-free base, and the next step of tornadogenesis is the formation of a rotating wall cloud. The vast majority of intense tornadoes occur with a wall cloud on the backside of a supercell.[62]

Evidence of a supercell comes from the storm's shape and structure, and cloud tower features such as a hard and vigorous updraft tower, a persistent, large overshooting top, a hard anvil (especially when backsheared against strong upper level winds), and a corkscrew look or striations. Under the storm and closer to where most tornadoes are found, evidence of a supercell and likelihood of a tornado includes inflow bands (particularly when curved) such as a "beaver tail", and other clues such as strength of inflow, warmth and moistness of inflow air, how outflow- or inflow-dominant a storm appears, and how far is the front flank precipitation core from the wall cloud. Tornadogenesis is most likely at the interface of the updraft and rear flank downdraft, and requires a balance between the outflow and inflow.[16]

Only wall clouds that rotate spawn tornadoes, and usually precede the tornado by five to thirty minutes. Rotating wall clouds may be a visual manifestation of a low-level mesocyclone. Barring a low-level boundary, tornadogenesis is highly unlikely unless a rear flank downdraft occurs, which is usually visibly evidenced by evaporation of cloud adjacent to a corner of a wall cloud. A tornado often occurs as this happens or shortly after; first, a funnel cloud dips and in nearly all cases by the time it reaches halfway down, a surface swirl has already developed, signifying a tornado is on the ground before condensation connects the surface circulation to the storm. Tornadoes may also occur without wall clouds, under flanking lines, and on the leading edge. Spotters watch all areas of a storm, and the cloud base and surface.[94]

Extremes


A map of the tornado paths in the Super Outbreak (April 3–4, 1974)

The most record-breaking tornado in recorded history was the Tri-State Tornado, which roared through parts of Missouri, Illinois, and Indiana on March 18, 1925. It was likely an F5, though tornadoes were not ranked on any scale in that era. It holds records for longest path length (219 miles, 352 km), longest duration (about 3.5 hours), and fastest forward speed for a significant tornado (73 mph, 117 km/h) anywhere on Earth. In addition, it is the deadliest single tornado in United States history (695 dead).[21] The tornado was also the costliest tornado in history at the time (unadjusted for inflation), but in the years since has been surpassed by several others if population changes over time are not considered. When costs are normalized for wealth and inflation, it ranks third today.[95]

The deadliest tornado in world history was the Daultipur-Salturia Tornado in Bangladesh on April 26, 1989, which killed approximately 1,300 people.[69] Bangladesh has had at least 19 tornadoes in its history kill more than 100 people, almost half of the total in the rest of the world.

The most extensive tornado outbreak on record was the April 25–28, 2011 tornado outbreak, which spawned 355 confirmed tornadoes over the southeastern United States - 211 of them within a single 24-hour period. The previous record was the Super Outbreak of 1974 which spawned nearly 148 tornadoes.

While direct measurement of the most violent tornado wind speeds is nearly impossible, since conventional anemometers would be destroyed by the intense winds and flying debris, some tornadoes have been scanned by mobile Doppler radar units, which can provide a good estimate of the tornado's winds. The highest wind speed ever measured in a tornado, which is also the highest wind speed ever recorded on the planet, is 301 ± 20 mph (484 ± 32 km/h) in the F5 Bridge Creek-Moore, Oklahoma, tornado which killed 36 people.[96] Though the reading was taken about 100 feet (30 m) above the ground, this is a testament to the power of the strongest tornadoes.[2]

Storms that produce tornadoes can feature intense updrafts, sometimes exceeding 150 mph (240 km/h). Debris from a tornado can be lofted into the parent storm and carried a very long distance. A tornado which affected Great Bend, Kansas, in November 1915, was an extreme case, where a "rain of debris" occurred 80 miles (130 km) from the town, a sack of flour was found 110 miles (180 km) away, and a cancelled check from the Great Bend bank was found in a field outside of Palmyra, Nebraska, 305 miles (491 km) to the northeast.[97] Waterspouts and tornadoes have been advanced as an explanation for instances of raining fish and other animals.[98]

Safety

Though tornadoes can strike in an instant, there are precautions and preventative measures that people can take to increase the chances of surviving a tornado. Authorities such as the Storm Prediction Center advise having a pre-determined plan should a tornado warning be issued. When a warning is issued, going to a basement or an interior first-floor room of a sturdy building greatly increases chances of survival.[99] In tornado-prone areas, many buildings have storm cellars on the property. These underground refuges have saved thousands of lives.[100]
Some countries have meteorological agencies which distribute tornado forecasts and increase levels of alert of a possible tornado (such as tornado watches and warnings in the United States and Canada). Weather radios provide an alarm when a severe weather advisory is issued for the local area, though these are mainly available only in the United States. Unless the tornado is far away and highly visible, meteorologists advise that drivers park their vehicles far to the side of the road (so as not to block emergency traffic), and find a sturdy shelter. If no sturdy shelter is nearby, getting low in a ditch is the next best option. Highway overpasses are one of the worst places to take shelter during tornadoes, as the constricted space can be subject to increased wind speed and funneling of debris underneath the overpass.[101]

Myths and misconceptions

Folklore often identifies a green sky with tornadoes, and though the phenomenon may be associated with severe weather, there is no evidence linking it specifically with tornadoes.[102] It is often thought that opening windows will lessen the damage caused by the tornado. While there is a large drop in atmospheric pressure inside a strong tornado, it is unlikely that the pressure drop would be enough to cause the house to explode. Some research indicates that opening windows may actually increase the severity of the tornado's damage.[citation needed] A violent tornado can destroy a house whether its windows are open or closed.[103][104]

The 1999 Salt Lake City tornado disproved several misconceptions, including the idea that tornadoes cannot occur in areas like Utah or in cities.

Another commonly held misconception is that highway overpasses provide adequate shelter from tornadoes. This belief is partly inspired by widely circulated video captured during the 1991 tornado outbreak near Andover, Kansas, where a news crew and several other people take shelter under an overpass on the Kansas Turnpike and safely ride out a tornado as it passes by.[105] However, a highway overpass is a dangerous place during a tornado: the subjects of the video remained safe due to an unlikely combination of events: the storm in question was a weak tornado, did not directly strike the overpass, and the overpass itself was of a unique design.[105] Due to the Venturi effect, tornadic winds are accelerated in the confined space of an overpass.[106] Indeed, in the 1999 Oklahoma tornado outbreak of May 3, 1999, three highway overpasses were directly struck by tornadoes, and at all three locations there was a fatality, along with many life-threatening injuries.[107] By comparison, during the same tornado outbreak, more than 2000 homes were completely destroyed, with another 7000 damaged, and yet only a few dozen people died in their homes.[101]

An old belief is that the southwest corner of a basement provides the most protection during a tornado. The safest place is the side or corner of an underground room opposite the tornado's direction of approach (usually the northeast corner), or the central-most room on the lowest floor. Taking shelter in a basement, under a staircase, or under a sturdy piece of furniture such as a workbench further increases chances of survival.[103][104]

Finally, there are areas which people believe to be protected from tornadoes, whether by being in a city, near a major river, hill, or mountain, or even protected by supernatural forces.[108] Tornadoes have been known to cross major rivers, climb mountains,[109] affect valleys, and have damaged several city centers. As a general rule, no area is safe from tornadoes, though some areas are more susceptible than others.[23][103][104]

Ongoing research


A Doppler on Wheels unit observing a tornado near Attica, Kansas

Meteorology is a relatively young science and the study of tornadoes is newer still. Although researched for about 140 years and intensively for around 60 years, there are still aspects of tornadoes which remain a mystery.[110] Scientists have a fairly good understanding of the development of thunderstorms and mesocyclones,[111][112] and the meteorological conditions conducive to their formation. However, the step from supercell (or other respective formative processes) to tornadogenesis and predicting tornadic vs. non-tornadic mesocyclones is not yet well known and is the focus of much research.[72]

Also under study are the low-level mesocyclone and the stretching of low-level vorticity which tightens into a tornado,[72] namely, what are the processes and what is the relationship of the environment and the convective storm. Intense tornadoes have been observed forming simultaneously with a mesocyclone aloft (rather than succeeding mesocyclogenesis) and some intense tornadoes have occurred without a mid-level mesocyclone.[113]

In particular, the role of downdrafts, particularly the rear-flank downdraft, and the role of baroclinic boundaries, are intense areas of study.[114]

Reliably predicting tornado intensity and longevity remains a problem, as do details affecting characteristics of a tornado during its life cycle and tornadolysis. Other rich areas of research are tornadoes associated with mesovortices within linear thunderstorm structures and within tropical cyclones.[115]

Scientists still do not know the exact mechanisms by which most tornadoes form, and occasional tornadoes still strike without a tornado warning being issued.[116] Analysis of observations including both stationary and mobile (surface and aerial) in-situ and remote sensing (passive and active) instruments generates new ideas and refines existing notions. Numerical modeling also provides new insights as observations and new discoveries are integrated into our physical understanding and then tested in computer simulations which validate new notions as well as produce entirely new theoretical findings, many of which are otherwise unattainable. Importantly, development of new observation technologies and installation of finer spatial and temporal resolution observation networks have aided increased understanding and better predictions.[117]

Research programs, including field projects such as the VORTEX projects (Verification of the Origins of Rotation in Tornadoes Experiment), deployment of TOTO (the TOtable Tornado Observatory), Doppler On Wheels (DOW), and dozens of other programs, hope to solve many questions that still plague meteorologists.[44] Universities, government agencies such as the National Severe Storms Laboratory, private-sector meteorologists, and the National Center for Atmospheric Research are some of the organizations very active in research; with various sources of funding, both private and public, a chief entity being the National Science Foundation.[92][118] The pace of research is partly constrained by the number of observations that can be taken; gaps in information about the wind, pressure, and moisture content throughout the local atmosphere; and the computing power available for simulation.[119]

Solar storms similar to tornadoes have been recorded, but it is unknown how closely related they are to their terrestrial counterparts.[120]

Gallery


Gene


From Wikipedia, the free encyclopedia


This stylistic diagram shows a gene in relation to the double helix structure of DNA and to a chromosome (right). The chromosome is X-shaped because it is dividing. Introns are regions often found in eukaryote genes that are removed in the splicing process (after the DNA is transcribed into RNA): Only the exons encode the protein. The diagram labels a region of only 55 or so bases as a gene. In reality, most genes are hundreds of times longer.

The chemical structure of a four-base fragment of a DNA double helix.

A gene is the molecular unit of heredity of a living organism. It is used extensively by the scientific community as a name given to some stretches of deoxyribonucleic acids (DNA) and ribonucleic acids (RNA) that code for a polypeptide or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains. Genes hold the information to build and maintain an organism's cells and pass genetic traits to offspring. All organisms have genes corresponding to various biological traits, some of which are instantly visible, such as eye color or number of limbs, and some of which are not, such as blood type, increased risk for specific diseases, or the thousands of basic biochemical processes that comprise life. The word gene is derived from the Greek word genesis meaning "birth", or genos meaning "origin" (see pangenesis).

A modern working definition of a gene is "a locatable region of genomic sequence, corresponding to a unit of inheritance, which is associated with regulatory regions, transcribed regions, and or other functional sequence regions ".[1][2] Colloquial usage of the term gene (e.g., "good genes", "hair color gene") may actually refer to an allele: a gene is the basic instruction— a sequence of nucleic acids (DNA or, in the case of certain viruses RNA), while an allele is one variant of that gene. Thus, when the mainstream press refers to "having" a "gene" for a specific trait, this is customarily inaccurate. In most cases, all people would have a gene for the trait in question, although certain people will have a specific allele of that gene, which results in the trait variant. Further, genes code for proteins, which might result in identifiable traits, but it is the gene (genotype), not the trait (phenotype), which is inherited.

Big genes are a class of genes whose nuclear transcript spans 500 kb (1 kb = 1,000 base pairs) or more of chromosomal DNA. The largest of the big genes is the gene for dystrophin, which spans 2.3 Mb. Many big genes have modestly sized mRNAs; the exons encoding these RNAs typically encompass about 1% of the total chromosomal gene region in which they occur.

History


Gregor Mendel

The existence of genes was first implied from the work of Gregor Mendel (1822–1884), who, between the years of 1857 to 1864 planted 8000 common edible pea plants and studied and tabulated the inheritance patterns in peaplants (Pisum) tracking inheritance of traits from parent to offspring and describing these mathematically as 2n combinations where n is the number of differing characteristics in the original peas. Although he did not use the term gene, he explained his results in terms of inherited characteristics. The notion of a gene[3] is evolving with the science of genetics, but began when Mendel noticed that biological variations are inherited from parent or grandparent organisms as specific, discrete traits and are transmitted thus unaltered from the original source. Prior to Mendel's work, the dominant theory of heredity was one of blending inheritance, pangenesis, which suggested that each parent contributed fluids to the fertilisation process and that in meiosis the traits of the parents blended and mixed to produce the offspring. Although Mendel's work was largely unrecognized after its first publication in 1866, it was 'rediscovered' in 1900 by three European scientists, Hugo de Vries, Carl Correns, and Erich von Tschermak, who claimed to have reached similar conclusions in their own research. However, these scientists were not yet aware of the identity of the 'discrete units' on which genetic material resides. The biological entity responsible for defining traits was later termed a gene, but the biological basis for inheritance remained unknown until DNA was identified as the genetic material in the 1940s. Mendel was also the first to show independent assortment, the distinction between dominant and recessive traits, the distinction between a heterozygote and homozygote, the phenomenon of discontinuing inheritance and what would later be described as genotype (the genetic material of an organism) and phenotype (the visible traits of that organism) and the conversion of one form into another within few generations.

Charles Darwin used the term gemmule to describe a microscopic unit of inheritance, and what would later become known as chromosomes had been observed separating out during cell division by Wilhelm Hofmeister as early as 1848. The idea that chromosomes are the carriers of inheritance was expressed in 1883 by Wilhelm Roux. Darwin also coined the word pangenesis by (1868).[4] The word pangenesis is made from the Greek words pan (a prefix meaning "whole", "encompassing") and genesis ("birth") or genos ("origin").

Mendel's concept was given a name by Hugo de Vries in 1889, in his book Intracellular Pangenesis; although probably unaware of Mendel's work at the time, he coined the term "pangen" for "the smallest particle [representing] one hereditary characteristic".[5] Danish botanist Wilhelm Johannsen coined the word "gene" ("gen" in Danish and German) in 1909 to describe the fundamental physical and functional units of heredity,[6] while the related word genetics was first used by William Bateson in 1905.[7] He derived the word from de Vries' "pangen". In the early 1900s, Mendel's work received renewed attention from scientists. In 1910, Thomas Hunt Morgan showed that genes reside on specific chromosomes. He later showed that genes occupy specific locations on the chromosome. With this knowledge, Morgan and his students began the first chromosomal map of the fruit fly Drosophila. In 1928, Frederick Griffith showed that genes could be transferred. In what is now known as Griffith's experiment, injections into a mouse of a deadly strain of bacteria that had been heat-killed transferred genetic information to a safe strain of the same bacteria, killing the mouse.

A series of subsequent discoveries led to the realization decades later that chromosomes within cells are the carriers of genetic material, and that they are made of DNA (deoxyribonucleic acid), a polymeric molecule found in all cells on which the 'discrete units' of Mendelian inheritance are encoded. In 1941, George Wells Beadle and Edward Lawrie Tatum showed that mutations in genes caused errors in specific steps in metabolic pathways. This showed that specific genes code for specific proteins, leading to the "one gene, one enzyme" hypothesis.[7] Oswald Avery, Colin Munro MacLeod, and Maclyn McCarty showed in 1944 that DNA holds the gene's information.[8] In 1952, Rosalind Franklin and Raymond Gosling produced a strikingly clear x-ray diffraction pattern indicating a helical form, and in 1953, James D. Watson and Francis Crick demonstrated the molecular structure of DNA. Together, these discoveries established the central dogma of molecular biology, which states that proteins are translated from RNA which is transcribed from DNA. This dogma has since been shown to have exceptions, such as reverse transcription in retroviruses.

In 1972, Walter Fiers and his team at the Laboratory of Molecular Biology of the University of Ghent (Ghent, Belgium) were the first to determine the sequence of a gene: the gene for Bacteriophage MS2 coat protein.[9] Richard J. Roberts and Phillip Sharp discovered in 1977 that genes can be split into segments. This led to the idea that one gene can make several proteins. Recently (as of 2003–2006), biological results let the notion of gene appear more slippery. In particular, genes do not seem to sit side by side on DNA like discrete beads. Instead, regions of the DNA producing distinct proteins may overlap, so that the idea emerges that "genes are one long continuum".[1] It was first hypothesized in 1986 by Walter Gilbert that neither DNA nor protein would be required in such a primitive system as that of a very early stage of the earth if RNA could perform as simply a catalyst and genetic information storage processor.

The modern study of genetics at the level of DNA is known as molecular genetics and the synthesis of molecular genetics with traditional Darwinian evolution is known as the modern evolutionary synthesis.

Mendelian inheritance and classical genetics


Crossing between two pea plants heterozygous for purple (B, dominant) and white (b, recessive) blossoms

According to the theory of Mendelian inheritance, variations in phenotype—the observable physical and behavioral characteristics of an organism—are due in part to variations in genotype, or the organism's particular set of genes, each of which specifies a particular trait. Different forms of a gene, which may give rise to different phenotypes, are known as alleles. Organisms such as the pea plants Mendel worked on, along with many plants and animals, have two alleles for each trait, one inherited from each parent. Alleles may be dominant or recessive; dominant alleles give rise to their corresponding phenotypes when paired with any other allele for the same trait, whereas recessive alleles give rise to their corresponding phenotype only when paired with another copy of the same allele. For example, if the allele specifying tall stems in pea plants is dominant over the allele specifying short stems, then pea plants that inherit one tall allele from one parent and one short allele from the other parent will also have tall stems. Mendel's work demonstrated that alleles assort independently in the production of gametes, or germ cells, ensuring variation in the next generation.

Physical definitions

RNA genes and genomes in the world

When proteins are manufactured, the gene is first copied into RNA as an intermediate product. In other cases, the RNA molecules are the actual functional products. For example, RNAs known as ribozymes are capable of enzymatic function, and microRNA has a regulatory role. The DNA sequences from which such RNAs are transcribed are known as RNA genes.

Some viruses store their entire genomes in the form of RNA, and contain no DNA at all. Because they use RNA to store genes, their cellular hosts may synthesize their proteins as soon as they are infected and without the delay in waiting for transcription. On the other hand, RNA retroviruses, such as HIV, require the reverse transcription of their genome from RNA into DNA before their proteins can be synthesized. In 2006, French researchers came across a puzzling example of RNA-mediated inheritance in mice. Mice with a loss-of-function mutation in the gene Kit have white tails. Offspring of these mutants can have white tails despite having only normal Kit genes. The research team traced this effect back to mutated Kit RNA.[10] While RNA is common as genetic storage material in viruses, in mammals in particular RNA inheritance has been observed very rarely.

Functional structure of a gene


Diagram of the "typical" eukaryotic protein-coding gene. Promoters and enhancers determine what portions of the DNA will be transcribed into the precursor mRNA (pre-mRNA). The pre-mRNA is then spliced into messenger RNA (mRNA) which is later translated into protein.

The vast majority of living organisms encode their genes in long strands of DNA (deoxyribonucleic acid). DNA consists of a chain made from four types of nucleotide subunits, each composed of: a five-carbon sugar (2'-deoxyribose), a phosphate group, and one of the four bases adenine, cytosine, guanine, and thymine. The most common form of DNA in a cell is in a double helix structure, in which two individual DNA strands twist around each other in a right-handed spiral. In this structure, the base pairing rules specify that guanine pairs with cytosine and adenine pairs with thymine. The base pairing between guanine and cytosine forms three hydrogen bonds, whereas the base pairing between adenine and thymine forms two hydrogen bonds. The two strands in a double helix must therefore be complementary, that is, their bases must align such that the adenines of one strand are paired with the thymines of the other strand, and so on.

Due to the chemical composition of the pentose residues of the bases, DNA strands have directionality. One end of a DNA polymer contains an exposed hydroxyl group on the deoxyribose; this is known as the 3' end of the molecule. The other end contains an exposed phosphate group; this is the 5' end. The directionality of DNA is vitally important to many cellular processes, since double helices are necessarily directional (a strand running 5'-3' pairs with a complementary strand running 3'-5'), and processes such as DNA replication occur in only one direction. All nucleic acid synthesis in a cell occurs in the 5'-3' direction, because new monomers are added via a dehydration reaction that uses the exposed 3' hydroxyl as a nucleophile.

The expression of genes encoded in DNA begins by transcribing the gene into RNA, a second type of nucleic acid that is very similar to DNA, but whose monomers contain the sugar ribose rather than deoxyribose. RNA also contains the base uracil in place of thymine. RNA molecules are less stable than DNA and are typically single-stranded. Genes that encode proteins are composed of a series of three-nucleotide sequences called codons, which serve as the words in the genetic language. The genetic code specifies the correspondence during protein translation between codons and amino acids. The genetic code is nearly the same for all known organisms.

All genes have regulatory regions in addition to regions that explicitly code for a protein or RNA product. A regulatory region shared by almost all genes is known as the promoter, which provides a position that is recognized by the transcription machinery when a gene is about to be transcribed and expressed. A gene can have more than one promoter, resulting in RNAs that differ in how far they extend in the 5' end.[11] Although promoter regions have a consensus sequence that is the most common sequence at this position, some genes have "strong" promoters that bind the transcription machinery well, and others have "weak" promoters that bind poorly. These weak promoters usually permit a lower rate of transcription than the strong promoters, because the transcription machinery binds to them and initiates transcription less frequently. Other possible regulatory regions include enhancers, which can compensate for a weak promoter. Most regulatory regions are "upstream"—that is, before or toward the 5' end of the transcription initiation site. Eukaryotic promoter regions are much more complex and difficult to identify than prokaryotic promoters.

Many prokaryotic genes are organized into operons, or groups of genes whose products have related functions and which are transcribed as a unit. By contrast, eukaryotic genes are transcribed only one at a time, but may include long stretches of DNA called introns which are transcribed but never translated into protein (they are spliced out before translation). Splicing can also occur in prokaryotic genes, but is less common than in eukaryotes.[12]

Chromosomes

The total complement of genes in an organism or cell is known as its genome, which may be stored on one or more chromosomes; the region of the chromosome at which a particular gene is located is called its locus. A chromosome consists of a single, very long DNA helix on which thousands of genes are encoded. Prokaryotesbacteria and archaea—typically store their genomes on a single large,circular chromosome, sometimes supplemented by additional small circles of DNA called plasmids, which usually encode only a few genes and are easily transferable between individuals. For example, the genes for antibiotic resistance are usually encoded on bacterial plasmids and can be passed between individual cells, even those of different species, via horizontal gene transfer. Although some simple eukaryotes also possess plasmids with small numbers of genes, the majority of eukaryotic genes are stored on multiple linear chromosomes, which are packed within the nucleus in complex with storage proteins called histones. The manner in which DNA is stored on the histone, as well as chemical modifications of the histone itself, are regulatory mechanisms governing whether a particular region of DNA is accessible for gene expression. The ends of eukaryotic chromosomes are capped by long stretches of repetitive sequences called telomeres, which do not code for any gene product but are present to prevent degradation of coding and regulatory regions during DNA replication. The length of the telomeres tends to decrease each time the genome is replicated in preparation for cell division; the loss of telomeres has been proposed as an explanation for cellular senescence, or the loss of the ability to divide, and by extension for the aging process in organisms.[13]

Whereas the chromosomes of prokaryotes are relatively gene-dense, those of eukaryotes often contain so-called "junk DNA", or regions of DNA that serve no obvious function. Simple single-celled eukaryotes have relatively small amounts of such DNA, whereas the genomes of complex multicellular organisms, including humans, contain an absolute majority of DNA without an identified function.[14] However it now appears that, although protein-coding DNA makes up barely 2% of the human genome, about 80% of the bases in the genome may be expressed, so the term "junk DNA" may be a misnomer.[2]

Gene expression

In all organisms, there are two major steps separating a protein-coding gene from its protein: First, the DNA on which the gene resides must be transcribed from DNA to messenger RNA (mRNA); and, second, it must be translated from mRNA to protein. RNA-coding genes must still go through the first step, but are not translated into protein. The process of producing a biologically functional molecule of either RNA or protein is called gene expression, and the resulting molecule itself is called a gene product.

Genetic code


Schematic diagram of a single-stranded RNA molecule illustrating the position of three-base codons.

The genetic code is the set of rules by which information encoded within a gene is translated into a functional protein. Each gene consists of a specific sequence of nucleotides encoded in DNA or RNA. The nucleotide being made up of a sugar, a phosphate molecule and a specific base (adenine, thymine, cytosine, guanine or sometimes uracil [thymine is replaced with uracil in some viruses[15]]); a correspondence between nucleotides, the basic building blocks of genetic material, and amino acids, the basic building blocks of proteins, must be established for genes to be successfully translated into functional proteins. Sets of three nucleotides, known as codons, each correspond to a specific amino acid or to a signal; three codons are known as "stop codons" and, instead of specifying a new amino acid, alert the translation machinery that the end of the gene has been reached, just as a specific set of 3 bases, "AUG", known as the "start codon", signifies the gene to start transcribing. There are 64 possible codons (four possible nucleotides at each of three positions, hence 43 possible codons) and only 20 standard amino acids; hence the code is redundant and multiple codons can specify the same amino acid. The correspondence between codons and amino acids is nearly universal among all known living organisms.

Transcription

The process of genetic transcription produces a single-stranded RNA molecule known as messenger RNA, whose nucleotide sequence is complementary to the DNA from which it was transcribed. The DNA strand whose sequence matches that of the RNA is known as the coding strand and the strand from which the RNA was synthesized is the template strand. Transcription is performed by an enzyme called an RNA polymerase, which reads the template strand in the 3' to 5' direction and synthesizes the RNA from 5' to 3'. To initiate transcription, the polymerase first recognizes and binds a promoter region of the gene. Thus a major mechanism of gene regulation is the blocking or sequestering of the promoter region, either by tight binding by repressor molecules that physically block the polymerase, or by organizing the DNA so that the promoter region is not accessible.

In prokaryotes, transcription occurs in the cytoplasm; for very long transcripts, translation may begin at the 5' end of the RNA while the 3' end is still being transcribed. In eukaryotes, transcription necessarily occurs in the nucleus, where the cell's DNA is sequestered; the RNA molecule produced by the polymerase is known as the primary transcript and must undergo post-transcriptional modifications before being exported to the cytoplasm for translation. The splicing of introns present within the transcribed region is a modification unique to eukaryotes; alternative splicing mechanisms can result in mature transcripts from the same gene having different sequences and thus coding for different proteins. This is a major form of regulation in eukaryotic cells.

Translation

Translation is the process by which a mature mRNA molecule is used as a template for synthesizing a new protein. Translation is carried out by ribosomes, large complexes of RNA and protein responsible for carrying out the chemical reactions to add new amino acids to a growing polypeptide chain by the formation of peptide bonds. The genetic code is read three nucleotides at a time, in units called codons, via interactions with specialized RNA molecules called transfer RNA (tRNA). Each tRNA has three unpaired bases known as the anticodon that are complementary to the codon it reads; the tRNA is also covalently attached to the amino acid specified by the complementary codon. When the tRNA binds to its complementary codon in an mRNA strand, the ribosome ligates its amino acid cargo to the new polypeptide chain, which is synthesized from amino terminus to carboxyl terminus. During and after its synthesis, the new protein must fold to its active three-dimensional structure before it can carry out its cellular function.

DNA replication and inheritance

The growth, development, and reproduction of organisms relies on cell division, or the process by which a single cell divides into two usually identical daughter cells. This requires first making a duplicate copy of every gene in the genome in a process called DNA replication. The copies are made by specialized enzymes known as DNA polymerases, which "read" one strand of the double-helical DNA, known as the template strand, and synthesize a new complementary strand. Because the DNA double helix is held together by base pairing, the sequence of one strand completely specifies the sequence of its complement; hence only one strand needs to be read by the enzyme to produce a faithful copy. The process of DNA replication is semiconservative; that is, the copy of the genome inherited by each daughter cell contains one original and one newly synthesized strand of DNA.[16]

After DNA replication is complete, the cell must physically separate the two copies of the genome and divide into two distinct membrane-bound cells. In prokaryotes - bacteria and archaea - this usually occurs via a relatively simple process called binary fission, in which each circular genome attaches to the cell membrane and is separated into the daughter cells as the membrane invaginates to split the cytoplasm into two membrane-bound portions. Binary fission is extremely fast compared to the rates of cell division in eukaryotes. Eukaryotic cell division is a more complex process known as the cell cycle; DNA replication occurs during a phase of this cycle known as S phase, whereas the process of segregating chromosomes and splitting the cytoplasm occurs during M phase. In many single-celled eukaryotes such as yeast, reproduction by budding is common, which results in asymmetrical portions of cytoplasm in the two daughter cells.

Molecular inheritance

The duplication and transmission of genetic material from one generation of cells to the next is the basis for molecular inheritance, and the link between the classical and molecular pictures of genes. Organisms inherit the characteristics of their parents because the cells of the offspring contain copies of the genes in their parents' cells. In asexually reproducing organisms, the offspring will be a genetic copy or clone of the parent organism. In sexually reproducing organisms, a specialized form of cell division called meiosis produces cells called gametes or germ cells that are haploid, or contain only one copy of each gene. The gametes produced by females are called eggs or ova, and those produced by males are called sperm. Two gametes fuse to form a fertilized egg, a single cell that once again has a diploid number of genes—each with one copy from the mother and one copy from the father.

During the process of meiotic cell division, an event called genetic recombination or crossing-over can sometimes occur, in which a length of DNA on one chromatid is swapped with a length of DNA on the corresponding sister chromatid. This has no effect if the alleles on the chromatids are the same, but results in reassortment of otherwise linked alleles if they are different. The Mendelian principle of independent assortment asserts that each of a parent's two genes for each trait will sort independently into gametes; which allele an organism inherits for one trait is unrelated to which allele it inherits for another trait. This is in fact only true for genes that do not reside on the same chromosome, or are located very far from one another on the same chromosome. The closer two genes lie on the same chromosome, the more closely they will be associated in gametes and the more often they will appear together; genes that are very close are essentially never separated because it is extremely unlikely that a crossover point will occur between them. This is known as genetic linkage.

Mutation

DNA replication is for the most part extremely accurate, with an error rate per site of around 10−6 to 10−10 in eukaryotes.[16] (Although in prokaryotes and viruses, the rate is much higher.) Rare, spontaneous alterations in the base sequence of a particular gene arise from a number of sources, such as errors in DNA replication and the aftermath of DNA damage. These errors are called mutations
The cell contains many DNA repair mechanisms for preventing mutations and maintaining the integrity of the genome; however, in some cases—such as breaks in both DNA strands of a chromosome—repairing the physical damage to the molecule is a higher priority than producing an exact copy. Due to the degeneracy of the genetic code, some mutations in protein-coding genes are silent, or produce no change in the amino acid sequence of the protein for which they code; for example, the codons UCU and UCC both code for serine, so the U→C mutation has no effect on the protein. Mutations that do have phenotypic effects are most often neutral or deleterious to the organism. Variants may confer benefits to the organism's fitness; it is commonly thought that mutations may produce beneficial variants. The most common mutations include point mutations in which a single codon is replaced, frame shift mutation where a single nucleotide base is inserted or deleted from the DNA strand so that all bases are shifted over, silent mutations where a single nucleotide base is replaced but without causing a change for the amino acid being coded for, and nonsense mutations, where a change in a single nucleotide base causes a codon to be turned into a stop codon hence terminating transcription at this point.
Mutations propagated to the next generation lead to variations within a species' population. Variants of a single gene are known as alleles, and differences in alleles may give rise to differences in traits. Although it is rare for the variants in a single gene to have clearly distinguishable phenotypic effects, certain well-defined traits are in fact controlled by single genetic loci. A gene's most common allele is called the wild type allele, and rare alleles are called mutants. However, this does not imply that the wild-type allele is the ancestor from which the mutants are descended. For the most part, these mutations are recessive and are phased out quickly. However, on occasion these mutations appear as dominant to other alleles, becoming predominant and increasing in the rate they are seen in a population.

Genome

Chromosomal organization

The total complement of genes in an organism or cell is known as its genome. In prokaryotes, the vast majority of genes are located on a single chromosome of circular DNA, while eukaryotes usually possess multiple individual linear DNA helices packed into dense DNA-protein complexes called chromosomes. Genes that appear together on one chromosome of one species may appear on separate chromosomes in another species. Many species carry more than one copy of their genome within each of their somatic cells. Cells or organisms with only one copy of each chromosome are called haploid; those with two copies are called diploid; and those with more than two copies are called polyploid. The copies of genes on the chromosomes are not necessarily identical. In sexually reproducing organisms, one copy is normally inherited from each parent.

Number of genes



The protein-encoding component of the human genome, categorized by function of each gene product, given both as number of genes and as percentage of all genes.[17]

Early estimates of the number of human genes that used expressed sequence tag data put it at 50 000–100 000.[18] Following the sequencing of the human genome and other genomes, it has been found that rather few genes (~20 000 in human, mouse and fly, ~13 000 in roundworm, >46,000 in rice[19]) encode all the proteins in an organism.[20] These protein-coding sequences make up 1–2% of the human genome.[21] A large part of the genome is transcribed however, to introns, retrotransposons and seemingly a large array of noncoding RNAs.[20][21] Total number of proteins (the Earth's proteome) is estimated to be 5 million sequences.[22]

Genetic and genomic nomenclature

Gene nomenclature has been established by the HUGO Gene Nomenclature Committee (HGNC) for each known human gene in the form of an approved gene name and symbol (short-form abbreviation). All approved symbols are stored in the HGNC Database. Each symbol is unique and each gene is only given one approved gene symbol. This also facilitates electronic data retrieval from publications. In preference each symbol maintains parallel construction in different members of a gene family and can be used in other species, especially the mouse.

Essential genes

Essential genes are those genes of an organism that are thought to be critical for its survival. Surprisingly few genes have been shown to be absolutely essential for the survival of bacteria, e.g. only about 10% of the ~4,200 genes of Escherichia coli.

Evolutionary concept of a gene

George C. Williams first explicitly advocated the gene-centric view of evolution in his 1966 book Adaptation and Natural Selection. He proposed an evolutionary concept of gene to be used when we are talking about natural selection favoring some genes. The definition is: "that which segregates and recombines with appreciable frequency." According to this definition, even an asexual genome could be considered a gene, insofar that it have an appreciable permanency through many generations.
The difference is: the molecular gene transcribes as a unit, and the evolutionary gene inherits as a unit.

Richard Dawkins' books The Selfish Gene (1976) and The Extended Phenotype (1982) defended the idea that the gene is the only replicator in living systems. This means that only genes transmit their structure largely intact and are potentially immortal in the form of copies. So, genes should be the unit of selection. In River Out of Eden, Dawkins further refined the idea of gene-centric selection by describing life as a river of compatible genes flowing through geological time. Scoop up a bucket of genes from the river of genes, and we have an organism serving as temporary bodies or survival machines. A river of genes may fork into two branches representing two non-interbreeding species as a result of geographical separation.

Gene targeting and implications

Gene targeting is commonly referred to techniques for altering or disrupting mouse genes and provides the mouse models for studying the roles of individual genes in embryonic development, human disorders, aging and diseases. The mouse models, where one or more of its genes are deactivated or made inoperable, are called knockout mice. Since the first reports in which homologous recombination among homologous chromosomes in embryonic stem cells was used to generate gene-targeted mice,[23] gene targeting has proven to be a powerful means of precisely manipulating the mammalian genome, producing at least ten thousand mutant mouse strains and it is now possible to introduce mutations that can be activated at specific time points, or in specific cells or organs, both during development and in the adult animal.[24][25]
Gene targeting strategies have been expanded to all kinds of modifications, including point mutations, isoform deletions, mutant allele correction, large pieces of chromosomal DNA insertion and deletion, tissue specific disruption combined with spatial and temporal regulation and so on. It is predicted that the ability to generate mouse models with predictable phenotypes will have a major impact on studies of all phases of development, immunology, neurobiology, oncology, physiology, metabolism, and human diseases. Gene targeting is also in theory applicable to species from which totipotent embryonic stem cells can be established, and therefore may offer a potential to the improvement of domestic animals and plants.[25][26]

Changing concept

The concept of the gene has changed considerably (see history section). From the original definition of a "unit of inheritance", the term evolved to mean a DNA-based unit that can exert its effects on the organism through RNA or protein products. It was also previously believed that one gene makes one protein; this concept was overthrown by the discovery of alternative splicing and trans-splicing.[7]

The definition of a gene is still changing. The first cases of RNA-based inheritance have been discovered in mammals.[27] Evidence is also accumulating that the control regions of a gene do not necessarily have to be close to the coding sequence on the linear molecule or even on the same chromosome. Spilianakis and colleagues discovered that the promoter region of the interferon-gamma gene on chromosome 10 and the regulatory regions of the T(H)2 cytokine locus on chromosome 11 come into close proximity in the nucleus possibly to be jointly regulated.[28] Even the coding sequence of a gene itself doesn't have to be all on the same chromosome: Marande and Burger showed that, in the mitochondria of the protist Diplonema papillatum, "genes are systematically fragmented into small pieces that are encoded on separate chromosomes, transcribed individually, and then concatenated into contiguous messenger RNA molecules".[29]

The concept that genes are clearly delimited is also being eroded. There is evidence for fused proteins stemming from two adjacent genes that can produce two separate protein products. While it is not clear whether these fusion proteins are functional, the phenomenon is more frequent than previously thought.[30] Even more ground-breaking than the discovery of fused genes is the observation that some proteins can be composed of exons from far away regions and even different chromosomes.[2][31] This new data has led to an updated, and probably tentative, definition of a gene as "a union of genomic sequences encoding a coherent set of potentially overlapping functional products".[7] This new definition categorizes genes by functional products, whether they be proteins or RNA, rather than specific DNA loci; all regulatory elements of DNA are therefore classified as gene-associated regions.[7]

Bayesian inference

From Wikipedia, the free encyclopedia https://en.wikipedia.org/wiki/Bayesian_inference Bayesian inference ( / ...