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Tuesday, February 3, 2015

Europa (moon)


From Wikipedia, the free encyclopedia

Europa
Europa-moon.jpg
Europa's trailing hemisphere in approximate natural color. The prominent crater in the lower right is Pwyll and the darker regions are areas where Europa's primarily water ice surface has a higher mineral content. Imaged on 7 September 1996 by Galileo spacecraft.
Discovery
Discovered by Galileo Galilei
Simon Marius
Discovery date 8 January 1610[1]
Designations
Jupiter II
Adjectives Europan
Orbital characteristics[3]
Epoch 8 January 2004
Periapsis 664862 km[a]
Apoapsis 676938 km[b]
Mean orbit radius
670900 km[2]
Eccentricity 0.009[2]
3.551181 d[2]
Average orbital speed
13.740 km/s[2]
Inclination 0.470° (to Jupiter's equator)[2]
Satellite of Jupiter
Physical characteristics
Mean radius
1560.8±0.5 km (0.245 Earths)[4]
3.09×107 km2 (0.061 Earths)[c]
Volume 1.593×1010 km3 (0.015 Earths)[d]
Mass (4.799844±0.000013)×1022 kg (0.008 Earths)[4]
Mean density
3.013±0.005 g/cm3[4]
1.314 m/s2 (0.134 g)[e]
2.025 km/s[f]
Synchronous[5]
0.1°[6]
Albedo 0.67 ± 0.03[4]
Surface temp. min mean max
Surface ≈ 50 K[7] 102 K (−171.15°C) 125 K
5.29 (opposition)[4]
Atmosphere
Surface pressure
0.1 µPa (10−12 bar)[8]

Europa Listeni/jʊˈrpə/[9] (Jupiter II), is the sixth-closest moon of the planet Jupiter, and the smallest of its four Galilean satellites, but still the sixth-largest moon in the Solar System. Europa was discovered in 1610 by Galileo Galilei[1] Progressively better observations of Europa have occurred over the centuries by Earth-bound telescopes, and by space probe flybys starting in the 1970s.
Slightly smaller than the Moon, Europa is primarily made of silicate rock and has a water-ice crust and probably an iron–nickel core. It has a tenuous atmosphere composed primarily of oxygen. Its surface is striated by cracks and streaks, whereas craters are relatively rare. It has the smoothest surface of any known solid object in the Solar System.[10] The apparent youth and smoothness of the surface have led to the hypothesis that a water ocean exists beneath it, which could conceivably serve as an abode for extraterrestrial life.[11] This hypothesis proposes that heat from tidal flexing causes the ocean to remain liquid and drives geological activity similar to plate tectonics.[12] On 8 September 2014, NASA reported finding evidence confirming earlier reports of plate tectonics in Europa's thick ice shell – the first sign of such geological activity on a world other than Earth.[13]
In December 2013, NASA reported the detection of "clay-like minerals" (specifically, phyllosilicates), often associated with "organic material" on the icy crust of Europa.[14] In addition, NASA announced, based on studies with the Hubble Space Telescope, that water vapor plumes were detected on Europa and were similar to water vapor plumes detected on Enceladus, moon of Saturn.[15]

The Galileo mission, launched in 1989, provided the bulk of current data on Europa. No spacecraft has yet landed on Europa, but its intriguing characteristics have led to several ambitious exploration proposals. The European Space Agency's Jupiter Icy Moon Explorer (JUICE) is a mission to Europa that is due to launch in 2022.[16] NASA is planning a robotic mission that would be launched in the "mid-2020s".[17]

Discovery and naming

Europa, along with Jupiter's three other large moons, Io, Ganymede, and Callisto, was discovered by Galileo Galilei on 8 January 1610,[1] and possibly independently by Simon Marius. The first reported observation of Io and Europa was made by Galileo Galilei on 7 January 1610 using a 20×-magnification refracting telescope at the University of Padua. However, in that observation, Galileo could not separate Io and Europa due to the low magnification of his telescope, so that the two were recorded as a single point of light. Io and Europa were seen for the first time as separate bodies during Galileo's observations of the Jupiter system the following day, 8 January 1610 (used as the discovery date for Europa by the IAU).[1] It is named after a Phoenician noblewoman in Greek mythology, Europa, who was courted by Zeus and became the queen of Crete.[18]

Like all the Galilean satellites, Europa is named after a lover of Zeus, the Greek counterpart of Jupiter, in this case Europa, daughter of the king of Tyre. The naming scheme was suggested by Simon Marius, who apparently discovered the four satellites independently, though Galileo accused Marius of plagiarism.[19][20] Marius attributed the proposal to Johannes Kepler.[19][20]

The names fell out of favor for a considerable time and were not revived in general use until the mid-20th century.[21] In much of the earlier astronomical literature, Europa is simply referred to by its Roman numeral designation as Jupiter II (a system also introduced by Galileo) or as the "second satellite of Jupiter". In 1892, the discovery of Amalthea, whose orbit lay closer to Jupiter than those of the Galilean moons, pushed Europa to the third position. The Voyager probes discovered three more inner satellites in 1979, so Europa is now considered Jupiter's sixth satellite, though it is still sometimes referred to as Jupiter II.[21]

Orbit and rotation


Animation showing Io's Laplace resonance with Europa and Ganymede

Europa orbits Jupiter in just over three and a half days, with an orbital radius of about 670,900 km. With an eccentricity of only 0.009, the orbit itself is nearly circular, and the orbital inclination relative to the Jovian equatorial plane is small, at 0.470°.[22] Like its fellow Galilean satellites, Europa is tidally locked to Jupiter, with one hemisphere of Europa constantly facing Jupiter. Because of this, there is a sub-Jovian point on Europa's surface, from which Jupiter would appear to hang directly overhead. Europa's prime meridian is the line intersecting this point.[23] Research suggests the tidal locking may not be full, as a non-synchronous rotation has been proposed: Europa spins faster than it orbits, or at least did so in the past. This suggests an asymmetry in internal mass distribution and that a layer of subsurface liquid separates the icy crust from the rocky interior.[5]

The slight eccentricity of Europa's orbit, maintained by the gravitational disturbances from the other Galileans, causes Europa's sub-Jovian point to oscillate about a mean position. As Europa comes slightly nearer to Jupiter, Jupiter's gravitational attraction increases, causing Europa to elongate towards and away from it. As Europa moves slightly away from Jupiter, Jupiter's gravitational force decreases, causing Europa to relax back into a more spherical shape, and creating tides in its ocean. The orbital eccentricity of Europa is continuously pumped by its mean-motion resonance with Io.[24] Thus, the tidal flexing kneads Europa's interior and gives it a source of heat, possibly allowing its ocean to stay liquid while driving subsurface geological processes.[12][24] The ultimate source of this energy is Jupiter's rotation, which is tapped by Io through the tides it raises on Jupiter and is transferred to Europa and Ganymede by the orbital resonance.[24][25]

Scientists analyzing the unique cracks lining the icy face of Europa found evidence showing that this moon of Jupiter likely spun around a tilted axis at some point in time. If this hypothesis is correct, this tilt would be an explanation for many of Europa's features. Europa's immense network of crisscrossing cracks serves as a record of the stresses caused by massive tides in the moon's global ocean. Europa's tilt could influence calculations of how much of the moon's history is recorded in its frozen shell, how much heat is generated by tides in its ocean, and even how long the ocean has been liquid. The moon's ice layer must stretch to accommodate these changes. When there is too much stress, it cracks. A tilt in the moon's axis could suggest that Europa's cracks may be much more recent than previously thought. The reason is that the direction of the spin pole may change by as much as a few degrees per day, completing one precession period over several months. A tilt also could affect the estimates of the age of Europa's ocean. Tidal forces are thought to generate the heat that keeps Europa's ocean liquid, and a tilt in the spin axis might suggest that more heat is generated by tidal forces. This heat might help the ocean to remain liquid longer. Scientists did not specify when the tilt would have occurred and measurements have not been made of the tilt of Europa's axis.[26]

Physical characteristics


Europa (lower left) compared to the Moon (top left) and Earth (right) to scale approximately. (montage)

Europa is slightly smaller than the Moon. At just over 3,100 kilometres (1,900 mi) in diameter, it is the sixth-largest moon and fifteenth largest object in the Solar System. Though by a wide margin the least massive of the Galilean satellites, it is nonetheless more massive than all known moons in the Solar System smaller than itself combined.[27] Its bulk density suggests that it is similar in composition to the terrestrial planets, being primarily composed of silicate rock.[28]

Internal structure

It is believed that Europa has an outer layer of water around 100 km (62 mi) thick; some as frozen-ice upper crust, some as liquid ocean underneath the ice. Recent magnetic field data from the Galileo orbiter showed that Europa has an induced magnetic field through interaction with Jupiter's, which suggests the presence of a subsurface conductive layer.[29] The layer is likely a salty liquid water ocean. Portions of the crust are estimated to have undergone a rotation of nearly 80°, nearly flipping over (see true polar wander), which would be unlikely if the ice were solidly attached to the mantle.[30] Europa probably contains a metallic iron core.[31]

Surface features


Approximate-natural-color (left) and enhanced-color (right) Galileo view of leading hemisphere

Europa is one of the smoothest objects in the Solar System when considering the lack of large scale features such as mountains or craters,[32] however on a smaller scale Europa's equator has been theorised to be covered in 10 metre tall icy spikes called penitentes caused by the effect of direct overhead sunlight on the equator melting vertical cracks.[33] The prominent markings crisscrossing Europa seem to be mainly albedo features, which emphasize low topography. There are few craters on Europa because its surface is tectonically active and young.[34][35] Europa's icy crust gives it an albedo (light reflectivity) of 0.64, one of the highest of all moons.[22][35] This would seem to indicate a young and active surface; based on estimates of the frequency of cometary bombardment that Europa probably endures, the surface is about 20 to 180 million years old.[36] There is currently no full scientific consensus among the sometimes contradictory explanations for the surface features of Europa.[37]

The radiation level at the surface of Europa is equivalent to a dose of about 5400 mSv (540 rem) per day,[38] an amount of radiation that would cause severe illness or death in human beings exposed for a single day.[39]

Lineae

Realistic-color Galileo mosaic of Europa's anti-Jovian hemisphere showing numerous lineae

Mosaic of Galileo images showing features indicative of tidal flexing: lineae, lenticulae (domes, pits) and Conamara Chaos.

Europa's most striking surface features are a series of dark streaks crisscrossing the entire globe, called lineae (English: lines). Close examination shows that the edges of Europa's crust on either side of the cracks have moved relative to each other. The larger bands are more than 20 km (12 mi) across, often with dark, diffuse outer edges, regular striations, and a central band of lighter material.[40] The most likely hypothesis states that these lineae may have been produced by a series of eruptions of warm ice as the Europan crust spread open to expose warmer layers beneath.[41] The effect would have been similar to that seen in Earth's oceanic ridges. These various fractures are thought to have been caused in large part by the tidal flexing exerted by Jupiter. Because Europa is tidally locked to Jupiter, and therefore always maintains the same approximate orientation towards Jupiter, the stress patterns should form a distinctive and predictable pattern. However, only the youngest of Europa's fractures conform to the predicted pattern; other fractures appear to occur at increasingly different orientations the older they are. This could be explained if Europa's surface rotates slightly faster than its interior, an effect that is possible due to the subsurface ocean mechanically decoupling Europa's surface from its rocky mantle and the effects of Jupiter's gravity tugging on Europa's outer ice crust.[42] Comparisons of Voyager and Galileo spacecraft photos serve to put an upper limit on this hypothetical slippage. The full revolution of the outer rigid shell relative to the interior of Europa occurs over a minimum of 12,000 years.[43] Studies of Voyager and Galileo images have revealed evidence of subduction on Europa's surface, suggesting that, just as the cracks are analogous to ocean ridges,[44][45] so plates of icy crust analogous to tectonic plates on Earth are recycled into the molten interior. Together, the evidence for crustal spreading at bands [44] and convergence at other sites [45] marks the first evidence for plate tectonics on any world other than Earth. [13]

Other geological features

Craggy, 250 m high peaks and smooth plates are jumbled together in a close-up of Conamara Chaos.

Other features present on Europa are circular and elliptical lenticulae (Latin for "freckles"). Many are domes, some are pits and some are smooth, dark spots. Others have a jumbled or rough texture. The dome tops look like pieces of the older plains around them, suggesting that the domes formed when the plains were pushed up from below.[46]

One hypothesis states that these lenticulae were formed by diapirs of warm ice rising up through the colder ice of the outer crust, much like magma chambers in Earth's crust.[46] The smooth, dark spots could be formed by meltwater released when the warm ice breaks through the surface. The rough, jumbled lenticulae (called regions of "chaos"; for example, Conamara Chaos) would then be formed from many small fragments of crust embedded in hummocky, dark material, appearing like icebergs in a frozen sea.[47]

An alternative hypothesis suggest that lenticulae are actually small areas of chaos and that the claimed pits, spots and domes are artefacts resulting from over-interpretation of early, low-resolution Galileo images. The implication is that the ice is too thin to support the convective diapir model of feature formation. [48] [49]

In November 2011, a team of researchers from the University of Texas at Austin and elsewhere presented evidence in the journal Nature suggesting that many "chaos terrain" features on Europa sit atop vast lakes of liquid water.[50][51] These lakes would be entirely encased in Europa's icy outer shell and distinct from a liquid ocean thought to exist farther down beneath the ice shell. Full confirmation of the lakes' existence will require a space mission designed to probe the ice shell either physically or indirectly, for example using radar.

Two possible models of Europa

Subsurface ocean

Scientists' consensus is that a layer of liquid water exists beneath Europa's surface, and that heat from tidal flexing allows the subsurface ocean to remain liquid.[12][52] Europa's surface temperature averages about 110 K (−160 °C; −260 °F) at the equator and only 50 K (−220 °C; −370 °F) at the poles, keeping Europa's icy crust as hard as granite.[7] The first hints of a subsurface ocean came from theoretical considerations of tidal heating (a consequence of Europa's slightly eccentric orbit and orbital resonance with the other Galilean moons). Galileo imaging team members argue for the existence of a subsurface ocean from analysis of Voyager and Galileo images.[52] The most dramatic example is "chaos terrain", a common feature on Europa's surface that some interpret as a region where the subsurface ocean has melted through the icy crust. This interpretation is extremely controversial. Most geologists who have studied Europa favor what is commonly called the "thick ice" model, in which the ocean has rarely, if ever, directly interacted with the present surface.[53] The different models for the estimation of the ice shell thickness give values between a few kilometers and tens of kilometers.[54] The best evidence for the thick-ice model is a study of Europa's large craters. The largest impact structures are surrounded by concentric rings and appear to be filled with relatively flat, fresh ice; based on this and on the calculated amount of heat generated by Europan tides, it is predicted that the outer crust of solid ice is approximately 10–30 km (6–19 mi) thick, including a ductile "warm ice" layer, which could mean that the liquid ocean underneath may be about 100 km (60 mi) deep.[36][55] This leads to a volume of Europa's oceans of 3 × 1018 m3, slightly more than two times the volume of Earth's oceans.

The thin-ice model suggests that Europa's ice shell may be only a few kilometers thick. However, most planetary scientists conclude that this model considers only those topmost layers of Europa's crust that behave elastically when affected by Jupiter's tides. One example is flexure analysis, in which Europa's crust is modeled as a plane or sphere weighted and flexed by a heavy load. Models such as this suggest the outer elastic portion of the ice crust could be as thin as 200 metres (660 ft). If the ice shell of Europa is really only a few kilometers thick, this "thin ice" model would mean that regular contact of the liquid interior with the surface could occur through open ridges, causing the formation of areas of chaotic terrain.[54]

In late 2008, it was suggested Jupiter may keep Europa's oceans warm by generating large planetary tidal waves on Europa because of its small but non-zero obliquity. This previously unconsidered kind of tidal force generates so-called Rossby waves that travel quite slowly, at just a few kilometers per day, but can generate significant kinetic energy. For the current axial tilt estimate of 0.1 degree, the resonance from Rossby waves would store 7.3×1017 J of kinetic energy, which is two thousand times larger than that of the flow excited by the dominant tidal forces.[56][57] Dissipation of this energy could be the principal heat source of Europa's ocean.

The Galileo orbiter found that Europa has a weak magnetic moment, which is induced by the varying part of the Jovian magnetic field. The field strength at the magnetic equator (about 120 nT) created by this magnetic moment is about one-sixth the strength of Ganymede's field and six times the value of Callisto's.[58] The existence of the induced moment requires a layer of a highly electrically conductive material in Europa's interior. The most plausible candidate for this role is a large subsurface ocean of liquid saltwater.[31] Spectrographic evidence suggests that the dark, reddish streaks and features on Europa's surface may be rich in salts such as magnesium sulfate, deposited by evaporating water that emerged from within.[59] Sulfuric acid hydrate is another possible explanation for the contaminant observed spectroscopically.[60] In either case, because these materials are colorless or white when pure, some other material must also be present to account for the reddish color, and sulfur compounds are suspected.[61]

Plumes


Water vapor plumes on Jupiter's moon Europa (artist's impression).[62]

Europa may have periodically occurring plumes of water 200 km (120 mi) high, or more than 20 times the height of Mt. Everest.[15][63][64] These plumes appear when Europa is at its farthest point from Jupiter, and are not seen when Europa is at its closest point to Jupiter, in agreement with tidal force modeling predictions.[65] The tidal forces are about 1,000 times stronger than the Moon's effect on Earth. The only other moon in the Solar System exhibiting water vapor plumes is Enceladus.[15] The estimated eruption rate at Europa is about 7000 kg/s[65] compared to about 200 kg/s for the plumes of Enceladus.[66][67]

Atmosphere

Observations with the Goddard High Resolution Spectrograph of the Hubble Space Telescope, first described in 1995, revealed that Europa has a thin atmosphere composed mostly of molecular oxygen (O2).[68][69] The surface pressure of Europa's atmosphere is 0.1 μPa, or 10−12 times that of the Earth.[8] In 1997, the Galileo spacecraft confirmed the presence of a tenuous ionosphere (an upper-atmospheric layer of charged particles) around Europa created by solar radiation and energetic particles from Jupiter's magnetosphere,[70][71] providing evidence of an atmosphere.

Magnetic field around Europa. The red line shows a trajectory of the Galileo spacecraft during a typical flyby (E4 or E14).

Unlike the oxygen in Earth's atmosphere, Europa's is not of biological origin. The surface-bounded atmosphere forms through radiolysis, the dissociation of molecules through radiation.[72] Solar ultraviolet radiation and charged particles (ions and electrons) from the Jovian magnetospheric environment collide with Europa's icy surface, splitting water into oxygen and hydrogen constituents. These chemical components are then adsorbed and "sputtered" into the atmosphere. The same radiation also creates collisional ejections of these products from the surface, and the balance of these two processes forms an atmosphere.[73] Molecular oxygen is the densest component of the atmosphere because it has a long lifetime; after returning to the surface, it does not stick (freeze) like a water or hydrogen peroxide molecule but rather desorbs from the surface and starts another ballistic arc. Molecular hydrogen never reaches the surface, as it is light enough to escape Europa's surface gravity.[74][75]

Observations of the surface have revealed that some of the molecular oxygen produced by radiolysis is not ejected from the surface. Because the surface may interact with the subsurface ocean (considering the geological discussion above), this molecular oxygen may make its way to the ocean, where it could aid in biological processes.[76] One estimate suggests that, given the turnover rate inferred from the apparent ~0.5 Gyr maximum age of Europa's surface ice, subduction of radiolytically generated oxidizing species might well lead to oceanic free oxygen concentrations that are comparable to those in terrestrial deep oceans.[77]

The molecular hydrogen that escapes Europa's gravity, along with atomic and molecular oxygen, forms a gas torus in the vicinity of Europa's orbit around Jupiter. This "neutral cloud" has been detected by both the Cassini and Galileo spacecraft, and has a greater content (number of atoms and molecules) than the neutral cloud surrounding Jupiter's inner moon Io. Models predict that almost every atom or molecule in Europa's torus is eventually ionized, thus providing a source to Jupiter's magnetospheric plasma. [78]

Exploration


Europa in 1979 by Voyager 1

Exploration of Europa began with the Jupiter flybys of Pioneer 10 and 11 in 1973 and 1974 respectively. The first closeup photos were of low resolution compared to later missions.

The two Voyager probes traveled through the Jovian system in 1979 providing more detailed images of Europa's icy surface. The images caused many scientists to speculate about the possibility of a liquid ocean underneath.

Starting in 1995, the Galileo spaceprobe began a Jupiter orbiting mission that lasted for eight years, until 2003, and provided the most detailed examination of the Galilean moons to date. It included the Galileo Europa Mission and Galileo Millennium Mission, with numerous close flybys of Europa.[79]
New Horizons imaged Europa in 2007, as it flew by the Jovian system while on its way to Pluto.

Jupiter and Europa as seen in 2007 by the New Horizons spacecraft

Future missions

Conjectures on extraterrestrial life have ensured a high profile for Europa and have led to steady lobbying for future missions.[80][81] The aims of these missions have ranged from examining Europa's chemical composition to searching for extraterrestrial life in its hypothesized subsurface oceans.[82][83] Robotic missions to Europa need to endure the high radiation environment around itself and Jupiter.[81] Europa receives about 5.40 Sv of radiation per day.[84]

In 2011, a Europa mission was recommended by the U.S. Planetary Science Decadal Survey.[85] In response, NASA commissioned Europa lander concept studies in 2011, along with concepts for a Europa flyby (Europa Clipper), and a Europa orbiter.[86][87] The orbiter element option concentrates on the "ocean" science, while the multiple-flyby element (Clipper) concentrates on the chemistry and energy science. On 13 January 2014, the House Appropriations Committee announced a new bipartisan bill that includes $80 million funding to continue the Europa mission concept studies.[88][89]
  • Europa Clipper — In July 2013 an updated concept for a flyby Europa mission called Europa Clipper was presented by the Jet Propulsion Laboratory (JPL) and the Applied Physics Laboratory (APL).[90] The aim of Europa Clipper is to explore Europa in order to investigate its habitability, and to aid selecting sites for a future lander. The Europa Clipper would not orbit Europa, but instead orbit Jupiter and conduct 45 low-altitude flybys of Europa during its envisioned mission. The probe would carry an ice-penetrating radar, short-wave infrared spectrometer, topographical imager, and an ion- and neutral-mass spectrometer.
  • Europa Orbiter — Its objective would be to characterize the extent of the ocean and its relation to the deeper interior. Instrument payload could include a radio subsystem, laser altimeter, magnetometer, Langmuir probe, and a mapping camera.[91][92]
  • Europa Lander — It would investigate the moon's habitability and assess its astrobiological potential by confirming the existence and determining the characteristics of water within and below Europa's icy shell.[93]
In 2012, Jupiter Icy Moon Explorer was selected by the European Space Agency (ESA) as a planned mission.[16][94] That mission includes some flybys of Europa, but is more focused on Ganymede.

Old proposals


Europa Lander Mission concept circa 2005 (NASA).

In the early 2000s, Jupiter Europa Orbiter led by NASA and the Jupiter Ganymede Orbiter led by the ESA were proposed together as an Outer Planet Flagship Mission to Jupiter's icy moons, and called Europa Jupiter System Mission with a planned launch in 2020.[95] In 2009 it was given priority over Titan Saturn System Mission.[96] At that time, there was competition from other proposals.[97] Japan proposed Jupiter Magnetospheric Orbiter. Russia expressed interest in sending Europa Lander as part of the international effort.[98] The overall plan collapsed in the early 2010s.[94]

Jovian Europa Orbiter was an ESA Cosmic Vision concept study from 2007. Another concept was Ice Clipper,[99] which would have used an impactor similar to the Deep Impact mission—it would make a controlled crash into the surface of Europa, generating a plume of debris that would then be collected by a small spacecraft flying through the plume.[100][101]

Artist's concept of the cryobot (a thermal drill, seen upper left) and its deployed 'hydrobot' submersible

Jupiter Icy Moons Orbiter (JIMO) was a partially developed fission-powered spacecraft with ion thrusters that was cancelled in 2006.[81][102] It was part of Project Prometheus.[102] The Europa Lander Mission proposed a small nuclear-powered Europa lander for JIMO.[103] It would travel with the orbiter, which would also function as a communication relay to Earth.[103]

The Europa Orbiter received a go-ahead in 1999 but was canceled in 2002. This orbiter featured a special radar that would allow it to scan below the surface.[32]

More ambitious ideas have been put forward including an impactor in combination with a thermal drill to search for biosignatures that might be frozen in the shallow subsurface.[104][105]

Another proposal put forward in 2001 calls for a large nuclear-powered "melt probe" (cryobot) that would melt through the ice until it reached an ocean below.[81][106] Once it reached the water, it would deploy an autonomous underwater vehicle (hydrobot) that would gather information and send it back to Earth.[107] Both the cryobot and the hydrobot would have to undergo some form of extreme sterilization to prevent detection of Earth organisms instead of native life and to prevent contamination of the subsurface ocean.[108] This proposed mission has not yet reached a serious planning stage.[109]

Potential for extraterrestrial life


A black smoker in the Atlantic Ocean. Driven by geothermal energy, this and other types of hydrothermal vents create chemical disequilibria that can provide energy sources for life.

Europa has emerged as one of the top locations in the Solar System in terms of potential habitability and the possibility of hosting extraterrestrial life.[110] Life could exist in its under-ice ocean, perhaps subsisting in an environment similar to Earth's deep-ocean hydrothermal vents. Life in such an ocean could possibly be similar to microbial life on Earth in the deep ocean.[82][111] So far, there is no evidence that life exists on Europa, but the likely presence of liquid water has spurred calls to send a probe there.[112]

Until the 1970s, life, at least as the concept is generally understood, was believed to be entirely dependent on energy from the Sun. Plants on Earth's surface capture energy from sunlight to photosynthesize sugars from carbon dioxide and water, releasing oxygen in the process, and are then consumed by oxygen-respiring animals, passing their energy up the food chain. Even life in the deep ocean, far below the reach of sunlight, was believed to obtain its nourishment either from the organic detritus raining down from the surface, or by eating animals that in turn depend on that stream of nutrients.[113] An environment's ability to support life was thus thought to depend on its access to sunlight.

This giant tube worm colony dwells beside a Pacific Ocean vent. Although the worms require oxygen (hence their blood-red color), methanogens and some other microbes in the vent communities do not.

However, in 1977, during an exploratory dive to the Galapagos Rift in the deep-sea exploration submersible Alvin, scientists discovered colonies of giant tube worms, clams, crustaceans, mussels, and other assorted creatures clustered around undersea volcanic features known as black smokers.[113] These creatures thrive despite having no access to sunlight, and it was soon discovered that they comprise an entirely independent food chain. Instead of plants, the basis for this food chain was a form of bacterium that derived its energy from oxidization of reactive chemicals, such as hydrogen or hydrogen sulfide, that bubbled up from Earth's interior. This chemosynthesis revolutionized the study of biology by revealing that life need not be sunlight-dependent; it only requires water and an energy gradient in order to exist. It opened up a new avenue in astrobiology by massively expanding the number of possible extraterrestrial habitats.

Water vapor plume on Europa (artist concept) (December 12, 2013).[15]

For comparison, the eruption of a natural repeating water geyser on Earth

Although the tube worms and other multicellular eukaryotic organisms around these hydrothermal vents respire oxygen and thus are indirectly dependent on photosynthesis, anaerobic chemosynthetic bacteria and archaea that inhabit these ecosystems provide a possible model for life in Europa's ocean.[77] The energy provided by tidal flexing drives active geological processes within Europa's interior, just as they do to a far more obvious degree on its sister moon Io. Although Europa, like the Earth, may possess an internal energy source from radioactive decay, the energy generated by tidal flexing would be several orders of magnitude greater than any radiological source.[114] However, such an energy source could never support an ecosystem as large and diverse as the photosynthesis-based ecosystem on Earth's surface.[115] Life on Europa could exist clustered around hydrothermal vents on the ocean floor, or below the ocean floor, where endoliths are known to inhabit on Earth. Alternatively, it could exist clinging to the lower surface of Europa's ice layer, much like algae and bacteria in Earth's polar regions, or float freely in Europa's ocean.[116] However, if Europa's ocean were too cold, biological processes similar to those known on Earth could not take place. Similarly, if it were too salty, only extreme halophiles could survive in its environment.[116] In September 2009, planetary scientist Richard Greenberg calculated that cosmic rays impacting on Europa's surface convert some water ice into free oxygen (O2), which could then be absorbed into the ocean below as water wells up to fill cracks. Via this process, Greenberg estimates that Europa's ocean could eventually achieve an oxygen concentration greater than that of Earth's oceans within just a few million years. This would enable Europa to support not merely anaerobic microbial life but potentially larger, aerobic organisms such as fish.[117]

In 2006, Robert T. Pappalardo, an assistant professor in the Laboratory for Atmospheric and Space Physics at the University of Colorado in Boulder said,
We've spent quite a bit of time and effort trying to understand if Mars was once a habitable environment. Europa today, probably, is a habitable environment. We need to confirm this ... but Europa, potentially, has all the ingredients for life ... and not just four billion years ago ... but today.[80]
In November 2011, a team of researchers presented evidence in the journal Nature suggesting the existence of vast lakes of liquid water entirely encased in Europa's icy outer shell and distinct from a liquid ocean thought to exist farther down beneath the ice shell.[50][51] If confirmed, the lakes could be yet another potential habitat for life.

A paper published in March 2013 suggests that hydrogen peroxide is abundant across much of the surface of Jupiter's moon Europa.[118] The authors argue that if the peroxide on the surface of Europa mixes into the ocean below, it could be an important energy supply for simple forms of life, if life were to exist there. The scientists think hydrogen peroxide is an important factor for the habitability of the global liquid water ocean under Europa's icy crust because hydrogen peroxide decays to oxygen when mixed into liquid water.

On December 11, 2013, NASA reported the detection of "clay-like minerals" (specifically, phyllosilicates), often associated with organic materials, on the icy crust of Europa.[14] The presence of the minerals may have been the result of a collision with an asteroid or comet according to the scientists.[14]

Life on Earth could have been blasted into space by asteroid collisions and arrived on the moons of Jupiter in a process called lithopanspermia.[119]

Hox gene


From Wikipedia, the free encyclopedia

Hox genes (also known as homeotic genes) are a group of related genes that control the body plan of an embryo along the anterior-posterior (head-tail) axis. After the embryonic segments have formed, the Hox proteins determine the type of segment structures (e.g. legs, antennae, and wings in fruit flies or the different types of vertebrae in humans) that will form on a given segment. Hox proteins thus confer segmental identity, but do not form the actual segments themselves.[1]
Hox genes are defined as having the following properties:
  • their protein product is a transcription factor
  • they contain a DNA sequence known as the homeobox
  • in many animals, the organization of the Hox genes of the chromosome is the same as the order of their expression along the anterior-posterior axis of the developing animal, and are thus said to display colinearity.[2]

Hox genes code for transcription factors

The products of Hox genes are Hox proteins. Hox proteins are transcription factors, which are proteins that are capable of binding to specific nucleotide sequences on the DNA called enhancers where they either activate or repress genes. The same Hox protein can act as a repressor at one gene and an activator at another. The ability of Hox proteins to bind DNA is conferred by a part of the protein referred to as the homeodomain. The homeodomain is a 60-amino-acid-long DNA-binding domain (encoded by its corresponding 180-base-pair DNA sequence, the homeobox). This amino acid sequence folds into a "helix-turn-helix" (i.e. homeodomain fold) motif that is stabilized by a third helix. The consensus polypeptide chain is (typical intron position noted with dashes):[3]
RRRKRTA-YTRYQLLE-LEKEFLF-NRYLTRRRRIELAHSL-NLTERHIKIWFQN-RRMK-WKKEN

The sequence and function of Hox genes is highly conserved

The homeodomain protein motif is highly conserved across vast evolutionary distances. In addition, homeodomains of individual Hox proteins usually exhibit greater similarity to homeodomains in other species than to proteins encoded by adjacent genes within their own Hox cluster. These two observations led to the suggestions that Hox gene clusters evolved from a single Hox gene via tandem duplication and subsequent divergence and that a prototypic Hox gene cluster containing at least seven different Hox genes was present in the common ancestor of all bilaterian animals.[4]

The functional conservation of Hox proteins can be demonstrated by the fact that a fly can function perfectly well with a chicken Hox protein in place of its own.[5] So, despite having a last common ancestor that lived over 670 million years ago,[6] the chicken and fly version of the same Hox gene can actually take each other's places when swapped.

Hox gene function in Drosophila


Homeobox gene expression in Drosophila melanogaster

Drosophila melanogaster is an important model for understanding body plan generation and evolution. The general principles of Hox gene function and logic elucidated in flies will apply to all Bilaterian organisms, including humans. Drosophila, like all insects, has 8 Hox genes. These are clustered into two complexes, both of which are located on chromosome 3. The Antennapedia complex (not to be confused with the Antp gene) consists of 5 genes: labial (lab), proboscipedia (pb), Deformed (Dfd) Sex combs reduced (Scr) and Antennapedia (Antp). The Bithorax complex, named after the Ultrabithorax gene, consists of the remaining 3 genes: Ultrabithorax (Ubx), abdominal-A (abd-A) and Abdominal-B (abd-B).

lab (labial)

The lab gene is the most anteriorly expressed gene. It is expressed in the head, primarily in the intercalary segment (an appendage-less segment between the antenna and mandible), and also in the midgut. Loss of function of lab results in the failure of the Drosophila embryo to internalize the mouth and head structures that initially develop on the outside of its body (a process called head involution). Failure of head involution disrupts or deletes the salivary glands, pharynx. The lab gene was initially so named because it disrupted the labial appendage; however, the lab gene is not expressed in the labial segment, and the labial appendage phenotype is likely a result of the broad disorganization resulting from the failure of head involution.[7]

pb (proboscipedia)

The pb gene is responsible for the formation of the labial and maxillary palps. There is evidence that pb interacts with Scr.[8]

Dfd (Deformed)

The Dfd gene is responsible for the formation of the maxillary and mandibular segments in the larval head.[9] The mutant phenotypes of Deformed are similar to those of labial. Loss of function of Deformed in the embryo results in a failure of head involution (see labial gene), with a loss of larval head structures. Mutations in the adult have either deletions of parts of the head or transformations of head to thoracic identity.[7]

Scr (Sex combs reduced)

The Scr gene is responsible for cephalic and thoracic development in Drosophila embryo and adult.[10]

Antp (Antennapedia)

The 2nd thoracic segment, or T2, develops a pair of legs and a pair of wings. The Antp gene specifies this identity by promoting leg formation and allowing (but not directly activating) wing formation. A dominant Antp mutation, caused by a chromosomal inversion, causes Antp to be expressed in the antennal imaginal disc, so that, instead of forming an antenna, the disc makes a leg, resulting in a leg coming out of the fly's head.

Wild type (left) Mutant (right)

Ubx (Ultrabithorax)

The third thoracic segment, or T3, bears a pair of legs and a pair of halteres (highly reduced wings that function in balancing during flight). Ubx patterns T3 largely by repressing genes involved in wing formation. The wing blade is composed of two layers of cells that adhere tightly to one another, and are supplied with nutrient by several wing veins. One of the many genes that Ubx represses is blistered, which activates proteins involved in cell-cell adhesion, and spalt, which patterns the placement of wing veins. In Ubx loss-of-function mutants, Ubx no longer represses wing genes, and the halteres develop as a second pair of wings, resulting in the famous four-winged flies. When Ubx is misexpressed in the 2nd thoracic segment, such as occurs in flies with the "Cbx" enhancer mutation, it represses wing genes, and the wings develop as halteres, resulting in a four-haltered fly.

abd-A (abdominal-A)

In Drosophila, abdominal-A (abd-A) is expressed along most of the abdomen, from abdominal segment 1 (A1) to A8. Expression of abdominal-A is necessary to specify the identity of most of the abdominal segments. A major function of abd-A in insects is to repress limb formation. In abd-A loss-of-function mutants, abdominal segments A2 through A8 are transformed into an identity more like A1. When abd-A is ectopically expressed throughout the embryo, all segments anterior of A4 are transformed to an A4-like abdominal identity.[7] The abd-A gene also affects the pattern of cuticle generation in the ectoderm, and pattern of muscle generation in the mesoderm.[8]

Abd-B (Abdominal-B)

Abd-B is transcribed in two different forms, a regulatory protein, and a morphogenic protein. Regulatory Abd-B suppress embryonic ventral epidermal structures in the 8th and 9th segment of the Drosophila abdomen. Both the regulatory protein, and the morphogenic protein are involved in the development of the tail segment.[8]

Classification of Hox proteins

Proteins with a high degree of sequence similarity are also generally assumed to exhibit a high degree of functional similarity, i.e. Hox proteins with identical homeodomains are assumed to have identical DNA-binding properties (unless additional sequences are known to influence DNA-binding). To identify the set of proteins between two different species that are most likely to be most similar in function, classification schemes are used. For Hox proteins, three different classification schemes exist: phylogenetic inference based, synteny based, and sequence similarity based.[11] The three classification schemes provide conflicting information for Hox proteins expressed in the middle of the body axis (Hox6-8 and Antp, Ubx and Abd-A). A combined approach used phylogenetic inference based information of the different species and plotted the protein sequence types onto the phylogenetic tree of the species. The approach identified the proteins that best represent ancestral forms (Hox7 and Antp) and the proteins that represent new, derived versions (or were lost in an ancestor and are now missing in numerous species).[12]

Genes regulated by Hox proteins

Hox genes act at many levels within developmental gene hierarchies: at the "executive" level they regulate genes that in turn regulate large networks of other genes (like the gene pathway that forms an appendage). They also directly regulate what are called realisator genes or effector genes that act at the bottom of such hierarchies to ultimately form the tissues, structures, and organs of each segment. Segmentation involves such processes as morphogenesis (differentiation of precursor cells into their terminal specialized cells), the tight association of groups of cells with similar fates, the sculpting of structures and segment boundaries via programmed cell death, and the movement of cells from where they are first born to where they will ultimately function, so it is not surprising that the target genes of Hox genes promote cell division, cell adhesion, apoptosis, and cell migration.[13]

Examples of targets
Organism Target gene Normal function of target gene Regulated by
Drosophila distal-less activates gene pathway for limb formation ULTRABITHORAX[14]
(represses distal-less)
distal-less activates gene pathway for limb formation ABDOMINAL-A[14]
(represses distal-less)
decapentaplegic triggers cell shape changes in the gut that are
required for normal visceral morphology
ULTRABITHORAX[15]
(activates decapentaplegic)
reaper Apoptosis: localized cell death creates the segmental
boundary between the maxilla and mandible of the head
DEFORMED[16]
(activates reaper)
decapentaplegic prevents the above cell changes in more posterior
positions
ABDOMINAL-B[15]
(represses decapentaplegic)
Mouse EphA7 Cell adhesion: causes tight association of cells in
distal limb that will form digit, carpal and tarsal bones
HOX-A13[13]
(activates EphA7)
Cdkn1a Cell cycle: differentiation of myelomonocyte cells into
monocytes (white blood cells), with cell cycle arrest
Hox-A10[17]
(activates Cdkn1a)

Enhancer sequences that are bound by homeodomains

The DNA sequence that is bound by the homeodomain protein contains the nucleotide sequence TAAT, with the 5' terminal T being the most important for binding.[18] This sequence is conserved in nearly all sites recognized by homeodomains, and probably distinguishes such locations as DNA binding sites. The base pairs following this initial sequence are used to distinguish between homeodomain proteins, all of which have similar recognition sites. For instance, the nucleotide following the TAAT sequence is recognized by the amino acid at position 9 of the homeodomain protein. In the maternal protein Bicoid, this position is occupied by lysine, which recognizes and binds to the nucleotide guanine. In Antennapedia, this position is occupied by glutamine, which recognizes and binds to adenine. If the lysine in Bicoid is replaced by glutamine, the resulting protein will recognize Antennapedia-binding enhancer sites.[19]

However, all homeodomain-containing transcription factors bind essentially the same DNA sequence. The sequence bound by the homeodomain of a Hox protein is only 6 nucleotides long, and such a short sequence would be found at random many times throughout the genome, far more than the number of actual functional sites. Especially for Hox proteins, which produce such dramatic changes in morphology when misexpressed, this raises the question of how each transcription factor can produce such specific and different outcomes if they all bind the same sequence. One mechanism that introduces greater DNA sequence specificity to Hox proteins is to bind protein cofactors. Two such Hox cofactors are Extradenticle (Exd) and Homothorax (Hth). Exd and Hth bind to Hox proteins and appear to induce conformational changes in the Hox protein that increase its specificity.[20]

Regulation of Hox genes

Just as Hox genes regulate realisator genes, they are in turn regulated themselves by gap genes and pair-rule genes, which are in their turn regulated by maternally-supplied mRNA. This results in a transcription factor cascade: maternal factors activate gap or pair-rule genes; gap and pair-rule genes activate Hox genes; then, finally, Hox genes activate realisator genes that cause the segments in the developing embryo to differentiate. Regulation is achieved via protein concentration gradients, called morphogenic fields. For example, high concentrations of one maternal protein and low concentrations of others will turn on a specific set of gap or pair-rule genes. In flies, stripe 2 in the embryo is activated by the maternal proteins Bicoid and Hunchback, but repressed by the gap proteins Giant and Kruppel. Thus, stripe 2 will only form wherever there is Bicoid and Hunchback, but not where there is Giant and Kruppel.[21]

MicroRNA strands located in hox clusters have been shown to inhibit more anterior hox genes ("posterior prevalence phenomenon"), possibly to better fine tune its expression pattern.[22]

Non-coding RNA (ncRNA) has been shown to be abundant in Hox clusters. In humans, 231 ncRNA may be present. One of these, HOTAIR, silences in trans (it is transcribed from the HOXC cluster and inhibits late HOXD genes) by binding to Polycomb-group proteins (PRC2).[23]

The chromatin structure is essential for transcription but it also requires the cluster to loop out of the chromosomal territory.[24]

In higher animals including humans, retinoic acid regulates differential expression of Hox genes along the anteroposterior axis.[25] Genes in the 3' ends of Hox clusters are induced by retinoic acid resulting in expression domains that extend more anteriorly in the body compared to 5' Hox genes that are not induced by retinoic acid resulting in expression domains that remain more posterior.

Quantitative PCR has shown several trends regarding colinearity: the system is in equilibrium and the total number of transcripts depends on the number of genes present according to a linear relationship.[26]

Collinearity of Hox genes

In some organisms, especially vertebrates, the various Hox genes are situated very close to one another on the chromosome in groups or clusters. Interestingly, the order of the genes on the chromosome is the same as the expression of the genes in the developing embryo, with the first gene being expressed in the anterior end of the developing organism. The reason for this colinearity is not yet completely understood. The diagram above shows the relationship between the genes and protein expression in flies.

Hox nomenclature

The Hox genes are named for the homeotic phenotypes that result when their function is disrupted, wherein one segment develops with the identity of another (e.g. legs where antenna should be). Hox genes in different phyla have been given different names, which has led to confusion about nomenclature. The complement of Hox genes in Drosophila is made up of two clusters, the Antennapedia complex and the Bithorax complex, which together were historically referred to as the HOM-C (for Homeotic Complex). Although historically HOM-C genes have referred to Drosophila homologues, while Hox genes referred to vertebrate homologues, this distinction is no longer made, and both HOM-C and Hox genes are called Hox genes.

Human genes

Humans have Hox genes in four clusters:
cluster chromosome genes
HOXA chromosome 7 HOXA1, HOXA2, HOXA3, HOXA4, HOXA5, HOXA6, HOXA7, HOXA9, HOXA10, HOXA11, HOXA13
HOXB chromosome 17 HOXB1, HOXB2, HOXB3, HOXB4, HOXB5, HOXB6, HOXB7, HOXB8, HOXB9, HOXB13
HOXC chromosome 12 HOXC4, HOXC5, HOXC6, HOXC8, HOXC9, HOXC10, HOXC11, HOXC12, HOXC13
HOXD chromosome 2 HOXD1, HOXD3, HOXD4, HOXD8, HOXD9, HOXD10, HOXD11, HOXD12, HOXD13

History

The Hox genes are so named because mutations in them cause homeotic transformations. Homeotic transformations were first identified and studied by William Bateson in 1894, who coined the term "homeosis". After the rediscovery of Mendel's genetic principles, Bateson and others realized that some examples of homeosis in floral organs and animal skeletons could be attributed to variation in genes.

Definitive evidence for a genetic basis of some homeotic transformations was obtained by isolating homeotic mutants. The first homeotic mutant was found by Calvin Bridges in Thomas Hunt Morgan's laboratory in 1915. This mutant shows a partial duplication of the thorax and was therefore named Bithorax (bx). It transforms the third thoracic segment (T3) toward the second (T2). Bithorax arose spontaneously in the laboratory and has been maintained continuously as a laboratory stock ever since.[27]

The genetic studies by Morgan and others provided the foundation for the systematic analyses of Edward B. Lewis and Thomas Kaufman, which provided preliminary definitions of the many homeotic genes of the Bithorax and Antennapedia complexes, and also showed that the mutant phenotypes for most of these genes could be traced back to patterning defects in the embryonic body plan.

Ed Lewis, Christiane Nüsslein-Volhard and Eric F. Wieschaus identified and classified 15 genes of key importance in determining the body plan and the formation of body segments of the fruit fly Drosophila melanogaster.[when?][citation needed] For their work, Lewis, Nüsslein-Volhard, and Wieschaus were awarded the Nobel Prize in Physiology or Medicine in 1995.

In 1983, the homeobox was discovered independently by researchers in two labs: Ernst Hafen, Michael Levine, and William McGinnis (in Walter Gehring's lab at the University of Basel, Switzerland) and Matthew P. Scott and Amy Weiner (in Thomas Kaufman's lab at Indiana University in Bloomington).

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