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Xylem
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Schematic cross section of part of a leaf, xylem shown as red circles at figure 8
Xylem is one of the two types of transport tissue in vascular plants, phloem being the other. The basic function of xylem is to transport water from roots to shoots and leaves, but it also transports nutrients.[1][2] The word "xylem" is derived from the Greek word ξύλον (xylon), meaning "wood"; the best-known xylem tissue is wood, though it is found throughout the plant.[3] The term was introduced by Carl Nägeli in 1858.[4][5]
Contents
[hide]
1 Structure
2 Primary and secondary xylem
3 Main function – upwards water transport
3.1 Cohesion-tension theory
3.2 Measurement of pressure
4 Evolution
5 Development
5.1 Protoxylem and metaxylem
5.2 Patterns of protoxylem and metaxylem
6 See also
7 References
7.1 General references
Structure[edit]
Cross section of some xylem cells
The most distinctive xylem cells are the long tracheary elements that transport water. Tracheids and vessel elements are distinguished by their shape; vessel elements are shorter, and are connected together into long tubes that are called vessels.[6]
Xylem also contains two other cell types: parenchyma and fibers.[7]
Xylem can be found:
in vascular bundles, present in non-woody plants and non-woody parts of woody plants
in secondary xylem, laid down by a meristem called the vascular cambium in woody plants
as part of a stelar arrangement not divided into bundles, as in many ferns.
In transitional stages of plants with secondary growth, the first two categories are not mutually exclusive, although usually a vascular bundle will contain primary xylem only.
The branching pattern exhibited by xylem follows Murray's law.[8]
Primary and secondary xylem[edit]
Primary xylem is formed during primary growth from procambium. It includes protoxylem and metaxylem. Metaxylem develops after the protoxylem but before secondary xylem. Metaxylem has wider vessels and tracheids than protoxylem.
Secondary xylem is formed during secondary growth from vascular cambium. Although secondary xylem is also found in members of the gymnosperm groups Gnetophyta and Ginkgophyta and to a lesser extent in members of the Cycadophyta, the two main groups in which secondary xylem can be found are:
conifers (Coniferae): there are some six hundred species of conifers. All species have secondary xylem, which is relatively uniform in structure throughout this group. Many conifers become tall trees: the secondary xylem of such trees is used and marketed as softwood.
angiosperms (Angiospermae): there are some quarter of a million to four hundred thousand species of angiosperms. Within this group secondary xylem is rare in the monocots.[9] Many non-monocot angiosperms become trees, and the secondary xylem of these is used and marketed as hardwood.
Main function – upwards water transport[edit]
The xylem, vessels and tracheids of the roots, stems and leaves are interconnected to form a continuous system of water conducting channels reaching all parts of the plants. It transports water and soluble mineral nutrients from the roots throughout the plant. It is also used to replace water lost during transpiration and photosynthesis. Xylem sap consists mainly of water and inorganic ions, although it can contain a number of organic chemicals as well. The transport is passive, not powered by energy spent by the tracheary elements themselves, which are dead by maturity and no longer have living contents. Transporting sap upwards becomes more difficult as the height of a plant increases and upwards transport of water by xylem is considered to limit the maximum height of trees.[10] Three phenomena cause xylem sap to flow:
Pressure flow hypothesis: Sugars produced in the leaves and other green tissues are kept in the phloem system, creating a solute pressure differential versus the xylem system carrying a far lower load of solutes- water and minerals. The phloem pressure can rise to several MPa,[11] far higher than atmospheric pressure. Selective inter-connection between these systems allows this high solute concentration in the phloem to draw xylem fluid upwards by negative pressure.
Transpirational pull: Similarly, the evaporation of water from the surfaces of mesophyll cells to the atmosphere also creates a negative pressure at the top of a plant. This causes millions of minute menisci to form in the mesophyll cell wall. The resulting surface tension causes a negative pressure or tension in the xylem that pulls the water from the roots and soil.
Root pressure: If the water potential of the root cells is more negative than that of the soil, usually due to high concentrations of solute, water can move by osmosis into the root from the soil. This causes a positive pressure that forces sap up the xylem towards the leaves. In some circumstances, the sap will be forced from the leaf through a hydathode in a phenomenon known as guttation. Root pressure is highest in the morning before the stomata open and allow transpiration to begin. Different plant species can have different root pressures even in a similar environment; examples include up to 145 kPa in Vitis riparia but around zero in Celastrus orbiculatus.[12]
The primary force that creates the capillary action movement of water upwards in plants is the adhesion between the water and the surface of the xylem conduits.[13][14] Capillary action provides the force that establishes an equilibrium configuration, balancing gravity. When transpiration removes water at the top, the flow is needed to return to the equilibrium.
Transpirational pull results from the evaporation of water from the surfaces of cells in the leaves. This evaporation causes the surface of the water to recess into the pores of the cell wall. By capillary action, the water forms concave menisci inside the pores. The high surface tension of water pulls the concavity outwards, generating enough force to lift water as high as a hundred meters from ground level to a tree's highest branches.
Transpirational pull requires that the vessels transporting the water be very small in diameter; otherwise, cavitation would break the water column. And as water evaporates from leaves, more is drawn up through the plant to replace it. When the water pressure within the xylem reaches extreme levels due to low water input from the roots (if, for example, the soil is dry), then the gases come out of solution and form a bubble – an embolism forms, which will spread quickly to other adjacent cells, unless bordered pits are present (these have a plug-like structure called a torus, that seals off the opening between adjacent cells and stops the embolism from spreading).
Cohesion-tension theory[edit]
The cohesion-tension theory is a theory of intermolecular attraction that explains the process of water flow upwards (against the force of gravity) through the xylem of plants. It was proposed in 1894 by John Joly and Henry Horatio Dixon.[15] Despite numerous objections,[16][17] this is the most widely accepted theory for the transport of water through a plant's vascular system based on the classical research of Dixon-Joly (1894), Askenasy (1895),[18] and Dixon (1914,1924).[19][20]
Water is a polar molecule. When two water molecules approach one another, the slightly negatively charged oxygen atom of one forms a hydrogen bond with a slightly positively charged hydrogen atom in the other. This attractive force, along with other intermolecular forces, is one of the principal factors responsible for the occurrence of surface tension in liquid water. It also allows plants to draw water from the root through the xylem to the leaf.
Water is constantly lost through transpiration from the leaf. When one water molecule is lost another is pulled along by the processes of cohesion and tension. Transpiration pull, utilizing capillary action and the inherent surface tension of water, is the primary mechanism of water movement in plants. However, it is not the only mechanism involved. Any use of water in leaves forces water to move into them.
Transpiration in leaves creates tension (differential pressure) in the cell walls of mesophyll cells. Because of this tension, water is being pulled up from the roots into the leaves, helped by cohesion (the pull between individual water molecules, due to hydrogen bonds) and adhesion (the stickiness between water molecules and the hydrophilic cell walls of plants). This mechanism of water flow works because of water potential (water flows from high to low potential), and the rules of simple diffusion.[21]
Over the past century, there has been a great deal of research regarding the mechanism of xylem sap transport; today, most plant scientists continue to agree that the cohesion-tension theory best explains this process, but multiforce theories that hypothesize several alternative mechanisms have been suggested, including longitudinal cellular and xylem osmotic pressure gradients, axial potential gradients in the vessels, and gel- and gas-bubble-supported interfacial gradients.[22][23]
Measurement of pressure[edit]
A diagram showing the setup of a pressure bomb
Until recently, the differential pressure (suction) of transpirational pull could only be measured indirectly, by applying external pressure with a pressure bomb to counteract it.[24] When the technology to perform direct measurements with a pressure probe was developed, there was initially some doubt about whether the classic theory was correct, because some workers were unable to demonstrate negative pressures. More recent measurements do tend to validate the classic theory, for the most part. Xylem transport is driven by a combination[citation needed] of transpirational pull from above and root pressure from below, which makes the interpretation of measurements more complicated.
Evolution[edit]
Xylem appeared early in the history of terrestrial plant life. Fossil plants with anatomically preserved xylem are known from the Silurian (more than 400 million years ago), and trace fossils resembling individual xylem cells may be found in earlier Ordovician rocks. The earliest true and recognizable xylem consists of tracheids with a helical-annular reinforcing layer added to the cell wall. This is the only type of xylem found in the earliest vascular plants, and this type of cell continues to be found in the protoxylem (first-formed xylem) of all living groups of plants. Several groups of plants later developed pitted tracheid cells, it seems, through convergent evolution. In living plants, pitted tracheids do not appear in development until the maturation of the metaxylem (following the protoxylem).
In most plants, pitted tracheids function as the primary transport cells. The other type of tracheary element, besides the tracheid, is the vessel element. Vessel elements are joined by perforations into vessels. In vessels, water travels by bulk flow, as in a pipe, rather than by diffusion through cell membranes. The presence of vessels in xylem has been considered to be one of the key innovations that led to the success of the angiosperms.[25] However, the occurrence of vessel elements is not restricted to angiosperms, and they are absent in some archaic or "basal" lineages of the angiosperms: (e.g., Amborellaceae, Tetracentraceae, Trochodendraceae, and Winteraceae), and their secondary xylem is described by Arthur Cronquist as "primitively vesselless". Cronquist considered the vessels of Gnetum to be convergent with those of angiosperms.[26] Whether the absence of vessels in basal angiosperms is a primitive condition is contested, the alternative hypothesis states that vessel elements originated in a precursor to the angiosperms and were subsequently lost.
Photos showing xylem elements in the shoot of a fig tree (Ficus alba): crushed in hydrochloric acid, between slides and cover slips
To photosynthesize, plants must absorb CO2 from the atmosphere. However, this comes at a price: while stomata are open to allow CO2 to enter, water can evaporate.[27] Water is lost much faster than CO2 is absorbed, so plants need to replace it, and have developed systems to transport water from the moist soil to the site of photosynthesis.[27] Early plants sucked water between the walls of their cells, then evolved the ability to control water loss (and CO2 acquisition) through the use of stomata. Specialized water transport tissues soon evolved in the form of hydroids, tracheids, then secondary xylem, followed by an endodermis and ultimately vessels.[27]
The high CO2 levels of Silurian-Devonian times, when plants were first colonizing land, meant that the need for water was relatively low. As CO2 was withdrawn from the atmosphere by plants, more water was lost in its capture, and more elegant transport mechanisms evolved.[27] As water transport mechanisms, and waterproof cuticles, evolved, plants could survive without being continually covered by a film of water. This transition from poikilohydry to homoiohydry opened up new potential for colonization.[27] Plants then needed a robust internal structure that held long narrow channels for transporting water from the soil to all the different parts of the above-soil plant, especially to the parts where photosynthesis occurred.
During the Silurian, CO2 was readily available, so little water needed expending to acquire it. By the end of the Carboniferous, when CO2 levels had lowered to something approaching today's, around 17 times more water was lost per unit of CO2 uptake.[27] However, even in these "easy" early days, water was at a premium, and had to be transported to parts of the plant from the wet soil to avoid desiccation. This early water transport took advantage of the cohesion-tension mechanism inherent in water. Water has a tendency to diffuse to areas that are drier, and this process is accelerated when water can be wicked along a fabric with small spaces. In small passages, such as that between the plant cell walls (or in tracheids), a column of water behaves like rubber – when molecules evaporate from one end, they pull the molecules behind them along the channels. Therefore, transpiration alone provided the driving force for water transport in early plants.[27] However, without dedicated transport vessels, the cohesion-tension mechanism cannot transport water more than about 2 cm, severely limiting the size of the earliest plants.[27] This process demands a steady supply of water from one end, to maintain the chains; to avoid exhausting it, plants developed a waterproof cuticle. Early cuticle may not have had pores but did not cover the entire plant surface, so that gas exchange could continue.[27] However, dehydration at times was inevitable; early plants cope with this by having a lot of water stored between their cell walls, and when it comes to it sticking out the tough times by putting life "on hold" until more water is supplied.[27]
A banded tube from the late Silurian/early Devonian. The bands are difficult to see on this specimen, as an opaque carbonaceous coating conceals much of the tube. Bands are just visible in places on the left half of the image – click on the image for a larger view. Scale bar: 20 μm
To be free from the constraints of small size and constant moisture that the parenchymatic transport system inflicted, plants needed a more efficient water transport system. During the early Silurian, they developed specialized cells, which were lignified (or bore similar chemical compounds)[27] to avoid implosion; this process coincided with cell death, allowing their innards to be emptied and water to be passed through them.[27] These wider, dead, empty cells were a million times more conductive than the inter-cell method, giving the potential for transport over longer distances, and higher CO2 diffusion rates.
The earliest macrofossils to bear water-transport tubes are Silurian plants placed in the genus Cooksonia.[28] The early Devonian pretracheophytes Aglaophyton and Horneophyton have structures very similar to the hydroids of modern mosses. Plants continued to innovate new ways of reducing the resistance to flow within their cells, thereby increasing the efficiency of their water transport. Bands on the walls of tubes, in fact apparent from the early Silurian onwards,[29] are an early improvisation to aid the easy flow of water.[30] Banded tubes, as well as tubes with pitted ornamentation on their walls, were lignified[31] and, when they form single celled conduits, are considered to be tracheids. These, the "next generation" of transport cell design, have a more rigid structure than hydroids, allowing them to cope with higher levels of water pressure.[27] Tracheids may have a single evolutionary origin, possibly within the hornworts,[32] uniting all tracheophytes (but they may have evolved more than once).[27]
Water transport requires regulation, and dynamic control is provided by stomata.[33] By adjusting the amount of gas exchange, they can restrict the amount of water lost through transpiration. This is an important role where water supply is not constant, and indeed stomata appear to have evolved before tracheids, being present in the non-vascular hornworts.[27]
An endodermis probably evolved during the Silu-Devonian, but the first fossil evidence for such a structure is Carboniferous.[27] This structure in the roots covers the water transport tissue and regulates ion exchange (and prevents unwanted pathogens etc. from entering the water transport system). The endodermis can also provide an upwards pressure, forcing water out of the roots when transpiration is not enough of a driver.
Once plants had evolved this level of controlled water transport, they were truly homoiohydric, able to extract water from their environment through root-like organs rather than relying on a film of surface moisture, enabling them to grow to much greater size.[27] As a result of their independence from their surroundings, they lost their ability to survive desiccation – a costly trait to retain.[27]
During the Devonian, maximum xylem diameter increased with time, with the minimum diameter remaining pretty constant.[30] By the middle Devonian, the tracheid diameter of some plant lineages (Zosterophyllophytes) had plateaued.[30] Wider tracheids allow water to be transported faster, but the overall transport rate depends also on the overall cross-sectional area of the xylem bundle itself.[30] The increase in vascular bundle thickness further seems to correlate with the width of plant axes, and plant height; it is also closely related to the appearance of leaves[30] and increased stomatal density, both of which would increase the demand for water.[27]
While wider tracheids with robust walls make it possible to achieve higher water transport pressures, this increases the problem of cavitation.[27] Cavitation occurs when a bubble of air forms within a vessel, breaking the bonds between chains of water molecules and preventing them from pulling more water up with their cohesive tension. A tracheid, once cavitated, cannot have its embolism removed and return to service (except in a few advanced angiosperms[verification needed] which have developed a mechanism of doing so). Therefore, it is well worth plants' while to avoid cavitation occurring. For this reason, pits in tracheid walls have very small diameters, to prevent air entering and allowing bubbles to nucleate. Freeze-thaw cycles are a major cause of cavitation. Damage to a tracheid's wall almost inevitably leads to air leaking in and cavitation, hence the importance of many tracheids working in parallel.[27]
Cavitation is hard to avoid, but once it has occurred plants have a range of mechanisms to contain the damage.[27] Small pits link adjacent conduits to allow fluid to flow between them, but not air – although ironically these pits, which prevent the spread of embolisms, are also a major cause of them.[27] These pitted surfaces further reduce the flow of water through the xylem by as much as 30%.[27] Conifers, by the Jurassic, developed an ingenious improvement, using valve-like structures to isolate cavitated elements. These torus-margo structures have a blob floating in the middle of a donut; when one side depressurizes the blob is sucked into the torus and blocks further flow.[27] Other plants simply accept cavitation; for instance, oaks grow a ring of wide vessels at the start of each spring, none of which survive the winter frosts. Maples use root pressure each spring to force sap upwards from the roots, squeezing out any air bubbles.
Growing to height also employed another trait of tracheids – the support offered by their lignified walls. Defunct tracheids were retained to form a strong, woody stem, produced in most instances by a secondary xylem. However, in early plants, tracheids were too mechanically vulnerable, and retained a central position, with a layer of tough sclerenchyma on the outer rim of the stems.[27] Even when tracheids do take a structural role, they are supported by sclerenchymatic tissue.
Tracheids end with walls, which impose a great deal of resistance on flow;[30] vessel members have perforated end walls, and are arranged in series to operate as if they were one continuous vessel.[30] The function of end walls, which were the default state in the Devonian, was probably to avoid embolisms. An embolism is where an air bubble is created in a tracheid. This may happen as a result of freezing, or by gases dissolving out of solution. Once an embolism is formed, it usually cannot be removed (but see later); the affected cell cannot pull water up, and is rendered useless.
End walls excluded, the tracheids of prevascular plants were able to operate under the same hydraulic conductivity as those of the first vascular plant, Cooksonia.[30]
The size of tracheids is limited as they comprise a single cell; this limits their length, which in turn limits their maximum useful diameter to 80 μm.[27] Conductivity grows with the fourth power of diameter, so increased diameter has huge rewards; vessel elements, consisting of a number of cells, joined at their ends, overcame this limit and allowed larger tubes to form, reaching diameters of up to 500 μm, and lengths of up to 10 m.[27]
Vessels first evolved during the dry, low CO2 periods of the late Permian, in the horsetails, ferns and Selaginellales independently, and later appeared in the mid Cretaceous in angiosperms and gnetophytes.[27] Vessels allow the same cross-sectional area of wood to transport around a hundred times more water than tracheids![27] This allowed plants to fill more of their stems with structural fibers, and also opened a new niche to vines, which could transport water without being as thick as the tree they grew on.[27] Despite these advantages, tracheid-based wood is a lot lighter, thus cheaper to make, as vessels need to be much more reinforced to avoid cavitation.[27]
Development[edit]
Patterns of xylem development: xylem in brown; arrows show direction of development from protoxylem to metaxylem.
Xylem development can be described by four terms: centrarch, exarch, endarch and mesarch. As it develops in young plants, its nature changes from protoxylem to metaxylem (i.e. from first xylem to after xylem). The patterns in which protoxylem and metaxylem are arranged is important in the study of plant morphology.
Protoxylem and metaxylem[edit]
As a young vascular plant grows, one or more strands of primary xylem form in its stems and roots. The first xylem to develop is called 'protoxylem'. In appearance protoxylem is usually distinguished by narrower vessels formed of smaller cells. Some of these cells have walls which contain thickenings in the form of rings or helices. Functionally, protoxylem can extend: the cells are able to grow in size and develop while a stem or root is elongating. Later, 'metaxylem' develops in the strands of xylem. Metaxylem vessels and cells are usually larger; the cells have thickenings which are typically either in the form of ladderlike transverse bars (scalariform) or continuous sheets except for holes or pits (pitted). Functionally, metaxylem completes its development after elongation ceases when the cells no longer need to grow in size.[34][35]
Patterns of protoxylem and metaxylem[edit]
There are four main patterns to the arrangement of protoxylem and metaxylem in stems and roots.
Centrarch refers to the case in which the primary xylem forms a single cylinder in the center of the stem and develops from the center outwards. The protoxylem is thus found in the central core and the metaxylem in a cylinder around it.[36] This pattern was common in early land plants, such as "rhyniophytes", but is not present in any living plants.[citation needed]
The other three terms are used where there is more than one strand of primary xylem.
Exarch is used when there is more than one strand of primary xylem in a stem or root, and the xylem develops from the outside inwards towards the center, i.e. centripetally. The metaxylem is thus closest to the center of the stem or root and the protoxylem closest to the periphery. The roots of vascular plants are normally considered to have exarch development.[34]
Endarch is used when there is more than one strand of primary xylem in a stem or root, and the xylem develops from the inside outwards towards the periphery, i.e. centrifugally. The protoxylem is thus closest to the center of the stem or root and the metaxylem closest to the periphery. The stems of seed plants typically have endarch development.[34]
Mesarch is used when there is more than one strand of primary xylem in a stem or root, and the xylem develops from the middle of a strand in both directions. The metaxylem is thus on both the peripheral and central sides of the strand with the protoxylem between the metaxylem (possibly surrounded by it). The leaves and stems of many ferns have mesarch development.[34]
See also[edit]
Wikimedia Commons has media related to Xylem.
Phloem
Soil plant atmosphere continuum
Stele
Suction
Tylosis
Vascular bundle
Vascular tissue
Xylem sap
References[edit]
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Jump up ^ Stewart, W.N.; Rothwell, G.W. (1993). Paleobiology and the evolution of plants. Cambridge University Press.
^ Jump up to: a b c d Foster, A.S.; Gifford, E.M. (1974). Comparative Morphology of Vascular Plants (2nd ed.). San Francisco: W.H. Freeman. pp. 55–56. ISBN 978-0-7167-0712-7.
Jump up ^ Taylor, T.N.; Taylor, E.L.; Krings, M. (2009). Paleobotany, the Biology and Evolution of Fossil Plants (2nd ed.). Amsterdam; Boston: Academic Press. pp. 207ff., 212ff. ISBN 978-0-12-373972-8.
Jump up ^ White, A. Toby; Kazlev, M. Alan. "Glossary". palaeos.com. Archived from the original on December 20, 2010.
General references[edit]
C. Wei; E. Steudle; M. T. Tyree; P. M. Lintilhac (May 2001). "The essentials of direct xylem pressure measurement". Plant, Cell and Environment. 24 (5): 549–555. doi:10.1046/j.1365-3040.2001.00697.x. is the main source used for the paragraph on recent research.
N. Michele Holbrook; Michael J. Burns; Christopher B. Field (November 1995). "Negative Xylem Pressures in Plants: A Test of the Balancing Pressure Technique". Science. 270 (5239): 1193–4. Bibcode:1995Sci...270.1193H. doi:10.1126/science.270.5239.1193. is the first published independent test showing the Scholander bomb actually does measure the tension in the xylem.
Pockman, W.T.; J.S. Sperry; J.W. O'Leary (December 1995). "Sustained and significant negative water pressure in xylem". Nature. 378 (6558): 715–6. Bibcode:1995Natur.378..715P. doi:10.1038/378715a0. is the second published independent test showing the Scholander bomb actually does measure the tension in the xylem.
Campbell, Neil A.; Jane B. Reece (2002). Biology (6th ed.). Benjamin Cummings. ISBN 978-0-8053-6624-2.
Kenrick, Paul; Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution Press. ISBN 1-56098-730-8.
Muhammad, A.F.; R. Sattler (1982). "Vessel Structure of Gnetum and the Origin of Angiosperms". American Journal of Botany. Botanical Society of America. 69 (6): 1004–21. doi:10.2307/2442898. JSTOR 2442898.
Melvin T. Tyree; Martin H. Zimmermann (2003). Xylem Structure and the Ascent of Sap (2nd ed.). Springer. ISBN 3-540-43354-6. recent update of the classic book on xylem transport by the late Martin Zimmermann
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text:
The Hunter (video game)
From Wikipedia, the free encyclopedia
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The Hunter
Logo for The Hunter
Developer(s)
Emote Games
Expansive Worlds (remake)
Avalanche Studios (remake)
Publisher(s)
Emote Games
Expansive Worlds (remake)
Avalanche Studios (remake)
Platform(s)
Microsoft Windows
Release
WW: March 2009
WW: 3 June 2014 (remake)
Genre(s)
First-person hunting
Mode(s)
Single-player, multiplayer
Three elk in a wooded area
The Hunter, stylized as theHunter, is an online first-person hunting video game released in March 2009. The Hunter: Primal, a standalone expansion featuring dinosaurs, was launched in March 2015.
Contents
[hide]
1 Gameplay
1.1 Payment
2 Development
3 Reception
4 Expansion and sequel
5 Notes
6 External links
Gameplay[edit]
The Hunter is a first-person hunting game that recreates hunting wild animals in as realistic manner as possible. The initial setting is on an open world island based on various locations, including the islands and surroundings of Washington state in the United States, Australian Outback, French Alps and more. It features five huntable animal types whitetail deer, european rabbit, cottontail rabbit, snowshoe hare and pheasant. The game is fully playable for no charge, but the game's full ability can be only unlocked with paid membership which allows to hunt all species and provides replenishing ammo for the obtained weapons.
To play the player must first create a character on the game's website, which includes social networking tools and messaging capabilities to allow players to connect and share in-game experiences. Creating a character requires selecting an avatar from pool of unique faces, and adding a unique name. Once a character is created and the tutorial completed, the player can hunt freely and activate missions and earn one of two types of in game currency called gm$. Completing the first 3 whitetail deer missions unlocks further missions from other animals in the game.
Players must use natural cover to avoid detection by the animals in the game. whitetail deer, European rabbit, cottontail rabbit, snowshoe hare and pheasant may be hunted for free, while mule deer, blacktail, Roosevelt elk, turkey, coyote, feral hog, black bear, moose, European wild boar, roe deer, red deer, red fox, brown bear, mallard, Canada goose, reindeer, alpine ibex, red kangaroo, bison, Sitka deer, snowshoe hare, gadwall, northern pintail, American black duck, polar bear, willow ptarmigan, rock ptarmigan, white-tailed ptarmigan, Big Horn Sheep, Rocky Mountain Elk and Grizzly Bear can be hunted by subscribers, or free users via the Free Rotation function added some time in 2016, Each animal species reacts to different stimuli in different ways. Scent detection for the quadrupeds is their keenest sense, which means the player has to be aware of wind direction or the animal might smell the player's scent and run off. Players must also pay careful attention to the soundscape, listening for subtle signals that indicate the presence of animals in the locality.[1]
The core gameplay revolves around tracking and harvesting (killing) animals with the weapons provided, although a digital camera is also provided for those who do not wish to shoot with a gun or a bow. Animal tracks can be found and identified with the "Huntermate" an in-game GPS-like device that identifies tracks, scat, rooting, bedding, blood trails, and animal calls and grunts.
In order to hunt certain species, The Hunter requires the player to utilize the correct weapon for each hunt. All bows are considered ethical for all species while rifles, guns and shotguns of different caliber must be selected according to the animal the player will hunt. Players that break this rule will not be given credit for their efforts and will receive a stern email from Doc the warden of all the Evergreen Hunting Reserves.
Hunting areas added since May 2011 are: Settler Creeks (rugged North American mountain forest with abandoned homesteads); Redfeather Falls (rough Canadian mountains and swamps); Hirschfelden (German highland farms and forests); Hemmeldal (Northern Swedish snowy mountains, swamps and icy lakes), which is also the first reserve to allow hunting during snowfall; Rougarou Bayou (Southern United States); Val des Bois (rugged mountains, meadows and woods in the Alps), Bushranger's Run (dry, rocky and sandy Australian Outback), and Whiterime Ridge (chilly arctic landscape surrounded by high mountains and icefields),Timbergold Trails (Rocky Mountains Inspired Reserve). Subscribers can choose to start anywhere in these hunting areas (provided they have a tent, otherwise a hunting lodge can be chosen to start in a list of two or three in each reserve), and at any time of day, between 5 AM and 6 PM.
Whitetail deer can be both typical or non-typical since the update in January, 2015 which introduced this feature.
Free-to-play players were initially restricted at launch to Whitehart Island. Then from May 2011 a large 2.3 square-mile spur, called Logger's Point, was made the free-to-play starting point. Access was barred from there to the main Whitehart Island, unless one had a paid subscription. As of 3 June 2013 all hunting reserves can be visited and played by free-to-play users of the game, but no animal can be shot without a license apart from whitetail deer, european rabbit, cottontail rabbit, snowshoe hare and pheasant.
The Hunter also features minor RPG elements. As players track and spot each animal their skill in that area improves. Weapon skills can also be improved by scoring successful hits on animals. Improving the hunting skills makes animals easier to track, and a weapon will not suffer as much "wobble" when aimed.
Skills, missions, and hunting statistics are recorded by the game and then displayed on the player's profile page on the website, although they cannot be seen while hunting in the game client.
Multiplayer was released on 3 June 2013 and allows full members both to host and to join maps with up to 8 players. Free members can only join multiplayer if the host allows it. A new set of animations has been also released, together with a new inventory mode, new clothes and some minor changes. Even the website has undergone a full restyle.
In July 2013 the red fox was released, followed by mallard ducks in the Rougarou Bayou swampy reserve.
Canada goose was the next on the list, on 27 March 2014, making Hirschfelden the first ever reserve with six species and the first ever with two species of game birds as well.
In September 2014 the alpine ibex was added on the brand new Val des Bois alpine reserve, as the first prey of the goat family. The reserve marks the release of the first new in-game activity: simplified climbing. Two species of rodents, cottontail rabbit and European rabbit followed in a few months. The following and first ever all-dry reserve is dedicated to the hot, dry outback of western Australia with a new species, red kangaroo.
Dogs were added for the retrieval of ducks during waterfowling and Canada goose hunting.
In 2015 the American buffalo, Sitka deer, Arctic fox, polar bear and snowshoe hare were added to Whiterime Ridge Reserve.This new reserve introduced extreme cold climate conditions that require the player to wear special protection against the elements or suffer hypothermia.
In November 2015, other 3 waterfowl bird in the Rougarou Bayou swampy reserve.
Some time in 2016 They added an animal Rotation which allows for players to hunt some other animals such as Mule Deer, Black Bears, and every other animal on the game via a Free Rotation.
Payment[edit]
The game is free-to-play, but licenses must be purchased to hunt any animal other than whitetail deer, european rabbit, cottontail rabbit, snowshoe hare and pheasant, but each week there are random animals that as a free player you can shoot as well and are featured on the front page of the Expansive Worlds web site. Any additional guns, ammo, and equipment can be purchased with in-game currency. Memberships allow the player to hunt all species and receive unlimited ammo for a defined period (3, 6 or 12 months according to the price).
Membership types each grant different set of weapons, and weapons can be bought in the store among many other things that makes the hunt more sophisticated. There are strict rules which weapon can be used to hunt which species, so buying different guns are needed in the long run to hunt all the species.
Development[edit]
The Hunter was first developed by UK-based Emote Games, in conjunction with the Swedish Avalanche Studios. On 18 February 2010, Avalanche Studios announced the acquisition of the full rights to The Hunter intellectual property previously owned by Emote Games.[2] In relation to the acquisition the new studio, Expansive Worlds was formed that will act as the online games division of Avalanche and continue the development of The Hunter.[3]
The Hunter uses Avalanche Studio's proprietary Avalanche Engine 2[4] which uses procedurally-generated content, and can thus pack a large and sophisticated game world into a relatively small download (which is 1.47Gb as of December 2013). The advanced engine enables, subtle lighting and shading, and natural weather effects.
Reception[edit]
Reception
Aggregate score
Aggregator
Score
GameRankings 79.75%[5]
Review score
Publication
Score
GamePro 4.5/5[6]
The Hunter gathered positive reviews during its launch period. However, it received "Mostly Negative" reviews on Steam due to the inability to hunt different kinds of animals without an in-game hunting license. A player can buy hunting licenses with in-game currency or real money to get a membership. The standalone expansion received a "Mixed" review for the same reason. Metacritic gave The Hunter: Primal a score of 49 out of 100.
Expansion and sequel[edit]
theHunter: Primal, a standalone expansion featuring dinosaurs, was released on Steam's early access on 15 December 2014, and fully launched on 31 March 2015.[7][8]
A sequel titled, theHunter: Call of the Wild, was released on Steam in February 2017.[9]
Notes[edit]
Jump up ^ Grimm, Michael (20 April 2009). "The Hunter Review". GamesRadar. Retrieved 22 April 2014.
Jump up ^ The Hunter Community Update as of 2010-02-18, http://www.thehunter.com/pub/newsarchive.php?id=2010-02-18
Jump up ^ New Swedish development studio formed by Avalanche Studios, http://www.avalanchestudios.se/start/10-03-23/New_Swedish_development_studio_formed_by_Avalanche_Studios.aspx
Jump up ^ "Making of The Hunter – Avalanche Studios".
Jump up ^ GameRankings.com as of 2009-07-18, http://www.gamerankings.com/pc/954010-the-hunter/index.html
Jump up ^ GamePro review https://web.archive.org/web/20110607135228/http://www.gamepro.com/article/reviews/209305/the-hunter-page-1-of-2/
Jump up ^ Makuch, Eddie (15 December 2014). ""Frighteningly Realistic Dinosaur Hunter" Goes Live on Steam". GameSpot. Retrieved 9 July 2015.
Jump up ^ Makuch, Eddie (31 March 2015). "Just Cause Dev's Dinosaur-Hunting Game Available Now, Adds Eye-Tracking". GameSpot. Retrieved 9 July 2015.
Jump up ^ "theHunter: Call of the Wild™". Steam. Retrieved 2017-11-05.
External links[edit]
Official website
Official website for theHunter: Call Of The Wild
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Other games
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Warner Bros. Interactive Entertainment
Retrieved from "https://en.wikipedia.org/w/index.php?title=The_Hunter_(video_game)&oldid=825367527"
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United States House Committee on Science, Space, and Technology
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The Committee on Science, Space and Technology is a committee of the United States House of Representatives. It has jurisdiction over non-defense federal scientific research and development. Specifically, the committee has partial or complete jurisdiction over the following federal agencies: NASA, the Department of Energy, EPA, ATSDR, NSF, FAA, NOAA, NIST, FEMA, the U.S. Fire Administration, and USGS.
Contents
[hide]
1 History
2 Members
2.1 115th Congress
3 Subcommittees
4 Committee chairmen, 1959-present
5 See also
6 References
7 External links
History[edit]
In the wake of the Soviet Sputnik program in the late 1950s, Congress created the Select Committee on Astronautics and Space Exploration in 1958, chaired by majority leader John William McCormack. This select committee drafted the National Aeronautics and Space Act that created the National Aeronautics and Space Administration (NASA). A staff report of the committee, the Space Handbook: Astronautics and its Applications, provided non-technical information about spaceflight to U.S. policy makers.[1]
The committee also chartered the permanent House Committee on Science and Astronautics, which officially began on January 3, 1959, and was the first new standing committee established in the House since 1946. The name was changed in 1974 to the House Committee on Science and Technology. The name was changed again in 1987 to the House Committee on Science, Space and Technology. After the Republican Party gained a majority in Congress in 1994, the name of the committee was changed to the House Committee on Science. With the return of control to the Democrats in 2007, the committee's name was changed back to the House Committee on Science and Technology.
In the 112th Congress, Committee Chairman Ralph Hall added “Space” back into the committee’s name: “The Committee on Science, Space, and Technology” – a nod to the committee’s history, broad jurisdiction, and the importance of space exploration in maintaining American innovation and competitiveness.[2]
On December 1, 2016, the committee's Twitter account posted a link to an article on Breitbart which argued that climate change was the result of natural weather processes.[3] The tweet was criticized by members of the scientific community on Twitter for promoting an unscientific and misleading article.[4] The Committee's Ranking Member Eddie Bernice Johnson also criticized the tweet, writing, "False news & false facts put us all in danger."[5]
Members[edit]
115th Congress[edit]
Majority[6]
Minority[7]
Lamar Smith, Texas, Chair
Dana Rohrabacher, California
Frank Lucas, Oklahoma, Vice Chair
Mo Brooks, Alabama
Randy Hultgren, Illinois
Bill Posey, Florida
Thomas Massie, Kentucky
Jim Bridenstine, Oklahoma
Randy Weber, Texas
Steve Knight, California
Brian Babin, Texas
Barbara Comstock, Virginia
Gary Palmer, Alabama
Barry Loudermilk, Georgia
Ralph Abraham, Louisiana
Darin LaHood, Illinois
Daniel Webster, Florida
Jim Banks, Indiana
Andy Biggs, Arizona
Roger Marshall, Kansas
Neal Dunn, Florida
Clay Higgins, Louisiana
Ralph Norman, South Carolina
Eddie Bernice Johnson, Texas, Ranking Member
Zoe Lofgren, California
Dan Lipinski, Illinois
Suzanne Bonamici, Oregon
Ami Bera, California
Elizabeth Esty, Connecticut
Marc Veasey, Texas
Don Beyer, Virginia, Vice Ranking Member
Jacky Rosen, Nevada
Jerry McNerney, California
Ed Perlmutter, Colorado
Paul Tonko, New York
Bill Foster, Illinois
Mark Takano, California
Colleen Hanabusa, Hawaii
Charlie Crist, Florida
Subcommittees[edit]
There are five subcommittees in the 115th Congress.
Subcommittee
Chair[8]
Ranking Member
Energy Randy Weber (R-TX) Marc Veasey (D-TX)
Environment Andy Biggs (R-AZ) Suzanne Bonamici (D-OR)
Oversight Darin Lahood (R-IL) Don Beyer (D-VA)
Research and Technology Barbara Comstock (R-VA) Dan Lipinski (D-IL)
Space Brian Babin (R-TX) Ami Bera (D-CA)
Committee chairmen, 1959-present[edit]
Chairmen since 1959.[2]
Overton Brooks, 1959–1961
George P. Miller, 1961–1973
Olin E. Teague, 1973–1978
Don Fuqua, 1979–1987
Robert A. Roe, 1987–1991
George Brown, Jr., 1991–1995
Robert S. Walker, 1995–1997
Jim Sensenbrenner, 1997–2001
Sherwood Boehlert, 2001–2007
Bart Gordon, 2007–2011
Ralph Hall, 2011–2013
Lamar Smith, 2013–present
See also[edit]
List of current United States House of Representatives committees
References[edit]
Jump up ^ "Space Handbook: Astronautics and its Applications". NASA.
^ Jump up to: a b "A BRIEF HISTORY OF THE COMMITTEE ON SCIENCE & TECHNOLOGY" (PDF). United States Government. November 7, 2007.
Jump up ^ Feltman, Rachel (December 1, 2016). "The House Committee on Science just tweeted a 'science' article from Breitbart". Popular Science. Retrieved December 2, 2016.
Jump up ^ "It's likely Earth's hottest year on record — and some people are talking about global cooling". Washington Post. Retrieved 2016-12-15.
Jump up ^ McCausland, Phil (December 1, 2016). "House Science Committee Tweets Climate-Change Denying Breitbart Article, Debunked by Scientists". NBC News. Retrieved December 2, 2016.
Jump up ^ H.Res. 6, H.Res. 51
Jump up ^ H.Res. 7, H.Res. 45, H.Res. 52, H.Res. 95
Jump up ^ Subcommittee assignments
Bush, Daniel (January 14, 2015). "New leaders named for energy, enviro subcommittees". Energy & Environment Daily. Environment & Energy Publishing. Retrieved January 14, 2015.
"Smith Announces Subcommittee Chairmen and Vice Chairmen for the 114th Congress". Committee on Science, Space and Technology. January 13, 2015. Retrieved January 14, 2015.
External links[edit]
Official web site (Archive)
House Science, Space, and Technology Committee. Legislation activity and reports, Congress.gov.
Republican Science Committee website
Democratic Science Committee website
A History of the Committee on Science and Technology: 85th-110th Congresses 1958-2008
Works by United States House Committee on Science, Space, and Technology at Project Gutenberg
Works by or about United States House Committee on Science, Space, and Technology at Internet Archive
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Chairmen of the United States House Committee on Science, Space, and Technology
Brooks
Miller
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Fuqua
Roe
Brown
Walker
Sensenbrenner
Boehlert
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Hall
Smith
Authority control
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Sail Away (David Gray song)
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"Sail Away"
Single by David Gray
from the album White Ladder
Released
16 July 2001
Format
CD, DVD, 12"
Genre
Folk rock
Label
IHT, EastWest
Songwriter(s)
David Gray
Producer(s)
David Gray, Marius de Vries
David Gray singles chronology
"This Year's Love"
(2001) "Sail Away"
(2001) "Say Hello Wave Goodbye"
(2001)
"This Year's Love"
(re-issue)
(2001) "Sail Away"
(2001) "Say Hello Wave Goodbye"
(2001)
"Sail Away" is a song by David Gray. It was released as the fourth single from White Ladder on 16 July 2001. Remixes by Rae and Christian and Biffco were also commissioned and featured on the single formats. The DVD single features live video footage taken from his concert DVD release David Gray: Live. The song is featured in the movie 15 Minutes.
The song is often played on the Expedia phone service while on hold,[citation needed] due to its message of escape.[citation needed]
Sail Away has been covered by Norwegian singer/songwriter Alejandro Fuentes, which was performed at the Hallelujah concerts.
Contents
[hide]
1 Track listings
2 Personnel
3 References
4 External links
Track listings[edit]
CD single[1]
"Sail Away" (Biffco Radio Edit) – 3:44
"Sail Away" (Biffco Club Mix) – 7:02
"Sail Away" (Rae and Christian Remix) – 5:35
DVD single[2]
"Sail Away" (Live at The Point) (Video) – 5:20
"Sail Away" (Biffco Radio Edit) (Audio) – 3:44
"This Year's Love" (Live at The Point) (Video Excerpt) – 0:30
"Please Forgive Me" (Live at The Point) (Video Excerpt) – 0:30
"Wisdom" (Live at The Point) (Video Excerpt) – 0:30
Promo 12"[3]
"Sail Away" (Rae and Christian Remix) – 5:35
"Sail Away" (Rae and Christian Instrumental) – 5:24
Personnel[edit]
Voice, guitar, piano, keyboard: David Gray
Drums, keyboard, bass, voice: Craig McClune
Programmers: Iestyn Polson, Marius de Vries, and Steve Sidylenk
References[edit]
Jump up ^ David Gray - Sail Away at Discogs
Jump up ^ David Gray - Sail Away (DVD) at Discogs
Jump up ^ David Gray - Sail Away (Rae & Christian Remixes) (Vinyl) at Discogs
External links[edit]
Lyrics of this song at MetroLyrics
[hide]
v
t
e
David Gray
Studio albums
A Century Ends
Flesh
Sell, Sell, Sell
White Ladder
Lost Songs 95–98
A New Day at Midnight
Life in Slow Motion
Draw the Line
Foundling
Mutineers
Compilations
The EPs 1992–1994
Shine: The Best of the Early Years
A Thousand Miles Behind
Greatest Hits
Singles
"Birds Without Wings"
"Shine"
"Wisdom"
"Faster Sooner Now"
"Late Night Radio"
"This Year's Love"
"Babylon"
"Please Forgive Me"
"Sail Away"
"Say Hello Wave Goodbye"
"Dead in the Water"
"The Other Side"
"Be Mine"
"The One I Love"
"Hospital Food"
"Alibi"
"You're the World to Me"
"Fugitive"
"Full Steam"
"A Moment Changes Everything"
Videos
David Gray: Live
Live in Slow Motion
Other musicians
Tim Bradshaw
Rob Malone
Neil MacColl
Craig McClune
David Nolte
Keith Prior
Related articles
Discography
Music videos
This Year's Love
Retrieved from "https://en.wikipedia.org/w/index.php?title=Sail_Away_(David_Gray_song)&oldid=757745738"
Categories:
David Gray (musician) songs
2001 singles
Songs written by David Gray (musician)
1998 songs
Song recordings produced by Marius de Vries
East West Records singles
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This page was last edited on 1 January 2017, at 14:12.
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