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List of men's footballers with 500 or more goals
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In top-level association football, 27 players have scored 500 or more goals over the course of their career, according to research by the Rec.Sport.Soccer Statistics Foundation. Only players who were active in top division football for all or most of their careers are considered. The ranking below takes into account goals scored in official matches played with national teams at all age levels and clubs in all divisions.[1]
Ranking[edit]
Key
Unverified data is excluded from players' data; goal and match totals may be higher.
Players in bold are currently active; data is accurate as on 26 November 2017.[2][3]
Rank
Player
Goals
Matches
Ratio
Years
1 Bican, JosefJosef Bican (CZE) 805 530 1.52 1931–1956
2 Romário (BRA) 772 994 0.78 1985–2007
3 Pelé (BRA) 767 831 0.92 1957–1977
4 Puskás, FerencFerenc Puskás (HUN) 746 754 0.99 1943–1966
5 Müller, GerdGerd Müller (GER) 735 793 0.93 1962–1981
6 Ronaldo, CristianoCristiano Ronaldo (POR) 637 916 0.7 2002–present
7 Messi, LionelLionel Messi (ARG) 611 781 0.78 2004–present
8 Deák, FerencFerenc Deák (HUN) 576 — 1940–1957
9 Seeler, UweUwe Seeler (GER) 575 — 1953–1978
Maravilha, TúlioTúlio Maravilha (BRA) 575 — 1988–2012
11 Friedenreich, ArthurArthur Friedenreich (BRA) 558 — 1909–1935
12 Willimowski, ErnstErnst Willimowski (POL) 554 — 1934–1955
13 Eusébio (POR) 552 — 1957–1979
14 McGrory, JimmyJimmy McGrory (SCO) 550 — 1922–1937
15 Binder, FranzFranz Binder (AUT) 546 — 1949–1976
16 Peyroteo, FernandoFernando Peyroteo (POR) 544 — 1937–1949
17 Sánchez, HugoHugo Sánchez (MEX) 541 — 1976–1997
18 Walter, FritzFritz Walter (GER) 539 — 1937–1959
19 Takács, JózsefJózsef Takács (HUN) 523 — 1917–1934
20 Zsengellér, GyulaGyula Zsengellér (HUN) 522 — 1935–1953
Zico (BRA) 522 — 1971–1994
22 Di Stéfano, AlfredoAlfredo Di Stéfano (ARG) 514 — 1945–1966
Krankl, HansHans Krankl (AUT) 514 — 1970–1989
24 Nordahl, GunnarGunnar Nordahl (SWE) 513 — 1937–1958
25 Dinamite, RobertoRoberto Dinamite (BRA) 512 — 1971–1993
26 Greaves, JimmyJimmy Greaves (ENG) 511 — 1957–1980
27 Bene, FerencFerenc Bene (HUN) 508 — 1961–1978
Source: Rec.Sport.Soccer Statistics Foundation[1]
See also[edit]
List of men's footballers with 50 or more international goals
List of footballers with most goals in World Cup qualifiers
List of footballers with the most official appearances
References[edit]
^ Jump up to: a b Kolos, Vladimir (21 January 2016). "Prolific Scorers Data: Best Goalscorers All-Time (Official Matches)". Rec.Sport.Soccer Statistics Foundation. Retrieved 22 March 2016.
Jump up ^ "Matches of Cristiano Ronaldo". Soccerway. Retrieved 16 April 2016.
Jump up ^ "Matches of L. Messi". Soccerway. Retrieved 16 April 2016.
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Lava dome
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Image of the rhyolitic lava dome of Chaitén Volcano during its 2008–2010 eruption.
One of the Mono Craters, an example of a rhyolite dome.
In volcanology, a lava dome or volcanic dome is a roughly circular mound-shaped protrusion resulting from the slow extrusion of viscous lava from a volcano. The geochemistry of lava domes can vary from basalt to rhyolite although most preserved domes tend to have high silica content.[1] The characteristic dome shape is attributed to high viscosity that prevents the lava from flowing very far. This high viscosity can be obtained in two ways: by high levels of silica in the magma, or by degassing of fluid magma. Since viscous basaltic and andesitic domes weather fast and easily break apart by further input of fluid lava, most of the preserved domes have high silica content and consist of rhyolite or dacite.
Existence of lava domes has been suggested for some domed structures on the Martian surface in the western part of Arcadia Planitia and within Terra Sirenum.[2][3]
Contents
[hide]
1 Dome dynamics
2 Related landforms
2.1 Cryptodomes
2.2 Lava coulées
3 Examples of lava domes
4 References
5 External links
Dome dynamics[edit]
Lava domes in the crater of Mount St. Helens
Lava domes evolve unpredictably, due to non-linear dynamics caused by crystallization and outgassing of the highly viscous lava in the dome's conduit.[4] Domes undergo various processes such as growth, collapse, solidification and erosion.
Lava domes grow by endogenic dome growth or exogenic dome growth. The former implies dome interior expansion to accommodate new lava and the latter refers to superficial piling up of lava.[1] It is the high viscosity of the lava that prevents it from flowing far from the vent from which it extrudes, creating a dome-like shape of sticky lava that then cools slowly in situ. Domes may reach heights of several hundred meters, and can grow slowly and steadily for months (e.g. Unzen volcano), years (e.g. Soufrière Hills volcano), or even centuries (e.g. Mount Merapi volcano). The sides of these structures are composed of unstable rock debris. Due to the intermittent buildup of gas pressure, erupting domes can often experience episodes of explosive eruption over time. If part of a lava dome collapses while it is still molten, it can produce pyroclastic flows,[5] one of the most lethal forms of volcanic event. Other hazards associated with lava domes are the destruction of property, forest fires, and lahars triggered by pyroclastic flows near mud, snow and ice. Lava domes are one of the principal structural features of many stratovolcanoes worldwide. Lava domes are prone to unusually dangerous explosions since they contain rhyolitic silica-rich lava.
Characteristics of lava dome eruptions include shallow, long-period and hybrid seismicity, which is attributed to excess fluid pressures in the contributing vent chamber. Other characteristics of lava domes include their hemispherical dome shape, cycles of dome growth over long periods, and sudden onsets of violent explosive activity.[6] The average rate of dome growth may be used as a rough indicator of magma supply, but it shows no systematic relationship to the timing or characteristics of lava dome explosions.[7]
Related landforms[edit]
Photo showing the bulging cryptodome of Mt. St. Helens on April 27, 1980.
Cryptodomes[edit]
A cryptodome (from Greek κρυπτός, kryptos, "hidden, secret") is a dome-shaped structure created by accumulation of viscous magma at a shallow depth. One example of a cryptodome was in the May 1980 eruption of Mount St. Helens, where the explosive eruption began after a landslide caused the side of the volcano to fall, leading to explosive decompression of the subterranean cryptodome.
Lava coulées[edit]
Chao dacite coulée flow-domes (left center), northern Chile, viewed from Landsat 8.
Coulées (or coulees) are lava domes that have experienced some flow away from their original position, thus resembling both lava domes and lava flows.[1]
The world's largest known dacite flow is the Chao dacite dome complex, a huge coulée flow-dome between two volcanoes in northern Chile. This flow is over 14 kilometres (8.7 mi) long, has obvious flow features like pressure ridges, and a flow front 400 metres (1,300 ft) tall (the dark scalloped line at lower left).[8] There is another prominent coulée flow on the flank of Llullaillaco volcano, in Argentina,[9] and other examples in the Andes.
Examples of lava domes[edit]
Main article: List of lava domes
Lava domes
Name of lava dome
Country
Volcanic area
Composition
Last eruption
or growth episode
Chaitén lava dome Chile Southern Volcanic Zone Rhyolite 2009
Ciomadul lava domes Romania Carpathians Dacite Pleistocene
Cordón Caulle lava domes Chile Southern Volcanic Zone Rhyodacite to Rhyolite Holocene
Galeras lava dome Colombia Northern Volcanic Zone Unknown 2010
Katla lava dome Iceland Iceland hotspot Rhyolite 1999 onwards[10]
Lassen Peak United States Cascade Volcanic Arc Dacite 1917
Bridge River Vent lava dome Canada Cascade Volcanic Arc Dacite 2350 BP
Mount Merapi lava dome Indonesia Sunda Arc Unknown 2010
Nea Kameni Greece South Aegean Volcanic Arc Dacite 1950
Novarupta lava dome Alaska (United States) Aleutian Arc Rhyolite 1912
Nevados de Chillán lava domes Chile Southern Volcanic Zone Dacite 1986
Puy de Dôme France Chaîne des Puys Trachyte ca. 5760 BC
Santa María lava dome Guatemala Central America Volcanic Arc Dacite 2009
Sollipulli lava dome Chile Southern Volcanic Zone Andesite to Dacite 1240 ± 50 years
Soufrière Hills lava dome Montserrat Lesser Antilles Andesite 2009
Mount St. Helens lava domes United States Cascade Volcanic Arc Dacite 2008
Torfajökull lava dome Iceland Iceland hotspot Rhyolite 1477
Tata Sabaya lava domes Bolivia Andes Unknown ~ Holocene
Tate-iwa Japan Japan Arc Dacite Miocene[11]
Valles lava domes United States Jemez Mountains Rhyolite 50,000-60,000 BP
Wizard Island lava dome United States Cascade Volcanic Arc Rhyodacite[12] 2850 BC
References[edit]
^ Jump up to: a b c Fink, Jonathan H., Anderson, Steven W. (2001), "Bernard Lewis", in Sigursson, Haraldur, Encyclopedia of Volcanoes, Academic Press, pp. 307–319.
Jump up ^ Rampey, Michael L.; Milam, Keith A.; McSween, Harry Y.; Moersch, Jeffrey E.; Christensen, Philip R. (28 June 2007). "Identity and emplacement of domical structures in the western Arcadia Planitia, Mars". Journal of Geophysical Research. 112 (E6). doi:10.1029/2006JE002750.
Jump up ^ Brož, Petr; Hauber, Ernst; Platz, Thomas; Balme, Matt (April 2015). "Evidence for Amazonian highly viscous lavas in the southern highlands on Mars". Earth and Planetary Science Letters. 415: 200–212. doi:10.1016/j.epsl.2015.01.033.
Jump up ^ Melnik, O; Sparks, R. S. J. (4 November 1999), "Nonlinear dynamics of lava dome extrusion" (PDF), Nature, 402 (6757): 37–41, Bibcode:1999Natur.402...37M, doi:10.1038/46950
Jump up ^ Parfitt, E.A.; Wilson, L (2008), Fundamentals of Physical Volcanology, Massachusetts, USA: Blackwell Publishing, p. 256
Jump up ^ Sparks, R.S.J. (August 1997), "Causes and consequences of pressurisation in lava dome eruptions", Earth and Planetary Science Letters, 150 (3–4): 177–189, Bibcode:1997E&PSL.150..177S, doi:10.1016/S0012-821X(97)00109-X
Jump up ^ Newhall, C.G.; Melson., W.G. (September 1983), "Explosive activity associated with the growth of volcanic domes", Journal of Volcanology and Geothermal Research, 17 (1–4): 111–131, Bibcode:1983JVGR...17..111N, doi:10.1016/0377-0273(83)90064-1 )
Jump up ^ Chao dacite dome complex at NASA Earth Observatory
Jump up ^ Coulées! by Erik Klemetti, an assistant professor of Geosciences at Denison University.
Jump up ^ Eyjafjallajökull and Katla: restless neighbours
Jump up ^ Goto, Yoshihiko; Tsuchiya, Nobutaka (July 2004). "Morphology and growth style of a Miocene submarine dacite lava dome at Atsumi, northeast Japan". Journal of Volcanology and Geothermal Research. 134 (4): 255–275. doi:10.1016/j.jvolgeores.2004.03.015.
Jump up ^ Map of Post-Caldera Volcanism and Crater Lake USGS Cascades Volcano Observatory. Retrieved 2014-01-31.
Global Volcanism Program: Lava Domes
USGS Photo glossary of volcano terms: Lava dome
External links[edit]
Wikimedia Commons has media related to Lava domes.
Wikimedia Commons has media related to Lava coulées.
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Resting potential
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The Na+
/K+
-ATPase, as well as effects of diffusion of the involved ions, are major mechanisms to maintain the resting potential across the membranes of animal cells.
Play media
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The relatively static membrane potential of quiescent cells is called the resting membrane potential (or resting voltage), as opposed to the specific dynamic electrochemical phenomena called action potential and graded membrane potential.
Apart from the latter two, which occur in excitable cells (neurons, muscles, and some secretory cells in glands), membrane voltage in the majority of non-excitable cells can also undergo changes in response to environmental or intracellular stimuli[citation needed]. In principle, there is no difference between resting membrane potential and dynamic voltage changes like action potential from a biophysical point of view: all these phenomena are caused by specific changes in membrane permeabilities for potassium, sodium, calcium, and chloride ions, which in turn result from concerted changes in functional activity of various ion channels, ion transporters, and exchangers. Conventionally, resting membrane potential can be defined as a relatively stable, ground value of transmembrane voltage in animal and plant cells.
Any voltage is a difference in electric potential between two points—for example, the separation of positive and negative electric charges on opposite sides of a resistive barrier. The typical resting membrane potential of a cell arises from the separation of potassium ions from intracellular, relatively immobile anions across the membrane of the cell. Because the membrane permeability for potassium is much higher than that for other ions (disregarding voltage-gated channels at this stage), and because of the strong chemical gradient for potassium, potassium ions flow from the cytosol into the extracellular space carrying out positive charge, until their movement is balanced by build-up of negative charge on the inner surface of the membrane. Again, because of the high relative permeability for potassium, the resulting membrane potential is almost always close to the potassium reversal potential. But in order for this process to occur, a concentration gradient of potassium ions must first be set up. This work is done by the ion pumps/transporters and/or exchangers and generally is powered by ATP.
In the case of the resting membrane potential across an animal cell's plasma membrane, potassium (and sodium) gradients are established by the Na+/K+-ATPase (sodium-potassium pump) which transports 2 potassium ions inside and 3 sodium ions outside at the cost of 1 ATP molecule. In other cases, for example, a membrane potential may be established by acidification of the inside of a membranous compartment (such as the proton pump that generates membrane potential across synaptic vesicle membranes).[citation needed]
Contents
[hide]
1 Electroneutrality
2 Generation of the resting potential
3 Membrane transport proteins
4 Equilibrium potentials
5 Resting potentials
6 Measuring resting potentials
7 Summary of resting potential values in different types of cells
8 History
9 See also
10 References
11 External links
Electroneutrality[edit]
In most quantitative treatments of membrane potential, such as the derivation of Goldman equation, electroneutrality is assumed; that is, that there is no measurable charge excess in any side of the membrane. So, although there is an electric potential across the membrane due to charge separation, there is no actual measurable difference in the global concentration of positive and negative ions across the membrane (as it is estimated below), that is, there is no actual measurable charge excess on either side. That occurs because the effect of charge on electrochemical potential is hugely greater than the effect of concentration so an undetectable change in concentration creates a great change in electric potential.[citation needed]
Generation of the resting potential[edit]
Cell membranes are typically permeable to only a subset of ions. These usually include potassium ions, chloride ions, bicarbonate ions, and others. To simplify the description of the ionic basis of the resting membrane potential, it is most useful to consider only one ionic species at first, and consider the others later. Since trans-plasma-membrane potentials are almost always determined primarily by potassium permeability, that is where to start.
A diagram showing the progression in the development of a membrane potential from a concentration gradient (for potassium). Green arrows indicate net movement of K+ down a concentration gradient. Red arrows indicate net movement of K+ due to the membrane potential. The diagram is misleading in that while the concentration of potassium ions outside of the cell increases, only a small amount of K+ needs to cross the membrane in order to produce a membrane potential with a magnitude large enough to counter the tendency of the potassium ions to move down the concentration gradient.
Panel 1 of the diagram shows a diagrammatic representation of a simple cell where a concentration gradient has already been established. This panel is drawn as if the membrane has no permeability to any ion. There is no membrane potential because despite there being a concentration gradient for potassium, there is no net charge imbalance across the membrane. If the membrane were to become permeable to a type of ion that is more concentrated on one side of the membrane, then that ion would contribute to membrane voltage because the permeant ions would move across the membrane with net movement of that ion type down the concentration gradient. There would be net movement from the side of the membrane with a higher concentration of the ion to the side with lower concentration. Such a movement of one ion across the membrane would result in a net imbalance of charge across the membrane and a membrane potential. This is a common mechanism by which many cells establish a membrane potential.
In panel 2 of the diagram, the cell membrane has been made permeable to potassium ions, but not the anions (An−) inside the cell. These anions are mostly contributed by protein. There is energy stored in the potassium ion concentration gradient that can be converted into an electrical gradient when potassium (K+) ions move out of the cell. Note that potassium ions can move across the membrane in both directions but by the purely statistical process that arises from the higher concentration of potassium ions inside the cell, there will be more potassium ions moving out of the cell. Because there is a higher concentration of potassium ions inside the cells, their random molecular motion is more likely to encounter the permeability pore (ion channel) that is the case for the potassium ions that are outside and at a lower concentration. An internal K+ is simply "more likely" to leave the cell than an extracellular K+ is to enter it. It is a matter of simple diffusion doing work by dissipating the concentration gradient. As potassium leaves the cell, it is leaving behind the anions. Therefore, a charge separation is developing as K+ leaves the cell. This charge separation creates a transmembrane voltage. This transmembrane voltage is the membrane potential. As potassium continues to leave the cell, separating more charges, the membrane potential will continue to grow. The length of the arrows (green indicating concentration gradient, red indicating voltage), represents the magnitude of potassium ion movement due to each form of energy. The direction of the arrow indicates the direction in which that particular force is applied. Thus, the building membrane voltage is an increasing force that acts counter to the tendency for net movement of potassium ions down the potassium concentration gradient.
In Panel 3, the membrane voltage has grown to the extent that its "strength" now matches the concentration gradients. Since these forces (which are applied to K+) are now the same strength and oriented in opposite directions, the system is now in equilibrium. Put another way, the tendency of potassium to leave the cell by running down its concentration gradient is now matched by the tendency of the membrane voltage to pull potassium ions back into the cell. K+ continues to move across the membrane, but the rate at which it enters and leaves the cell are the same, thus, there is no net potassium current. Because the K+ is at equilibrium, membrane potential is stable, or "resting" (EK).
The resting voltage is the result of several ion-translocating enzymes (uniporters, cotransporters, and pumps) in the plasma membrane, steadily operating in parallel, whereby each ion-translocator has its characteristic electromotive force (= reversal potential = 'equilibrium voltage'), depending on the particular substrate concentrations inside and outside (internal ATP included in case of some pumps). H+ exporting ATPase render the membrane voltage in plants and fungi much more negative than in the more extensively investigated animal cells, where the resting voltage is mainly determined by selective ion channels.
In most neurons the resting potential has a value of approximately −70 mV. The resting potential is mostly determined by the concentrations of the ions in the fluids on both sides of the cell membrane and the ion transport proteins that are in the cell membrane. How the concentrations of ions and the membrane transport proteins influence the value of the resting potential is outlined below.
The resting potential of a cell can be most thoroughly understood by thinking of it in terms of equilibrium potentials. In the example diagram here, the model cell was given only one permeant ion (potassium). In this case, the resting potential of this cell would be the same as the equilibrium potential for potassium.
However, a real cell is more complicated, having permeabilities to many ions, each of which contributes to the resting potential. To understand better, consider a cell with only two permeant ions, potassium, and sodium. Consider a case where these two ions have equal concentration gradients directed in opposite directions, and that the membrane permeabilities to both ions are equal. K+ leaving the cell will tend to drag the membrane potential toward EK. Na+ entering the cell will tend to drag the membrane potential toward the reversal potential for sodium ENa. Since the permeabilities to both ions were set to be equal, the membrane potential will, at the end of the Na+/K+ tug-of-war, end up halfway between ENa and EK. As ENa and EK were equal but of opposite signs, halfway in between is zero, meaning that the membrane will rest at 0 mV.
Note that even though the membrane potential at 0 mV is stable, it is not an equilibrium condition because neither of the contributing ions is in equilibrium. Ions diffuse down their electrochemical gradients through ion channels, but the membrane potential is upheld by continual K+ influx and Na+ efflux via ion transporters. Such situation with similar permeabilities for counter-acting ions, like potassium and sodium in animal cells, can be extremely costly for the cell if these permeabilities are relatively large, as it takes a lot of ATP energy to pump the ions back. Because no real cell can afford such equal and large ionic permeabilities at rest, resting potential of animal cells is determined by predominant high permeability to potassium and adjusted to the required value by modulating sodium and chloride permeabilities and gradients.
In a healthy animal cell Na+ permeability is about 5% of the K+ permeability or even less, whereas the respective reversal potentials are +60 mV for sodium (ENa)and −80 mV for potassium (EK). Thus the membrane potential will not be right at EK, but rather depolarized from EK by an amount of approximately 5% of the 140 mV difference between EK and ENa. Thus, the cell's resting potential will be about −73 mV.
In a more formal notation, the membrane potential is the weighted average of each contributing ion's equilibrium potential. The size of each weight is the relative conductance of each ion. In the normal case, where three ions contribute to the membrane potential:
E m = g K + g t o t E K + + g N a + g t o t E N a + + g C l − g t o t E C l − {\displaystyle E_{m}={\frac {g_{K^{+}}}{g_{tot}}}E_{K^{+}}+{\frac {g_{Na^{+}}}{g_{tot}}}E_{Na^{+}}+{\frac {g_{Cl^{-}}}{g_{tot}}}E_{Cl^{-}}} ,
where
Em is the membrane potential, measured in volts
EX is the equilibrium potential for ion X, also in volts
gX is the relative conductance of ion X in arbitrary units (e.g. siemens for electrical conductance)
gtot is the total conductance of all permeant ions, in this case gK+ + gNa+ + gCl−
Membrane transport proteins[edit]
For determination of membrane potentials, the two most important types of membrane ion transport proteins are ion channels and ion transporters. Ion channel proteins create paths across cell membranes through which ions can passively diffuse without direct expenditure of metabolic energy. They have selectivity for certain ions, thus, there are potassium-, chloride-, and sodium-selective ion channels. Different cells and even different parts of one cell (dendrites, cell bodies, nodes of Ranvier) will have different amounts of various ion transport proteins. Typically, the amount of certain potassium channels is most important for control of the resting potential (see below). Some ion pumps such as the Na+/K+-ATPase are electrogenic, that is, they produce charge imbalance across the cell membrane and can also contribute directly to the membrane potential. Most pumps use metabolic energy (ATP) to function.
Equilibrium potentials[edit]
For most animal cells potassium ions (K+) are the most important for the resting potential.[1] Due to the active transport of potassium ions, the concentration of potassium is higher inside cells than outside. Most cells have potassium-selective ion channel proteins that remain open all the time. There will be net movement of positively charged potassium ions through these potassium channels with a resulting accumulation of excess negative charge inside of the cell. The outward movement of positively charged potassium ions is due to random molecular motion (diffusion) and continues until enough excess negative charge accumulates inside the cell to form a membrane potential which can balance the difference in concentration of potassium between inside and outside the cell. "Balance" means that the electrical force (potential) that results from the build-up of ionic charge, and which impedes outward diffusion, increases until it is equal in magnitude but opposite in direction to the tendency for outward diffusive movement of potassium. This balance point is an equilibrium potential as the net transmembrane flux (or current) of K+ is zero. A good approximation for the equilibrium potential of a given ion only needs the concentrations on either side of the membrane and the temperature. It can be calculated using the Nernst equation:
E e q , K + = R T z F ln  [ K + ] o [ K + ] i , {\displaystyle E_{eq,K^{+}}={\frac {RT}{zF}}\ln {\frac {[K^{+}]_{o}}{[K^{+}]_{i}}},}
where
Eeq,K+ is the equilibrium potential for potassium, measured in volts
R is the universal gas constant, equal to 8.314 joules·K−1·mol−1
T is the absolute temperature, measured in kelvins (= K = degrees Celsius + 273.15)
z is the number of elementary charges of the ion in question involved in the reaction
F is the Faraday constant, equal to 96,485 coulombs·mol−1 or J·V−1·mol−1
[K+]o is the extracellular concentration of potassium, measured in mol·m−3 or mmol·l−1
[K+]i is likewise the intracellular concentration of potassium
Potassium equilibrium potentials of around −80 millivolts (inside negative) are common. Differences are observed in different species, different tissues within the same animal, and the same tissues under different environmental conditions. Applying the Nernst Equation above, one may account for these differences by changes in relative K+ concentration or differences in temperature.
For common usage the Nernst equation is often given in a simplified form by assuming typical human body temperature (37 °C), reducing the constants and switching to Log base 10. (The units used for concentration are unimportant as they will cancel out into a ratio). For Potassium at normal body temperature one may calculate the equilibrium potential in millivolts as:
E e q , K + = 61.54 m V log  [ K + ] o [ K + ] i , {\displaystyle E_{eq,K^{+}}=61.54mV\log {\frac {[K^{+}]_{o}}{[K^{+}]_{i}}},}
Likewise the equilibrium potential for sodium (Na+) at normal human body temperature is calculated using the same simplified constant. You can calculate E assuming an outside concentration, [K+]o, of 10mM and an inside concentration, [K+]i, of 100mM. For chloride ions (Cl−) the sign of the constant must be reversed (−61.54 mV). If calculating the equilibrium potential for calcium (Ca2+) the 2+ charge halves the simplified constant to 30.77 mV. If working at room temperature, about 21 °C, the calculated constants are approximately 58 mV for K+ and Na+, −58 mV for Cl− and 29 mV for Ca2+. At physiological temperature, about 29.5 °C, and physiological concentrations (which vary for each ion), the calculated potentials are approximately 67 mV for Na+, −90 mV for K+, −86 mV for Cl− and 123 mV for Ca2+.
Resting potentials[edit]
The resting membrane potential is not an equilibrium potential as it relies on the constant expenditure of energy (for ionic pumps as mentioned above) for its maintenance. It is a dynamic diffusion potential that takes this mechanism into account—wholly unlike the equilibrium potential, which is true no matter the nature of the system under consideration. The resting membrane potential is dominated by the ionic species in the system that has the greatest conductance across the membrane. For most cells this is potassium. As potassium is also the ion with the most negative equilibrium potential, usually the resting potential can be no more negative than the potassium equilibrium potential. The resting potential can be calculated with the Goldman-Hodgkin-Katz voltage equation using the concentrations of ions as for the equilibrium potential while also including the relative permeabilities of each ionic species. Under normal conditions, it is safe to assume that only potassium, sodium (Na+) and chloride (Cl−) ions play large roles for the resting potential:
E m = R T F ln  ( P N a + [ N a + ] o + P K + [ K + ] o + P C l − [ C l − ] i P N a + [ N a + ] i + P K + [ K + ] i + P C l − [ C l − ] o ) {\displaystyle E_{m}={\frac {RT}{F}}\ln {\left({\frac {P_{Na^{+}}[Na^{+}]_{o}+P_{K^{+}}[K^{+}]_{o}+P_{Cl^{-}}[Cl^{-}]_{i}}{P_{Na^{+}}[Na^{+}]_{i}+P_{K^{+}}[K^{+}]_{i}+P_{Cl^{-}}[Cl^{-}]_{o}}}\right)}}
This equation resembles the Nernst equation, but has a term for each permeant ion. Also, z has been inserted into the equation, causing the intracellular and extracellular concentrations of Cl− to be reversed relative to K+ and Na+, as chloride's negative charge is handled by inverting the fraction inside the logarithmic term. *Em is the membrane potential, measured in volts *R, T, and F are as above *Ps is the relative permeability of ion s *[s]Y is the concentration of ion s in compartment Y as above. Another way to view the membrane potential, considering instead the conductance of the ion channels rather than the permeability of the membrane, is using the Millman equation (also called the Chord Conductance Equation):
E m = g K + E e q , K + + g N a + E e q , N a + + g C l − E e q , C l − g K + + g N a + + g C l − {\displaystyle E_{m}={\frac {g_{K^{+}}E_{eq,K^{+}}+g_{Na^{+}}E_{eq,Na^{+}}+g_{Cl^{-}}E_{eq,Cl^{-}}}{g_{K^{+}}+g_{Na^{+}}+g_{Cl^{-}}}}}
or reformulated
E m = g K + g t o t E e q , K + + g N a + g t o t E e q , N a + + g C l − g t o t E e q , C l − {\displaystyle E_{m}={\frac {g_{K^{+}}}{g_{tot}}}E_{eq,K^{+}}+{\frac {g_{Na^{+}}}{g_{tot}}}E_{eq,Na^{+}}+{\frac {g_{Cl^{-}}}{g_{tot}}}E_{eq,Cl^{-}}}
where gtot is the combined conductance of all ionic species, again in arbitrary units. The latter equation portrays the resting membrane potential as a weighted average of the reversal potentials of the system, where the weights are the relative conductances of each ion species (gX/gtot). During the action potential, these weights change. If the conductances of Na+ and Cl− are zero, the membrane potential reduces to the Nernst potential for K+ (as gK+ = gtot). Normally, under resting conditions gNa+ and gCl− are not zero, but they are much smaller than gK+, which renders Em close to Eeq,K+. Medical conditions such as hyperkalemia in which blood serum potassium (which governs [K+]o) is changed are very dangerous since they offset Eeq,K+, thus affecting Em. This may cause arrhythmias and cardiac arrest. The use of a bolus injection of potassium chloride in executions by lethal injection stops the heart by shifting the resting potential to a more positive value, which depolarizes and contracts the cardiac cells permanently, not allowing the heart to repolarize and thus enter diastole to be refilled with blood.
Although the GHK voltage equation and Millman's equation are related, they are not equivalent. The critical difference is that Millman's equation assumes the current-voltage relationship to be ohmic, whereas the GHK voltage equation takes into consideration the small, instantaneous rectifications predicted by the GHK flux equation caused by the concentration gradient of ions. Thus, a more accurate estimate of membrane potential can be calculated using the GHK equation than with Millman's equation.[2]
Measuring resting potentials[edit]
In some cells, the membrane potential is always changing (such as cardiac pacemaker cells). For such cells there is never any "rest" and the "resting potential" is a theoretical concept. Other cells with little in the way of membrane transport functions that change with time have a resting membrane potential that can be measured by inserting an electrode into the cell.[3] Transmembrane potentials can also be measured optically with dyes that change their optical properties according to the membrane potential.
Summary of resting potential values in different types of cells[edit]
Cell types
Resting potential
Hair cell (Cochlea) -15 to -40mV[4]
Skeletal muscle cells −95 mV[5]
Smooth muscle cells –60 mV
Astroglia –80 to –90 mV
Neurons –60 to –70 mV[6]
Erythrocytes –8.4 mV[7]
Chondrocytes -8mV[6]
Aorta Smooth muscle tissue -45mV[6]
Photoreceptor cells –40 mV
History[edit]
Resting currents in nerves were measured and described by Julius Bernstein in 1902 where he proposed a "Membrane Theory" that explained the resting potential of nerve and muscle as a diffusion potential.[8]
See also[edit]
Action potential
Depolarization
Hyperpolarization (biology)
Membrane potential
References[edit]
Jump up ^ An example of an electrophysiological experiment to demonstrate the importance of K+ for the resting potential. The dependence of the resting potential on the extracellular concentration of K+ is shown in Figure 2.6 of Neuroscience, 2nd edition, by Dale Purves, George J. Augustine, David Fitzpatrick, Lawrence C. Katz, Anthony-Samuel LaMantia, James O. McNamara, S. Mark Williams. Sunderland (MA): Sinauer Associates, Inc.; 2001.
Jump up ^ Hille, Bertil (2001) Ion Channels of Excitable Membranes, 3 ed.
Jump up ^ An illustrated example of measuring membrane potentials with electrodes is in Figure 2.1 of Neuroscience by Dale Purves, et al. (see reference #1, above).
Jump up ^ Ashmore, J. F.; Meech, R. W. (1986-07-24). "Ionic basis of membrane potential in outer hair cells of guinea pig cochlea". Nature. 322 (6077): 368–371. doi:10.1038/322368a0.
Jump up ^ "Muscles". users.rcn.com. 2015-01-24. Archived from the original on 2015-11-07. Retrieved 2016-06-01.
^ Jump up to: a b c Lewis, Rebecca; Asplin, Katie E.; Bruce, Gareth; Dart, Caroline; Mobasheri, Ali; Barrett-Jolley, Richard (2011-11-01). "The role of the membrane potential in chondrocyte volume regulation". Journal of Cellular Physiology. 226 (11): 2979–2986. doi:10.1002/jcp.22646. ISSN 1097-4652. PMC 3229839  . PMID 21328349.
Jump up ^ Cheng, K; Haspel, HC; Vallano, ML; Osotimehin, B; Sonenberg, M (1980). "Measurement of membrane potentials (psi) of erythrocytes and white adipocytes by the accumulation of triphenylmethylphosphonium cation". J. Membr. Biol. 56 (3): 191–201. doi:10.1007/bf01869476. PMID 6779011.
Jump up ^ Seyfarth, Ernst-August (2006-01-01). "Julius Bernstein (1839-1917): pioneer neurobiologist and biophysicist". Biological Cybernetics. 94 (1): 2–8. doi:10.1007/s00422-005-0031-y. ISSN 0340-1200. PMID 16341542.
External links[edit]
Neuroscience - online textbook by Purves, et al.
Basic Neurochemistry Molecular, Cellular, and Medical Aspects by Siegel, et al.
Bertil Hille Ion channels of excitable membranes, 3rd ed., Sinauer Associates, Sunderland, MA (2001). ISBN 0-87893-321-2
Wright, SH (2004). "Generation of resting membrane potential". Adv Physiol Educ. 28 (1–4): 139–42. doi:10.1152/advan.00029.2004. PMID 15545342.
Resting Membrane Potential - Online lecture notes on the resting membrane potential
The Origin of the Resting Membrane Potential - Online interactive tutorial (Flash)
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Cynthia Watros
From Wikipedia, the free encyclopedia
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This biography of a living person needs additional citations for verification. Please help by adding reliable sources. Contentious material about living persons that is unsourced or poorly sourced must be removed immediately, especially if potentially libelous or harmful. (March 2013) (Learn how and when to remove this template message)
Cynthia Watros
Watros in Iraq
Born
Cynthia Michele Watros
September 2, 1968
Lake Orion, Michigan, U.S.
Occupation
Actress
Years active
1994–present
Spouse(s)
Curtis Gilliland (m. 1996)
Children
2
Cynthia Michele Watros is an American actress. Watros is known for her roles as Libby Smith on the ABC TV series Lost, Kellie in The Drew Carey Show, Erin in Titus, and Annie Dutton in Guiding Light.
Contents
1 Career
2 Personal life
3 Filmography
3.1 Film
3.2 Television
4 Awards and nominations
5 References
6 External links
Career[edit]
Watros initially became known for her role as Annie Dutton on the CBS soap opera Guiding Light from 1994-98. Her character, a nurse, began as a heroine, but she gained notice and critical acclaim as her character gradually descended into madness. In 1998, Watros won the Daytime Emmy Award for Outstanding Lead Actress in a Drama Series for her role on Guiding Light. In 1998, she briefly filled in for Jensen Buchanan as Victoria Hudson McKinnon on Another World while Buchanan was on maternity leave.[citation needed]
Watros appeared as Erin Fitzpatrick on Titus from 2000 to 2002 and gained recognition as a comedic actress, playing the fiancée of the title character. According to the DVD commentary, Erin was originally intended to be neurotic and socially awkward, but the character was revised a week before the pilot was shot. After Titus was canceled, she took on the role of Kellie Newmark on The Drew Carey Show, as a replacement for Christa Miller. Watros played the role from 2002-04.[citation needed]
From 2005 to 2006, Watros was a cast member on the ABC series Lost, playing psychologist Libby, a member of the "Tailies", a group of survivors of the plane crash who were in the tail section of the plane (and not seen during Season One). She was a romantic interest to Hurley and was very mysterious showing up in flashbacks. In the episode "Two for the Road", Libby was shot and killed by Michael Dawson. Libby briefly reappeared in the 2008 episode "Meet Kevin Johnson". She returned in 2010 in the episode "Everybody Loves Hugo" in the final season of Lost, in which she played a large part in the story. In addition, she appeared in the season finale, albeit without speaking lines.
After her departure from Lost in 2006, Watros filmed a pilot for a show titled My Ex-Life for CBS. She was slated to play the ex-wife of a character played by Tom Cavanagh. However, the pilot was not chosen by CBS.[1] Watros was also the lead in the 2007 CBS television pilot for The Rich Inner Life of Penelope Cloud, a comedy about a former literary genius who, after an awakening, decides to pursue optimism instead of cynicism.
In 2009, Watros filmed a pilot for Valley Girls, a spin-off of the CW series Gossip Girl set in the 1980s, in which she played CeCe Rhodes, the socialite ex-wife of Rick Rhodes, and mother of the series' protagonists. In a May 2009 episode of the USA series In Plain Sight, "A Stand-Up Triple", she guest starred as a mother of three children in the witness protection program. Beginning in April 2010, Watros joined the cast of the TV medical drama House for seven episodes as Samantha Carr, one of Dr. James Wilson's ex-wives.
Watros appeared in the 2012 indie film, Electrick Children. In May 2012, she guest-starred in episode 8.22 of Grey's Anatomy. In June 2013, she guest-starred in episode 4.17 of Warehouse 13. She played Mary Matrix, coach of the FPS Varsity team in the online series Video Game High School. Her character first appears in season two of the series, which started July 2013. Watros starred as Avery Jenning's self-obsessed aunt in Dog with a Blog on December 6, 2013.
In 2013, Watros joined the cast of the CBS soap opera The Young and the Restless but left the show in early 2014 with her character being recast. In July 2014, she joined the cast of the MTV series Finding Carter.[2]
Personal life[edit]
Watros was born in Lake Orion, Michigan. She attended Macomb Community College in Clinton Township, Michigan,[3] and received a Bachelor of Fine Arts degree in Theatre from Boston University,[4] where she was part of Boston University's Professional Actors Training Program.[5]
Watros and Bonnie-Jill Laflin posing with a serviceman on a USO tour
Watros has been married to Curtis Gilliland since 1996. They have twin daughters, born in 2001.[6]
In 2008, Watros participated in a USO tour to Iraq, visiting several bases, including COP Callahan in Baghdad's East Adamiyah area.
Filmography[edit]
Film[edit]
Year
Title
Role
Notes
1995 Cafe Society Dianna Harris
1997 His and Hers Pam
2000 Mercy Streets Sam
2001 The Yellow Bird Alma Tutwiler Short film
2002 P.S. Your Cat Is Dead Kate
2005 Just Pray Perry Ann Lewis Short film
2007 Frank Jennifer York
2008 Duane Incarnate Connie
2008 American Crude Jane
2009 Calvin Marshall Karen
2010 Mars Allison Guthrie
2012 Electrick Children Gay Lynn
2013 Park City Nina
2014 Blood and Circumstance Mrs. Stabler
Television[edit]
Year
Title
Role
Notes
1994 New York Undercover Reporter Episode: "Missing"
1994–98 Guiding Light Annie Dutton / Dee
1997 Spin City Gayley Episode: "Striptease"
1998 Another World Vicky Hudson 9 episodes
1998 Profiler Helen Jefferies Episode: "Perfect Helen"
2000–02 Titus Erin Fitzpatrick Main role, 54 episodes
2002 A Nero Wolfe Mystery Phoebe Gunther "The Silent Speaker"
2002–04 The Drew Carey Show Kellie Newmark Main role (Season 8–9), 52 episodes
2005 Washington Street Maggie Movie
2005–06;
2008, 2010 Lost Elizabeth "Libby" Smith Main role (Season 2)
Special guest star (Seasons 4 & 6)
24 episodes
2007 Avenging Angel Maggie Movie
2007 The Rich Inner Life of Penelope Cloud Eva Movie
2007 Raines Sarah Carver Episode: "Closure"
2007 Law & Order: Criminal Intent Beth Hoyle Episode: "Offense"
2008 Fear Itself Meredith Kane Episode: "Spooked"
2008 The Bill Engvall Show AJ Episodes: "Susan's Best Friend"
"Bill Talks a Good Game"
2009 Family Guy Security System Voice Episode: "Ocean's Three and a Half"
2009 In Plain Sight Maureen Stewart / Maureen Sullivan Episode: "A Stand-Up Triple"
2009 Gossip Girl Young Celia Rhodes Episode: "Valley Girls"
2009 CSI: Crime Scene Investigation Barbie Aubrey Episode: "All In"
2009 The Closer Robin Milano Episode: "Identity Theft"
2009 Criminal Minds Heather Vanderwaal Episode: "Reckoner"
2009 Men of a Certain Age Erica Episode: "Mind's Eye"
2010 The Secret Life of the American Teenager Nicky Episode: "The Second Time Around"
2010 House M.D. Sam Carr 7 episodes
2010 Desperate Housewives Tracy Miller Episode: "Pleasant Little Kingdom"
2012 A Smile as Big as the Moon Dr. Deborah Barnhart Movie
2012 Retribution Karen Movie
2012 Grey's Anatomy Mrs. Connor Episode: "Let the Bad Times Roll"
2012 Hawaii Five-0 Katie Burgess Episode: "Lana I Ka Moana"
2013 Cynthia Watros Gets Lost As herself 4 pilot episodes on YouTube
2013 Warehouse 13 Janice Malloy Episode: "What Matters Most"
2013–14 Video Game High School Mary Matrix 12 episodes
2013 Dog with a Blog Aunt Sigourney Episode: "Twas the Flight Before Christmas"
2013–14 The Young and the Restless Kelly Andrews
2014–15 Finding Carter Elizabeth Wilson Main role
2015 Stolen from the Suburbs Katherine Movie
2017 Washed Away Movie
2017 Deadly Exchange Detective Hardy Movie
Awards and nominations[edit]
Year
Award
Work
Result
Ref.
1998 Daytime Emmy Award for Outstanding Lead Actress in a Drama Series Guiding Light Won [7]
Soap Opera Digest Award for Outstanding Villainess in a Drama Series – Daytime Nominated [8]
2006 Screen Actors Guild Award for Outstanding Performance by an Ensemble in a Drama Series Lost Won [9]
References[edit]
Jump up ^ My Ex Life at TV.com
Jump up ^ "Cynthia Watros - Finding Carter". mtv.com. MTV. Retrieved 12 August 2014.
Jump up ^ [1], Retrieved 11-10-2015.
Jump up ^ [2]
Jump up ^ "Emily Deschanel Biography". TV.com. Retrieved August 15, 2011.
Jump up ^ "The Young and the Restless ACTOR - Cynthia Watros".
Jump up ^ Margulies, Lee (May 18, 1998). "'All My Children' Tops Daytime Emmys". Los Angeles Times. Los Angeles, California: Tribune Company. Archived from the original on June 28, 2013. Retrieved May 6, 2013.
Jump up ^ "Event: Soap Opera Digest Awards (1998)". Internet Movie Database. February 4, 1993. Retrieved February 27, 2013.
Jump up ^ "12th Annual Screen Actors Guild Award Recipients". Archived from the original on 2007-08-19. Retrieved 2007-08-28.
External links[edit]
Wikimedia Commons has media related to Cynthia Watros.
Cynthia Watros on IMDb
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Daytime Emmy Award for Outstanding Lead Actress in a Drama Series
Elizabeth Hubbard (1974)
Susan Flannery (1975)
Helen Gallagher (1976)
Helen Gallagher (1977)
Laurie Heineman (1978)
Irene Dailey (1979)
Judith Light (1980)
Judith Light (1981)
Robin Strasser (1982)
Dorothy Lyman (1983)
Erika Slezak (1984)
Kim Zimmer (1985)
Erika Slezak (1986)
Kim Zimmer (1987)
Helen Gallagher (1988)
Marcy Walker (1989)
Kim Zimmer (1990)
Finola Hughes (1991)
Erika Slezak (1992)
Linda Dano (1993)
Hillary B. Smith (1994)
Erika Slezak (1995)
Erika Slezak (1996)
Jess Walton (1997)
Cynthia Watros (1998)
Susan Lucci (1999)
Susan Flannery (2000)
Martha Byrne (2001)
Susan Flannery (2002)
Susan Flannery (2003)
Michelle Stafford (2004)
Erika Slezak (2005)
Kim Zimmer (2006)
Maura West (2007)
Jeanne Cooper (2008)
Susan Haskell (2009)
Maura West (2010)
Laura Wright (2011)
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Heather Tom (2013)
Eileen Davidson (2014)
Maura West (2015)
Mary Beth Evans (2016)
Gina Tognoni (2017)
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2000: The West Wing, season 1/season 2
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Lowe
Moloney
Schiff
Sheen
Spencer
Whitford
2001: The West Wing, season 2/season 3
Hill
Janney
Lowe
Moloney
Schiff
Sheen
Spencer
Whitford
2002: Six Feet Under, season 2
Ambrose
Conroy
Griffiths
Hall
Krause
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2003: Six Feet Under, season 3
Ambrose
Conroy
Foster
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Hall
Krause
Macdissi
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Taylor
Wilson
2004: CSI: Crime Scene Investigation,
season 4/season 5
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2006: Grey's Anatomy, season 2/season 3
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Complete list
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Authority control
WorldCat Identities
ISNI: 0000 0000 4822 4661
LCCN: no2004107526
VIAF: 16944189
Retrieved from "https://en.wikipedia.org/w/index.php?title=Cynthia_Watros&oldid=848567849"
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