1021:
cell, and gentle suction is applied through the microelectrode to draw a piece of the cell membrane (the 'patch') into the microelectrode tip; the glass tip forms a high resistance 'seal' with the cell membrane. This configuration is the "cell-attached" mode, and it can be used for studying the activity of the ion channels that are present in the patch of membrane. If more suction is now applied, the small patch of membrane in the electrode tip can be displaced, leaving the electrode sealed to the rest of the cell. This "whole-cell" mode allows very stable intracellular recording. A disadvantage (compared to conventional intracellular recording with sharp electrodes) is that the intracellular fluid of the cell mixes with the solution inside the recording electrode, and so some important components of the intracellular fluid can be diluted. A variant of this technique, the "perforated patch" technique, tries to minimize these problems. Instead of applying suction to displace the membrane patch from the electrode tip, it is also possible to make small holes on the patch with pore-forming agents so that large molecules such as proteins can stay inside the cell and ions can pass through the holes freely. Also the patch of membrane can be pulled away from the rest of the cell. This approach enables the membrane properties of the patch to be analyzed pharmacologically. Patch-clamp may also be combined with RNA sequencing in a technique known as
127:
2334:
1059:
Recording in this way is in general called "single-unit" recording. The action potentials recorded are very much like the action potentials that are recorded intracellularly, but the signals are very much smaller (typically about 1 mV). Most recordings of the activity of single neurons in anesthetized and conscious animals are made in this way. Recordings of single neurons in living animals have provided important insights into how the brain processes information. For example,
1099:
1474:(information) and data required to meet a specific aim or aims in a clinical study. The "Minimum Information about a Neuroscience investigation" (MINI) family of reporting guideline documents aims to provide a consistent set of guidelines in order to report an electrophysiology experiment. In practice a MINI module comprises a checklist of information that should be provided (for example about the protocols employed) when a data set is described for publication.
1212:
906:
995:
1034:
much smaller so that there is very little ion exchange between the intracellular fluid and the electrolyte in the pipette. The electrical resistance of the micropipette electrode is reduced by filling with 2-4M KCl, rather than a salt concentration which mimics the intracellular ionic concentrations as used in patch clamping. Often the tip of the electrode is filled with various kinds of dyes like
1237:
875:
silver wire inserted into the pipette connects the electrolyte electrically to the amplifier and signal processing circuit. The voltage measured by the electrode is compared to the voltage of a reference electrode, usually a silver chloride-coated silver wire in contact with the extracellular fluid around the cell. In general, the smaller the electrode tip, the higher its
1406:
of modeling systems that are large enough and over sufficient timescales to be considered reproducing the macroscopic properties of the systems themselves. While atomistic simulations may access timescales close to, or into the microsecond domain, this is still several orders of magnitude lower than even the resolution of experimental methods such as patch-clamping.
1211:
1249:
930:, which open only when the membrane voltage is within a certain range. Voltage clamp measurements of current are made possible by the near-simultaneous digital subtraction of transient capacitive currents that pass as the recording electrode and cell membrane are charged to alter the cell's potential.
1145:
uses a carbon electrode to record changes in the chemical composition of the oxidized components of a biological solution. Oxidation and reduction is accomplished by changing the voltage at the active surface of the recording electrode in a process known as "scanning". Because certain brain chemicals
833:
Optical electrophysiological techniques were created by scientists and engineers to overcome one of the main limitations of classical techniques. Classical techniques allow observation of electrical activity at approximately a single point within a volume of tissue. Classical techniques singularize a
1298:
electrode (composed of the SSM and the absorbed vesicles) is so mechanically stable that solutions may be rapidly exchanged at its surface. This property allows the application of rapid substrate/ligand concentration jumps to investigate the electrogenic activity of the protein of interest, measured
1084:
If the electrode tip is slightly larger, then the electrode might record the activity generated by several neurons. This type of recording is often called "multi-unit recording", and is often used in conscious animals to record changes in the activity in a discrete brain area during normal activity.
1058:
An electrode introduced into the brain of a living animal will detect electrical activity that is generated by the neurons adjacent to the electrode tip. If the electrode is a microelectrode, with a tip size of about 1 micrometre, the electrode will usually detect the activity of at most one neuron.
1995:
Moschopoulou G.; Vitsa, K.; Bem, F.; Vassilakos, N.; Perdikaris, A.; Blouhos, P.; Yialouris, C.; Frossiniotis, D.; Anthopoulos, I.; Maggana, O.; Nomikou, K.; Rodeva, V.; Kostova, D.; Grozeva, S.; Michaelides, A.; Simonian, A.; Kintzios, S. (2008) Engineering of the membrane of fibroblast cells with
1405:
The benefit of such methods is the high level of detail of the active conduction mechanism, given by the inherently high resolution and data density that atomistic simulation affords. There are significant drawbacks, given by the uncertainty of the legitimacy of the model and the computational cost
1307:
The bioelectric recognition assay (BERA) is a novel method for determination of various chemical and biological molecules by measuring changes in the membrane potential of cells immobilized in a gel matrix. Apart from the increased stability of the electrode-cell interface, immobilization preserves
1033:
In situations where one wants to record the potential inside the cell membrane with minimal effect on the ionic constitution of the intracellular fluid a sharp electrode can be used. These micropipettes (electrodes) are again like those for patch clamp pulled from glass capillaries, but the pore is
1106:
and postsynaptic neuron. This is meant to represent a large population of synapses and neurons. When the synapse releases glutamate onto the postsynaptic cell, it opens ionotropic glutamate receptor channels. The net flow of current is inward, so a current sink is generated. A nearby electrode
1075:
To prepare the brain for such electrode insertion, delicate slicing devices like the compresstome vibratome, leica vibratome, microtome are often employed. These instruments aid in obtaining precise, thin brain sections necessary for electrode placement, enabling neuroscientists to target specific
874:
Today, most microelectrodes used for intracellular recording are glass micropipettes, with a tip diameter of < 1 micrometre, and a resistance of several megohms. The micropipettes are filled with a solution that has a similar ionic composition to the intracellular fluid of the cell. A chlorided
215:
Intracellular activity may also be observed using a specially formed (hollow) glass pipette containing an electrolyte. In this technique, the microscopic pipette tip is pressed against the cell membrane, to which it tightly adheres by an interaction between glass and lipids of the cell membrane.
1089:
and is suitable in areas where there are identified types of cells with well defined spike characteristics. If the electrode tip is bigger still, in general the activity of individual neurons cannot be distinguished but the electrode will still be able to record a field potential generated by the
1375:
A recent advance is the development of a technique called molecular identification through membrane engineering (MIME). This technique allows for building cells with defined specificity for virtually any molecule of interest, by embedding thousands of artificial receptors into the cell membrane.
1020:
who received the Nobel Prize in 1991. Conventional intracellular recording involves impaling a cell with a fine electrode; patch-clamp recording takes a different approach. A patch-clamp microelectrode is a micropipette with a relatively large tip diameter. The microelectrode is placed next to a
949:
by injecting current into a cell through the recording electrode. Unlike in the voltage clamp mode, where the membrane potential is held at a level determined by the experimenter, in "current clamp" mode the membrane potential is free to vary, and the amplifier records whatever voltage the cell
909:
The voltage clamp uses a negative feedback mechanism. The membrane potential amplifier measures membrane voltage and sends output to the feedback amplifier. The feedback amplifier subtracts the membrane voltage from the command voltage, which it receives from the signal generator. This signal is
199:
Neuronal electrophysiology is the study of electrical properties of biological cells and tissues within the nervous system. With neuronal electrophysiology doctors and specialists can determine how neuronal disorders happen, by looking at the individual's brain activity. Activity such as which
882:
Maintaining healthy brain slices is pivotal for successful electrophysiological recordings. The preparation of these slices is commonly achieved with tools such as the
Compresstome vibratome, ensuring optimal conditions for accurate and reliable recordings. Nevertheless, even with the highest
1316:
by changing the cell membrane potential. In this way, when a positive sample is added to the sensor, a characteristic, "signature-like" change in electrical potential occurs. BERA is the core technology behind the recently launched pan-European FOODSCAN project, about pesticide and food risk
2092:
Kokla A, Blouchos P., Livaniou E., Zikos C., Kakabakos S.E., Petrou P.S., Kintzios, S. (2013) Visualization of the membrane-engineering concept: evidence for the specific orientation of electroinserted antibodies and selective binding of target analytes. Journal of
Molecular Recognition 26:
1985:
Perdikaris, A.; Alexandropoulos, N; Kintzios, S. (2009) Development of a Novel, Ultra-rapid
Biosensor for the Qualitative Detection of Hepatitis B Virus-associated Antigens and Anti-HBV, Based on "Membrane-engineered" Fibroblast Cells with Virus-Specific Antibodies and Antigens. Sensors 9:
1236:
1217:
Schematic drawing of the classical patch clamp configuration. The patch pipette is moved to the cell using a micromanipulator under optical control. Relative movements between the pipette and the cell have to be avoided in order to keep the cell-pipette connection
231:
The electrophysiologist may choose not to insert the tip into a single cell. Instead, the electrode tip may be left in continuity with the extracellular space. If the tip is small enough, such a configuration may allow indirect observation and recording of
2082:
Ferentinos K.P., C.P. Yialouris, P. Blouchos, G. Moschopoulou, V. Tsourou, Kintzios, S. (2013) Pesticide
Residue Screening Using a Novel Artificial Neural Network Combined with a Bioelectric Cellular Biosensor. BioMed Research International. Article ID
251:. Still larger electrodes, such as uninsulated needles and surface electrodes used by clinical and surgical neurophysiologists, are sensitive only to certain types of synchronous activity within populations of cells numbering in the millions.
310:) often carries the specific meaning of intracardiac electrogram, which is like an electrocardiogram but with some invasive leads (inside the heart) rather than only noninvasive leads (on the skin). Electrophysiological recording for clinical
2072:
Moschopoulou, G., Valero, T., Kintzios, S. (2012) Superoxide determination using membrane-engineered cells: An example of a novel concept for the construction of cell sensors with customized target recognition properties. Sens. Actuat.175:
1225:
2062:
Moschopoulou G., Kintzios S. (2006) Application of "membrane-engineering" to bioelectric recognition cell sensors for the detection of picomole concentrations of superoxide radical: a novel biosensor principle. Anal. Chimica Acta 573–74:
879:. So an electrode is a compromise between size (small enough to penetrate a single cell with minimum damage to the cell) and resistance (low enough so that small neuronal signals can be discerned from thermal noise in the electrode tip).
1293:
monolayer. Because the painted membrane is supported by the electrode, it is called a solid-supported membrane. Mechanical perturbations, which usually destroy a biological lipid membrane, do not influence the life-time of an SSM. The
1038:
to fill the cells recorded from, for later confirmation of their morphology under a microscope. The dyes are injected by applying a positive or negative, DC or pulsed voltage to the electrodes depending on the polarity of the dye.
1174:
containing a microstructured aperture. A single cell is then positioned on the hole by suction and a tight connection (Gigaseal) is formed. The planar geometry offers a variety of advantages compared to the classical experiment:
216:
The electrolyte within the pipette may be brought into fluid continuity with the cytoplasm by delivering a pulse of negative pressure to the pipette in order to rupture the small patch of membrane encircled by the pipette rim (
1126:. The diagram to the right shows hippocampal synaptic field potentials. At the right, the lower trace shows a negative wave that corresponds to a current sink caused by positive charges entering cells through postsynaptic
986:). It then instructs a parallel circuit that has a large current source behind it (the electrical mains) and adjusts the resistance of that parallel circuit to give the same output voltage, but across a lower resistance.
969:
electronics. The amplifier increases the current behind the signal while decreasing the resistance over which that current passes. Consider this example based on Ohm's law: A voltage of 10 mV is generated by passing 10
862:
won the Nobel Prize in
Physiology or Medicine for their contribution to understanding the mechanisms underlying the generation of action potentials in neurons. Their experiments involved intracellular recordings from the
838:
and fluorescing proteins. After introducing one or more such compounds into tissue via perfusion, injection or gene expression, the 1 or 2-dimensional distribution of electrical activity may be observed and recorded.
834:
distributed phenomenon. Interest in the spatial distribution of bioelectric activity prompted development of molecules capable of emitting light in response to their electrical or chemical environment. Examples are
2014:
Mavrikou, S, Flampouri, E, Moschopoulou, G, Mangana, O, Michaelides, A, Kintzios, S (2008) Assessment of organophosphate and carbamate pesticide residues in cigarette tobacco with a novel cell biosensor. Sensors 8:
950:
generates on its own or as a result of stimulation. This technique is used to study how a cell responds when electric current enters a cell; this is important for instance for understanding how neurons respond to
1341:), and plant viruses (tobacco and cucumber viruses) in a specific, rapid (1–2 minutes), reproducible, and cost-efficient fashion. The method has also been used for the detection of environmental toxins, such as
2033:
Larou, E., Yiakoumettis, I., Kaltsas, G., Petropoulos, A., Skandamis, P., Kintzios, S. (2012) High throughput cellular biosensor for the ultra-sensitive, ultra-rapid detection of aflatoxin M1. Food
Control 29:
965:, sometimes referred to as a "unity gain amplifier"; its main purpose is to reduce the electrical load on the small signals (in the mV range) produced by cells so that they can be accurately recorded by low-
2052:
Apostolou T, Pascual N, Marco M-P, Moschos A, Petropoulos A, Kaltsas G, Kintzios S (2014) Extraction-less, rapid assay for the direct detection of 2,4,6-trichloroanisole (TCA) in cork samples. Talanta 125:
1248:
1071:
of the anesthetized cat, and showed how single neurons in this area respond to very specific features of a visual stimulus. Hubel and Wiesel were awarded the Nobel Prize in
Physiology or Medicine in 1981.
1085:
Recordings from one or more such electrodes that are closely spaced can be used to identify the number of cells around it as well as which of the spikes come from which cell. This process is called
850:
involves measuring voltage and/or current across the membrane of a cell. To make an intracellular recording, the tip of a fine (sharp) microelectrode must be inserted inside the cell, so that the
1281:, or membrane fragments containing the channel or transporter of interest are adsorbed to a lipid monolayer painted over a functionalized electrode. This electrode consists of a glass support, a
1146:
lose or gain electrons at characteristic voltages, individual species can be identified. Amperometry has been used for studying exocytosis in the nervous and endocrine systems. Many monoamine
220:). Alternatively, ionic continuity may be established by "perforating" the patch by allowing exogenous pore-forming agents within the electrolyte to insert themselves into the membrane patch (
1976:
Kintzios S., E. Pistola, P. Panagiotopoulos, M. Bomsel, N. Alexandropoulos, F. Bem, I. Biselis, R. Levin (2001) Bioelectric recognition assay (BERA). Biosensors and
Bioelectronics 16: 325–36
1242:
In planar patch configuration, the cell is positioned by suction. Relative movements between cell and aperture can then be excluded after sealing. An antivibration table is not necessary.
2005:
Flampouri E, Mavrikou S, Kintzios S, Miliaids G, Aplada-Sarli P (2010). Development and
Validation of a Cellular Biosensor Detecting Pesticide Residues in Tomatoes. Talanta 80: 1799–804.
854:
can be measured. Typically, the resting membrane potential of a healthy cell will be -60 to -80 mV, and during an action potential the membrane potential might reach +40 mV. In 1963,
153:) in biological tissues and, in particular, to the electrical recording techniques that enable the measurement of this flow. Classical electrophysiology techniques involve placing
1170:
Planar patch clamp is a novel method developed for high throughput electrophysiology. Instead of positioning a pipette on an adherent cell, cell suspension is pipetted on a
130:"Current Clamp" is a common technique in electrophysiology. This is a whole-cell current clamp recording of a neuron firing due to its being depolarized by current injection
1130:, while the upper trace shows a positive wave that is generated by the current that leaves the cell (at the cell body) to complete the circuit. For more information, see
2217:
Gibson, Frank; Overton, Paul G.; Smulders, Tom V.; Schultz, Simon R.; Eglen, Stephen J.; Ingram, Colin D.; Panzeri, Stefano; Bream, Phil; Sernagor, Evelyne (2008).
1025:
by extracting the cellular contents following recording in order to characterize the electrophysiological properties relationship to gene expression and cell-type.
2043:
Varelas, V., Sanvicens N, Marco MP, Kintzios S (2010) Development of a cellular biosensor for the detection of 2, 4, 6- trichloroanisole (TCA). Talanta 84: 936–40
1777:
Halliwell J., Whitaker M., Ogden D. (1994) Using microelectrodes. Microelectrode techniques: the
Plymouth Workshop handbook. ed. Ogden, D. Available online at
1424:
is the study of the electrical properties which govern heart rhythm and activity. Cardiac electrophysiology can be used to observe and treat disorders such as
2024:
Lokka K., Skandamis P., Kintzios S. (2013) Screening of Total Organophosphate Pesticides in Agricultural Products with a Cellular Biosensor CellBio 2: 131–37.
240:. Depending on the preparation and precise placement, an extracellular configuration may pick up the activity of several nearby cells simultaneously, termed
1428:(irregular heartbeat). For example, a doctor may insert a catheter containing an electrode into the heart to record the heart muscle's electrical activity.
1384:
While not strictly constituting an experimental measurement, methods have been developed to examine the conductive properties of proteins and biomembranes
212:
electrical activity of a single cell. However, this invasive setup reduces the life of the cell and causes a leak of substances across the cell membrane.
1162:(5-HT) are oxidizable. The method can also be used with cells that do not secrete oxidizable neurotransmitters by "loading" them with 5-HT or dopamine.
1118:
are local current sinks or sources that are generated by the collective activity of many cells. Usually, a field potential is generated by the
961:
Most current-clamp amplifiers provide little or no amplification of the voltage changes recorded from the cell. The "amplifier" is actually an
1847:
1756:
1420:
Clinical electrophysiology is the study of how electrophysiological principles and technologies can be applied to human health. For example,
247:
As electrode size increases, the resolving power decreases. Larger electrodes are sensitive only to the net activity of many cells, termed
1549:
2284:
200:
portions of the brain light up during any situations encountered. If an electrode is small enough (micrometers) in diameter, then the
180:
1693:"Microglia contribute to neuronal synchrony despite endogenous ATP-related phenotypic transformation in acute mouse brain slices"
17:
294:), and the specific types of electrophysiological recording are usually called by specific names, constructed on the pattern of
201:
1224:
2253:
1917:
1398:
is subjected to an externally applied voltage. Studies using these setups have been able to study dynamical phenomena like
2105:"Ion Leakage through Transient Water Pores in Protein-Free Lipid Membranes Driven by Transmembrane Ionic Charge Imbalance"
1534:
2162:"Computational Electrophysiology: The Molecular Dynamics of Ion Channel Permeation and Selectivity in Atomistic Detail"
1504:
1421:
1102:
A schematic diagram showing a field potential recording from rat hippocampus. At the left is a schematic diagram of a
1111:
electrode placed inside the cell body (#1) records the change in membrane potential that the incoming current causes.
217:
2218:
2277:
2353:
1467:
1254:
Scanning electron microscope image of a planar patch clamp chip. Both the pipette and the chip are made from
1115:
978:
of resistance. The electrometer changes this "high impedance signal" to a "low impedance signal" by using a
2358:
1996:
virus-specific antibodies: a novel biosensor tool for virus detection. Biosensors Bioelectron. 24: 1033–36.
1648:"Healthy Brain-pituitary Slices for Electrophysiological Investigations of Pituitary Cells in Teleost Fish"
205:
161:
Simple solid conductors, such as discs and needles (singles or arrays, often insulated except for the tip),
167:
Hollow, often elongated or 'pulled', tubes filled with an electrolyte, such as glass pipettes filled with
2436:
1509:
1415:
116:
2270:
1791:
1529:
1499:
1191:
982:
circuit. A voltage follower reads the voltage on the input (caused by a small current through a big
927:
315:
204:
may choose to insert the tip into a single cell. Such a configuration allows direct observation and
2451:
1514:
1432:
2405:
2323:
1519:
1368:
621:
282:, "electrical recording"), with the record thus produced being an electrogram. However, the word
1792:"Receptive fields, binocular interaction and functional architecture in the cat's visual cortex"
2390:
1350:
1330:
1313:
1119:
1002:
uses a micropipette attached to the cell membrane to allow recording from a single ion channel.
859:
416:
248:
164:
Tracings on printed circuit boards or flexible polymers, also insulated except for the tip, and
112:
1942:
Schulz, Patrick; Garcia-Celma, Juan J.; Fendler, Klaus (2008). "SSM-based electrophysiology".
1839:
1748:
2446:
2441:
1484:
1452:
1131:
1123:
1053:
876:
835:
546:
527:
237:
120:
2410:
2173:
2116:
1704:
1579:
1448:
1103:
966:
758:
708:
584:
565:
554:
474:
439:
241:
221:
8:
2456:
1494:
1489:
655:
504:
338:
291:
149:
Electrophysiology is the branch of physiology that pertains broadly to the flow of ions (
2177:
2120:
1725:
1708:
1692:
1583:
157:
into various preparations of biological tissue. The principal types of electrodes are:
2194:
2161:
2137:
2104:
1887:
1870:
1816:
1668:
1647:
1603:
1391:
1338:
1127:
946:
922:
crosses a cell's membrane at any given voltage. This is important because many of the
915:
855:
851:
637:
573:
485:
168:
1570:
Scanziani, Massimo; Häusser, Michael (2009). "Electrophysiology in the age of light".
2318:
2199:
2142:
1959:
1892:
1821:
1730:
1673:
1607:
1595:
1544:
1278:
1147:
884:
817:
602:
384:
311:
126:
2348:
2313:
2250:
2229:
2189:
2181:
2160:
Kutzner, Carsten; Grubmüller, Helmut; De Groot, Bert L.; Zachariae, Ulrich (2011).
2132:
2124:
1951:
1910:
1882:
1811:
1807:
1803:
1720:
1712:
1663:
1655:
1587:
1022:
979:
951:
864:
458:
272:
233:
104:
81:
2233:
1308:
the viability and physiological functions of the cells. BERA is used primarily in
270:
Electrophysiological recording in general is sometimes called electrography (from
115:, may also be referred to as electrophysiological recordings. They are useful for
2415:
2374:
2308:
2257:
2219:"Minimum Information about a Neuroscience Investigation (MINI) Electrophysiology"
1539:
1456:
1399:
1353:
in cork and wine, as well as the determination of very low concentrations of the
470:
428:
278:
2128:
1955:
1716:
1621:
1524:
1151:
1064:
1035:
405:
367:
350:
108:
73:
2333:
2185:
1634:
1435:. In this medical specialty, doctors measure the electrical properties of the
887:
induces rapid and robust phenotype changes of the brain's major immune cells,
2430:
2395:
1395:
1317:
assessment in Europe. BERA has been used for the detection of human viruses (
1201:
1180:
1086:
1068:
939:
900:
209:
34:
1098:
2203:
2146:
1963:
1896:
1825:
1734:
1677:
1633:
Movie featuring Alan Hodgkin recording action potentials from a squid axon
1599:
1255:
1017:
962:
955:
871:), and were among the first applications of the "voltage clamp" technique.
96:
1440:
1322:
1318:
1295:
1184:
1142:
1060:
1013:
1007:
999:
923:
801:
776:
259:
255:
225:
150:
85:
1591:
2293:
1778:
1425:
1354:
1342:
1290:
975:
971:
307:
287:
154:
69:
1471:
1386:
1346:
1309:
1197:
1159:
888:
2262:
1282:
1274:
1155:
983:
732:
27:
Study of the electrical properties of biological cells and tissues.
2159:
1659:
914:
The voltage clamp technique allows an experimenter to "clamp" the
1455:(1924–2006) are considered to have greatly advanced the field of
1171:
910:
amplified and returned into the cell via the recording electrode.
891:, which must be taken into consideration when using this model.
645:
610:
592:
100:
88:
77:
1299:
via capacitive coupling between the vesicles and the electrode.
905:
450:(specifically the cerebral cortex), with intracranial electrodes
1444:
1326:
748:
697:
680:
663:
516:
466:
994:
918:
at a chosen value. This makes it possible to measure how much
107:
activity. Recordings of large-scale electric signals from the
1436:
1334:
766:
447:
424:
401:
346:
92:
64:
57:
47:
37:
2216:
139:
1286:
784:
716:
828:
189:
dissociated cells from excised tissue (acute or cultured),
1941:
535:
512:
493:
265:
99:, it includes measurements of the electrical activity of
84:
or manipulations on a wide variety of scales from single
1691:
Peter, Berki; Csaba, Cserep; Zsuzsanna, Környei (2024).
254:
Other classical electrophysiological techniques include
1302:
1268:
1067:
recorded the activity of single neurons in the primary
1470:
or reporting guidelines specify the minimum amount of
1312:
in order to assay analytes that can interact with the
1230:
Scanning electron microscope image of a patch pipette.
2102:
1645:
72:
that studies the electrical properties of biological
1690:
1183:, which enables automatic compound application for
1379:
683:flow via streaming potential detected through skin
2103:Gurtovenko, Andrey A.; Vattulainen, Ilpo (2007).
1789:
1646:Fontaine, R.; Hodne, K.; Weltzien, F. A. (2018).
1431:Another example of clinical electrophysiology is
2428:
1840:"The Nobel Prize in Physiology or Medicine 1981"
1749:"The Nobel Prize in Physiology or Medicine 1991"
1569:
1462:
1402:of membranes and ion translocation by channels.
1273:With this electrophysiological approach, proteo
2278:
1868:
1409:
1367:The electronic read-out device with embedded
1028:
314:purposes is included within the category of
1635:https://www.youtube.com/watch?v=k48jXzFGMc8
1550:Transcutaneous electrical nerve stimulation
1394:simulations in which a model system like a
224:). Finally, the patch may be left intact (
144:
2285:
2271:
1790:D. H. Hubel; Wiesel, TN (1 January 1962).
1684:
1042:
842:
408:), with intracardiac electrodes (invasive)
353:), with cutaneous electrodes (noninvasive)
318:. The various "ExG" modes are as follows:
76:and tissues. It involves measurements of
2193:
2136:
1886:
1815:
1724:
1667:
171:solution or another electrolyte solution.
140:Classical electrophysiological techniques
1190:The system is accessible for optical or
1097:
1047:
993:
989:
945:The current clamp technique records the
904:
302:(abbreviation ExG). Relatedly, the word
125:
1107:(#2) detects this as a negativity. An
1079:
829:Optical electrophysiological techniques
192:artificially grown cells or tissues, or
134:
14:
2429:
1459:, enabling its clinical applications.
1364:The consumable biorecognition elements
266:Electrographic modalities by body part
2266:
1850:from the original on 23 December 2017
1329:viruses), veterinary disease agents (
1165:
2292:
1779:http://plymsea.ac.uk/id/eprint/7954/
1759:from the original on 10 October 2017
1303:Bioelectric recognition assay (BERA)
1269:Solid-supported membrane (SSM)-based
175:The principal preparations include:
1869:Papouin, T.; Haydon, P. G. (2018).
1535:Multiscale Electrophysiology Format
1093:
186:excised tissue (acute or cultured),
24:
2251:Book chapter on Planar Patch Clamp
1923:from the original on 31 March 2010
1505:Clinical cardiac electrophysiology
1468:Minimum Information (MI) standards
1422:clinical cardiac electrophysiology
306:(not being needed for those other
25:
2468:
2244:
1652:Journal of Visualized Experiments
2332:
1263:
1247:
1235:
1223:
1210:
1012:This technique was developed by
933:
926:in the membrane of a neuron are
894:
538:—via the corneoretinal potential
2210:
2153:
2096:
2086:
2076:
2066:
2056:
2046:
2037:
2027:
2018:
2008:
1999:
1989:
1979:
1970:
1935:
1903:
1862:
1380:Computational electrophysiology
629:stomach and bowel smooth muscle
431:), with extracranial electrodes
1871:"Obtaining Acute Brain Slices"
1832:
1808:10.1113/jphysiol.1962.sp006837
1783:
1771:
1741:
1639:
1627:
1614:
1563:
1137:
1116:Extracellular field potentials
883:standards of tissue handling,
13:
1:
2359:Physiological plant disorders
2354:Plant perception (physiology)
1556:
1463:Clinical reporting guidelines
1360:A BERA sensor has two parts:
1179:It allows for integration of
1076:brain regions for recording.
954:that act by opening membrane
469:throughout the body (usually
579:4—not applicable clinically
7:
2129:10.1529/biophysj.106.094797
1956:10.1016/j.ymeth.2008.07.002
1477:
1357:anion in clinical samples.
333:Prevalence in clinical use
236:from a single cell, termed
10:
2473:
1717:10.1038/s41467-024-49773-1
1510:Clinical electrophysiology
1416:Clinical electrophysiology
1413:
1410:Clinical electrophysiology
1051:
1005:
937:
928:voltage-gated ion channels
898:
222:perforated patch recording
179:living organisms (example
58:
48:
38:
2383:
2367:
2341:
2330:
2301:
2186:10.1016/j.bpj.2011.06.010
1796:The Journal of Physiology
1530:History of bioelectricity
1500:Cardiac electrophysiology
1029:Sharp electrode recording
622:electrogastroenterography
316:electrodiagnostic testing
91:to whole organs like the
1515:Clinical neurophysiology
1433:clinical neurophysiology
1289:layer, and an octadecyl
1090:activity of many cells.
938:Not to be confused with
395:intracardiac electrogram
256:single channel recording
145:Principle and mechanisms
56:, "nature, origin"; and
2406:Evolutionary physiology
2324:Physiology of dinosaurs
1911:"Automated patch clamp"
1520:Electrophysiology study
1369:artificial intelligence
1120:simultaneous activation
1043:Extracellular recording
848:Intracellular recording
843:Intracellular recording
378:electroventriculography
206:intracellular recording
18:Intracellular recording
2391:Comparative physiology
2234:10101/npre.2009.1720.2
1351:2,4,6-trichloroanisole
1331:foot and mouth disease
1310:biosensor applications
1204:side can be performed.
1112:
1003:
911:
860:Andrew Fielding Huxley
836:voltage sensitive dyes
726:electropancreatography
417:electroencephalography
249:local field potentials
131:
113:electroencephalography
103:, and, in particular,
2256:31 March 2010 at the
1697:Nature Communications
1485:Automated patch clamp
1453:Nathaniel A. Buchwald
1447:. Scientists such as
1132:local field potential
1124:synaptic transmission
1101:
1054:single-unit recording
1048:Single-unit recording
997:
990:Patch-clamp recording
908:
877:electrical resistance
691:electroblepharography
576:in arthropod antennae
528:electronystagmography
238:single-unit recording
195:hybrids of the above.
129:
2411:Molecular physiology
1622:U.S. patent 4425922A
1449:Duchenne de Boulogne
1104:presynaptic terminal
1080:Multi-unit recording
974:of current across 1
759:electroneuronography
742:electrohysterography
674:electroarteriography
585:electrocochleography
566:electroantennography
555:olfactory epithelium
547:electroolfactography
440:electrocorticography
242:multi-unit recording
218:whole-cell recording
135:Definition and scope
2178:2011BpJ...101..809K
2166:Biophysical Journal
2121:2007BpJ....92.1878G
2109:Biophysical Journal
1709:2024NatCo..15.5402B
1592:10.1038/nature08540
1584:2009Natur.461..930S
1495:Bioelectromagnetics
1490:Bioelectrochemistry
1390:. These are mainly
1128:glutamate receptors
1122:of many neurons by
867:of Atlantic squid (
811:electrovomerography
777:electropneumography
656:electropalatography
638:electroglottography
603:electrogastrography
574:olfactory receptors
505:electroretinography
404:(specifically, the
349:(specifically, the
339:electrocardiography
202:electrophysiologist
68:) is the branch of
1392:molecular dynamics
1166:Planar patch clamp
1113:
1004:
998:The cell-attached
947:membrane potential
912:
856:Alan Lloyd Hodgkin
852:membrane potential
795:electrospinography
709:electrodermography
632:2—somewhat common
616:2—somewhat common
597:2—somewhat common
541:2—somewhat common
522:2—somewhat common
499:2—somewhat common
486:electrooculography
453:2—somewhat common
434:2—somewhat common
411:2—somewhat common
361:electroatriography
292:electrophotography
169:potassium chloride
132:
2437:Electrophysiology
2424:
2423:
2401:Electrophysiology
2319:Insect physiology
2226:Nature Precedings
1545:Slice preparation
1339:blue tongue virus
1314:immobilized cells
1148:neurotransmitters
952:neurotransmitters
885:slice preparation
826:
825:
818:vomeronasal organ
787:(chest movements)
666:contact of tongue
234:action potentials
31:Electrophysiology
16:(Redirected from
2464:
2349:Plant physiology
2336:
2314:Human physiology
2287:
2280:
2273:
2264:
2263:
2238:
2237:
2223:
2214:
2208:
2207:
2197:
2157:
2151:
2150:
2140:
2100:
2094:
2090:
2084:
2080:
2074:
2070:
2064:
2060:
2054:
2050:
2044:
2041:
2035:
2031:
2025:
2022:
2016:
2012:
2006:
2003:
1997:
1993:
1987:
1983:
1977:
1974:
1968:
1967:
1939:
1933:
1932:
1930:
1928:
1922:
1915:
1907:
1901:
1900:
1890:
1866:
1860:
1859:
1857:
1855:
1836:
1830:
1829:
1819:
1787:
1781:
1775:
1769:
1768:
1766:
1764:
1745:
1739:
1738:
1728:
1688:
1682:
1681:
1671:
1643:
1637:
1631:
1625:
1624:
1618:
1612:
1611:
1578:(7266): 930–39.
1567:
1451:(1806–1875) and
1251:
1239:
1227:
1214:
1154:(noradrenalin),
1094:Field potentials
980:voltage follower
459:electromyography
321:
320:
117:electrodiagnosis
105:action potential
82:electric current
61:
60:
51:
50:
41:
40:
21:
2472:
2471:
2467:
2466:
2465:
2463:
2462:
2461:
2452:Neurophysiology
2427:
2426:
2425:
2420:
2416:Neurophysiology
2379:
2375:Cell physiology
2363:
2337:
2328:
2309:Fish physiology
2297:
2291:
2258:Wayback Machine
2247:
2242:
2241:
2221:
2215:
2211:
2158:
2154:
2101:
2097:
2091:
2087:
2081:
2077:
2071:
2067:
2061:
2057:
2051:
2047:
2042:
2038:
2032:
2028:
2023:
2019:
2013:
2009:
2004:
2000:
1994:
1990:
1984:
1980:
1975:
1971:
1940:
1936:
1926:
1924:
1920:
1913:
1909:
1908:
1904:
1867:
1863:
1853:
1851:
1838:
1837:
1833:
1788:
1784:
1776:
1772:
1762:
1760:
1747:
1746:
1742:
1689:
1685:
1644:
1640:
1632:
1628:
1620:
1619:
1615:
1568:
1564:
1559:
1554:
1540:Neurophysiology
1480:
1465:
1457:neurophysiology
1418:
1412:
1400:electroporation
1382:
1305:
1271:
1266:
1259:
1252:
1243:
1240:
1231:
1228:
1219:
1215:
1168:
1140:
1096:
1082:
1056:
1050:
1045:
1031:
1010:
992:
943:
942:in electronics.
936:
903:
897:
845:
831:
473:, occasionally
429:cerebral cortex
268:
226:patch recording
147:
142:
137:
28:
23:
22:
15:
12:
11:
5:
2470:
2460:
2459:
2454:
2449:
2444:
2439:
2422:
2421:
2419:
2418:
2413:
2408:
2403:
2398:
2393:
2387:
2385:
2384:Related topics
2381:
2380:
2378:
2377:
2371:
2369:
2365:
2364:
2362:
2361:
2356:
2351:
2345:
2343:
2339:
2338:
2331:
2329:
2327:
2326:
2321:
2316:
2311:
2305:
2303:
2299:
2298:
2290:
2289:
2282:
2275:
2267:
2261:
2260:
2246:
2245:External links
2243:
2240:
2239:
2209:
2152:
2115:(6): 1878–90.
2095:
2085:
2075:
2065:
2055:
2045:
2036:
2026:
2017:
2007:
1998:
1988:
1978:
1969:
1934:
1902:
1861:
1844:nobelprize.org
1831:
1782:
1770:
1753:nobelprize.org
1740:
1683:
1638:
1626:
1613:
1561:
1560:
1558:
1555:
1553:
1552:
1547:
1542:
1537:
1532:
1527:
1525:Hille equation
1522:
1517:
1512:
1507:
1502:
1497:
1492:
1487:
1481:
1479:
1476:
1464:
1461:
1414:Main article:
1411:
1408:
1381:
1378:
1373:
1372:
1365:
1304:
1301:
1270:
1267:
1265:
1262:
1261:
1260:
1253:
1246:
1244:
1241:
1234:
1232:
1229:
1222:
1220:
1216:
1209:
1206:
1205:
1195:
1192:scanning probe
1188:
1167:
1164:
1152:norepinephrine
1139:
1136:
1095:
1092:
1081:
1078:
1065:Torsten Wiesel
1052:Main article:
1049:
1046:
1044:
1041:
1036:Lucifer yellow
1030:
1027:
1006:Main article:
991:
988:
935:
932:
916:cell potential
899:Main article:
896:
893:
844:
841:
830:
827:
824:
823:
820:
815:
812:
808:
807:
804:
799:
796:
792:
791:
788:
782:
779:
773:
772:
769:
764:
761:
755:
754:
751:
746:
743:
739:
738:
735:
730:
727:
723:
722:
719:
714:
711:
705:
704:
701:
695:
692:
688:
687:
684:
678:
675:
671:
670:
667:
661:
658:
652:
651:
648:
643:
640:
634:
633:
630:
627:
624:
618:
617:
614:
608:
605:
599:
598:
595:
590:
589:ECOG or ECochG
587:
581:
580:
577:
571:
568:
562:
561:
558:
552:
549:
543:
542:
539:
533:
530:
524:
523:
520:
510:
507:
501:
500:
497:
491:
488:
482:
481:
480:1—very common
478:
464:
461:
455:
454:
451:
445:
442:
436:
435:
432:
422:
419:
413:
412:
409:
406:cardiac muscle
399:
396:
392:
391:
388:
387:cardiac muscle
382:
379:
375:
374:
371:
370:cardiac muscle
365:
362:
358:
357:
356:1—very common
354:
351:cardiac muscle
344:
341:
335:
334:
331:
328:
325:
267:
264:
197:
196:
193:
190:
187:
184:
173:
172:
165:
162:
146:
143:
141:
138:
136:
133:
109:nervous system
26:
9:
6:
4:
3:
2:
2469:
2458:
2455:
2453:
2450:
2448:
2445:
2443:
2440:
2438:
2435:
2434:
2432:
2417:
2414:
2412:
2409:
2407:
2404:
2402:
2399:
2397:
2396:Ecophysiology
2394:
2392:
2389:
2388:
2386:
2382:
2376:
2373:
2372:
2370:
2366:
2360:
2357:
2355:
2352:
2350:
2347:
2346:
2344:
2340:
2335:
2325:
2322:
2320:
2317:
2315:
2312:
2310:
2307:
2306:
2304:
2300:
2295:
2288:
2283:
2281:
2276:
2274:
2269:
2268:
2265:
2259:
2255:
2252:
2249:
2248:
2235:
2231:
2227:
2220:
2213:
2205:
2201:
2196:
2191:
2187:
2183:
2179:
2175:
2172:(4): 809–17.
2171:
2167:
2163:
2156:
2148:
2144:
2139:
2134:
2130:
2126:
2122:
2118:
2114:
2110:
2106:
2099:
2089:
2079:
2069:
2059:
2049:
2040:
2030:
2021:
2011:
2002:
1992:
1982:
1973:
1965:
1961:
1957:
1953:
1950:(2): 97–103.
1949:
1945:
1938:
1919:
1912:
1906:
1898:
1894:
1889:
1884:
1880:
1876:
1872:
1865:
1849:
1845:
1841:
1835:
1827:
1823:
1818:
1813:
1809:
1805:
1802:(1): 106–54.
1801:
1797:
1793:
1786:
1780:
1774:
1758:
1754:
1750:
1744:
1736:
1732:
1727:
1722:
1718:
1714:
1710:
1706:
1702:
1698:
1694:
1687:
1679:
1675:
1670:
1665:
1661:
1660:10.3791/57790
1657:
1653:
1649:
1642:
1636:
1630:
1623:
1617:
1609:
1605:
1601:
1597:
1593:
1589:
1585:
1581:
1577:
1573:
1566:
1562:
1551:
1548:
1546:
1543:
1541:
1538:
1536:
1533:
1531:
1528:
1526:
1523:
1521:
1518:
1516:
1513:
1511:
1508:
1506:
1503:
1501:
1498:
1496:
1493:
1491:
1488:
1486:
1483:
1482:
1475:
1473:
1469:
1460:
1458:
1454:
1450:
1446:
1442:
1438:
1434:
1429:
1427:
1423:
1417:
1407:
1403:
1401:
1397:
1396:lipid bilayer
1393:
1389:
1388:
1377:
1370:
1366:
1363:
1362:
1361:
1358:
1356:
1352:
1349:in food, and
1348:
1344:
1340:
1336:
1332:
1328:
1324:
1320:
1315:
1311:
1300:
1297:
1292:
1288:
1284:
1280:
1276:
1264:Other methods
1257:
1250:
1245:
1238:
1233:
1226:
1221:
1213:
1208:
1207:
1203:
1202:intracellular
1199:
1196:
1193:
1189:
1186:
1182:
1181:microfluidics
1178:
1177:
1176:
1173:
1163:
1161:
1157:
1153:
1149:
1144:
1135:
1133:
1129:
1125:
1121:
1117:
1110:
1109:intracellular
1105:
1100:
1091:
1088:
1087:spike sorting
1077:
1073:
1070:
1069:visual cortex
1066:
1062:
1055:
1040:
1037:
1026:
1024:
1019:
1015:
1009:
1001:
996:
987:
985:
981:
977:
973:
968:
964:
959:
957:
953:
948:
941:
940:Current clamp
934:Current clamp
931:
929:
925:
921:
920:ionic current
917:
907:
902:
901:Voltage clamp
895:Voltage clamp
892:
890:
886:
880:
878:
872:
870:
869:Loligo pealei
866:
861:
857:
853:
849:
840:
837:
821:
819:
816:
813:
810:
809:
805:
803:
800:
797:
794:
793:
789:
786:
783:
780:
778:
775:
774:
770:
768:
765:
762:
760:
757:
756:
752:
750:
747:
744:
741:
740:
736:
734:
731:
728:
725:
724:
720:
718:
715:
712:
710:
707:
706:
702:
699:
696:
693:
690:
689:
685:
682:
679:
676:
673:
672:
668:
665:
662:
659:
657:
654:
653:
649:
647:
644:
641:
639:
636:
635:
631:
628:
625:
623:
620:
619:
615:
613:smooth muscle
612:
609:
606:
604:
601:
600:
596:
594:
591:
588:
586:
583:
582:
578:
575:
572:
569:
567:
564:
563:
559:
556:
553:
550:
548:
545:
544:
540:
537:
534:
531:
529:
526:
525:
521:
518:
514:
511:
508:
506:
503:
502:
498:
496:—entire globe
495:
492:
489:
487:
484:
483:
479:
476:
472:
468:
465:
462:
460:
457:
456:
452:
449:
446:
443:
441:
438:
437:
433:
430:
427:(usually the
426:
423:
420:
418:
415:
414:
410:
407:
403:
400:
397:
394:
393:
389:
386:
383:
380:
377:
376:
372:
369:
366:
363:
360:
359:
355:
352:
348:
345:
342:
340:
337:
336:
332:
329:
326:
323:
322:
319:
317:
313:
309:
305:
301:
297:
293:
289:
285:
284:electrography
281:
280:
275:
274:
263:
261:
257:
252:
250:
245:
243:
239:
235:
229:
227:
223:
219:
213:
211:
210:intracellular
207:
203:
194:
191:
188:
185:
182:
178:
177:
176:
170:
166:
163:
160:
159:
158:
156:
152:
128:
124:
122:
118:
114:
110:
106:
102:
98:
94:
90:
87:
83:
79:
75:
71:
67:
66:
55:
45:
36:
32:
19:
2447:Neuroimaging
2442:Ion channels
2400:
2225:
2212:
2169:
2165:
2155:
2112:
2108:
2098:
2088:
2078:
2068:
2058:
2048:
2039:
2029:
2020:
2010:
2001:
1991:
1981:
1972:
1947:
1943:
1937:
1925:. Retrieved
1905:
1881:(2): e2699.
1878:
1875:Bio-Protocol
1874:
1864:
1852:. Retrieved
1843:
1834:
1799:
1795:
1785:
1773:
1761:. Retrieved
1752:
1743:
1700:
1696:
1686:
1651:
1641:
1629:
1616:
1575:
1571:
1565:
1466:
1430:
1419:
1404:
1385:
1383:
1374:
1359:
1325:viruses and
1306:
1272:
1256:borosilicate
1169:
1141:
1114:
1108:
1083:
1074:
1057:
1032:
1018:Bert Sakmann
1011:
963:electrometer
960:
956:ion channels
944:
924:ion channels
919:
913:
881:
873:
868:
847:
846:
832:
763:ENeG or ENoG
519:specifically
444:ECoG or iEEG
327:Abbreviation
303:
299:
295:
283:
277:
271:
269:
253:
246:
230:
214:
198:
174:
148:
97:neuroscience
63:
53:
46:, "amber" ;
43:
30:
29:
1703:(1): 5402.
1441:spinal cord
1319:hepatitis B
1277:, membrane
1194:techniques.
1185:ion channel
1143:Amperometry
1138:Amperometry
1061:David Hubel
1014:Erwin Neher
1008:Patch clamp
1000:patch clamp
972:nanoamperes
822:3—uncommon
806:3—uncommon
802:spinal cord
790:3—uncommon
771:3—uncommon
753:3—uncommon
737:3—uncommon
721:3—uncommon
703:3—uncommon
686:3—uncommon
669:3—uncommon
650:3—uncommon
560:3—uncommon
390:3—uncommon
385:ventricular
373:3—uncommon
304:electrogram
290:(including
260:amperometry
151:ion current
86:ion channel
80:changes or
2457:Biophysics
2431:Categories
2294:Physiology
1927:17 January
1557:References
1426:arrhythmia
1355:superoxide
1347:mycotoxins
1343:pesticides
1296:capacitive
1291:mercaptane
1187:screening.
865:giant axon
557:in mammals
343:ECG or EKG
312:diagnostic
286:has other
181:in insects
155:electrodes
121:monitoring
111:, such as
70:physiology
1608:205218803
1472:meta data
1387:in silico
1285:layer, a
1275:liposomes
1198:Perfusion
1160:serotonin
1023:patch-seq
967:impedance
889:microglia
330:Body part
2254:Archived
2204:21843471
2147:17208976
2093:627–232.
1986:2176–86.
1964:18675360
1918:Archived
1897:29552595
1848:Archived
1826:14449617
1757:Archived
1735:38926390
1726:11208608
1678:30176004
1600:19829373
1478:See also
1283:chromium
1279:vesicles
1156:dopamine
1150:; e.g.,
984:resistor
733:pancreas
681:arterial
471:skeletal
324:Modality
296:electro-
273:electro-
89:proteins
44:ēlektron
2302:Animals
2195:3175076
2174:Bibcode
2138:1861780
2117:Bibcode
2083:813519.
2053:336–40.
2015:2818–32
1944:Methods
1888:5856250
1817:1359523
1705:Bibcode
1669:6126815
1654:(138).
1580:Bibcode
1333:virus,
1218:intact.
1200:of the
664:palatal
646:glottis
611:stomach
593:cochlea
467:muscles
300:-graphy
279:-graphy
208:of the
101:neurons
78:voltage
2342:Plants
2202:
2192:
2145:
2135:
2063:90–96.
2034:208–12
1962:
1895:
1885:
1824:
1814:
1733:
1723:
1676:
1666:
1606:
1598:
1572:Nature
1445:nerves
1443:, and
1337:, and
1335:prions
1327:herpes
1258:glass.
1158:, and
767:nerves
749:uterus
700:muscle
698:eyelid
517:retina
475:smooth
368:atrial
308:senses
288:senses
95:. In
65:-logia
59:-λογία
54:physis
33:(from
2368:Cells
2296:types
2222:(PDF)
2073:88–94
1921:(PDF)
1914:(PDF)
1854:5 May
1763:5 May
1604:S2CID
1437:brain
785:lungs
448:brain
425:brain
402:heart
347:heart
298:+ +
93:heart
74:cells
49:φύσις
39:ἥλεκτ
35:Greek
2200:PMID
2143:PMID
1960:PMID
1929:2010
1893:PMID
1856:2018
1822:PMID
1765:2018
1731:PMID
1674:PMID
1596:PMID
1345:and
1321:and
1287:gold
1172:chip
1063:and
1016:and
858:and
717:skin
626:EGEG
258:and
119:and
2230:hdl
2190:PMC
2182:doi
2170:101
2133:PMC
2125:doi
1952:doi
1883:PMC
1812:PMC
1804:doi
1800:160
1721:PMC
1713:doi
1664:PMC
1656:doi
1588:doi
1576:461
814:EVG
798:ESG
781:EPG
745:EHG
729:EPG
713:EDG
694:EBG
677:EAG
660:EPG
642:EGG
607:EGG
570:EAG
551:EOG
536:eye
532:ENG
513:eye
509:ERG
494:eye
490:EOG
463:EMG
421:EEG
398:EGM
381:EVG
364:EAG
228:).
2433::
2228:.
2224:.
2198:.
2188:.
2180:.
2168:.
2164:.
2141:.
2131:.
2123:.
2113:92
2111:.
2107:.
1958:.
1948:46
1946:.
1916:.
1891:.
1877:.
1873:.
1846:.
1842:.
1820:.
1810:.
1798:.
1794:.
1755:.
1751:.
1729:.
1719:.
1711:.
1701:15
1699:.
1695:.
1672:.
1662:.
1650:.
1602:.
1594:.
1586:.
1574:.
1439:,
1134:.
976:MΩ
958:.
276:+
262:.
244:.
183:),
123:.
62:,
52:,
42:,
2286:e
2279:t
2272:v
2236:.
2232::
2206:.
2184::
2176::
2149:.
2127::
2119::
1966:.
1954::
1931:.
1899:.
1879:8
1858:.
1828:.
1806::
1767:.
1737:.
1715::
1707::
1680:.
1658::
1610:.
1590::
1582::
1371:.
1323:C
515:—
477:)
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
