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+*jj %C<\#255><\#255><\#253><\#178><\#252>}0<\#252>} <\#253><\#250>@Figure 3.1 Fluid mosaic. Transmembrane proteins and lipids are kept together by non-covalent interactions (ionic and hydrophobic). 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V<\#172><\#172>V<\#129><\#129><\#129>+<\#253><\#129>+<\#129>+<\#172>+<\#129><\#228><\#129><\#226><\#129><\#235>V+<\#234>V<\#227><\#129><\#226>++<\#236>V+<\#234>V<\#227><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#255><\#255><\#254>OIonic Fluxes Across the Neuronal Plasma Membrane<\#255><\#254>`!<\#221><\#239><\#144><\#255><\#255>:<\#153><\#225>.3<\#192>HG<\#210><\#135>e<\#146><\#130>b <\#237>KPS@`<\#228>P"_ _<\#160>%.<\#128>jB<\#151>N<\#209>NV<\#255><\#206> O!n1n B(B(B<\#168>p<\#160>` @<\#255><\#206>f<\#160>"n2<\#168>=@N^ _O<\#239> N<\#208>NV<\#255><\#206> O1n<\#160>=@ N^ _T<\#143>N<\#208>Q<\#193>`P<\#193>NV<\#255><\#206> O!n 1n"n !Q$Bh,B<\#168>.Jf<\#160>`<\#160>=@"n "<\#168>(N^"_O<\#239> N<\#209>NV<\#255><\#176> O!n1nB(<\#160>fBh<\#160> fC<\#232> "<\#238>"<\#174> <\#160> =@N^"_<\#218>wx<\#175><\#204><\#176><\#196><\#204><\#237><\#179><\#206><\#233><\#181><\#182><\#183><\#239><\#184><\#253><\#185><\#250><\#129><\#129><\#187><\#188> <\#254><\#190><\#191><\#192><\#193><\#194><\#195> <\#240><\#197><\#198><\#199><\#200><\#201><\#202><\#203><\#255><\#255><\#253>\The neuronal plasma membrane delimits the whole neuron, from the dendritic spines to the axon terminals. It is a barrier between the intracellular and extracellular environments. The general structure of xtracellular environments. The general structure of the neuronal plasma membrane is similar to that of other plasma membranes. It is made up of proteins inserted in a lipid bilayer, forming as a whole a <\#212>fluid mosaic<\#213> (Figure 3.1). However, insofar as there are functions that are exclusively neuronal, the n<\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147><\#129><\#147>BV<\#129><\#252>V<\#252>VV<\#129>V<\#252>V+VV<\#253>V+<\#236><\#129>+<\#244>VV<\#244><\#172><\#247><\#255>+<\#129><\#240><\#129><\#254>V+<\#253>V+<\#149><\#144><\#129>++<\#172><\#254><\#129><\#255><\#253>+<\#129>V++<\#129>+<\#129>++++<\#129>+V++<\#253>++++<\#254>+VV<\#254>+ ++++<\#129>++<\#254>V++<\#172>+++<\#254><\#255><\#129><\#172>+++++++<\#172>+++<\#254>V<\#129>+V<\#254>++<\#254>+<\#162><\#133>+VV<\#253><\#172><\#253>VV<\#253>VV+<\#129>V+VV<\#129>+VV+<\#129>+<\#129><\#255>+<\#254><\#253>V<\#172><\#254>V <\#129>+<\#172>+V<\#129>V<\#172>V<\#129>+<\#249>V,+<\#129><\#129>+<\#172>V+<\#172><\#172><\#129>+V+VV+<\#129>VV<\#129>VV+<\#129>VV<\#172><\#255>++<\#129>V<\#129><\#172><\#252>V<\#172>+<\#172>V+<\#172>+<\#172><\#129>VV<\#129><\#129><\#129><\#161><\#142>+V<\#172><\#253><\#129><\#253><\#129>+++<\#255>V<\#129><\#172><\#129>+<\#129>+<\#172>VV<\#129>+V<\#129><\#172><\#172><\#253>++V<\#172>V<\#172>VV<\#129>+VVVV<\#172><\#129>+<\#172><\#129><\#129>+<\#255>V<\#129><\#172><\#129>++<\#129>+<\#172><\#129>+<\#254><\#129>VV<\#254>-<\#172><\#129>+<\#172><\#255>V<\#129><\#172><\#129>+<\#129><\#172>V<\#180>require energy. Ions being charged particles, ionic fluxes are ionic currents across the membrane and therefore have an immediate effect on membrane potential (electrical signalling). The aim of this chapter is to look at which proteins Na+, K+, Ca2+ and CVVV<\#172><\#129><\#129>+<\#129>++<\#129>+<\#129>++<\#129><\#172><\#172><\#129>+<\#129><\#161><\#137><\#129><\#129><\#253><\#172><\#253>+<\#129>VV<\#129><\#253>V+<\#129>+<\#252>V<\#172><\#172><\#253><\#129>++V<\#129>+<\#254>V<\#129>+V+<\#129>VV<\#129><\#172>V+<\#129><\#254>V<\#254>V<\#172><\#172><\#172><\#254>V+<\#254><\#129><\#129><\#129><\#129>VV+<\#172><\#172>V++V<\#172><\#129>V+VV<\#252>V<\#9>VV<\#129>+<\#129><\#129>VV<\#161><\#140>V<\#129><\#254>V+<\#172><\#252><\#129><\#129>V<\#129>V<\#129><\#129>V<\#129>+<\#129>+<\#129>V<\#129>V<\#129>+V<\#129><\#129><\#172>V<\#172><\#253>0<\#129>V<\#129><\#172><\#172>VV+<\#129><\#129><\#172>VVV<\#172><\#129>+V<\#129><\#129>V<\#129>+V<\#129><\#129>V+<\#129><\#129>VV<\#129><\#129><\#172><\#186>ross neuronal plasma membrane The neuronal plasma membrane delimits the intracellular and extracellular compartments. There is an unequal distribution of ions across the plasma membrane: regardless of the animal<\#213>s environment (seawater, freshwater or air), potassium (K+) ions are the predominant cations in the intracellular fluid and sodium (Na+) ions are the predominant cations in the extracellular fluid. The main anions of the intracellular fluid are organic molecules (P<\#208>): negatively charged amino acids (glutamate and aspartate), proteins, nucleic acids, etc. In the extracellular fluid the predominant anions are chloride (Cl<\#208>) ions. A marked difference between cytosolic and extracellular Ca2+ concentrations is also observed (Figure 3.2). Insert Figure 3.Spatial distribution of Ca2+ ions inside the cell deserves a more detailed description. Ca2+ ions are present in the cytosol as <\#212>free<\#213> Ca2+ ions at a very low concentration (10<\#208>8 to 10<\#208>7m) and as bound Ca2+ ions (bound to Ca2+-binding proteins). They ay are also distributed in organelles able to sequester calcium, which include endoplasmic reticulum, calciosome and mitochondria, where they constitute the intracellular Ca2+ stores. Free intracellular Ca2+ ions present in the cytosol act as second messengers<\#172><\#129><\#129>V<\#172>VV<\#254><\#172>V<\#172>V<\#129><\#129>VV<\#129><\#129><\#172><\#172><\#172>V<\#129><\#255>V<\#129>+<\#129>++<\#253><\#129>++<\#254><\#129>V<\#129><\#129><\#172>+<\#255><\#160> <\#129><\#250>V<\#129><\#129><\#160><\#255>Apple 12" RGB StandardApple 12" RGB Standardint#0int#0int#1int#1n two compartments is called a <\#212>concentration gradient<\#213>. Measurements of Na+, K+, Ca2+ and Cl<\#208> concentrations have shown that concentration gradients for ions are constant in the external and cytosolic compartments, at the macroscopic level, during the entire neuronal life. How are concentration gradients kept constant? At least two hypotheses can explain this constancy: n<\#9>Na+, K+, Ca2+ and Cl<\#208> ions cannot cross the plasma membrane: plasma membrane is impermeable to these inorganic ions. n<\#9>Plasma membrane<\#129>j<\#129>jn<\#230><\#255> <\#255><\#255><\#255><\#255><\#230>n n<\#230> 4<\#255><\#255><\#255><\#255><\#152><\#128><\#230>n<\#230>HH~<\#235>~<\#235>x<\#128><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#204><\#204><\#255><\#255><\#255><\#255><\#153><\#153><\#255><\#255><\#255><\#255>ff<\#255><\#255><\#255><\#255>33<\#255><\#255><\#255><\#255><\#255><\#255><\#204><\#204><\#255><\#255><\#255><\#255><\#204><\#204><\#204><\#204><\#255><\#255><\#204><\#204><\#153><\#153><\#255><\#255><\#204><\#204>ff<\#255><\#255><\#204><\#204>33<\#255><\#255><\#204><\#204><\#255><\#255><\#153><\#153><\#255><\#255><\#255><\#255><\#153><\#153><\#204><\#204><\#255><\#255><\#153><\#153><\#255><\#255><\#253>(ioisotopes, it was shown that inorganic ions such as Na+, K+, Ca2+ and Cl<\#208> can effectively cross the plasma membrane: there is a transport of Na+, K+, Ca2+ or Cl<\#208> ions between the extracellular and cytosolic compartments. In contrast, organic ions such as proteins cannot diffuse through the membrane: the membrane is impermeable to them. The first demonstrations of ionic fluxes across plasma membrane by Hodgkin and Keynes (1955) were based on the use of radioisotopes of K+ or Na+ ions. These experiments were conducted on the isolated squid giant axon. When this axon is immerged in a bath containing a control concentration of radioactive *Na+ (24Na+) instead of cold Na+ (22Na+), *Na+ ions constantly appear in the cytoplasm. This *Na+ influx is not affected by dinitrophenol (DNP), a blocker of ATP synthesis in mitochondria. It does not require energy expenditure. This is passive transport. When the reverse experiment is conducted, the isolated squid giant axon is passively loaded with radioactive *Na+ by performing the above experiment, and is then transferred to a bath containing cold Na+. Measuring the quantity of *Na+ that appears in the bath per unit of time (d*Na+/dt, expressed in counts per minute) allows quantification of the efflux of *Na+ (Figure 3.3a). In the presence of dinitrophenol (DNP) this *Na+ efflux quickly diminishes to nearly zero. The process can be started up again by intracellular injection of ATP. Therefore the *Na+ efflux is active transport. The movement of Na+ from the cytosol to the outeuronal membrane differs from other plasma membranes by the nature, density and spatial distribution of the proteins of which it is composed. Electrical signalling in the nervous system involves the movement of ions. Among the ions present in the nervous sassive exchange of all principal ions? Ionic composition of cytosol and extracellular compartments are maintained at the expense of a continuous basal metabolism that provides energy (ATP) utilized to actively transport ions and thus to compensate for their passive movements (Figure 3.3b). 3.1.3 There is a difference of potential between the two faces of the membrane, called membrane potential Membrane potential (Vm) is by convention the difference between the potential of the internal and external faces of the membrane (Vm=Vi<\#208>Ve). Intracellular and extracellular media are neutral ionic solutions: in each medium, the concentration of positive ions is equal to that of negative ions. However, there is a very small excess of positive and negative ions accumulated on each side of the membrane. At <\#212>rest<\#213>, for example, a small excess of negative ions is accumulated at the internal side of the membrane. An equal number of positive ions is accumulated at the external side. This creates a difference of potential <\#208> the internal face is charged more negatively than the external face: Vm at rest is equal to <\#208>80 to <\#208>50mV, depending of the neuronal type (see Figure 3.2). What is particular to membrane of neurons (and of all excitable cells) is that Vm varies (Figure 3.4). It can be more negative or hyperpolarized or less negative (depolarized) or even positive (also depolarized, the internal face is positive compared to the external face). Vm varies between the extreme values <\#208>90mV and +30mV. The origin of the resting membrane potential is explained in Chapter 4, and variations of the membrane potential in the following chapters. Meanwhile, it is clear that a variable difference of potential exists between the two sides of the membrane, although the underlying mechanisystem fluids, Na+, K+, Ca2+ and Cl<\#208> ions seem to be responsible for almost all of the action. These ions cross the membrane through specific transmembrane proteins called ionic channels. These ionic fluxes are passive since they do not since they do not h the membrane either passively or actively; and (iii) there is a difference of potential between the two faces of the neuronal membrane. The remainder of this chapter examines the following questions: n<\#9>Where and how do ions passively cross the plasma l<\#208> ions passively use to cross the membrane, when and in which direction, the functional consequences of these passive ionic fluxes, and how they are maintained and controlled. 3.1 Observations and questions 3.1.1 There is an unequal distribution of ions a of the passive movements of ions? (Section 3.5) 3.2 Na+, K+, Ca2+ and Cl<\#208> ions passively cross the plasma membrane through transmembrane proteins <\#208> the channels When proteins are absent from a synthetic lipid bilayer, no movements of ions occur across this purely lipidic membrane. Owing to its central hydrophobic region, the lipid bilayer has a low permeability to hydrophilic substances such as ions, water and polar molecules; i.e. the lipid bilayer is a barrier for the diffusion of ions and most polar molecules. Therefore, Na+, K+, Ca2+ and Cl<\#208> cross the membrane at the level of specialized transmembrane proteins. The passive transport of ions occurs through proteins, called ionic channels (or simply <\#212>channels<\#213>). 3.2.1 Channels are a particular class of trnsmembrane proteins Transmembrane proteins span the entire width of the lipid bilayer (see Figure 3.1). They have hydrophobic regions containing a high fraction of non-polar amino acids and hydrophilic regions containing a high fraction of polar amino acids (see Appendix 3.1). Certain hydrophobic regions organize themselves inside the bilayer as transmembrane a-helices while more hydrophilic regions are in contact with the aqueous intracellular and extracellular environments. Interaction energies are very high between hydrophobic regions of the protein and hydrophobic regions of the lipid bilayer, as well as between hydrophilic regions of the protein and the extracellular and intracellular environments. These interactions strongly stabilize transmembrane proteins within the bilayer, thus preventing their extracellular and cytoplasmic regions from flipping back and forth. Ionic channels have a three-dimensional structure that delimits an aqueous pore through which certain ions can pass. They provide the ions with a passage through the membrane. Each channel may be regarded as an excitable molecule as it is specifically responsive to a stimulus and can be in at least two different states: closed and open. Channel opening, a switch from the closed to the open state, is tightly controlled (see Table 3.1) by: n<\#9>a change in the membrane potential <\#208> these are voltage-gated channels (Section 3.2.2); n<\#9>the binding of an extracellular ligand, such as a neurotransmitter <\#208> these are ligand-gated channels, also called receptor channels or ionotropic receptors (Section 3.2.3); n<\#9>the binding of an intracellular ligand such as Ca2+ ions or a cyclic nucleotide (Section 3.2.4); n<\#9>mechanical stimuli such as stretch <\#208> these are mechanoreceptors (Section 3.2.5). Insert Table 3.1 The channel<\#213>s response to its specific stimuli, called gating, is a simple opening or closing of the pore. The pore has the important property of selective permeability, allowing some restricted class of small ions to flow passively down their electrochemical gradients (see Section 3.3). These gated ion fluxes through pores make signals for the nervous sytem. 3.2.2 Voltage-gated channels open in response to a change in membrane potential Channels opened by voltage changes have a three-dimensional structure which contains an aqueous pore and a voltage <\#212>sensitive<\#213> region. Depending on the membrane potential, these channels may be in a closed state (ions cannot pass through the aqueous pore) or an open state (ions can pass through the aqueous pore); they change conformation <\#208> switching from one state to the other <\#208> depending on membrane potential changes. Most of these channels open transiently in response to membrane depolarization (Figure 3.5a), but some open in response to hyperpolarization. The aqueous pore of voltage-sensitive channels is essentially permeable to one ionic species. Voltage-gated channels are usually named after their most important permeant ion. However, no channel is perfectly selective. Thus, the voltage-dependent Na+ channel is also faairly permeable to NH4+. The voltage-gated Na+ channel consists of various subunits but only the principal subunit (a subunit) is required for functionality. The a subunit consists of four internally homologous domains (labelled I to IV))(labelled I to IV)) each of which is approximately 300 amino acids long and contains six transmembrane segments (Figure 3.5b). These four domains are said to be homologous because they share a high percentage of identical amino acid residues. They are separated from each other by non-homologous regions of variable lengths, of which the S5<\#208>S6 linkers form the P (pore-lining) segment. As depicted in Figure 3.5c, the protein wraps around a central pore. P segments of each domain come together to form the pore. They are responsible for the ionic selectivity (these regions exhibit exquisite conservation within a given channel family of like selectivity, but not among families with different selectivities). The general strategy for activation gating is the presence of a fourth transmembrane segment (S4) with positively charged residues (Figure 3.5b) that moves in response to a change of membrane potential (DV), somehow opening the channel. This structure is similar to the predicted structure of voltage-gated Ca2+ channels and the functional tetrameric form of K+ channels. The voltage-sensitive Na+ channel is responsible for the initial inward current of Na+ ions during the depolarizing phase of action potentials. It thus underlies the generation and propagation of Na+ action potentials (see Chapter 5). The multiple types of voltage-sensitive Ca2+ channels control the entry of Ca2+ ions into the cell. Ca2+ current underlies the generation of Ca2+ action potentials (see Chapters 6 and 19), and intracellular Ca2+ triggers several Ca2+-dependent intracellular processes, including the release of neurotransmitters from presynaptic terminals (see Chapter 8). The multiple types of voltage-sensitive K+ channels do not have a single function that can be assigned to them as a group. In general,ple types of voltage-sensitive K+ channels do not have a single function that can be assigned to them as a group. In general, they participate in resting membrane potential and action potential repolarization (see Chapters 4, 5 and 6). 3.2.3 Ligand-gated channels opened by extracellular ligands, and receptor channels opened by neurotransmitters These channels open as a result of thnels open as a result of thr ligands, and receptor channels opened by neurotransmitters These channels open as a result of the binding of an extracellular ligand (e.g. a neurotransmitter). Their common characteristic is that the receptor sites for the ligand and the ionic channel that they control are part of the same protein. Receptor channels are made up of several subunits (lco /Bg<\#169><\#238> - N<\#208>"_ 8<\#142>f<\#160>U.<\#128>N<\#209><\#192><\#184>.<\#128>N<\#209>NVYO/<\#160>"n B g" @ P02.g<\#178>@bpSAg<\#209><\#192>`<\#246>R@<\#160>.N^ _P<\#143>N<\#208>a"_NuNV<\#254><\#166>=|<\#255><\#133><\#255><\#252>Jn<\#255><\#252>fN<\#186><\#255><\#228>"<\#140>/ N<\#186><\#255><\#220>(Q?8 Z?,r<\#169><\#152>0.g<\#218>k<\#214>W@g<\#208>j<\#204>R@j<\#192>Hn<\#255><\#170><\#168><\#139> np<\#208>h=@<\#255><\#168>=h<\#255><\#166>0.<\#255><\#172><\#208>n<\#255><\#176><\#145>n<\#255><\#166>?.<\#255><\#168>?.<\#255><\#166><\#168><\#147>/.<\#168><\#163>=|<<\#255><\#254>Hn<\#255><\#178> receptor channel changes conformation and transiently switches to a state in which the aqueous pore is open (Figure 3.6a). Thus, receptor channels ensure fast synaptic transmission by triggering a rapid increase of ionic permeability in response to the binding of a neurotransmitter. It should be noted that these channels may also present receptor sites for other endogenous molecules such as allosteric agonists which modulate their activity. The best-known members of this group open in rrhis group open in response to neurotransmitters (Table 3.1): acetylcholine (nicotinic receptors), g-aminobutyric acid (GABAA receptors) and glutamate (NMDA and non-NMDA receptors). Nicotinic receptors (nAChR; see Chapter 9) and glutamate receptor channels (see Chapter 11) are permeable to cations Na+, K+ and Ca2+. Conversely, GABAA receptors (see Chapter 10) are permeable to Cl<\#208> ions. The primary structures of the nAChR, GABAa and glutamate receptor channels show many similarities. The nAChR, for example, is a pentamer of 16 known different types of subunits a, b, g, d and e. These different subunits show very high levels of sequence and hydrophobicity profile homologies: they contain a large N-terminal hydrophilic domain exposed to the synaptic cleft, followed by three transmembrane segments (M1<\#208>M3), a large intracellular loop and a C-terminal transmembrane segment (M4) (Figure 3.6b). Subunits are regularly distributed around an axis of quasi-symmetry delineating the ion channel (Figure 3.6c). The ion channel is lined by the M2 segment from each of the five subunits. The neurotransmitter receptor sites have been identified in the large extracellular domain of two identical subunits (the two a subunits of the nAChR receptor and the two b-subunits of the GABAA receptor). 3.2.4 Ligand-gated channels opened by intracellular ligands These are channels gated by GTP-dependent proteins (G proteins) or by second messengers such as Ca2+, IP3 or cyclic nucleotides. Channels that are directly opened by G proteins are, for example, the G protein-gated inwardly rectifying K+ (GIRK) channels (Figure 3.7). GIRK channels are present in the membrane of heart cells where they are coupled to a muscarinic acetylcholine receptor (m2AChR) and are involved in mediating a slowing of the heart rate (as a result of the efflux of K+ ions) in response to acetylcholine release from the vagus nerve. They are also present in mammalian central and peripheral neurons where they are coupled via G proteins to many different neurotransmitter receptors and act to decrease firing rate. The coupling between these G-coupled receptors (called metabotropic receptors) and GIRK channels is achieved by a high diversity of G proteins which are peripheral proteins present on the cytoplasmic side of the membrane and are formed by three different subunits (a, b, g). Activation of the metabotropic receptor catalyses the release of the Gbg subunit from the G protein (Gai/obg) and Gbg directly interacts with GIRK channels to increase their open probability (Figure 3.7a). Insert FCa2+-gated channels are K+, Cl<\#208> or cation permeable channels. A local increase in intracellular Ca2+ concentration favours the probability of the binding of Ca2+ ions to the Ca2+ receptor site(s) located on the cytoplasmic side of these channelsse channels. This in turn triggers the opening of the channels and the passage of ions across the membrane. Some of these channels are also voltage-gated. Their general role is to transduce a local increase in intracellular Ca2+ concentration in a change of membrane potential. Cyclic nucleotide-gated (CNG) channels are cation permeable channels. Despite the fact that CNG channels are gated by the binding of a ligand, cGMP or cAMP (Figure 3.8a), and not by voltage, they are structurally homologous to the voltage-gated channels since their primary structure presents six hypothetic transmembrane domains (Figure 3.8b). Consistent with this interpretation, immunocytochemistry studies have shown that the amino terminal domain is located on the cytoplasmic side of the membran and transduce electrical activity in neurons into biochemical events such as exocytosis. Ca2+ ions bound to cytosolic proteins or present in organelle stores are not active Ca2+ ions; only <\#212>free<\#213> Ca2+ ions have a role. A difference of concentration betwnary connection between voltage-gated channels and CNG channels is suggested also by the presence of a cGMP-binding region in some voltage-gated channels (such as the H-channel, a hyperpolarization-activated cation channel). CNG channels may exist as tetramers (Figure 3.8c). The role of CNG channels of photoreceptors (cGMP-gated) and olfactory receptor cells (cAMP-gated; see Chapter 15) is to convert external sensory stimuli into changes in membrane potential. Consequently, sensory transduction in these cases is very rapid. 3.2.5 Mechanically gated channels opened by mechanical stimuli Sensory receptors underlying the transduction of pressure, stretching, auditory and vestibular stimuli are ionic channels directly opened by the corresponded by the corresponding sensory stimulus. Consequently, transduction in these cases is very rapid. Mechanically gated ion channels, such as stretch-activated channels, transduce mechanical stimuli into changes of membrane potential of specialized sensory cells (see Chapter 12). These channels are K+ selective, Cl<\#208> selective or non-selective cation channels. Mechanosensory transduction underlies a wide range of senses, including proprioception, touch, balance and hearing. 3.2.6 Other channels: junctional channels or gap junctions These channels are different from all the channels mentioned so far because they are built with transmembrane proteins from two apposed plasma membranes. At the point of apposition the extracellular space is very narrow. These structures are called gap junctions because they connect the cytoplasms of two adjacent cells. These are poorly selective channels but are more permeable to cations than to anions. They are also permeable to a number of small molecules (with molecular weights lower than 1200D), such as the second messengers cAMP, inositol trisphosphate, but not to proteins or nucleic acids. Junctional channels are usually open at the resting membrane potential. Their opening can be modulated by either the transjunctional potential (the voltage difference between the two internal sides of the coupled membranes), by intracellular or extracellular factors, or by second messengers. Thus, depending on the cell type they may open or close as a function of membrane potential, intracellular pH, intracellular concentration of Ca2+ ions and the presence of cyclic nucleotides (Figure 3.9a). Connexins are the principal protein component of gap junctions. Genes coding for connexins were first sequenced from rat and human heart and liver cellheart and liver cells and the amino acid sequences deduced. Hydrophobicity profiles and specific antibody binding to certain regions of the protein have led to a transmembrane model of the connexins. Each connexin has four putative transmembrane a-helices and both NH2 and COOH terminals are oriented to the cytoplasmic side, accessible to intracellular regulatory elements (Figure 3.9b). Junctional channels have a hexameric structure with a central aqueous pore 15nm long, 2nm wide at the ends and approximately 1.5nm wide at t or present in organelle stores are not active Ca2+ ions; only <\#212>free<\#213> Ca2+ ions have a role. A difference of concentration betweeN^"_\<\#143>N<\#209>NV<\#255><\#206> O1n <\#160>=@"n"<\#168>N^"_\<\#143>N<\#209>4< N<\#250>4< "o oB@BA/H/I _???/<\#173><\#237> o?<<\#169><\#238> o <\#128>N<\#250> o /Bg<\#169><\#238> _POranes (Figure 3.9c). There is a high density of gap junctions at electrical synapses. The function of these gap junctions is electrical and metabolic coupling as well as transfer of second messengers between the associated cells. Electrical coupling allows the rapid propagation of action potentials between neurons. This would give synchrony among a population of active neurons for the purpose of escape behaviours, for example (electrical coupling of crayfish cord neurons). In the heart, the primary function of gap junctions is to allow current flow from the pacemaker cells to ventricular muscle cells leading to their synchronized rhythmic contraction. Metabolic coupling is particularly important in the case of non-vascularized cells such as those of the eye is permeable to Na+, K+, Ca2+ and Cl<\#208> ions but there are mechanisms that continuously maintain the unequal distribution of ions. 3.1.2 Neuronal plasma membrane is permeable to ions, allowing both passive and active transport of ions With the use of radad-sensitive Na+ channels underlie the generation and propagation of sodium and sodium<\#208>calcium action potentials. They are present in high densities at the initial segment and nodes of Ranvier of the axon (in the case of myelinated axons) or all along the axside (efflux) can be switched off reversibly by the use of metabolic inhibitors. Observation of active transport of ions provides a general answer to the question: how does the cell maintain its ionic composition in the face of continuous p of continuous pms are not explained in this chapter. 3.1.4 Questions In summary, we have the following facts: (i) there is an unequal distribution of ions across the neuronal plasma membrane, which stays constant during the entire neuronal life; (ii) ions move througrougmembrane? What are the respective roles of the concentration gradient and membrane potential? (Sections 3.2 and 3.3) n<\#9>How and where do ions actively cross the plasma membrane and thus compensate for the passive movements? (Section 3.4) n<\#9>What are the rolepropagation of calcium (dendrites) or sodium<\#208>calcium (soma) action potentials. Neurotransmitter receptors are located at postsynaptic regions <\#208> postsynaptic membrane of the dendritic spines, the dendritic branches, the soma or the axon (the latter in the case of axo-axonic synapses). They underlie synaptic transmission. They can also be found in presynaptic regions where they play a role in the control of neurotransmitter release. Sensory receptors are located exclusively in membranes of sensory receptor cs<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255> <\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>inked together by non-covalent bonds) which form the central aqueous pore. In general, the receptor sites of the neurotransmitter are present in the extracellular domains of two identical subunits. When the neurotransmitter binds to its receptor sites, the<\#164><\#249><\#154><\#200> 5{0 6/`Z0<\#160> <\#158>8<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255> 5{0<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>structure of the channel pore determines the type of ion(s) that diffuse passively through the channel The pores of ion channels select their permeant ions. The structural basis for ion channel selectivity has been studied in a bacterial K+ channel called the KcsA channel (it is a voltage-independent K+ channel). All K+ channels show a selectivity sequence K+=Rb+>Cs+, whereas permeability for the smallest alkali metal ions Na+ and Li+ is extremely low. Potassium is at least 10000 times more permeant than Na+, a feature that is essential to the function of K+ channels. Each subunit of the K+ channel consists of an N-terminal cytoplasmic domain, followed by six transmembrane helices (or two transmembrane helices for KcsA), and a C-terminal globular domain in the cytoplasm (Figures 3.11a,b). The P loop (P for pore) situated between transmembrane helices 5 and 6 or 1 and 2, depending on the K+ channel considered, is the region primarily responsible for ion selectivity. The KcsA channel i1 The KcsA channel is overexpressed in bacteria and the three-dimensional structure of its pore is investigated by the use of X-ray crystallography. The KcsA channel is a tetramer with fourfold symmetry around a central pore. The pore is constructed of an inverted teepee withe and the beginning of the P region is located on the extracellular side of the membrane. The pore of CNG channels is permeable to both monovalent and divalent cations. The cyclic nucleotide binding site is situtated near the carboxy terminus. An evolutiony predominantly non-polar side-chains pointing to the pore axis. The diameter of this region is sufficiently wide to allow the passage of fully hydrated cations. This long entry way then opens to a wider water-filled cavity (1nm across). Beyond this vestibule is the 1.2nm long selectivity filter. After this, the pore opens widely to the extracellular side of the membrane. What are the respective roles of the parts of the pore? Electrostatic calculations show that when an ion is moved along a narrow pore through a membrane it must cross an energy barrier that is maximum at the membrane centre. A K+ ion can move throughout the inner pore and cavity and still remain mostly hydrated, owing to the large diameter of these regions. The role of the inner pore and the cavity is to lower the electrostatic barrier. The cavity overcomes the electrostatic destabilization from the low dielectric bilayer by simply surrounding an ion with polarizable water. Another feature that contributes to the stabilization of the cation at the bilayer centre are the four pore helices which point directly at the centre of the cavity. The amino to carboxyl orientation of these helices imposes a negative electrostatic (cation attractive) potential via the helix dipole effect. These two mechanisms (large aqueous cavity and oriented helices) serve to stabilize a cation in the hydrophobic membrane interior. The selectivity filter that follows, in contrast, is lined exclusively by polar main-chain atoms. They create a stack of sequential carbonyl oxygen rings which provide multiple closely spaced binding sites for cations separated by 0.3<\#208>0.4nm. This selectivity filter attracts K+ ions and allows them to move. Why are cations permeant and not anions? As might have been anticipated for a cation he centre. Each hexamer belongs to the membrane of one of the coupled cells and constitutes a connexon formed by six connexins. A junctional channel is formed by the apposition of two connexons and its axis is perpendicular to the plane of the plasma membermeant than Na+ ions? The selectivity filter is so narrow that a K+ ion evidently dehydrates to enter into it and only a single K+ ion can pass through at one time. To compensate for the energy cost of dehydration, the carbonyl oxygen atoms come in very close contact with the ion and act like surrogate water <\#208> they substitute for the hydration waters of K+. This filter is too large to accommodate a Na+ ion with its smaller radius (main chain oxygens are spatially inflexible and their relative distances tolens. The exchange of second messengers (cyclic AMP, cyclic GMP, inositol trisphosphate, Ca2+, etc.) allows the transfer of information between coupled cells. 3.2.7 Distribution of the various channels in the neuronal plasma membrane (Figure 3.10) Voltage<Hn<\#255><\#184>/,<\#138> . <\#169><\#231>Jg0J.<\#253><\#176>gp`p?<N<\#186><\#232>X<\#143>/</,~ ln h<\#212>N<\#144>P<\#143>`Vp.<\#253><\#176>=@<\#255><\#206>Hn<\#253><\#176>/</,~N<\#186>,O<\#239> .<\#253><\#176>f Jn<\#255><\#206>gp`p?<N<\#186><\#231><\#176>X<\#143>/</,~ ln h<\#212>N<\#144>P<\#143> l<\#138> PJhLV<\#192>DH<\#128>??</,~ ln h<\#212>N<\#144>P<\#143> ln h<\#146> <\#176><\#172>zgB'/ N<\#186><\#231><\#218>\<\#143>`<\#169><\#152>(_p`<\#169><\#152>(_L<\#223> <\#224>N^Nu<\#234><\#156><\#237><\#237>p<\#237>p<\#255><\#244><\#255><\#244><\#237>^part near the entry and exit point of the selectivity filter. Binding at adjacent sites may provide the repulsive force for ion flow through the selectivity filter: two K+ ions at close proximity in the selectivity filter repel each other. The repulsion overcomes the strong interaction between ion and protein and allows rapid conduction in the setting of high selectivity. This leads to a rate of diffusion of around 108 ions per second. 3.3.2 The electrochemical gradient for a particular ion determines the direction of the passive diffusion of this ion through an open channel To predict the direction of diffusion of ions through an open channel, both the concentration gradient of the ion and the membrane potential have to be known. The resultant of these two forces is called the electrochemical gradient. To understand what the electrochemical gradient is for a particular ion, the concentration gradient and membrane potential will first be explained separately. The passive diffusion of ions down their concentr.<\#255><\#254><\#145>n<\#255><\#224>`Rn]Zp<\#134> P0.<\#255><\#224><\#176>hLm<\#254><\#182> l<\#134> PJhLg<\#150>Jn<\#255><\#224>lRn<\#255><\#224> l<\#134> P0.<\#255><\#224><\#176>hLfSn<\#255><\#224>Bn<\#255><\#254>Bn<\#255><\#252>B'<Hn<\#255><\#252>/,<\#134>?<<<\#169><\#231>Jg0.<\#255><\#224><\#176>n<\#255><\#252>gB'/.<\#255><\#252>/,<\#134>?<\<\#169><\#231></.<\#255><\#224>/,<\#134>?<\<\#169><\#231>/,<\#134>?<<\#169><\#231>/</,z ln h<\#212>N<\#144>P<\#143>`<\#169><\#152>(_p` 9n<\#208><\#169><\#152>(_&_N^Nu<\#253>D<\#253>v<\#245><\#192><\#252>@<\#248>&<\#253>v<\#253>D<\#241>t<\#244><\#162><\#244>2<\#244>2<\#255><\#226><\#255><\#226><\#244><\#244><\#245><\#245>p<\#255><\#218>NV<\#255><\#240>H<\#231>0..ation gradient The concentration gradient of a particular ion is the difference of concentration of this ion between the two sides of the plasma membrane. A difference of concentration allows passive diffusion of ions through an open channel, since ions move from the medium where their concentration is high to the medium where their concentration is lower. Suppose that membrane potential is null (Vm=0mV); there is no difference of potential between the two faces of the membrane, so ions will diffuse according to their concentration gradient only (Figure 3.12a). Since the extracellular concentrations of Na+, Ca2+ and Cl<\#208> are higher than the respective intracellular ones, these ions will diffuse passively towards the intracellular medium (when Na+, Ca2+ or Cl<\#208> permeable channels are open) as a result of their concentration gradient. In contrast, K+ will move from the intracellular medium to the extracellular one (when K+ permeable channels are open). The force that makes ions move down takes ions move down their concentration gradient is constant for a given ion since it depends only on the difference of concentration of this ion, which is itself continuously controlled to a constant value by active transport (pumps and transporters). However, this is not always true; during intense neuronal activity, concentration of ions may change (K+ concentration in particular) owing to the small volume of the external medium. At the microscopic level this is not true also; intracellular Ca2+ concentration, for example, on (in the case of non-myelinated axons). Na+ channels are also found in the membrane of the cell body, but they are generally absent or present at a very low density in dendritic membranes. In fact, dendrites rarely generate or propagate sodiuction pient cannot reverse by itself. The passive diffusion of ions according to potential gradient Membrane potential is a potential gradient that forces ions to move in one direction: positive ions are attracted by the <\#212>negative<\#213> side of the membrane and negative ions by the <\#212>positive<\#213> one. If we suppose that there is no concentration gradient for any ions (there is the same concentration of each ion in the extracellular and intracellular media), ions will diffuse according to membrane potential only. At a membrane potential Vm=<\#208>30mV (Figure 3.12b), positively charged ions, the cations Na+, Ca2+ and K+, will move from the extracellular medium to the intracellular one according to membrane potential. In contrast, anions (Cl<\#208>) will move from the intracellular medium to the extracellular one. The passive diffusion of ions according to both the concentration gradient and membrane potential: the electrochemical gradient In physiological conditions, the direction and amplitude of ion diffusion through an open channel is determined by both the concentration gradient and membrane potential. Since concentration gradient is constant for each ion, the direction and amplitude of diffusion varies with membrane potential. Figure 3.12 shows that at a membrane potential of <\#208>30mV, concentration gradient and membrane potential drive Na+ and Ca2+ ions in the same direction, toward the intracellular medium, whereas they drive K+ and Cl<\#208> in reverse directions (compare the two parts of Figure 3.12). The resultant of these two forces, concentration and potential gradients, is the electrochemical gradient. To know how to express the electrochemical gradient, the equilibrium potential must first be explained. The equilibrium potential, Eion All systems are moving toward equilibrium. The value of membrane potential where the concentration force that tends to move a particular ion in one direction is exactly balanced by the electrical force that tends to move the same ion in the reverse direction is called the <\#212>equilibrium potential<\#213> of the ion (Eion). The equilibrium potential for a particular ion is the membrane potential for which the net flux of this ion (fnet) through an open channel is null: when Vm=Eion, fnet=0mols<\#208>1. Eion can be calculated using the Nernst equation (see Appendim action potentials. The distribution of voltage-dependent K+ channels will not be analysed here because of the large diversity of these channels. Voltage-gated Ca2+ channels are located at high density at presynaptic terminals <\#208> axon terminal membranes, presynaptic dendrites or somatic presynaptic regions <\#208> where they ensure the Ca2+ inflow essential for the release of neurotransmitters. They can also be present in the dendritic or somatic membranes where they are the basis of the generation and tion and (140/14)=+58mV <\#9>EK=(58/1)log10(3/160)=<\#208>84mV <\#9>ECa=(58/2)log10(1/10<\#208>4)=+116mV <\#9>ECl=(58/<\#208>1)log10(150/14)=<\#208>58mV. These equations have the following meanings. If the channels open in a membrane are for example all permeable to K+ ions, the efflux of K+ ions will hyperpolarize the membrane until Vm= EK=<\#208>84mV, a potential where K+ ions have the same tendency to diffuse towards the intracellular medium according to their concentration gradient than to move in the reverse direction according to membrane potential. At that potential the efflux of K+ will stop and the membrane potential will stay stable if K+ channels stay open (see Chapter 4). Now, if only Na+ channels are open, the membrane potential will move toward Vm=+58mV, the potential at which the net flux of Na+ is null. Similarly when Vm=ECl=<\#208>60mV, Cl<\#208> ions have the same tendency to move down their concentration gradient than to move in the reverse direction according to membrane potential; the net flux of Cl<\#208> is null. In contrast, when Vm is different from ECl, the net flux of Cl<\#208> is not null. This holds true for all the other ions: when Vm is different from Eion there is a net flux of this ion. The electrochemical gradient We have seen that when Vm=Eion (i.e. Vm<\#208>Eion=0), there is no diffusion of this particular ion (fnet=0). In contrast, when Vm is different from Eion there is a passive diffusion of this ion through an open channel. The difference (Vm<\#208>Eion) is called the electrochemical gradient. It is the force that makes the ion move through the open channel. 3.3.3 The passive diffusion of ions through an open channel is a current To know the direction of passive diffusion of a particular ion and how many of these ions diffuse per unit of time, the direction and intensity of the net flux of ions (number of moles per second) through an open channel have to be measured. Usually the net flux (fnet) is not measured; the electrical counterpart of this net flux, the ionic current, is measured instead. Passive diffusion of ions through an open channel is a movement of charges through a resistance (resistance here is a measure of the difficulty of ions moving through the channel pore). Movement of charges through a resistance is a current. Through a single channel the current is called <\#212>single-channel current<\#213> or <\#212>unitary current<\#213>, iion. The relation between fnet and iion is: iion=fnetzF. The amplitude of iion is expressed in amp<\#143>res (A) which are coulombs per seconds (Cs<\#208>1). F is the Faraday constant (96500C); ells where they underlie sensory signal transduction. 3.3 The diffusion of ions through an open channel: What is an electrochemical gradient and an ionic current? It has been stated above that a channel is said to be in a closed state (C) when its ionic pf potential between the two ends of the resistance. For currents carried by ions (and not by electrons as in copper wires), I is called iion, the current that passes through the resistance of the channel pore which has a resistance R (called rion). But whaa Q<\#187><\#243><\#144><\#200><\#130> <\#195><\#227>P<\#168>H <\#200><\#130> <\#200><\#130>`<\#195><\#227>0<\#168>H.l<\#254>?<\#177><<\#200><\#130> <\#9>OJPCZ<\#250><\#237>f7<\#155>r7<\#155>r+G~+G~<\#243><\#138><\#243><\#138><\#159><\#150><\#159><\#150>K<\#162>K<\#161><\#249><\#247><\#173><\#249><\#247><\#173><\#237><\#163><\#186>V<\#171><\#185><\#225>O<\#197><\#225>O<\#197><\#212><\#251><\#209><\#212><\#251><\#209><\#200><\#167><\#221><\#200><\#167><\#221><\#188>S<\#233><\#188>S<\#249><\#155>s <\#240><\#200><\#143><\#228>t!vw-vw-j#9j#9]<\#207>E]<\#207>Eore does not allow ions to pass. In contrast, when the channel is said to be in the open state (O), ions can diffuse through the ionic pore. C s O This diffusion of ions through an open channel is a passive transport since it does not require energy exxxitary conductance. It is a measure of the ease of flow of ions (flow of current) through the channel pore. Whereas resistance is expressed in ohms (W), conductance is expressed in siemens (S). By convention iion is negative when it represents an inward flux of cations and positive when it represents an outward flux of cations. It is generally of the order of pico-amp<\#143>res (1pA=10<\#208>12A). At physiological concentrations, gion varies between 10 and 150 pico-siemens (pS), according to the channel type. Total current, Iion In physiological situations, several channels of the same type are open at the same time in the neuronal membrane. Suppose that only one type of channel is open in the membrane, for example Na+ channels. The total current INa that crosses the membrane at time t is the sum of the unitary currents iNa at time t: INa=NpoiNa, where N is the number of Na+ channels present in the membrane; po is the probability of Na+ channels being open at time t (Npo is therefore the number of open Na+ channe3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202>``<\#255><\#255>. Iion and iion can be measured experimentally. The latter is the current measured from a patch of membrane where only one channel of a particular type is present. Iion is the current measured from a whole cell membrane where N channels of the same type are present. Roles of ionic currents Ionic currents have two main functions: n<\#9>Ionic currents change the membrane potential: either they depolarize the membrane or repolarize it or hyperpolarize it, depending on the charge carrier. These terms are in reference to resting potential (see Figure 3.3). A depolarization can be an action potential (see Chapters 5 and 6) or postsynaptic excitatory potentials (EPSPs; see Chapters 9, 11 and 12). These changes of membrane potential are essential to neuronal communication. n<\#9>Ionic currents increase the concentration of a particular ion in the intracellular medium. Calcium current, for example, is always inward. It transiently and locally increases the intracellular concentration of Ca2+ ions and contributes to the triggering of Ca2+-dependent events such as secretion or contraction. 3.4 Active transport of Na+, K+, Ca2+ and Cl<\#208> ions by pumps and transporters maintain the unequal distribution of ions Passive movements of Na+, K+, Ca2+ or Cl<\#208> ions across the membrane would normally cause concentration changes in the extracellular and intracellular compartments if they were not constantly regulated during the entire life of the neuron by transport of ions in the reverse direction, against passive diffusion; i.e. against ele the extracellular side corresponding to the base of the teepee (Figure 3.11c). The overall length of the pore is 4.5nm and its diameter varies along its distance. From inside the cell the pore begins as a water-filled tunnel of 1.8nm length (isurroundedr transporters. Pumps obtain energy from the hydrolysis of ATP, whereas transporters use the energy of an ionic gradient, for example the sodium driving force. 3.4.1 Pumps are ATPases that actively transport ions Pumps have ATPase activity (they hydrolyse ATP). This ATPase activity is generally the easiest way of identifying them. Pumps are membrane-embedded enzymes that couple the hydrolysis of ATP to active translocation of ions across the membrane. The central issue of ion motive ATPases is to couple thediameter varies along its distance. From inside the cell the pore begins as a water-filled tunnel of 1.8nm length (inner pore) surrounded b!Q$Bh,B<\#168>.Jf<\#160>`<\#160>=@"n "<\#168>(N^"_O<\#239> N<\#209>NV<\#255><\#176> O!n1nB(<\#160>fBh<\#160> fC<\#232> "<\#238>"<\#174> <\#160> =@N^"_O<\#239>N<\#209>NV<\#255><\#206> O!n the centre of the pore cannot readily be changed). It is proposed that a K+ ion fits in the filter so precisely that the energetic costs and gains are well balanced. What drives K+ ions to move on? K+ ions bind simultaneously at two binding sites 0.75nmic costs and gains are well balanced. What drives K+ ions to move on? K+ ions bind simultaneously at two binding sites 0.75nm a<\#208>"_ 8<\#142>f<\#160>U.<\#128>N<\#209><\#192><\#184>.<\#128>N<\#209>NVYO/<\#160>"n B g" @ P02.g<\#178>@bpSAg<\#209><\#192>`<\#246>R@<\#160>.N^ _P<\#143>N<\#208>a"_NuNV<\#254><\#166>=|<\#255><\#133><\#255><\#252>Jn<\#255><\#252>fN<\#186><\#255><\#228>ty for the ion transported, whether the binding sites are accessible from the cytoplasmic or the extracellular sides. For example, when the Na+ binding sites are accessible from the cytoplasm, the protein is in a conformation with a high affinity (KA=1mm) for intracellular Na+ ions, and so Na+ ions bind to the three sites. In contrast, when the three Na+ have been translocated to the extracellular side, the protein is in a conformation with a low affinity for Na+ ions so that the three Na+ are released in the extracellular space. These conformational changes are energy-dependent. The Na/K-ATPase has a catalytic a-subunit comprising about 1000 amino acids with a molecular weight of approximately 100kDa. It comprises the catalytic site for ATP hydrolysis. can increase locally by a factor of between 100 and 1000 but stay stable in the entire cytosol. However, these increases of ion concentration do not change the direction of the concentration gradient for this ion since, as we will see, a concentration grargy to produce electric signals (action potentials, synaptic potentials) as well as to actively transport other molecules. The Ca-ATPase pump The function of Ca-ATPases is to maintain (with the Na<\#208>Ca transporter) the intracellular Ca2+ concentration at verd<\#164><\#249><\#154><\#200> /<\#247>` 0<\#170>y<\#239> <\#158>8<\#254><\#160>/,<\#130><\#169><\#131> l /<\#247>`<\#146> <\#172>z/,<\#250><\#168>} l<\#250><\#160>`2 n<\#255><\#255><\#176>fB,HBn<\#255><\#170>`<\#130> L0.<\#255><\#170><\#208><\#192><\#208><\#192>=h.<\#255><\#200> L<\#208><\#192><\#208><\#192>=h><\#255><\#198> L<\#208><\#192><\#208><\#192>"L<\#210><\#192><\#210><\#192>3h&. L<\#208><\#192><\#208><\#192>"L<\#210><\#192><\#210><\#192>3h6> L<\#208><\#192><\#208><\#192>1n<\#255><\#200>& L<\#208><\#192><\#208><\#192>1n<\#255><\#198>6Jn<\#255><\#200>g L0.<\#255><\#170>H<\#192><\#231><\#136><\#209><\#192>HhN?.<\#255><\#198>?.<\#255><\#200>N<\#186> P<\#143>Rn<\#255><\#170> n<\#255><\#170>m<\#255>xHlNHl6Hl&N<\#186><\#238><\#148>O<\#239> N<\#186><\#240>2Jl&release of Ca2+ from intracellular stores. Maintaining a low intracellular Ca2+ concentration is critical since Ca2+ ions control several intracellular reactions and are toxic at a high concentration. Ca-ATPases are located in the plasma membrane and in the membrane of the reticulum. The former extrude Ca2+ from the cytoplasm whereas the latter sequester Ca2+ inside the reticulum (see also Figure 8.8). Ca-ATPases have been isolated from the sarcoplasmic reticulum of rabbit muscle cells. They have only one subunit, called a, which presents a transmembrane organization similar to that of the a-subunit of the Na/K-ATPase. There are several sequence homologies between these two subunits. Most homologous regions are located in the large cytoplasmic domain implicax 3.2): Eion=(RT/zF)ln([ion]e/[ion]i), where R is the constant of an ideal gas (8.314 VCK<\#208>1mol<\#208>1), T is the absolute temperature in kelvins (273.16 + th<\#161>C); F is the Faraday constant (96500Cmol<\#208>1); z is the valence of the ion; and [ion]. is the rature in kelvins (273.16 + the temperature in <\#161>C); F is the Faraday constant (96500Cmol<\#208>1); z is the valence of the ion; and [ion]. is the co<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>neurons. The Na<\#208>Ca transporter This transporter uses the energy of the Na+ gradient to actively carry Ca2+ ions towards the extracellular environment. It is situated in the neuronal plasma membrane and operates in synergy with the Ca-ATPase and with transpncentration of the ion in the extracellular or intracellular medium. This gives: <\#9>Eion=(58/z)log10([ion]e/[ion]i),<\#9>(1) From the equation and concentrations of Figure 3.2, the equilibrium potentials for each ion can be calculated: <\#9>ENa=(58/1)log10e protein has 12 hypothetic transmembrane segments. Neurotransmitter transporters Inactivation of most neurotransmitters present in the synaptic cleft is achieved by rapid reuptake into the presynaptic neural element and astrocytic glial cells. This is i<\#255><\#255>s performed by specific neurotransmitter transporters, transmembrane proteins that couple neurotransmitter transport to the movement of ions down their concentration gradient. Certain neurotransmitter precursors are also taken up by this type of active transport (glutamine and choline, for instance). Once in the cytoplasm, neurotransmitters are concentrated inside synaptic vesicles by distinct transport systems driven by the H+ concentration gradient (maintained by the vesicular H+-ATPase). 3.5 Summary Passage of ions through the membrane is a regulated process and the flow of ions across the neuronal plasma membrane is not a simple and anarchic diffusion through a lipid bilayer. Instead, it is restricted through transmembrane proteins whose opening (channel proteins) or activation (pumps or transporters) is tightly controlled by different factors. The following are the answers to the questions of Section 3.1. Where and how do ions passively cross the plasma membrane? Ions move passively across the plasma membrane through ionic channels that are specifically permeable to one or several ions. They move down their electrochemical gradient. This passive movement of charges is a current that can be recorded. Through a single channel it is a unitary current iion, and through N channels it is a macroscopic current or total current Iion. n<\#9>The type of ion that moves through an open channel (ionic selectivity of the channel pore) is determined by the structure of the channel itself. This ionic selectivity gives the name to the channel. For example, a Na+ channel is permeable to Na+ ions; a cationic channel is permeable to cations, Na+, K+ and sometimes also Ca2+. n<\#9>The direction of ion diffusion through a single channel depends on the electrochemical gradient or driving force for this particular ion (Vm<\#208>Eion). n<\#9>The number of charges that diffuse through an open channel per unit of time (iion) depends on the electrochemical gradient (Vm<\#208>Eion) but also on how easily ions move through the pore of the channel (expressed as the conductance gion of the channel). How and where do ions actively cross the plasma membrane and thus compensate for the passive movements? Active movements of Na+, K+, Ca2+ or Cl<\#208> ions across the membrane occur through pumps or transporters. Pumps z is the valence of the ion (+1 for Na+ and K+, <\#208>1 for Cl<\#208>, +2 for Ca+); and fnet is the net flux of the ion in mols<\#208>1. In general, currents are expressed following Ohm<\#213>s law: U=RI, where I is the current through a resistance R and U is the difference ot is U in biological systems? U is the force that makes ions move in a particular direction; it is the electrochemical gradient for the considered ion and is also called the driving force: U=Vm<\#208>Eion (Figure 3.13). Unitary current, iion According to Ohm<\#213>s law, the current iion through a single channel is derived from (Vm<\#208>Eion) = rion<\#180> iion. So: iion=(1/rion)(Vm<\#208>Eion)=gion (Vm<\#208>Eion). gion is the reciprocal of resistance; it is called the conductance of the channel, or unf the channel, or unctrochemical gradients. This type of transport is described as active since it requires energy in order to oppose the electrochemical gradient of the transported ions. Ions cross the membrane actively through specialized proteins known as pumps oas pumps olular Ca2+-dependent events such as secretion or contraction. Appendix 3.1Hydrophobicity profile of a transmembrane protein The hydrophobicity profile of a protein represents the degree of attraction to water of different regions of the molecule. By conv hydrolysis of ATP (and their auto-phosphorylation) to the translocation of ions. The Na/K-ATPase pump Na/K-ATPases maintain the unequal distribution of Na+ and K+ ions across the membrane. Na+ and K+ ions cross the membrane through different Na+ and K+ petein This determination is possible when the primary structure of the protein is known. Each one of the 20 amino acids has been given a value for its hydropathic character that approximately reflects the hydrophilic or hydrophobic nature of its side-chain;rmeable channels (voltage-sensitive Na+ and K+ channels plus receptor channels). This pump operates continuously at a rhythm of 100 ions per second (compared with 106<\#208>108 ions per second for a channel), adjusting its activity to the electrical activity ohigh percentage of non-polar amino acids (isoleucine or valine) are hydrophobic. Computer programs exist that are capable of evaluating the hydrophilic or hydrophobic character all along a peptide sequence. By convention this analysis iiion this analysis is performed from the NH2 towards the COOH terminal. The length of each segment analysed is chosen by the experimenter, usually between 7 and 20 amino acids. The results can be expressed graphically: the hydrophobicity index of each segment is plotted againf the neuron. It actively transports three Na+ ions towards the extracellular space for each two K+ ions that it carries into the cell. The energy of ATP hydrolysis is needed for the conformational changes that allow the pump to change its affinis affinire hydrophobic regions, usually characterized by stretches of approximately 20 amino acids with positive values, are localized in the interior of the lipid bilayer, in the form of transmembrane a-helices. On the other hand, the more hydrophilic regions, characterized by uninterrupted negative values, are in contact with aqueous environments, thus constituting the hydrophilic extracellular or intracellular domains of the protein (see Figures 3.5<\#208>3.8). Appendix 3.2The Nernst equation The material in this appendix is adapted from Katz B (ed) (1966) Nerve, Muscle and Synapse (New York: McGraw-Hill). When Vm=Eion, a particular ion has an equal tendency to diffuse in one direction according to its concentration gradient as to move in the reverse direction according to membrane potential. The net flux of this ion is null, so the current carried by this ion is null. Vm=Eion means that: <\#9>osmotic work (Wo)=electrical work (We).<\#9>(a) The osmotic work required to move one mole of a particular ion from a compartment where its concentration is low to a compartment when its concentration is high is equal to the electrical work needed to move one mole of this ion against the membrane potential in the opposite direction. Here, active diffusion of ions is considered It presents ten hypothetic transmembrane domains. A second b-subunit is closely associated with it. They generally form a tetramer (ab)2. The steady unequal distribution of Na+ and K+ ions constitutes a reserve of energy for a cell. The neuron uses this enovalent anions such as Cl<\#208>, and to +2 for divalent cations such as Ca2+. F is the Faraday constant. F for hydrogen is the charge of one hydrogen atom: F=Ne. Here N is the Avogadro number, which is 6.022<\#180>1023mol<\#208>1 (one mole of hydrogen atoms contains er (ab)2. The steady unequal distribution of Na+ and K+ ions constitutes a reserve of energy for a cell. The neuron uses this ene<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>penditure. n<\#9>Which type(s) of ions will move through a given open channel: cations, anions? n<\#9>In which direction will these ions move, from the external medium to the cytosol or the reverse? n<\#9>How many of these ions will move per unit of time? 3.3.1 The e concentration is high can be compared to the work done in compressing 1g equivalent of an ideal gas. The gas is contained in a cylinder with a movable piston. Mechanical work to move the piston is W, calculated from force times distance of displacement H<\#138><\#233>]<\#162>D<\#130><\#226> P<\#253>/<\#253>"$<\#204><\#130>"TH<\#138><\#169>_<\#162><\#132>H<\#132>BoD  #<\#196><\#248><\#132>"T_<\#202><\#169>c<\#163>P<\#143><P/<\#253>/<\#253>"<\#204><\#132>"(H<\#138><\#169>}<\#162><\#132>`<\#128><\#194>or  "<\#204><\#132>"(J<\#138><\#169>]<\#162>DP<\#132>"/<\#253>"<\#248><\#158><\#194>(G <\#233>c<\#167>dH<\#142>Bct   <\#128>A <\#160><\#149><\#130>ra?<\#255>?<\#255> <\#128><\#226><\#254>@<\#9><\#241>? <\#132><\#226>ge <\#128>@ <\#128>er<\#255><\#255> @LN@fc<\#176><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#227><\#160>of the piston (dl). The force exerted is equal to the pressure p of the gas multiplied by the surface area S of the piston. So the work dW done to displace the piston is pSdl, which equals pdv. Therefore the work done in compressing a gas from a volume v1 to a volume v2 is: <\#9>W= #v2v1 pdv.<\#9>(c) The gas law tells us that pv=RT (hence p=RT/v), with R the constant of an ideal gas (8.314VCK<\#208>1mol<\#208>1) and T is the absolute temperature. Equation (c) can be changed to: <\#9>W=RT (1/v)dv=RT(lnv1<\#208>lnv2ted in ATP recognition and hydrolysis. 3.4.2 Transporters use the energy stored in the transmembrane electrochemical gradient of Na+, H+ or When transporters carry Na+ or H+ ions (along their electrochemical gradient) in the same direction as the transporNa+, H+ or other ions When transporters carry Na+ or H+ ions (along their electrochemical gradient) in the same direction as the transporte<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>y low levels by active expulsion of Ca2+. In fact, the intracellular Ca2+ concentration is 10000 times lower than the extracellular concentration despite the inflow of Ca2+ (through receptor channels and voltage-gated Ca2+ channels) and the intracellular ort mechanisms of the smooth sarcoplasmic reticulum to maintain the intracellular Ca2+ concentration at a very low level. Molecular cloning, expression and deduced amino acid sequence of the Na<\#208>Ca exchanger of the cardiac sarcolemmal membrane shows that th (C). So F=96500Cmol<\#208>1. Therefore zF with z = 1 is the charge of 1mol of protons or 1mol of monovalent cations (Na+, K+). The charge of one mole of monovalent anions (Cl<\#208>) is <\#208>F (z=<\#208>1); the charge of 1mol of divalent cations (Ca2+) is 2F (z=2)channel, both the intracellular and extracellular entryways are negatively charged by acidic amino acids, an effect that would raise the local concentration of cations while lowering the concentration of anions. Why are K+ ions at least 10000 times morepls in the membrane at time t); and iNa is the unitary Na+ current. More generally: Iion=Npoiion. By analogy, the total conductance of the membrane for a particular ion is: Gion=Npogion; and from iion= gion(Vm<\#208>Eion) above: Iion= Gion(Vm<\#208>Eion)obtain energy from the hydrolysis of ATP, whereas transporters use the energy of an ionic gradient, for example the sodium driving force. These transports require energy since they operate against the electrochemical gradient of the transo. They maintain ig force. These transports require energy since they operate against the electrochemical gradient of the transported ions or molecules. They maintain ionic concentrations at constant values in the extracellular and intracellular compartments despite the continuous passive movements of ions across the membrane. What are the roles of electrochemical gradients and passive movements of ions? The electrochemical gradients of ions are a reserve of energy: they allow the existence of ionic currents and drive sollow the existence of ionic currents and drive some active transports. Ionic currents have two main functions: (i) they evoke transient changes of membrane potential which are electrical signals of the neuron (action potentials or postsynaptic potentials or sensory potentials) essential to neuronal communication; and (ii) they locally increase the concentration of a particular ion in the intracellular medium, for example Ca2+ ions, and thus trigger intracelfor example Ca2+ ions, and thus trigger intracelention, the more hydrophilic regions are denoted <\#208> and the more hydrophobic ones are denoted +. Hydrophilicity and hydrophobicity are the two extremes of the hydrophobicity scale. Determination of the hydrophilic and hydrophobic characters of a proof a pro i.e. its tendency to remain in contact with water (hydrophilicity) or its tendency to repel it (hydrophobicity) (Table A3.1). Protein segments containing a high percentage of polar or charged amino acids are hydrophilic, whereas segments containing a g a st the position of the segment in the protein. Thus, one obtains the hydrophobicity profile (Figure A3.1). Determination of the transmembrane organization of the protein In the case of transmembrane proteins one assumes that the moe moe assumes that the moinstead of passive diffusion. The electrical work required to move 1mole of an ion against a potential difference Eion is: <\#9>We=zFEion,<\#9>(b) where z is the valence of the transported ion, equal to +1 for monovalent cations such as Na+ or K+, to <\#208>1 for mo<\#250>4< "o <\#171><\#254>B@BA/H/I _???/<\#173><\#237> o?<<\#169><\#238> o <\#128>N<\#250> o /Bg<\#169><\#238> _PON<\#208>"_ 8<\#142>f<\#160>U.<\#128>N<\#209><\#192><\#184>.<\#128>N<\#209>NVYO/<\#160>"n B g" @ P02.g<\#178>@bpSAg<\#209><\#192>`<\#246>R@<\#160>.N^ _P<\#143>N<\#208>a"_NuNV<\#254><\#166>=|<\#255><\#133><\#255><\#252>Jn<\#255><\#252>fN<\#186><\#255><\#228>"<\#140>/ N<\#186><\#255><\#220>(Q?8 Z?,r<\#169><\#152>0.g<\#218>k<\#214>W@g<\#208>j<\#204>R@j<\#192>6<\#180>1023 protons and the same number of electrons), and e is the elementary charge of a proton, which is 1.602<\#180>10<\#208>19 coulombs (C). So F=96500Cmol<\#208>1. Therefore zF with z = 1 is the charge of 1mol of protons or 1mol of monivalent cations (Na+, K+)d ion or molecule the process is called symport. When the movements occur in opposite directions the process is called antiport. We shall study only transporters implicated in the electrical or secretory activity of he electrical or secretory activity of ; etc. The osmotic work required to move 1mole of ions from a compartment where its concentration is low to a compartment where th. The osmotic work required to move 1mol of ions from a compartment where its concentration is low to a compartment where thR=8.314VCK<\#208>1mol<\#208>1) and T is the absolute temperature. Equation (c) can be changed to: <\#9>W=RT #v2v1 (1/v)dv=RT(lnv1<\#208>lnv2) <\#9>=RTln(v1/v2).<\#9>(d) By analogy the osmotic work is: <\#9>Wo=RTln([ion]e/[ion]i).<\#9>(e) From equation (a), Wo=<\#208>We, so from equations (b) and (e) the Nernst equation is obtained: <\#9>RTln([ion]e/[ion]i)=zFEion <\#9>Eion=(RT/zF)ln([ion]e/[ion]i).<\#9>(Nernb) and (e) the Nernst equation is obtained: <\#9>RTln([ion]e/[ion]i)=zFEion <\#9>Eion=(RT/zF)ln([ion]e/[ion]i).<\#9>(Nernst) At 20<\#161>C, RT/F is about 25mV, and moving from Neperian logarithms to decimal ones a factor of 2.3 is needed. Hence: <\#9>Eion=(58/z)log10([ion]e/[ion]i). Further reading Blaustein MP, Lederer WJ (1999) Sodium/calcium exchange: its physiological implications. Physiol. Rev. 79, 763<\#208>854. Carafoli E, Brini M (2000) Calcium pumps: structural basis for and mechanism of calcium transmembrane transport. Curr. Opin. Chem. Biol. 4, 152<\#208>161. <\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#252><\#205><\#207><\#208><\#209>J<\#210><\#211><\#212><\#213><\#214><\#215><\#216><\#238><\#217><\#218><\#219><\#220><\#221><\#255><\#222><\#236><\#223><\#224><\#225><\#226><\#227><\#228><\#229><\#130><\#230><\#231><\#131><\#232><\#234><\#235><\#236><\#236><\#237><\#238><\#239><\#240><\#241><\#242><\#243><\#244><\#245><\#247><\#246><\#245><\#247><\#248><\#249><\#255><\#251><\#252><\#253><\#236><\#254><\#255><\#236>"<\#9> %& 2<\#249><=<\#247>HK<\#254><\#135><\#254><\#235>W<\#129>Z<\#140> !<\#145>#$[<\#129>\<\#129>'()*+,-./<\#235>01`b3456789:;f<\#129>g<\#146>i>?@ABCDEFGtIuv<\#236>LMNO<\#146>QRSTUVw<\#246>XYy{~<\#248>]^_<\#132><\#129><\#134><\#129>a<\#142>cde<\#140><\#129><\#141><\#139><\#158><\#216>h<\#143>j<\#251>klmnopqrs<\#147><\#129><\#148><\#149><\#206><\#150><\#151><\#208>x<\#152><\#248>z<\#153><\#252>|<\#236>}<\#154><\#231><\#128><\#253><\#129><\#130><\#131><\#155><\#156><\#133><\#157><\#129><\#144><\#159><\#131><\#136><\#137><\#138><\#139><\#132><\#160><\#161><\#138><\#162><\#163><\#188>,<\#152>)s& )_%)%)%)%)<\#136>")#%<\#255><\#255><\#253>U)%)%)%)<\#175>O)<\#239>%)L%)<\#130>%)<\#157>%)B%)"& )%)>%)-%)%%)%))%)%)<\#195>%)%)<\#147>%)R%)%)<\#131>%)%)%)%)<\#181>/)5 )%)%)%)%)` )#%)%)%)%)\i)J%)%)%)%)F%)%)%)%)9%)%)<\#164>%)%)%)%)%)%)%)%)1%)<\#139>")|%)Y%)%),")%)")L%)& )2%)<\#131>%) ")%)!& )j)" )l" )")" )")" )+" )H& )R" ")& )<\#174>" )u)<\#147> )"<\#9>)<\#135> )")X )D    X)s%)%)%)%")<\#133>")@)G& ) ) ")<\#157>&<\#9>) )X )<\#174> )2%). )<\#189>Q<\#9><\#247>0 <\#9>-<\#9>'/<\#9>)%)F )- )<\#168>& )& )<\#175>& )< ")h%)1%)#%)<%)p%)N%)!%)%)'%)@%)%)<\#166>%)<\#205>o)<\#227>" )<\#165>& )<\#145>&<\#9>)( ) )<\#176>%)%)%) )/%)01)<\#135> ) )0& )e& )<\#204> ), ) ) <)c%) )<\#131>%)& )| )_%) )I )=- ) )L& )%)%)%)C%):%)%)%)<\#207>& )<\#163>& )<\#164>& )?)<\#178>%)%)<\#162>;)^<\#181>.> )z ) )k& )<\#153>& ))F%)B[)%)%)%)f%)<\#169>%)<\#221>n)<\#235>9)y )P )b )4q)<\#136>%)?%)%)'%)%)%)<%)%)K%)2%) %)<\#185>&)h%)<\#161>& )98)<\#172>%)-%)))<\#9>1),%)>%)<\#228>%).%)<\#180>%)[ )%)<\#227>%)<\#250>%)<\#142>)<\#163>" )<\#219>B)<\#129>" )<\#154>& )/%)%)%)%)%)%)Y%)I%)c"<\#9>)(%)<\#144>%)=>)<\#177>" ) & )*%)%)%){%)D<\#128>)+& )e%)%)R%)%)1& )<\#214>)5" )p" )(" )")" )" ) %)" )A" )")") ) ) ")+%)%)")2")")&%)")% )\" )") ) ) )0& )=" ) )" ) )" ) )%) " ) )<\#135>%)%)," )")%)<\#218>%);%)4%)<" )4%)" )" ) %)<\#160>%)" )" )%);" )" )")" )")" )" )" )5" )" )" )S" )" )lI)" )<\#183>" )" )" )" )" )")")" )@%)")%")%%)%) %) %)" )$%):")")")%")")<\#144>") " )\")<\#9>" )")")<\#158>")" )" )& ))$" )+" )" ) ) )" )" )" " " " " )" " )3" )<\#170> ):" )<\#174>%)& )M)<\#192>%)" ")"")$" )<\#9>")" )" " )<\#9>")%)#" )%)")" )#%)"")" )%)" )" " )")L" )"  )") " ) )" )" )<\#9>" " " " " )" )<\#149>" ):")()) )<\#209>& )<\#221> )<\#214>%)-%)4    J)%)%)%)%)<\#222>")<\#206>5)})4%)%)%)%)-%)%))%)<\#203><\#202><\#202>Cellular and Molecular Neurobiologyl <\#214>s``<\#255><\#255>%)x%)%)<\#130>%)5%)!%)i" ))%)%)=%)n%)%)} )<\#204> )I<\#205>5<\#163>582655 and not the rapid industrialisation and state control which Stalin proposed. They were sacked from the leadership in 1926. By 1929 Stalin was undisputed leader of the USSR. Stalin carries ou ) )'%)%)<\#197>)[%);%)%)b%)0%).%)>%)#%)<\#190>%)3%) & )<\#255><\#255><\#253>/<\#164>(<\#207><\#137><\#208><\#212><\#203> <\#216><\#203><\#211><\#133>h#!# !E <\#245> + <\#9>   (   <\#9>  <\#217>Table 3.1 Examples of ionic channels Channels <\#9>Voltage-gated <\#9><\#9> Ligand-gated<\#9><\#9>Mechanically gated  Opened by <\#9>Depolarization<\#9>Exlar ligand<\#9>Intracellular ligand<\#9><\#9>Mechanical stimuliHyperpolarization <\#9> Localization <\#9>Plasma membrane <\#9>Plasma membrane<\#9>tracellular ligand<\#9> Intracellular ligand<\#9><\#9>Mechanical stimuliHyperpolarization <\#9> Localization <\#9>Plasma membrane <\#9>Plasma membrane<\#9>Plasma membrane <\#9>Organelle membrane <\#9>Plasma membrane Examples <\#9>Na+ channels<\#9>nAChR<\#9>G protein-gated channels| )E )<\#209>\??)%)%)|")H")b*<\#234><\#222>)W%)6%)  )<\#160>;)"" )")7" ) )<\#217>%)%)5%)%)%) )"<\#9>)}" )<\#255><\#255><\#252><\#164>ndent action <\#9>IPSP<\#9>IPSP<\#9><\#9>potentials<\#9>[Ca2+]i increase <\#9>Action potential <\#9><\#9>[Ca2+]i increase <\#9><\#9><\#202><\#202>repolarization<\#9><\#9><\#9>Receptor potentiall<\#168><\#247>P 7x8 <\#155>`<\#168><\#247>@ 7x8"<\#175><\#141> <\#154><\#250>`<\#175><\#141><\#141><\#175><\#175><\#144><\#168><\#247> 7x8 <\#155><\#160><\#168><\#247> 7x8;<\#178><\#178> <\#154><\#250>`0A0<\#172> A <\#172><\#9>IP3-gated Ca2+ <\#9>Stretch-activated Ca2+ channels<\#9>iGluR<\#9>Ca2+-gated channels<\#9><\#202><\#202>channel<\#9><\#202><\#202>channels K+ channels<\#9>5-HT3<\#9>CNG channels<\#9>Ca2+-gated Ca2+ Cationic channels <\#9>GABAA<\#9>ATP-gated channels <\#9><\#202><\#202>channelGlyR<\#9><\#9><\#9><\#9> Closed by <\#9>Inactivation<\#9>Desensitization<\#9><\#9><\#9><\#157><\#214><\#215><\#157>`' +2<(+*N3*j3*<\#153>B[Figure 3.4 Variations of the membrane potential of neurons (Vm). When the membrane potential is less negative than resting membrane potential (Vrest), the membrane is said to be depolarized. In contrast, when the membrane potential is more negative than Vrest, the membrane is said to be hyperpolarized. 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(a) Effect of dinitrophenol on the outflux of *Na+ as a function of time. The axon is previously loaded with *Na+. At t=1, the axon is transferred in a bath devoid of *Na+. The ordinate (logarithmic) axis is the quantity of *Na+ ions that appear in the bath (that leave the axon) as a function of time. At t=100min, DNP (0.2mm) is added to the bath for 90min. The efflux, which previously decreased linearly with time, is totally blocked" ) )")B" )+" )" )a )h)%)%)%)%)<\#255>O)<\#243>%))%)4@)6I)m& )<\#131> )E& )?)<\#255> )<\#242>&)!)A")" )" )" <\#161> after one hour of DNP. This blockade is reversible. (b) Passive and active Na+ fluxes are in opposite directions. Plot (a) adapted from Hodgkin AL, Keynes RD (1955) Active transport of cations in giant axons from sepia and loligo, J. Physiol. 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Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#224><\#165><\#166> <\#218><\#224><\#160>' +7*G*7*(* 4*7*<\#199>*3*7*$<\#188>s people? 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<\#129><\#250>V<\#129><\#172><\#129><\#129><\#227><\#228><\#255><\#248><\#250> <\#128>mj<\#254>Li<\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160>X<\#255><\#255>sHATMS<\#254><\#199><\#252>X<\#255><\#255>s<\#254>tonn<\#254><\#253><\#255><\#248><\#255><\#248><\#250> <\#128>k<\#192><\#254>N<\#190><\#250><9<\#252><\#254>) #&O&568#(&<\#255><\#248><\#255><\#248><\#250> <\#128>h<\#144><\#208><\#254><\#134><\#253><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160>P<\#255><\#255>lHATMS<\#254><\#199><\#252>P<\#255><\#255>l<\#254>nnes<\#254><\#253><\#255><\#248><\#255><\#248><\#250> <\#128>dX<\#232><\#254><\#137>h<\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160>K<\#255><\#255>h<\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5Q582655T<\#160><\#160><\#255><\#250>+EATMS<\#254><\#199><\#253><\#255><\#250>+<\#254>50l<\#254><\#247><\#255><\#248><\#254><\#128><\#252><\#152><\#210><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#255><\#250>0EATMS<\#254><\#199><\#253><\#255><\#250>0<\#254>0li<\#254><\#247><\#255><\#248><\#254>F<\#144><\#252>3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#144><\#240> <\#219><\#144>`' +7*m* 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(a) The opening of the G protein-gated inward rectifier K channels (GIRK) requires the binding of the G protein bg subunits. 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Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5<\#192>582655k(<\#156>z>[<\#153>@t&><\#139><\#155><\#152>u '=3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#225> STUV<\#225><\#168>' +7*<\#172>4*7*37*z3*m3*7*:3*f3 7*#<\#190><\#255><\#254>@Figure 3.9 Gap junction channels. (a) These channels allow bidirectional fluxes of ions and molecules of low molecular weight to cross them. Their opening is regulated, among other factors, by intracellular Ca2+ ions and pH. (b) Putative transmembrane organization of connexins. (c) Diagram of the structure of gap junctions in a lipid bilayer. These junctions are made up of 12 connexins molecules organized into two hexamers, called connexons. Each connexon has two domains, a transmembrane and a cytoplasmic domain. The aqueous pore situated at the centre of the connexon spans both plasma membranes. Drawing (b) adapted from Milks LC, Kumar NM, Houghten N et al. (1988) Topology of the 32-kD liver gap junction protein determined by site-directed antibody localizations, EMBO J. 7, 2967<\#208>2975; (c) from Makowski L, Caspar DLD, Phillips WC et al. (1984) Gap junction structures: VI. Variation and conservation in connexon conformation and packing, Biophys. 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(a) CNG-gated channel open when four molecules of a cyclic nucleotide are bound to the receptor sites located on the intracellular face of the protein. 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Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5<\#9><\#236>582655`<\#255><\#255>`3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#153> <\#223><\#210><\#195><\#196><\#199><\#152>' +>*<\#250>4*4* *5*4**<\#149>**(IP<\#255><\#255>))Y" )&)")&) @ZO<\#209><\#219>8 /4  w <\#147><\#9>if1<\#205><\#194>j<\#137><\#9><\#238>fferences originating in local circuit neurons from the spinal cord. General localization of voltage-dependent Na+ and Ca2+ channels (Vdep channels) and receptor channels (nAChR, Glu and GABAA receptors). Voltage-gated K+ channels and GABAB channels have been omitted. One channel symbolizes a population of channels of the illustrated type. nAChR, nicotinic acetylcholine receptor; GABA, g-aminobutyric acid; GABAA, type-A GABA receptor; Glu, glutamate. h6/ ln/(<\#154>N<\#186><\#195>:?.<\#255><\#226>0.<\#255><\#224><\#208>l6?<\#168><\#147> ln/(<\#154><\#168><\#132>B'?< ln/(<\#150>##~<\#255> <\#255><\#255><\#255><\#255>~ ~ 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+VVV+<\#129><\#129>+<\#254><\#129><\#129>V<\#129>VV<\#129>V<\#129>+<\#129>+<\#129>V<\#129>V<\#129>+V<\#254><\#129>V<\#172><\#253>0<\#129>V<\#129><\#172><\#172>VV+V<\#129><\#172>VVV<\#172><\#129>+V<\#129><\#129>+<\#129>+VV<\#129>V+V<\#129>VV<\#129><\#129><\#172><\#129><\#129><\#129>V<\#172>VV<\#254>"<\#172>V<\#172>V<\#129><\#129>VV<\#129><\#129><\#172><\#172><\#129>V<\#129><\#255>V<\#129>+<\#129>+VV<\#129>V<\#129>++<\#254><\#129>V<\#129>V<\#172>+<\#255><\#129><\#170><\#129><\#224>V+<\#129><\#129><\#129><\#170><\#255><\#129><\#129><\#170><\#255>explain why Lenin changed the economic policy of Russia from War Communism to NEP in 1921. (8 marks) <\#255><\#255>& V&568#(&N^NuNV)nBgHxHlN<\#186><\#243>B0,N^NuNV)nBgHxPHlN<\#186><\#243>"N^NuNV)n&)n *)n"BgHxJHlN<\#186><\#242><\#250>N^NuNV)n9n j)n.)n&)n*BgHxKHlN<\#186><\#242><\#198>0,N^NuNV)n&BgHxLHlN<\#186><\#242><\#166>N^NuNV<\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5 X58265561<\#172>#3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#160> <\#224><\#160>P' +*'7*'7*3*  <\#128>from War Communism to NEP in 1921.<\#9>(8 marks) Source clearly shows that production had not reached 1913 levels /-/ <\#212>War<\#9><\#146> <\#168> x Eq3<\#9>@a<\#186> Y <\#175> c SjQ[<\#252>Io<\#228><\#159><\#137><<\#130>?<\#9><\#255><\#255><\#245><\#146>C<\#166>C<\#166><\#230><\#255> <\#255><\#255><\#255><\#255><\#230> <\#230> 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<\#234>&568#(&<\#254>) #<\#152>T<\#160><\#160>X<\#132>o<\#160>DATMS<\#254><\#199><\#252>X<\#132>o<\#160><\#254>d<\#162>:<\#254><\#247><\#255><\#248><\#254><\#140>`<\#216>f<\#252><\#135><\#168><\#250><9<\#252><\#254>+M, "<\#146><\#193><\#152>T<\#160><\#160><\#132><\#185><\#155><\#216>DATMS<\#254><\#199><\#252><\#132><\#185><\#155><\#216><\#254>Mosc<\#254><\#202><\#202>Cellular and Molecular 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#<\#152>T<\#160><\#160><\#132><\#203><\#155><\#231>DATMS<\#254><\#199><\#252><\#132><\#203><\#155><\#231><\#254>owd<\#254><\#247><\#255><\#248><\#254><\#211><\#190><\#248><\#146><\#252><\#134><\#253><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#132><\#207>3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>O ZO@' +2+ 7*7*2<\#254><\#199><\#252><\#235><\#175><\#204><\#254>ack <\#254><\#247><\#255><\#248><\#254><\#184>w<\#224><\#249><\#252><\#136>}<\#250> }<\#252><\#254>) 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#<\#152>T<\#160><\#160><\#200>o<\#223><\#137>DATMS<\#254><\#199><\#252><\#200>o<\#223><\#137><\#254>rial<\#254><\#247><\#255><\#248><\#254>wd<\#232><\#214><\#252>N<\#190><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#200>& <\#238>&568#(&<9<\#252><\#254>) #<\#152>T<\#160><\#160><\#200>s<\#223><\#143>DATMS<\#254><\#199><\#252><\#200>s<\#223><\#143><\#254>al <\#254><\#247><\#255><\#248><\#254>{<\#156><\#208><\#214><\#252><\#133><\#211><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#200>w<\#223><\#145>CATMS<\#254><\#199><\#252><\#200>w<\#223><\#145><\#254>l <\#253><\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5 <\#240>582655<\#132><\#254>eas<\#253><\#247><\#255><\#248><\#254>p$<\#136><\#223><\#252><\#133><\#211><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#209>l<\#232><\#136>DATMS<\#254><\#199><\#252><\#209>l<\#232><\#136><\#254>asr<\#254><\#247><\#255><\#248><\#254>tS <\#223><\#252><\#133><\#211><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#209>3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>& <\#205>&568#(&<\#160><\#160><\#145><\#200><\#168><\#228>BATMS<\#254><\#199><\#252><\#145><\#200><\#168><\#228><\#254>st<\#252><\#247><\#255><\#248><\#254><\#208><\#159><\#216><\#159><\#252>t<\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#145><\#204><\#168><\#230>DATMS<\#254><\#199><\#252><\#145><\#204><\#168><\#230><\#254>tr<\#254><\#247><\#255><\#248><\#254><\#212>@p<\#159><\#252>N<\#190><\#250><9<\#252><\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5 <\#207>582655<\#168><\#252>N<\#190><\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#154><\#182><\#177><\#207>DATMS<\#254><\#199><\#252><\#154><\#182><\#177><\#207><\#254>icul<\#254><\#247><\#255><\#248><\#254><\#190>x<\#168><\#252>8?<\#250><9<\#252><\#254>) #<\#152>T<\#160><\#160><\#154><\#183><\#177><\#211>DATMS<\#254><\#199><\#252><\#154><\#183><\#177>3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>) <\#227> <\#228><\#253> <\#244>,<\#136>' +@+-*4*g4*<\#168>4*94*L3*7*X<\#209><\#204>(0<\#168><\#252><\#137>h<\#250><9<\#252><\#254>) #<\#152>T( <\#185> <#w<\#221><\#9>5} #<\#137>p<\#242> <\#145>pV  <\#153> <\#160> ;r K <\#152> <\#254> h' OT@6 IH 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V<\#129>V<\#129><\#129>V<\#129><\#129>V<\#129><\#237><\#253>V<\#129>V<\#129><\#129>V<\#129><\#239>+<\#129>+<\#129><\#129>+<\#129>V<\#129>+<\#129><\#129>+<\#241><\#172>V<\#172>VV<\#129>V<\#129>+<\#129><\#129>+<\#241><\#172><\#252><\#172>+<\#129><\#129>+<\#254>V<\#241><\#172><\#254><\#172>V<\#129>+<\#129><\#129>+<\#254>V<\#241>VV<\#129><\#254>V+<\#129><\#129>+<\#254>V<\#204><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133><\#129><\#129><\#133>N<\#249> +<\#255>++<\#129>++<\#129>+<\#253>++<\#253>+<\#254><\#129>V<\#252><\#129>+<\#172><\#252><\#129>+<\#236>VV<\#244><\#172><\#243><\#129><\#247><\#255>+<\#172><\#241>VV<\#254><\#129><\#252><\#129>+<\#129><\#151><\#152><\#249>V<\#129>V<\#129><\#129><\#172><\#129>++<\#129><\#129><\#253><\#129>+V++VV<\#129>++VV+<\#254>+<\#252>V<\#254>+++V+<\#254>++<\#254>+<\#129>++<\#254>+++<\#172>+++<\#254><\#172><\#172><\#129>++++<\#254>+<\#129>++++VV+<\#253><\#254>++<\#129><\#163><\#154><\#249> 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<\#172>VV<\#172>VV<\#129><\#129>+<\#254>VV<\#129>VV+<\#253>VV<\#172><\#129>+<\#254>V%V+<\#172>V+<\#129>V++V<\#129><\#172>VV<\#172>VV+VV<\#172><\#172>VV<\#255><\#172><\#253>(<\#129><\#129>+<\#129>V<\#129>V<\#129>V<\#129>+VV+<\#172><\#129>V+VV<\#129><\#129><\#129>VV<\#129><\#172>V+<\#253>V+<\#129><\#163><\#154><\#250>V<\#129>+<\#129><\#254>VVV++<\#129><\#254>VV<\#253><\#172><\#254><\#172><\#172>+<\#129>+<\#172><\#172>VV<\#129>+<\#172>VV<\#129>+<\#254>V<\#129><\#172><\#172><\#253>V%VV<\#129>VV<\#172>VVVV<\#172><\#172>+<\#129>VV<\#129>+<\#172><\#172>VV<\#172><\#129><\#253>,<\#172>+<\#129><\#172>VV++<\#172>VV<\#129>+<\#129><\#172>VV<\#129>+<\#172><\#172><\#129><\#129>+<\#172><\#129>V+<\#129>V<\#129><\#172><\#129><\#162><\#154><\#250><\#172><\#254><\#172><\#129>V<\#129><\#129>V<\#129><\#129>+V<\#253>+<\#172>V<\#129>+V<\#129><\#254>V+<\#129><\#129>VV<\#129>V+<\#129><\#129>V<\#172>VV<\#254>V<\#129>V<\#129>V<\#129>+<\#129><\#129>V<\#172>+<\#172>V+<\#129><\#254><\#129><\#129>V<\#129>VV<\#129>V<\#129>+<\#129>V<\#255><\#172>+<\#129>V<\#172>+<\#172><\#253><\#129><\#129><\#129><\#254><\#129><\#129><\#129><\#253>V VV<\#129>+V<\#129><\#129>V<\#129>V<\#129><\#254>V +V<\#129>VV<\#129>V<\#172>+<\#129>V<\#129><\#162><\#129><\#235>++<\#129>+<\#129>+<\#129><\#162><\#255><%8<\#193><\#210><\#232><%D<\#132><\#232><%D<\#132><\#232><%D<\#132><\#238>2UD<\#132><\#238>2UD<\#132><\#238>2UD<\#132><\#238>2U8<\#193><\#210><\#238>2U8<\#193><\#210>5 <\#229> <\#202> <\#203>5X#!# <\#177>: ! !! + $<\#232>%&"'*<\#153><\#158><\#244>P<\#224>4i %* )!*\* )-+t+&t<\#149><\#153> <\#160> ;r K <\#152> y index  Isoleucine<\#9>4.5Valine<\#9>4.2Leucine<\#9>3.8Phenylalanine<\#9>2.8Cysteine/cystine<\#9>2.5Methionine<\#9>1.9Alanine<\#9>1.8Glycine<\#9><\#208>0.4Threonine<\#9><\#208>0.7Tryptophane<\#9><\#208>0.9Serine<\#9><\#208>0.8Tyrosine<\#9><\#208>1.3Proline<\#9><\#208>1.6Histidine<\#9><\#208>3.2Glutamic acid<\#9><\#208>3.5Glutamine<\#9><\#208>3.5Aspartic acid<\#9><\#208>3.5Asparagine<\#9><\#208>3.5Lysine<\#9><\#208>3.9Arginine<\#9><\#208>4.5P<\#191><\#224> 7x8 <\#172><\#243>8<\#192><\#243>9P<\#191><\#192> 7x8<\#243>8<\#192>P<\#191><\#176> 7x8<\#243>8<\#192>P<\#191><\#160><\#226><\#183>p<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#226><\#183>p<\#243>8<\#192>P<\#191><\#128> 7x8<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#226><\#183>p<\#243>8<\#192>P<\#191>` 7x8<\#243>8<\#192>P<\#191>P 7x8<\#243>8<\#192>P<\#191>@ 7x8<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#226><\#183>p<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>& <\#209>&568#(&<\#186><\#255><\#255><\#254><\#144>5<\#148>5|<\#253><\#250>@<\#153><\#138>!F<\#218>L<\#153><\#224>!F<\#210>$<\#244><\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>5 <\#210>582655<\#158><\#255><\#255><\#254>X<<\#252>5T<\#253><\#250>@<\#153><\#217>(<\#218>L<\#153><\#195><\#135><\#215> %3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>& <\#211>&568#(&5(5 <\#253><\#250>@<\#153>g<\#177>c<\#218>L<\#153><\#145>`<\#194><\#215> %<\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255>3<\#202>Ionic Fluxes Across the Neuronal Plasma Membrane<\#202><\#202><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#202><\#202>Cellular and Molecular Neurobiology<\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255><\#255> (in their extracellular domains). Drawings (b) and (c) adapted from Changeux JP, Edelstein SJ (1998) Allosteric receptors after 30 years Neuron 21, 959<\#208>980, with permission. <\#152> <\#174>5Z5]-;<\#144> )!*\* )-+t+&t<\#149>.B<\#225>. Y x<\#169><\#180>  28<\#208>60, with permission. <\#195>0)$<\#144>h&<\#218>@ L<\#208><\#195><\#208><\#195>"L<\#210><\#195><\#210><\#195>3h$& Cl8 L<\#208><\#195><\#208><\#195>"L<\#210><\#195><\#210><\#195>0(&<\#176>i(f" L<\#208><\#195><\#208><\#195>"L<\#210><\#195><\#210><\#195>0((<\#176>i*f SE L<\#208><\#195><\#208><\#195>Sh& Co8 L<\#208><\#195><\#208><\#195>"L<\#210><\#195><\#210><\#195>0(&<\#176>i$f" L<\#208><\#195><\#208><\#195>"L<\#210><\#195><\#210><\#195>0($<\#176>i"f RE L<\#208><\#195><\#208><\#195>Rh& L H<\#192><\#231><\#136><\#209><\#192>HhN L<\#208><\#195><\#208><\#195>?(6 L<\#208><\#195><\#208><\#195>?(&N<\#186>P<\#143>HlNHl6Hl&N<\#186><\#241><\#158>O<\#239> N<\#186><\#243>< L<\#208><\#195><\#208><\#195>0h&/ lD<\#213><\#213>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128>\ <\#243><\#172> <\#243><\#172>'<\#128>GlyR<\#9><\#9><\#9><\#9> Closed by <\#9>Inactivation<\#9>Desensitization<\#9><\#9><\#9>AdaptationRepolarization<\#9>Ligand recapture or degradation<\#9><\#9>End of stimulus Roles <\#9>Na+ and Ca2+-<\#9>EPSP<\#9>EPSP<\#9>[Ca2+]i increase <\#9>Receptor potential depe<\#9>EPSP<\#9>EPSP<\#9>[Ca2+]i increase <\#9>receptor potential depe  <\#139>    J  !  2@%F<\#129>f x <\#207>$6!Z<\#181>l<\#237><\#220><\#228><\#202><\#176>trically around a central aqueous pore. Drawings (b) and (c) adapted from Marban E, Yamagishi T, Tomaselli G (1998) Structure and function of voltage-gated sodium channels, J. Physiol. (Lond.) 508, 647<\#208>657, with permission. P0P brane a-helices (M1 and M2) and the P region. (c) Top view of the putative tetrameric arrangement of the GIRK subunits around the central pore. $ 5q5555)5n the top. Aqueous inner pore and cavity are in green. The selectivity filter is in grey. See text for further explanations. O: oxygen. Adapted from Doyle DA, Cabral JM, Pfuetzner RA et al. (1998) The structure of the phannel: molecular basis of K+ conductd channel of rods. The region between S5 and S6, the P segment, is thought to line the ion conducting pore. The cyclic nucleotide binding domain is shown in green. (c) Top view of the putative tetrameric arrangement of the subunits around the central pore.gement of the subunits around the central pore. Drawings (b) and (c) adapted from Zagotta WN, Siegelbaum SA (1996) Structure and function of cyclic nucleotide-gated channels, Ann. Rev. Neurosci. 19, 135<\#208>263. -@<\#253><\#170>/*<\#160><\#169><\#216>J<\#174><\#253><\#170>g n<\#253><\#170><\#160>) n<\#253><\#170>p<\#208><\#144>)@<\#248>?<<\#255><\#255> ln hrFigure 3.10 Localization of channels in the plasma membrane of neurons. The diagram illustrates a motoneuron innervating a striated muscle cell and receiving excitatory (glutamate) and inhibitory (GABAergic) aatory (glutamate) and inhibitory (GABAergic) aFigure 3.12 Passive diffusion. Passive diffusion of ions according to (a) their concentration gradient only, or (b) to membrane potential only at Vm=<\#208>30mV. <\#255><\#255>otassium channel: molecular basis of K+ conduction and selectivity, Science 280, 69<\#208>77; and Choe S, Robinson R (1998) An ingenious filter: the structural basis for ion channel selectivity. Neuron 20, 821<\#208>823. 5<\#139>  -<\#253><\#250>@<\#153><\#239>.LS<\#225>.<\#240>T<\#169>3<\#244>*< <\#243><\#172> <\#243><\#172><\#229><\#235>xl_XQAAWDBN<\#255><\#255><\#146>6Figure A3.1 Hydrophobicity profile of the a-subunit of two voltage-sensitive channels. The voltage-activated Na+ channel underlying the sodium action potential, and the voltage-activated dihydropyridine-sensitive Ca2+ channel. One can distinguish four hydrophobic domains (I to IV) in each protein, each composed of six segments (1 to 6). The hydrophilic COOH terminal domain of the Na+ channel is approximately twice as long as that of the Ca2+ channel. From Alsobrook JP, Stevens CF (1988) Cloning the calcalcTable A3.1 Hydrophobicity index of the side chains of each of the 20 amino acids. From Kyte J, Doolittle RF (1982). A simple method for displaying the hydropathic character of a protein. J. Mol. Biol. 157, 107<\#208>132, with permission. 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Y x<\#169><\#180> <\#255><\#255><\#255><\#231>E<\#255><\#255><\#252><\#166> <\#168>| Q<\#234><\#164><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#166><\#166>US<\#132>O<\#249>|>(, <\#243><\#172>J<\#255><\#255><\#255><\#254><\#255><\#166> <\#195><\#168><\#254><\#252>} @<\#153>x<\#213><\#192><\#189><\#131><\#237>x<\#213><\#192><\#151>+K<\#255><\#255><\#255><\#253><\#255><\#166>9<\#131><\#144><\#253><\#255>}@<\#153>eA<\#235><\#179>6<\#148>eA<\#235><\#243> L<\#255><\#255><\#255><\#252><\#255><\#166>P<\#228><\#228><\#252><\#254>}@<\#153><\#197><\#137>t<\#185><\#138><\#216><\#197><\#137>t<\#249><\#187><\#174>M<\#255><\#255><\#255><\#251><\#255><\#166> <\#170>x<\#251><\#249>}@ <\#153>V<\#199>G<\#250>"V<\#199><\#176> <\#137>N<\#255><\#255><\#255><\#250><\#255><\#166>:GT<\#250><\#248>}@<\#153><\#197><\#215><\#255>\9<\#245><\#197><\#215><\#255><\#155><\#205><\#181>O<\#255><\#255><\#255><\#249><\#255><\#166> <\#166>0<\#249><\#251> }@<\#153>x<\#213><\#192><\#195><\#133><\#202>x<\#213><\#192><\#157>P<\#255><\#255><\#255><\#248><\#255><\#166>9<\#130>4<\#248><\#250>}@<\#153>x<\#213><\#192>v<\#212><\#229>x<\#213><\#192>Pm# Q<\#255><\#255><\#255><\#247>P<\#236>`<\#247><\#245>}@<\#153><\#165>4<\#226><\#169><\#176>8<\#165>4<\#226><\#135><\#195>4R<\#255><\#255><\#255><\#246>P<\#225><\#128><\#246><\#244>}@ <\#153><\#139>O<\#152><\#169><\#176>8<\#139>O<\#152><\#135><\#195>4S<\#255><\#255><\#255><\#245>:G<\#216><\#245><\#247>}@<\#153>N<\#142>C<\#169><\#176>8N<\#142>C<\#135><\#195>4T<\#255><\#255><\#255><\#244>P<\#226><\#224> <\#244><\#246>}@<\#153>><\#158><\#179><\#169><\#176>8><\#158><\#179><\#135><\#195>4U<\#255><\#255><\#255><\#243>P<\#236><\#243><\#241>}@<\#153>!<\#243><\#169><\#176>8!<\#243><\#135><\#195>4 V<\#255><\#255><\#255><\#242>P<\#226><\#184><\#242><\#240>}@<\#153> <\#9><\#161><\#169><\#176>8 <\#9><\#161><\#135><\#195>4<\#128>W<\#255><\#255><\#253>0P<\#229>8P<\#226><\#216><\#242><\#251><\#254><\#249>@<\#255><\#254><\#255><\#254><\#255><\#254> !<\#255><\#254><\#153><\#243>"I<\#168><\#176>8<\#231>(<\#164><\#136><\#195>4 <\#243><\#172><\#249><\#214><\#249><\#214><\#249><\#214><\#249><\#214>P<\#231>DPDXQAAW6BN<\#255><\#255>t<\#248>5G3 HD:Current jobs:DRG:Cellular & molecular:table 3.1&fX<\#255><\#255><\#255><\#255> " <\#168>tQ<\#234><\#176><\#254><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#243><\#196>`<\#168><\#176>8 <\#180>%<\#135><\#146> <\#243><\#172>G<\#255><\#255><\#253>* <\#210><\#224> # <\#166>l<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>LAH<\#216><\#138><\#196><\#230>3+<\#128> <\#243><\#172>F<\#255><\#255><\#254>BQ<\#227>HP<\#232>,<\#250><\#250> $@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>XR "m>(,<\#255>P<\#232>xn<\#230><\#179><\#254><\#228><\#255>M-<<\#153>iEPSF<\#182>Z<\#162><\#227><\#254><\#151><\#255><\#186> %<\#161><\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.04D<\#255><\#255><\#252><\#160> <\#170>@ <\#210><\#136><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>X<(,w<\#130><\#221>8&]<\#255> <\#170><\#240><\#216><\#234> & <\#244><\#193>0<\#254><\#233><\#254><\#209>-<<\#153>iEPSF<\#182><\#250><\#255>1<\#255><\#228> <\#207><\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.03<\#128><\#255><\#255><\#255><\#253> <\#170>L9<\#129>X '<\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128><\#255>8 <\#243><\#172> <\#243><\#172>Ph&<\#128><\#9><\#255><\#255><\#252>_ <\#210><\#244> <\#166>,<\#253><\#250>@ (<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172><\#128> <\#255><\#255><\#252>]P<\#224>:E|<\#253><\#250>@ )<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172><\#128> <\#255><\#255><\#255><\#252> <\#170>L9<\#130>X<\#253><\#250>@ *<\#153>S<\#206><\#227><\#168><\#176>8<\#232>1<\#244><\#136><\#195>48 <\#243><\#172> <\#243><\#172>P`$I<\#255><\#255><\#255><\#255>P<\#225><\#196>9<\#131><\#180><\#252><\#250><\#243>"I<\#168><\#176>8<\#231>(<\#164><\#136><\#195>4 +H<\#255><\#255><\#255><\#255>9<\#131><\#204> <\#170>|<\#252><\#250><\#243>"I<\#168><\#176>8<\#231>(<\#164><\#136><\#195>48<\#9> ,<\#132>$6!Z<\#181>l<\#237><\#220>:E(<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184> <\#171>)<\#255>a.3<\#134> <\#132> <\#243><\#172> Y <\#138><\#188><\#148>.B<\#225>. Y x<\#169><\#180>  -<\#255><\#255><\#255><\#248>Z<\#255><\#255><\#252>[P<\#217><\#184> <\#195><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#149>z<\#212>S<\#132><\#247>43+<\#128> <\#243><\#172>Y<\#255><\#255><\#252> .W <\#196>\P<\#229><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#239>.lS<\#132><\#155>v<\#184>3+<\#128><\#255>1L<\#200><\#171><\#228>C<\#247><\#200>3 <\#254><\#206><\#254><\#225>-<<\#153> /iEPSF<\#181><\#230>qA<\#254><\#160><\#255>Mj<\#250><\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.05<\#128><\#255><\#255><\#255><\#251> <\#170>L9<\#128><\#144><\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128> 0*<\#150>8 <\#243><\#172> <\#243><\#172>P[6<\#128><\#255><\#255><\#251>:F(P<\#229><\#204><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172> 1<\#128><\#255><\#255><\#251>P<\#234><\#132> <\#207>T<\#253><\#250>@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172> 2<\#128><\#255><\#255><\#255><\#250> <\#170>L <\#170><\#204><\#253><\#250>@<\#153>S<\#206><\#227><\#168><\#176>8<\#243>P<\#136><\#195>4.<\#153>8 <\#243><\#172> <\#243><\#172>P 3X<\#132>$6!Z<\#181>l<\#237><\#220> <\#207><\#232>*<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184> <\#204><\#196>+<\#255>a.3<\#134> <\#132> <\#243><\#172>Y <B<\#225>. Y x<\#169><\#180> e1 4<\#255><\#255><\#255><\#246>_<\#255><\#255><\#251>9<\#129>89<\#133><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>\<\#189>j<\#168><\#176>8<\#152><\#129><\#212><\#136><\#195>4 <\#243><\#172> 5^<\#255><\#255><\#251> P<\#218>l9<\#130>d<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>X<\#168><\#176>8<\#128><\#154>|<\#136><\#195>4<\#255> <\#206><\#152>w<\#206><\#248><\#207><\#175><\#254><\#229><\#255>Q-< 6<\#153>iEPSF<\#182>Vy5<\#254><\#251><\#255>Ms<\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.07\<\#255><\#255><\#250>eP<\#219> <\#170><<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>o<\#169><\#229>S<\#132> 7<\#239>(s3+<\#128> <\#243><\#172>[<\#255><\#255><\#250>_ <\#170><\#180> <\#170><\#200><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>M<\#212><\#218>S<\#132><\#170>oV3+<\#128> 8<\#255>9<\#128><\#140>J<\#208> <\#133> <\#141><\#141><\#255>s<\#254>u-<<\#153>iEPSF<\#182>V<\#174><\#255>,<\#255>Nx<\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.06<\#128> 9<\#255><\#255><\#255><\#249> <\#170>L <\#208><\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128>Cn8 <\#243><\#172> <\#243><\#172>PS4<\#128><\#255><\#255><\#249><\#177>P<\#231><\#156>P<\#221><\#212> :<\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172><\#128><\#255><\#255><\#249><\#175>P<\#219>| <\#199>(<\#253><\#250> ;@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172><\#128><\#255><\#255><\#255><\#248> <\#170>L <\#165><\#252><\#253><\#250>@ <<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4L8 <\#243><\#172> <\#243><\#172>PNF<\#132>$6!Z<\#181>l<\#237><\#220>1D,<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>P<\#230>P-<\#255> =a.3<\#134> <\#132> <\#243><\#172> Y <\#173><\#160><\#163><B<\#225>. Y x<\#169><\#180> #<\#228>2<\#255><\#255><\#255><\#244>e<\#255><\#255><\#249><\#173>:G`P<\#220>H<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254> ><\#153><\#249>1<\#168><\#176>8<\#149>|<\#226><\#136><\#195>4 <\#243><\#172>d<\#255><\#255><\#249><\#168> <\#167><\#164>P<\#230>\<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>6 ?<\#228>tm<\#132>N'<\#237><\#228><\#136><\#195>4<\#255> <\#205><\#140><\#224><\#232><\#254>#<\#255>-<<\#153>jEPSF<\#181><\#227><\#172>H<\#255> <\#255><\#171>(<\#190><\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular & molecular:Cel @lular Artwork:03.09cc<\#255><\#255><\#249>:D<\#244> <\#208><\#140><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>6<\#228>t<\#168><\#176>8?<\#213><<\#136><\#195>4<\#255>P<\#224><\#208><\#158>-< A<\#153>jEPSF<\#182><\#250><\#143><\#254><\#226><\#255><\#215><\#255><\#231>u<\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.09bb<\#255><\#255><\#248><\#178> <\#200><\#140> <\#208>p<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254> B<\#153>X<\#168><\#176>8V<\#197><\#162><\#136><\#195>4<\#255>:B<\#248>*\<\#215><\#9><\#169><\#178><\#221><\#252><\#255>$<\#255>-<<\#153>jEPSF<\#181><\#227><\#171><\#166><\#254><\#246><\#255><\#155><\#255><\#252><\#198><\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular C & molecular:Cellular Artwork:03.09aa<\#255><\#255><\#248>(9<\#128>$ <\#208>h<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#164>srS<\#132><\#250>&3+<\#128> <\#243><\#172> D`<\#255><\#255><\#248>$P<\#218>`:F<<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>XS<\#132><\#216>u<\#128>:<\#215>}<\#255> <\#167>@<\#150><\#223><\#162><\#255>^-< E<\#153>iEPSF<\#182><\#188><\#189><\#254><\#220><\#255>Ms,<\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.08<\#128>"<\#255><\#255><\#255><\#247> <\#170>L9<\#131><\#200><\#253><\#250>@ F<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128>V8 <\#243><\#172> <\#243><\#172><\#9>PIx<\#128>$<\#255><\#255><\#247>B <\#202> <\#204><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> G <\#243><\#172><\#128>&<\#255><\#255><\#247>@ <\#202><\#212> <\#204><\#204><\#253><\#250>@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 H <\#243><\#172><\#128>(<\#255><\#255><\#255><\#246> <\#170>L:C<\#144><\#253><\#250>@<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4]o8 <\#243><\#172> I <\#243><\#172> PD<\#170><\#132>$6!Z<\#181>l<\#237><\#220> <\#202>8.<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>:D<\#196>/<\#255>a.3<\#134> <\#132> <\#243><\#172> Y N<\#250>4 B<\#225>. <\#244><\#9> J<\#255><\#255><\#255><\#248>g<\#255><\#255><\#247>> <\#206>9<\#128>P<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>F<\#203><\#253>S<\#132><\#185>W<\#230>3+<\#128> <\#243><\#172> Kf<\#255><\#255><\#247>: <\#205><\#192> <\#206><\#204><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>XS<\#132>K<\#200><\#232>3+<\#128><\#255>:C<\#160>~6<\#204>z-< L<\#153>iEPSF<\#182>Vy<\#183><\#254>^<\#255><\#173>u+<\#255><\#255><\#163>l 0BG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.10<\#128>+<\#255><\#255><\#255><\#245> <\#170>L9<\#128>(<\#253><\#250>@ M<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128>`8 <\#243><\#172> <\#243><\#172> P?<\#188><\#128>-<\#255><\#255><\#246>9<\#130><\#152>P<\#231><\#228><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> N <\#243><\#172><\#128>/<\#255><\#255><\#246>:D4:C<\#216><\#253><\#250>@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 O <\#243><\#172><\#128>1<\#255><\#255><\#255><\#244> <\#170>L <\#204><\#136><\#253><\#250>@<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4aI8 <\#243><\#172> P <\#243><\#172> P<<\#136><\#132>$6!Z<\#181>l<\#237><\#220> <\#168><\#156>0<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>P<\#236>01<\#255>a.3<\#134> <\#132> <\#243><\#172>Y <B<\#225>. Y x<\#169><\#180> Q <\#199>1<\#255><\#255><\#255><\#245>~<\#255><\#255><\#246>9<\#131><\#176>9<\#129><\#224><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#208>s{<\#168><\#176>8<\#246>N<\#136><\#195>4 <\#243><\#172> R}<\#255><\#255><\#246>:C:C<\#252><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#196><\#157><\#179><\#140><\#211>=0<\#136><\#195>4<\#255>P<\#234>|<\#230><\#255><\#255><\#255>-< S<\#153>jEPSF<\#182>V}<\#254><\#249><\#255><\#189><\#255><\#250><\#163><\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.12b|<\#255><\#255><\#245><\#208> <\#205>P<\#227><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254> T<\#255><\#254><\#153><\#246>4<\#168><\#176>8<\#211>=0<\#129><\#225><\#232><\#255>9<\#129>`<\#224><\#239><\#144>%<\#254><\#220>-<<\#153>jEPSF<\#181><\#230>u<\#223><\#254><\#153><\#255><\#194><\#255><\#181><\#163><\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG: UCellular & molecular:Cellular Artwork:03.12a<\#128>l<\#255><\#255><\#245><\#148>:C<\#228>P<\#234><\#160><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#176>gS<\#132><\#248>*3+<\#128> <\#243><\#172> V<\#128>j<\#255><\#255><\#245><\#138>P<\#236>P<\#237>`<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>2<\#133>ZS<\#202>Nv:$O<\#255> <\#168><\#193><\#247>lo$<\#9><\#138><\#167><\#253>_-< W<\#153>pEPSF<\#182>[<\#250><\#254><\#255><\#255><\#179><\#255><\#194><\#170><\#255><\#253><\#244><\#128> 0Ibook2000:A Customers:Academic Press:6734 Cellular:Cellular Artwork:03.11c<\#128>i<\#255><\#255><\#245>:EH9<\#131><\#156><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254> X<\#255><\#254><\#153><\#153><\#168>6S<\#132>:<\#226>f3+<\#128><\#255>9<\#130><\#128><\#232>^@<\#251><\#174>4<\#230><\#255>-<<\#153>pEPSF<\#182>V{,<\#255><\#255><\#164><\#255><\#251><\#193><\#255><\#253><\#244><\#128> 0Ibook2000:A Customers: YAcademic Press:6734 Cellular:Cellular Artwork:03.11bh<\#255><\#255><\#244><\#194>:GXP<\#231><\#172><\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>K<\#167><\#253>S<\#132>x3+<\#128><\#255>9<\#130><\#144><\#165><\#230>wv<\#228><\#246>: Z<\#253><\#203>-<<\#153>pEPSF<\#182>Vz<\#158><\#255><\#255><\#220><\#255><\#185><\#196><\#255><\#253><\#244><\#128> 0Ibook2000:A Customers:Academic Press:6734 Cellular:Cellular Artwork:03.11a<\#128>4<\#255><\#255><\#255><\#243> <\#170>L:F<\#224><\#253><\#250> [@<\#153><\#250>8S<\#132><\#240>T<\#169>3+<\#128>s<\#221>8 <\#243><\#172> <\#243><\#172> P7<\#186><\#128>6<\#255><\#255><\#244><\#170>:D<\#156>P<\#224><\#200><\#253><\#250>@ \<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172><\#128>n<\#255><\#255><\#244><\#168>P<\#227>,:G<\#128><\#253><\#250>@<\#153> ]0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172><\#128>p<\#255><\#255><\#255><\#242> <\#170>L0<\#244><\#253><\#250>@<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4 ^x<\#216>8 <\#243><\#172> <\#243><\#172>P2<\#204><\#132>$6!Z<\#181>l<\#237><\#220>P<\#229>|2<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>9<\#131><\#188>3<\#255>a.3<\#134> <\#132> <\#243><\#172>Y _<B<\#225>. Y x<\#169><\#180> <\#199>1<\#255><\#255><\#255><\#249><\#134><\#255><\#255><\#244><\#166>P<\#229>x9<\#130><\#132><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#208>>7<\#168><\#176>8 <\#231><\#147><\#191><\#224> ` <\#243><\#172><\#133><\#255><\#255><\#244><\#164>P<\#230><\#184>:Ep<\#250><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#244><\#168><\#176>8<\#213>;"<\#147><\#191><\#224><\#255>P<\#236><\#228><\#151> a09bG<\#179><\#254>P-<<\#153>jEPSF<\#181><\#230>vI<\#255><\#255><\#237><\#255><\#235><\#133><\#255><\#255><\#163>l<\#152>CG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.13b<\#132><\#255><\#255><\#244>t:CpP<\#233>T<\#250><\#250> b@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>B<\#216> <\#168><\#176>8s<\#159><\#208><\#147><\#191><\#224><\#255>9<\#130>l <\#229><\#251><\#228><\#228><\#254><\#227><\#255>-<<\#153>jEPSF<\#182><\#191>q<\#254><\#217><\#255><\#212> c<\#255><\#250><\#186><\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular & molecular:Cellular Artwork:03.13a<\#128>t<\#255><\#255><\#255><\#241> <\#170>LP<\#233>P<\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128><\#133>8 <\#243><\#172> d <\#243><\#172>P-<\#254><\#128>v<\#255><\#255><\#244> <\#167><\#212>9<\#130><\#136><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172> e<\#128>x<\#255><\#255><\#244>9<\#128><\#220>P<\#224><\#220><\#253><\#250>@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172> f<\#128>z<\#255><\#255><\#255><\#240> <\#170>LP<\#229>p<\#253><\#250>@<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4<\#151><\#128>8 <\#243><\#172> <\#243><\#172>P*<\#202><\#132>$6 g!Z<\#181>l<\#237><\#220>Q<\#234> 4<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>P<\#235><\#224>5<\#255>a.3<\#134> <\#132> <\#243><\#172>Y .B<\#225>. Y x<\#169><\#180> <\#255><\#255><\#255><\#250><\#128><\#129><\#255><\#255><\#255><\#239> h<\#170>LP<\#231> <\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128><\#161>`8 <\#243><\#172> <\#243><\#172>P%<\#220><\#128><\#131><\#255><\#255><\#244> <\#210><\#184>P<\#237><<\#253><\#250> i@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172><\#128><\#136><\#255><\#255><\#244>P<\#233>xP<\#235> <\#253><\#250>@ j<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172><\#128><\#138><\#255><\#255><\#255><\#238> <\#170>LP<\#235>@<\#253><\#250>@ k<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4<\#179><\#231>8 <\#243><\#172> <\#243><\#172>P$B<\#132>$6!Z<\#181>l<\#237><\#220>9<\#129>D6<\#255>a.3<\#134> <\#132> <\#243><\#172>$l<\#237><\#220>!Z<\#181><\#163><\#219><\#184>P<\#232><\#244>7<\#255>a.3<\#134> <\#132> l <\#243><\#172>Y XB<\#225>. Y x<\#169><\#180> <\#163><\#197><\#171><\#183><\#229><\#242>a<\#255><\#255><\#255><\#243><\#168><\#255><\#255><\#244>P<\#220>$P<\#220>4<\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254> m<\#153><\#173>b,S<\#132><\#247>43+<\#128> <\#243><\#172><\#167><\#255><\#255><\#244>  <\#168>l <\#166><\#172><\#250><\#250>@ <\#243><\#172><\#255><\#254><\#255><\#254><\#255><\#254><\#153><\#162>u<\#128>S n<\#132><\#180><\#199>Q3+<\#128><\#255>:E( |-<\#208><\#190>!<\#221>Z<\#228><\#254><\#168><\#255>-<<\#153>jEPSF<\#182><\#9>}<\#255> <\#255><\#136>-<\#255><\#255><\#163>l 0CG3 HD:Current jobs:DRG:Cellular & molecular:Cellula or Artwork:A03.01<\#153><\#255><\#255><\#255><\#254><\#255><\#166>:FH<\#254><\#252>}@<\#153><\#157>-Oh<\#157>1<\#245><\#240><\#166><\#255><\#255><\#255><\#253>P<\#227>8<\#253><\#255>}@ p<\#153><\#168>xdI<\#216><\#168>xd2*<\#210><\#162><\#255><\#255><\#255><\#252>P<\#236><\#252><\#252><\#254>}@<\#153><\#143>'MH<\#253><\#197><\#143>'M2 <\#191><\#128><\#163><\#255><\#255><\#243>7:D<\#228>:F<\#208> q<\#248><\#250><\#254><\#249>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>XH6<\#148><\#240><\#195>3+<\#128> <\#243><\#172><\#249><\#214><\#249><\#214><\#249><\#214><\#249><\#214>P<\#231>pPDXQAAW6BN<\#255><\#255>t<\#248>5G3 HD:Current jobs:DR rG:Cellular & molecular:table 3.1&f<\#164><\#255><\#255><\#255><\#255>Q<\#232><\#156>:F<\#232><\#253><\#250>@<\#255><\#254><\#255><\#254><\#255><\#254><\#255><\#254><\#153>X<\#162>H6<\#144>L<\#188>2<\#148>u <\#243><\#172> s<\#128><\#141><\#255><\#255><\#255><\#237> <\#170>LP<\#227><\#253><\#250>@<\#153>S<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128><\#197>!8 <\#243><\#172> <\#243><\#172>Pt<\#128> t<\#143><\#255><\#255><\#243>3 <\#166><\#220>P<\#227><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> <\#243><\#172><\#128><\#145><\#255><\#255><\#243>1 uP<\#226><\#144>1 <\#253><\#250>@<\#153>0<\#242>U<\#168><\#176>8FA<\#190><\#136><\#195>4 <\#243><\#172><\#128><\#147><\#255><\#255><\#255><\#236> <\#170>LQ<\#232>H<\#253><\#250> v@<\#153>S<\#206><\#227><\#168><\#176>8<\#240>T<\#169><\#136><\#195>4<\#201>(8 <\#243><\#172> <\#243><\#172>:5<\#149><\#255><\#255><\#255><\#255>Q<\#233><\#160>0<\#228><\#252><\#250> wXG<\#253><\#197><\#148><\#240><\#195>3+<\#128> <\#148><\#255><\#255><\#255><\#255>:EP <\#166><\#148><\#252><\#250>XG<\#253><\#197><\#148><\#240><\#195>3+<\#128> x D$6!Z<\#181>l<\#237><\#220>P<\#232><\#200>8<\#255>a.3<\#134> <\#132> <\#243><\#172>Y <\#255><\#255><\#255><\#252><\#128><\#171><\#255><\#255><\#255><\#235> <\#170>L <\#166><\#136><\#253><\#250>@<\#153> yS<\#206><\#227>S<\#132><\#240>T<\#169>3+<\#128><\#217>[8 <\#243><\#172> <\#243><\#172>:0N<\#128><\#173><\#255><\#255><\#243>/P<\#229>P<\#236><\#248><\#253><\#250>@<\#153>0<\#242>US<\#132>FA<\#190>3+<\#128> z <\#243><\#172><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158><\#158> {xQ<\#231><\#212><\#255><\#255><\#242><\#132>t <\#129>