1) Drop off in the Bi 150 box outside Baxter 331 or to the head TA (jcolas).
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1 Bi/CNS/NB 150 Problem Set 3 Due: Tuesday, Oct. 27, at 4:30 pm Instructions: 1) Drop off in the Bi 150 box outside Baxter 331 or to the head TA (jcolas). 2) Submit with this cover page. 3) Use a separate sheet of paper for each problem. 4) Type all answers if possible. 5) Use complete, grammatically correct sentences. 6) Include your name and the page number on every page. 7) Note that late problem sets receive a 10% deduction for every day past the due date. Name: Time and date submitted: Total pages (including cover page): Comments: Problem 1 grade: Problem 1 comments: Problem 2 grade: Problem 2 comments: Total grade: 1
2 Problem 1 (1.5 points): Synaptic integration Problem 1.A (0.9 points): Inhibition Many inhibitory synapses in the CNS utilize GABA A and glycine receptors. Both GABA A and glycine receptor channels are permeable to Cl - ions. 1.A.a. Calculate the Nernst potential for Cl - ions, E Cl, at 25 C. Consult the ionic concentrations listed on the fourth slide from the second half of the first lecture. E Cl = -25*ln(110/10) = -60mV (0.1 pt) E Cl = -58*log(110/10) = -60mV 1.A.b. The resting membrane potential of neurons is sometimes close to E Cl. When this is the case, how does the membrane voltage change if only GABA A receptors are activated? There is no change in membrane potential. When the membrane potential is equal to E Cl, no current flows. (0.2 pt) 2
3 1.A.c. Consider a neuron receiving two excitatory inputs as shown below. The traces at the right show the membrane potential in the soma without inhibition. i) Excitatory input 1 is stimulated alone. ii) Excitatory inputs 1 and 2 are stimulated simultaneously. Excitatory input 1 Excitatory input 2 dendrite A Inhibitory input i) ii) B 1 alone 1+2 soma V V time For each trace, redraw the trace and then superimpose the trace if a GABAergic input were active at location A during the excitatory stimulation. Repeat for location B. Explain the reasoning behind each trace. Plots: Blue: EPSP traces for input 1 alone Green: EPSP traces for inputs 1 and 2 The left plot shows EPSPs without inhibition. The middle plot shows EPSPs with inhibition at location A. The right plot shows EPSPs with inhibition at location B. Inhibition at location A efficiently blocks input 2 but has little effect on input 1. On the other hand, inhibition at location B blocks both inputs 1 and 2 efficiently. (0.15 pt for each of 4 location/stimulus combinations) 3
4 Problem 1.B (0.6 points): Active dendrites 1.B.a. Panel B in the figure below shows simultaneous recordings of the membrane potential from the soma and a dendrite when an action potential is induced by injection of current into the soma. Panel A shows the recording sites. A B Soma Dendrite V time Redraw the dendritic traces and then superimpose the dendritic voltage trace during action potentials in a mutant animal that lacks voltage-gated Na + channels at the dendrites. The action potentials are induced in the same way as before by injecting current at the soma. Assume that the only difference is in the dendrites and their lack of voltage-gated Na + channels. Explain the reasoning behind the traces. Plot: The amplitude of the dendritic potential is much lower because, given the lack of Na + channels, there is no regenerative current. However, there should still be passive spread. (0.2 pt) 4
5 1.B.b. In the case of a normal neuron, how might backpropagating spikes due to dendritic Na + channels influence the coincidence-detection properties of NMDA receptors? Backpropagation of action potentials from soma to dendrites is important for removing Mg 2+ to open the NMDA channel, which is critical for coincidence detection. (0.2 pt) A more detailed analysis: In a passive membrane it does not matter much whether synaptic inputs are clustered or distributed in dendrites as long as the total input strength is the same. That is, each synaptic potential will be summated approximately linearly. However, in active dendrites with Na + channels, the membrane potential summation can be nonlinear. For example, Na + channels may open enough only with clustered inputs because of greater local membrane depolarization relative to the case of distributed inputs. Na + channels opening further increases the membrane potential known as dendritic spiking, which can be regenerative leading to a significant difference between clustered and distributed inputs. 1.B.c. Describe the roles played by Na +, Ca 2+, and Mg 2+ ions during coincidence detection. The Na + ions depolarize the membrane potential and allow Mg 2+ ions to leave their binding (blocking) sites. This in turn allows Ca 2+ ions to flow into the postsynaptic cell, acting as a second messenger. (0.2 pt) 5
6 Problem 2 (1.5 points): Synaptic transmission Problem 2.A (0.3 points): Chronology Write the following events in the correct sequence as they occur at the neuromuscular junction of the California olive-tree rat (Rattus pasadensis) during normal synaptic transmission. If an event overlaps in time with any of the others, explain this in your answer. Begin from event 9. 1) Action potential arrives at the presynaptic terminal. 2) Muscle fiber action potential. 3) Na + enters, and K + exits through the same channels. 4) There is a local increase in intracellular Ca 2+ concentration near the Ca 2+ channels. 5) ACh is hydrolyzed to acetate and choline. 6) Choline is transported into the presynaptic terminal. 7) ACh molecules fill the synaptic vesicles. 8) ACh is released into the synaptic cleft. 9) The synaptic vesicles become acidic. (Begin here.) 10) Na + is pumped out, and K + pumped into the presynaptic terminal and muscle fiber. 11) ACh binds to ACh-gated ion channels on the postsynaptic membrane. 12) ACh diffuses within the synaptic cleft. 13) Muscle fiber reaches threshold depolarization. 14) ACh-gated ion channels open. 15) ACh-filled vesicles fuse with the presynaptic membrane. 16) Ca 2+ enters the presynaptic terminal through voltage gated Ca 2+ channels. 9 The synaptic vesicles become acidic. 7 ACh molecules fill the synaptic vesicles. 1 Action potential arrives at the presynaptic terminal. 16 Ca 2+ enters the presynaptic terminal through voltage gated Ca 2+ channels. 4 There is a local increase in intracellular Ca 2+ concentration near the Ca 2+ channels. 15 ACh-filled vesicles fuse with the presynaptic membrane. 8 ACh is released into the synaptic cleft. 12 ACh diffuses within the synaptic cleft. 11 ACh binds to ACh-gated ion channels on the postsynaptic membrane. 14 ACh-gated ion channels open. 3 Na + enters, and K + exits through the same channels. 13 Muscle fiber reaches threshold depolarization. 2 Muscle fiber action potential. 5 ACh is hydrolyzed to acetate and choline. 6 Choline is transported into the presynaptic terminal. 10 Na + is pumped out, and K + pumped into the presynaptic terminal and muscle fiber. 6
7 However, note the following: Event 10 overlaps with all events because it occurs not only after the action potential, but also at rest to balance the leaky Na + and K + currents. Events 5 and 6 take place all along with events 8, 12, 11, 14, 3, 13, and 2. (0.02 pt for each of 15 events following event 9) Problem 2.B (1.2 points): Excitatory postsynaptic potentials You are in the lab recording from neuron pairs each connected by a glutamatergic synapse in four different mutant animals and in a wild-type animal. In each animal a different protein involved in neurotransmitter release is mutated. You stimulate the presynaptic cell and record from the postsynaptic neuron of each pair. For each mutation below, describe the following: i) After the first stimulus, how does the excitatory postsynaptic potential (EPSP) you record from the postsynaptic neuron differ from the EPSP recorded from a wild-type animal? Does it increase, decrease, stay comparatively the same, or fail to be generated at all? ii) After five stimulations of the presynaptic cell in quick succession, how does the EPSP you record from the mutant animal differ from that in the same experiment on a wild-type animal? Explain your reasoning in each case. The mutations are as follows: 2.B.a. Mutation of the voltage-gated Ca 2+ channels of the presynaptic terminal. The mutant channels have a lower opening threshold. i) Increases the EPSP. (0.15 pt) - Lower threshold of Ca 2+ channels. - Same stimulus leads to greater Ca 2+ influx. - Ca 2+ binds to synaptotagmin on docked vesicles. - More vesicles fuse. - More neurotransmitter is released. - More post-synaptic channels open. - Greater EPSP. 7
8 ii) Both of the following answers can be valid depending on the reasoning. (0.15 pt) First possibility: Slightly increases the EPSP. - More residual Ca 2+ in the presynaptic terminal due to previous stimulations. - Ca 2+ /calmodulin activated. - Synapsin phosphorylated. - Storage vesicles released from cytoskeleton. - More vesicles docked and ready to go at membrane. - More fusion. - More neurotransmitter release. - More postsynaptic channel opening. - Greater EPSP. Second possibility: Slightly decreases the EPSP. - All vesicles docked and ready to go have been used by previous stimuli. - Fewer vesicles ready at the membrane. - Less neurotransmitter release. - Less postsynaptic channel opening. - Lesser EPSP. 2.B.b. Mutation of the Ca 2+ binding sites of synaptotagmin. The mutant protein does not bind Ca 2+. i) No EPSP due to synaptotagmin-independent asynchronous release. (0.15 pt) ii) No EPSP due to synaptotagmin-independent asynchronous release. (0.15 pt) - Synaptotagmin is unable to act as a Ca2+ sensor. - Vesicles do not fuse with the membrane. - Neurotransmitter is not released. - Postsynaptic receptors are not opened. - No EPSP. 2.B.c. Mutation of the glutamate transporter to inhibit its function. Several redundant processes maintain the intracellular glutamate concentration of ~10 mm. Contrast this with the concentration of ~1 μm in cerebrospinal fluid. i) Increases the EPSP. (0.15 pt) - Glutamate is not being taken back into the presynaptic terminal. - Glutamate spends more time in the synaptic cleft. - Increasing opening events of postsynaptic channels. - Longer/increased EPSP. 8
9 ii) Increases the EPSP. (0.15 pt) - Glutamate is not being taken back into the presynaptic terminal, so it simply diffuses. - Redundant mechanisms maintain intracellular glutamate so that vesicles can still be refilled and released. - Glutamate spends more time in the synaptic cleft. - Increasing opening events of postsynaptic channels. - Longer/increased EPSP. 2.B.d. Knockout of N-ethylmaleimide-sensitive factor (NSF). i) Neither increases nor decreases the EPSP. (0.15 pt) - Docked vesicles are available. - NSF is only necessary for recycling. - Therefore, no effect during the first stimulation. ii) Decreases the EPSP. (0.15 pt) - Due to the lack of NSF, SNARE complex proteins cannot be recycled. - New vesicles cannot be docked. - When stimulus comes in, fewer vesicles will be able to respond to Ca 2+ influx. - Less neurotransmitter will be released. - Lesser EPSP. 9
1) Drop off in the Bi 150 box outside Baxter 331 or to the head TA (jcolas).
Bi/CNS/NB 150 Problem Set 3 Due: Tuesday, Oct. 27, at 4:30 pm Instructions: 1) Drop off in the Bi 150 box outside Baxter 331 or e-mail to the head TA (jcolas). 2) Submit with this cover page. 3) Use a
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