© 2013 Pearson Education, Inc. Figure Synapses. Axodendritic synapses Dendrites Cell body Axoaxonal synapses Axon Axosomatic synapses Axon Axosomatic synapses Cell body (soma) of postsynaptic neuron

© 2013 Pearson Education, Inc. Varieties of Synapses: Electrical Synapses Less common than chemical –Electrically coupled (joined by gap junctions that connect cytoplasm of adjacent neurons) Very rapid Unidirectional or bidirectional Synchronize activity –More abundant in: Embryonic nervous tissue

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Presynaptic neuron Action potential arrives at axon terminal. Mitochondrion Axon terminal Synaptic cleft Synaptic vesicles Postsynaptic neuron Postsynaptic neuron Presynaptic neuron 1

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Presynaptic neuron Action potential arrives at axon terminal. Voltage-gated Ca 2+ channels open and Ca 2+ enters the axon terminal. Mitochondrion Axon terminal Synaptic cleft Synaptic vesicles Postsynaptic neuron Postsynaptic neuron Presynaptic neuron 1 2

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Presynaptic neuron Action potential arrives at axon terminal. Voltage-gated Ca 2+ channels open and Ca 2+ enters the axon terminal. Ca 2+ entry causes synaptic vesicles to release neurotransmitter by exocytosis Mitochondrion Axon terminal Synaptic cleft Synaptic vesicles Postsynaptic neuron Postsynaptic neuron Presynaptic neuron 1 2 3

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Presynaptic neuron Action potential arrives at axon terminal. Voltage-gated Ca 2+ channels open and Ca 2+ enters the axon terminal. Ca 2+ entry causes synaptic vesicles to release neurotransmitter by exocytosis Neurotransmitter diffuses across the synaptic cleft and binds to specific receptors on the postsynaptic membrane. Mitochondrion Axon terminal Synaptic cleft Synaptic vesicles Postsynaptic neuron Postsynaptic neuron Presynaptic neuron

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Ion movement Graded potential Binding of neurotransmitter opens ion channels, resulting in graded potentials. 5

© 2013 Pearson Education, Inc. Figure Chemical synapses transmit signals from one neuron to another using neurotransmitters. Enzymatic degradation Diffusion away from synapse Neurotransmitter effects are terminated by reuptake through transport proteins, enzymatic degradation, or diffusion away from the synapse. Reuptake 6

© 2013 Pearson Education, Inc. Synaptic Delay Time for NT to be released, diffuse across synapse, and bind to receptors –0.3–5.0 ms Synaptic delay is rate-limiting step of neural transmission

© 2013 Pearson Education, Inc. Postsynaptic Potentials NT receptors cause graded potentials that vary in strength with: –Amount of NT released –Time nt stays in area

© 2013 Pearson Education, Inc. Table 11.2 Comparison of Graded Potentials and Action Potentials (1 of 4)

© 2013 Pearson Education, Inc. Table 11.2 Comparison of Graded Potentials and Action Potentials (2 of 4)

© 2013 Pearson Education, Inc. Table 11.2 Comparison of Graded Potentials and Action Potentials (3 of 4)

© 2013 Pearson Education, Inc. Table 11.2 Comparison of Graded Potentials and Action Potentials (4 of 4)

© 2013 Pearson Education, Inc. Postsynaptic Potentials Types of postsynaptic potentials –EPSP—excitatory postsynaptic potentials –IPSP—inhibitory postsynaptic potentials

© 2013 Pearson Education, Inc.

Excitatory Synapses and EPSPs NT binding opens chemically gated channels Allows simultaneous flow of Na + and K + in opposite directions Na + influx greater than K + efflux  net depolarization called EPSP (not AP) EPSP help trigger AP if EPSP is of threshold strength – Can spread to axon hillock, trigger opening of voltage-gated channels, and cause AP to be generated

© 2013 Pearson Education, Inc. Figure 11.18a Postsynaptic potentials can be excitatory or inhibitory. An EPSP is a local depolarization of the postsynaptic membrane that brings the neuron closer to AP threshold. Neurotransmitter binding opens chemically gated ion channels, allowing Na + and K + to pass through simultaneously. Threshold Stimulus –55 –70 Time (ms) Membrane potential (mV) Excitatory postsynaptic potential (EPSP)

© 2013 Pearson Education, Inc. Inhibitory Synapses and IPSPs Reduces postsynaptic neuron's ability to produce an action potential –Makes membrane more permeable to K + or Cl – If K + channels open, it moves out of cell If Cl - channels open, it moves into cell –Therefore neurotransmitter hyperpolarizes cell Inner surface of membrane becomes more negative AP less likely to be generated

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Figure 11.18b Postsynaptic potentials can be excitatory or inhibitory. Threshold Stimulus –55 –70 Time (ms) Membrane potential (mV) An IPSP is a local hyperpolarization of the postsynaptic membrane that drives the neuron away from AP threshold. Neurotransmitter binding opens K + or Cl – channels. Inhibitory postsynaptic potential (IPSP)

© 2013 Pearson Education, Inc. Synaptic Integration: Summation A single EPSP cannot induce an AP EPSPs can summate to influence postsynaptic neuron IPSPs can also summate Most neurons receive both excitatory and inhibitory inputs from thousands of other neurons –Only if EPSP's predominate and bring to threshold  AP

© 2013 Pearson Education, Inc. Two Types of Summation Temporal summation –One or more presynaptic neurons transmit impulses in rapid-fire order Spatial summation –Postsynaptic neuron stimulated simultaneously by large number of terminals at same time

© 2013 Pearson Education, Inc. Figure 11.19a Neural integration of EPSPs and IPSPs. Threshold of axon of postsynaptic neuron Resting potential 0 Membrane potential (mV) –55 –70 E1E1 E1E1 E1E1 No summation: 2 stimuli separated in time cause EPSPs that do not add together. Excitatory synapse 1 (E 1 ) Excitatory synapse 2 (E 2 ) Inhibitory synapse (I 1 ) Time

© 2013 Pearson Education, Inc. Figure 11.19b Neural integration of EPSPs and IPSPs. Threshold of axon of postsynaptic neuron Resting potential 0 Membrane potential (mV) –55 –70 E1E1 Excitatory synapse 1 (E 1 ) Excitatory synapse 2 (E 2 ) Inhibitory synapse (I 1 ) Time E1E1 E1E1 Temporal summation: 2 excitatory stimuli close in time cause EPSPs that add together.

© 2013 Pearson Education, Inc. Figure 11.19c Neural integration of EPSPs and IPSPs. Threshold of axon of postsynaptic neuron Resting potential 0 Membrane potential (mV) –55 –70 E1E1 Excitatory synapse 1 (E 1 ) Excitatory synapse 2 (E 2 ) Inhibitory synapse (I 1 ) Time E 1 + E 2 Spatial summation: 2 simultaneous stimuli at different locations cause EPSPs that add together. E2E2

© 2013 Pearson Education, Inc. Figure 11.19d Neural integration of EPSPs and IPSPs. Threshold of axon of postsynaptic neuron Resting potential 0 Membrane potential (mV) –55 –70 E1E1 Excitatory synapse 1 (E 1 ) Excitatory synapse 2 (E 2 ) Inhibitory synapse (I 1 ) Time Spatial summation of EPSPs and IPSPs: Changes in membane potential can cancel each other out. l1l1 E 1 + l 1 l1l1

© 2013 Pearson Education, Inc. Integration: Synaptic Potentiation Repeated synapse increases ability of presynaptic cell to excite postsynaptic neuron –Ca 2+ conc increases in presynaptic terminal and postsynaptic neuron Brief high-frequency stimulation partially depolarizes postsynaptic neuron –Chemically gated channels allow Ca 2+ entry –Ca 2+ promotes more effective responses to subsequent stimuli

© 2013 Pearson Education, Inc. Integration: Presynaptic Inhibition Excitatory nt release by one neuron inhibited by another neuron via an axoaxonic synapse Less neurotransmitter released Smaller EPSPs formed

© 2013 Pearson Education, Inc. Neurotransmitters Language of nervous system 50 or more nt have been identified Most neurons make two or more nts –Neurons can exert several influences Usually released at different stimulation frequencies Classified by chemical structure and by function

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Acetylcholine (ACh) –First identified; best understood –Released at neuromuscular junctions,by some ANS neurons, by some CNS neurons –Synthesized from acetic and choline by enzyme choline acetyltransferase –Degraded by enzyme acetylcholinesterase (AChE)

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Biogenic amines Catecholamines –Dopamine, norepinephrine (NE), and epinephrine –Synthesized from amino acid tyrosine Indolamines –Serotonin and histamine –Serotonin synthesized from amino acid tryptophan; histamine synthesized from amino acid histidine Broadly distributed in brain –Play roles in emotional behaviors and biological clock Some ANS motor neurons (especially NE) Imbalances associated with mental illness

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Amino acids Glutamate Aspartate Glycine GABA—gamma (  )-aminobutyric acid

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Peptides (neuropeptides) Substance P –Mediator of pain signals Endorphins –Beta endorphin, dynorphin and enkephalins –Act as natural opiates; reduce pain perception Gut-brain peptides –Somatostatin and cholecystokinin

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Purines –ATP! –Adenosine Potent inhibitor in brain Caffeine blocks adenosine receptors –Act in both CNS and PNS –Produce fast or slow responses –Induce Ca 2+ influx in astrocytes

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure Gases and lipids - gasotransmitters Nitric oxide (NO), carbon monoxide (CO), hydrogen sulfide gases (H 2 S) Bind with G protein–coupled receptors in the brain Lipid soluble Synthesized on demand NO involved in learning and formation of new memories; brain damage in stroke patients, smooth muscle relaxation in intestine H 2 s acts directly on ion channels to alter function

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Chemical Structure –Endocannabinoids Act at same receptors as THC (active ingredient in marijuana) –Most common G protein-linked receptors in brain Lipid soluble Synthesized on demand Believed involved in learning and memory May be involved in neuronal development, controlling appetite, and suppressing nausea

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Function Great diversity of functions Can classify by –Effects – excitatory versus inhibitory –Actions – direct versus indirect

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Function Effects - excitatory versus inhibitory –Neurotransmitter effects can be excitatory (depolarizing) and/or inhibitory (hyperpolarizing) –Effect determined by receptor to which it binds GABA and glycine usually inhibitory Glutamate usually excitatory Acetylcholine and NE bind to at least two receptor types with opposite effects –ACh excitatory at neuromuscular junctions in skeletal muscle –ACh inhibitory in cardiac muscle

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Direct versus Indirect Actions Direct action –Neurotransmitter binds to and opens ion channels –Promotes rapid responses by altering membrane potential –Examples: ACh and amino acids

© 2013 Pearson Education, Inc. Classification of Neurotransmitters: Direct versus Indirect Actions Indirect action –NT acts through second messengers, usually G protein pathways –Broader, longer-lasting effects similar to hormones –Biogenic amines, neuropeptides, and dissolved gases

© 2013 Pearson Education, Inc. Figure Indirect neurotransmitter receptor mechanism: G protein-inked receptors a 5b 5c Ligand (1st messenger) ReceptorG proteinEnzyme 2nd messenger Recall from Chapter 3 that G protein signaling mechanisms are like a molecular relay race. Neurotransmitter (1st messenger) binds and activates receptor. Receptor G protein Adenylate cyclase Closed ion channelOpen ion channel GDP Receptor activates G protein. G protein activates adenylate cyclase. Adenylate cyclase converts ATP to cAMP (2nd messenger). Active enzyme cAMP changes membrane permeability by opening or closing ion channels. cAMP activates enzymes. cAMP activates specific genes. Nucleus

© 2013 Pearson Education, Inc. Channel-Linked (Ionotropic) Receptors: Mechanism of Action Ligand-gated ion channels Action is immediate and brief Excitatory receptors are channels for small cations –Na + influx contributes most to depolarization Inhibitory receptors allow Cl – influx that causes hyperpolarization

© 2013 Pearson Education, Inc. Basic Concepts of Neural Integration Neurons function in groups Groups contribute to broader neural functions There are billions of neurons in CNS –Must be integration so the individual parts fuse to make a smoothly operating whole

© 2013 Pearson Education, Inc. Organization of Neurons: Neuronal Pools Functional groups of neurons –Integrate incoming information received from receptors or other neuronal pools –Forward processed information to other destinations Simple neuronal pool –Single presynaptic fiber branches and synapses with several neurons in pool –Discharge zone—neurons most closely associated with incoming fiber –Facilitated zone—neurons farther away from incoming fiber

© 2013 Pearson Education, Inc. Presynaptic (input) fiber Facilitated zone Discharge zoneFacilitated zone Figure Simple neuronal pool.

© 2013 Pearson Education, Inc. Types of Circuits Circuits –Patterns of synaptic connections in neuronal pools Four types of circuits –Diverging –Converging –Reverberating –Parallel after-discharge

© 2013 Pearson Education, Inc. Figure 11.23a Types of circuits in neuronal pools. Input Diverging circuit One input, many outputs An amplifying circuit Example: A single neuron in the brain can activate 100 or more motor neurons in the spinal cord and thousands of skeletal muscle fibers Many outputs

© 2013 Pearson Education, Inc. Figure 11.23b Types of circuits in neuronal pools. Converging circuit Many inputs, one output A concentrating circuit Example: Different sensory stimuli can all elicit the same memory Input 1 Input 2Input 3 Output

© 2013 Pearson Education, Inc. Figure 11.23c Types of circuits in neuronal pools. Reverberating circuit Signal travels through a chain of neurons, each feeding back to previous neurons An oscillating circuit Controls rhythmic activity Example: Involved in breathing, sleep-wake cycle, and repetitive motor activities such as walking Output Input

© 2013 Pearson Education, Inc. Figure 11.23d Types of circuits in neuronal pools. Output Input Parallel after-discharge circuit Signal stimulates neurons arranged in parallel arrays that eventually converge on a single output cell Impulses reach output cell at different times, causing a burst of impulses called an after-discharge Example: May be involved in exacting mental processes such as mathematical calculations

© 2013 Pearson Education, Inc. Patterns of Neural Processing: Serial Processing Input travels along one pathway to a specific destination System works in all-or-none manner to produce specific, anticipated response Example – spinal reflexes –Rapid, automatic responses to stimuli –Particular stimulus always causes same response –Occur over pathways called reflex arcs Five components: receptor, sensory neuron, CNS integration center, motor neuron, effector

© 2013 Pearson Education, Inc. Figure A simple reflex arc. Interneuron Spinal cord (CNS) Stimulus Receptor Sensory neuron Integration center Motor neuron Effector Response

© 2013 Pearson Education, Inc. Patterns of Neural Processing: Parallel Processing Input travels along several pathways Different parts of circuitry deal simultaneously with the information –One stimulus promotes numerous responses Important for higher-level mental functioning Example: a sensed smell may remind one of an odor and any associated experiences

© 2013 Pearson Education, Inc. Developmental Aspects of Neurons Nervous system originates from neural tube and neural crest formed from ectoderm The neural tube becomes CNS –Neuroepithelial cells of neural tube proliferate to form number of cells needed for development –Neuroblasts become amitotic and migrate –Neuroblasts sprout axons to connect with targets and become neurons

© 2013 Pearson Education, Inc. Axonal Growth: Finding the Target Growth cone at tip of axon interacts via: –Cell surface adhesion proteins provide anchor points –Neurotropins that attract or repel the growth cone –Nerve growth factor (NGF) keeps neuroblast alive Must find target and right place to form synapse –Astrocytes provide physical support About 2/3 of neurons die before birth –If do not form synapse with target –Many cells also die due to apoptosis (programmed cell death) during development

© 2013 Pearson Education, Inc. Figure A neuronal growth cone.