ANIMAL PHYSIOLOGY BIOL 3151: Principles of Animal Physiology Dr. Tyler Evans Email: tyler.evans@csueastbay.edu Phone: 510-885-3475 Office Hours: M,W 10:30-12:00 or appointment Website: http://evanslabcsueb.weebly.com/ LAST LECTURE TRANSMISSION AT THE NEUROMUSCULAR JUNCTION • when an action potential reaches the axon terminal of the neuromuscular junction it triggers calcium (Ca+2) channels to open • the concentration of Ca+2 inside the neuron is much lower than outside, so Ca+2 moves into the neuron along its concentration gradient • this increase in internal Ca+2 concentration triggers the release of SYNAPTIC VESICLES, synaptic vesicles contain neurotransmitters, which are then released across the synapse Textbook Fig 4.16 pg 162 LAST LECTURE Diversity in Neurophysiology • although all neurons have the same basic components, each of these components has been modified by evolution to better perform specific tasks • all neurons have DENDRITES, a CELL BODY (SOMA) and an AXON, but details of each structure are variable EXAMPLES OF NEURON DIVERSITY textbook Fig 4.18 pg 166 LAST LECTURE Diversity in Neurophysiology • OHM’S LAW describes the speed of an action potential traveling down an axon • speed or CURRENT (I) of the signal depends on two variables: voltage and resistance V= IR voltage current or V I= R resistance • essentially, the strength of the signal along an axon (current) is greatest when voltage (input energy) is high and resistance is low LAST LECTURE Diversity in Neurophysiology Altering myelination and axon diameter • fastest conduction occurs in either large diameter axons or myelinated axons textbook Table 4.3 pg. 172 LAST LECTURE Diversity in Neurophysiology Altering myelination and axon diameter • squid giant axons can obtain rapid conduction speed in the absence of myelination by increasing the diameter of its axons • squid giant axons are hundreds of times larger in diameter than mammalian axons • squid axons can be 1 mm in diameter, while most mammalian axons are 5µm in diameter Diversity in Neurophysiology Altering myelination and axon diameter • in squid, giant axons are found in the neurons that stimulate muscle contraction around the mantle cavity • squid mantle cavity is used for jet propulsion locomotion • squid can expand and contract the mantle, drawing water into the mantle cavity and rapidly expelling it through the siphon water intake mantle cavity Siphon (output) NEUROPHYSIOLOGY: Diversity in neural signaling Altering myelination and axon diameter • this form of locomotion allows the squid to move very fast, but in order for this jet propulsion to work properly, muscle fibers throughout the entire mantle must contract at THE SAME TIME! • however some parts of the mantle are much further away from the squid’s central nervous system than others CNS • in order for signals to reach all parts of the mantle at the same time, action potentials must be conducted faster in order to reach distal parts of the mantle at the same time as action potentials controlling the mantle muscles close to the CNS NEUROPHYSIOLOGY: Diversity in neural signaling How do squid ensure that signals needing to travel only a short distance reach target muscles AT THE SAME TIME as those traveling longer distances CNS NEUROPHYSIOLOGY: Diversity in neural signaling ANSWER: • axons that activate muscles at the far end of the mantle have VERY LARGE DIAMETERS • axons that activate muscles in the region of the mantle closest to the central nervous system have smaller diameters • combining axons of various diameters allows the near-simultaneous contraction of the mantle Mantle Contraction in the squid Loligo Textbook Fig 4.24 pg 177 NEUROPHYSIOLOGY: Diversity in neural signaling Why do large diameter axons conduct signals faster? • the effects of RESISTANCE explain why larger diameter axons, including squid giant axons, conduct signals more rapidly that small diameter axons • recall from last lecture: • OHM’S LAW describes the relationship between current, voltage and resistance V= IR voltage current or V I= R resistance • essentially, the strength of the signal along an axon (current) is greatest when voltage (input energy) is high and RESISTANCE IS LOW NEUROPHYSIOLOGY: Diversity in neural signaling • membrane resistance and intracellular resistance both decrease as axon diameter increases (inversely proportional) • so large diameter axons conduct signals faster by reducing resistance • resistance is caused by ions “leaking” across membranes and less “leaking” occurs in large diameter axons. V I= R • by keeping R low, large diameter axons increase conduction speed NEUROPHYSIOLOGY: Diversity in neural signaling Consequences of Increasing Axon diameter • although increasing axon diameter provides faster conduction velocity, there are two main disadvantages to using large axons to increase conduction velocity: 1. Large axons take up more space and this may limit the number of neurons that can be packed into the nervous system • mammals have nervous systems densely packed with neurons and accessory cells and cannot afford to lose additional space textbook Fig 4.19 pg. 168 NEUROPHYSIOLOGY: Diversity in neural signaling Consequences of Increasing Axon diameter • although increasing axon diameter provides faster conduction velocity, there are two main disadvantages to using large axons to increase conduction velocity: 2. Large diameter axons have a much larger volume of cytoplasm per unit length, making them energetically expensive to make and maintain. • cytoplasm contains all of the organelles (e.g. mitochondria) and many energy demanding processes occur in the cytoplasm textbook Fig 4.25 pg. 179 NEUROPHYSIOLOGY: Diversity in neural signaling Consequences of Increasing Axon diameter • as a result, giant axons are typically used when extremely high speed conduction is necessary for survival. • in squid the axons control the rapid contraction of the mantle used during escape and prey capture behaviors • Not all muscles in the mantle will contract during basic swimming NEUROPHYSIOLOGY: Diversity in neural signaling Myelinated Axons • myelinated neurons are found only in vertebrates, though lampreys and hagfish (very old lineage) lack myelin sheaths • although some invertebrates have “wrappings” that act similar to vertebrate myelin sheaths, these membrane wrappings are not as effective in signal conduction NEUROPHYSIOLOGY: Diversity in neural signaling Myelinated Axons • • • • true myelin sheaths were an important evolutionary innovation allowed rapid signal conduction in a small amount of space helped to provide conditions for complex vertebrate nervous systems more complex nervous systems allowed animals to evolve more complex behavior, physiology, social systems, etc. Mammals (lots of myelin) Lampreys/Hagfish (no myelin) NEUROPHYSIOLOGY: Diversity in neural signaling Myelinated Axons • nervous systems that rely on large diameter axons and less effective axon wrappings are simple relative to complex nervous systems seen in vertebrates • invertebrates have simple nervous systems that typically lack a central processing center like a brain NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • recall that an action potential in a neuromuscular axon terminates by triggering the release of the neurotransmitter ACETYLCHOLINE, which ultimately triggers muscle contraction. Textbook Fig 4.17 pg 163 NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • however, SYNAPTIC TRANSMISSION is incredibly diverse • animals use different types of transmission to accomplish different tasks • some neurons don’t even use neurotransmitters in signaling, but are instead are directly connected to their target cells via GAP JUNCTIONS • synapses where pre- and post-synaptic cells are connected by gap junctions are called ELECTRICAL SYNAPSES NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • in ELECTRICAL SYNAPSES, signals are sent directly from the presynaptic cell to the postsynaptic cell via GAP JUNCTIONS • in CHEMICAL SYNPASES, the signal is transmitted via chemical neurotransmitters, which cross the SYNAPTIC CLEFT and bind to a receptor on the postsynaptic cell to induce a response. textbook Fig 4.26 pg 181 NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • ELECTRICAL SYNAPSES and CHEMICAL SYNAPSES differ in a number of ways: 1. DIRECTION OF FLOW OF INFORMATION • in chemical synapses, information flows in a single direction, that is from the presynaptic cell to the post-synaptic cell e.g. vertebrate motor neuron (Fig 4.2 pg. 145) NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • ELECTRICAL SYNAPSES and CHEMICAL SYNAPSES differ in a number of ways: 1. DIRECTION OF FLOW OF INFORMATION • in electrical synapses, information can flow in BOTH DIRECTIONS • because pre- and post-synaptic cells are directly connected textbook Fig 4.26 pg 181 NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • ELECTRICAL SYNAPSES and CHEMICAL SYNAPSES differ in a number of ways: 2. SPEEED OF TRANSMISSION • electrical synapses are fast • chemical synapses are slower because of delays associated with the diffusion of neurotransmitters across the synaptic cleft and their binding to receptors on the post-synaptic neuron textbook Fig 4.26 pg 181 What type of signal would be involved in reflexes? Chemical OR Electrical NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • there is large diversity in the types of neurotransmitters (see Table 4.4 pg 184) • collectively, these features allow for more complex signaling events to occur • the proportions of electrical to chemical synapses differs among organisms What type of synaptic transmission would you expect mammals to use most often? NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • major benefit to the use of chemical signals is the diversity of signals that can be conducted: • there is a lot of variation in both the structure of chemical synapses and in the types of neurotransmitters TYPES OF SYNAPSE • axon terminal, axon vericosities, en passant synapse and spine synapse all refer to different synapse structures and therefore different functions in cells textbook Fig 4.27 pg 182 NEUROPHYSIOLOGY: Diversity in neural signaling Variation in synaptic transmission • each of these types of synapse can also be found in different locations along the neuron LOCATIONS OF SYNAPSE • dendritic synapse, axoaxonic synapse, axosomatic synapse and axodendritic synapse all refer to different synapse locations textbook Fig 4.27 pg 182 NEUROPHYSIOLOGY: Diversity in neural signaling Neurons are Unique to Animals • neurons are only found in animals, though plants do have sensory receptors, ion channels, action potentials, and can process information. • the Venus Flytrap catches prey when sensory hairs are stimulated • triggers an action potential which spreads through cells via PLASMODESMATA very similar to GAP JUNCTIONS, causing the trap to shut LECTURE SUMMARY • increases in axon diameter and axon myelination increase the speed of conduction • squid use giant axons to ensure mantle tissue contracts simultaneously • large axons take up more space and require more energy to maintain • vertebrates use myelination to increase conduction speed an innovation that helped to develop complex nervous systems • neural signal are also varied by differences in synaptic transmission • animals can alter the types of synaptic transmission (electrical or chemical), the structure of the synapse, the location of the synapse and the type of neurotransmitters • this diversity in neural signaling is necessary for the complex nervous systems found in vertebrates NEXT LECTURE: Cellular Movement and Muscles (Chapter 5)