Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1

Appendix Basics of the Nervous System

Chapter Outline: I. Introduction to the Nervous System II. Overvie of the Nervous System A. Overa!! Organi"ation B. #he Brain $. #he %ore&rain i. 'iencepha!on ii. (o&es of the Brain III. #he Neuron: $onnecting 'ifferent parts of the Brain A. Structure B. %unction i. #he )esting Potentia! ii. #he Action Potentia! I*. #he Synapse: +a,ing the Brain %unction and (earn A. Structure B. %unction i. Preparation ii. )e!ease iii. Binding iv. )emova! *. $omp!eting the $irc!e: -enerating a Ne Action Potentia! A. Summation *I. #he $ortex *II. Sensory $oding or .o does the Brain a&!e to (et /s 0no 1hat is -oing on Out #here2 A. $oding &y a sing!e ce!! B. $oding across many ce!!s

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.3

Introduction to the Nervous System #his text is a&out the fascinating and often surprising or!d of our sensory systems and ho they operate to a!!o us the the rich and varied experiences of the or!d that e en4oy. +uch of the operation of sensory systems is a!so the tai! of the operation of the nervous system. A &asic ,no !edge of the structure and operation of the nervous system i!! &e he!pfu! in understanding some of the topics discussed in the main chapters of the text and this appendix i!! provide the needed &ac,ground. So hat is the nervous system2 #he nervous system is the part of our &ody made up of neurons and their associated support ce!!s. #here are5 &y some estimates5 over a 1 &i!!ion neurons 6)E%E)EN$E7 and perhaps over 18 times as many support ce!!s. $o!!ective!y5 the nervous system contro!s and coordinates the functions of the rest of the &ody. As organisms &ecame more comp!ex5 there needed to &e a ay for the organism to &e a&!e to have a!! of its disparate parts function together5 for examp!e5 move in a common direction. #he nervous system evo!ved to a!!o the organism to achieve this goa!. Overview of the Nervous System -iven the comp!exity of the human nervous system5 it is a onder that e can understand as much as e a!ready do. %ortunate!y5 the nervous system is high!y organi"ed and not a random co!!ection of interconnecting neurons. 1ithout these patterns of organi"ation5 the nervous system ou!d pro&a&!y e!ude comprehension5 and pro&a&!y ou!d not function as e!! either. #he next fe sections i!! give an overvie of the organi"ation of the nervous system. Overall Organization #he nervous system can &e su&divided into severa! main divisions as sho n in %igure AP 1.1. Brief!y5 the nervous system is divided into t o ma4or divisions: the centra! nervous system 6$NS7 hich is contained ithin &ony cases5 in particu!ar the s,u!! and spine5 and the periphera! nervous system 6PNS7 hich spreads through the rest of the &ody. #he centra! nervous system primari!y processes information and coordinates the operation of the rest of the &ody. #he periphera! nervous system primari!y functions to carry signa!s to the &ody from the &rain and from the &ody to the &rain. In other ords5 the periphera! nervous system carries the decisions of the centra! nervous system to the part of the &ody to carry out the action and ,eeps the centra! nervous system informed of the state of the &ody and the environment so that the &rain can &est function.

Figure AP 1.1. The major divisions of the nervous system. See the text for details.
#he centra! nervous system can &e further divided into t o parts5 the &rain5 in the s,u!!5 and the spina! chord in the spine. #he &rain has the great ma4ority of the neurons of the nervous system and primari!y functions to process information and is the seat of the most comp!ex processing. #he spina! chord dua!!y functions to carry signa!s to and from the &rain and performs many !ess comp!icated operations itse!f. #he periphera! nervous system can a!so &e divided into t o further divisions5 the somatic and autonomic. #he somatic division connects to striated or s,e!eta! musc!es. #he information coming from the sensory systems is a!so considered part

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.9

of the somatic division of the periphera! nervous system. Sometimes the somatic division is ca!!ed the vo!untary nervous system as it contro!s the s,e!eta! musc!es e can vo!untari!y move and receives input from the sensory systems that e are conscious!y attuned to. #he autonomic &ranch is so ca!!ed &e cause it contro!s so:ca!!ed automatic functions such as heart rate and &reathing that e do not conscious!y contro!. #here are t o &ranches of the autonomic nervous system5 the sympathetic and parasympathetic. #he sympathetic or,s to increase energy uti!i"ation. It is part of the emergency response system of our &ody. It increases heart rate5 &reathing and re!ated functions hi!e inhi&iting digestion and other systems that impede our emergency responses. #he parasympathetic in most cases operates on the same systems &ut has the opposite effect. %or examp!e5 it s!o s heart rate and &reathing and stimu!ates digestion. %rom these o&servations one may conc!ude that the parasympathetic system operates to ca!m systems and or, to increase energy savings. A!so5 since the parasympathetic system is not near!y as interconnected as the sympathetic system it can he!p to modu!ate or ad4ust the operation of the sympathetic nervous system hich tends to operate as a comp!ete system. #he parasympathetic nervous system cou!d ,eep &reathing from increasing as much as the heart rate depending upon the situation and in this ay a!!o the operation of the sympathetic system to &e more sensitive to particu!ar situations. 1ith this &ac,ground5 !et us !oo, a &it more c!ose!y at the &rain. The Brain %igure AP 1.3 sho s the &rain in t o vie s5 from the side and !oo,ing at the midd!e of a &rain that has &een cut in ha!f. #he &rain is a !arge mushrooming of the nervous system at the top of the spina! chord. #he adu!t &rain eighs a&out 15988 to 15;88 grams hich is a fair a&out of eight for the de!icate spina! chord to carry. #o he!p support and protect the &rain5 it f!oats in a sea of cere&ro:spina! f!uid. #he f!uid cushions the &rain against norma! &!o s to the head that might other ise cause damage. Additiona! protection is provided &y the s,u!! and three !ayers of mem&ranes ,no n co!!ective!y as the meninges.

Figure AP 1.2. The Brain. On the left is the view from side. On the right is the view loo ing at the !rain that has !een "ut in half. #he centra! nervous system is typica!!y divided into three main regions as sho n in %igure AP 1.9. #hese divisions are derived from studying the deve!opment of the &rain. Starting at the &ase of the &rain5 the hind &rain is made up of the medu!!a5 the pons and the cere&e!!um. #he medu!!a is5 in appearance5 a s e!!ing of the spina! chord. It is !arge!y tracts of fi&ers carrying information to and from the rest of the &rain. .o ever5 there are important centers5 ca!!ed nuc!ei 6or a nucleus in singu!ar7. )e!ated to sensation an perception there are centers re!ated to hearing5 e.g5 the coch!ear nuc!eus5 taste5 e.g. the nuc!eus of the so!itary tract5 and the !emnisca! path ays of the somatosensory system. #he pons rests a&ove the medu!!a. #o ards the &ac, of the heard from &oth the medu!!a and the pons and the medu!!a is the cere&e!!um. #he cere&e!!um is a comp!ex figure that seems invo!ved in some forms of !earning and in some e!ements of motor contro!. Sti!!5 e i!! not run into it in the course of our discussions a&out sensation and perception5 though it ou!d not surprise me if in the future e did not need to ,no more a&out this structure.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.;

Figure AP 1.#. The three major divisions of the !rain.

#he mid&rain is a very sma!! region of the &rain as sho n in %igure 1.9. It is made up of t o parts5 the tectum and tegmentum. #he tegmentum p!ays some important ro!es in movement. #he tectum is divided into t o parts the superior co!!icu!us and inferior co!!icu!us. #hey form t o &umps at the &ac, of the &rain stem as sho n in %igure 1.;. 1hi!e the mid&rain is <uite sma!! it these t o structures are <uite important e!ements of our sensory systems. #he superior co!!icu!us p!ays a ro!e in the visua! system. In fact in anima!s that !ac, a fore&rain5 such as repti!es5 amphi&ians5 and &irds5 the superior co!!icu!us5 ca!!ed the optic tectum in these anima!s5 is the primary visua! center in the &rain. #he inferior co!!icu!us is an important processing center for our auditory systems.

Figurer AP 1.$. The %nferior and Su&erior 'olli"ulus of the (id!rain.

The Fore rain A&ove the mid&rain5 the &rain expands into the fore&rain. #he fore&rain can &e divided into severa! su&:regions. In this section5 I i!! focus on those structures that are most re!evant to the sensory systems and their geographica! re!ationship to each other. The !iencephalon" At the top of the mid&rain are three structures that are sti!! considered part of the &rain stem 60o!& = 1hisha 5 1>>?7. $o!!ective!y they are ca!!ed the diencepha!on5 and individua!!y they are the hypotha!amus5 thalamus5 and

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.@

pinea! &ody 6%igure AP 1.@7. #he pinea! &ody p!ays a ro!e in regu!ating our dai!y cyc!es such as our s!eepA a,e cyc!es. #he hypotha!amus is vita! in contro!!ing many of our &asic &io!ogica! functions inc!uding eating. In addition5 the autonomic system is !arge!y contro!!ed from the hypotha!amus. %rom the perspective of the sensory systems5 the tha!amus is the most important of these three three structures. #he t o tha!ami5 one in each hemisphere5 are made up of numerous different nuc!ei5 areas for synapses to occur. #hese different nuc!ei connect to and receive connections from most regions of the cortex of the fore&rain. Our sensory systems a!! connect ith the tha!amus prior to proceeding to the cortex. So the visua! system connects to the !atera! genicu!ate nuc!eus of the tha!amus &efore reaching the visua! cortex5 and the auditory system connects to the media! genicu!ate nuc!eus of the tha!amus &efore reaching the cortex.

Figure AP 1.). The *ien"e&halon showing the hy&othalamus+ thalamus+ and &ineal !ody. ,%-,A.T A /0(A. B1A%. FO1 T/2 T23T
The #o es of the Brain" #he fore&rain is divided into four genera! regions of the &rain to assist us in ,no ing here a part of the fore&rain is to &e found. #hese !o&es are the fronta! !o&e5 in the front of the &rain5 the tempora! !o&es on the side5 the parieta! !o&e5 found on top5 and the occipita! found on the &ac, 6%igure AP 1.?7. #hese !o&es are names after the &ones of the s,u!! that they are under. #he top !ayer of the !o&es is the cortex here most of the synaptic connections of the fore&rain are to &e found. #he occipita! !o&e contains the visua! cortex. In fact5 the occipita! !o&e is primari!y visua! in function. #he tempora! !o&e has the auditory cortex a!ong the top hori"onta! &ump. #he somatosensory cortex is in the front of the parieta! !o&e. #aste and sme!! are found near&y. %urther discussion of the cortex i!! &e saved for !ater. No it is time to examine the ce!!s that ma,e up the &rain and ho they interact ith each other.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.?

%llustration 18 Figure AP 1.9. -o!es of the !rain. % .22* TO O:21-A; T/2 :%S0A-+ A0*%TO1;+ A.* SO(ATOS2.SO1; 'O1T23 O. T/%S %(A62.
The Neuron: Connecting !ifferent parts of the Brain Structure #he neuron $to glossary% is a ce!! of the &ody. #hat may seem a trivia! statement &ut this statement revea!s an important feature a&out theories. #he more genera! the theory the &etter e !i,e it. #hat &odies are made of ce!!s is a theory. An incredi&!y e!! supported theory and one of the most genera! theories in &io!ogy. It defines the ay e !oo, at !iving organisms. It is not the point of this text5 or this appendix for that matter5 to discuss ce!!s5 &ut ,no ing that a neuron is a ce!! te!!s us a !ot a&out neurons. #hey have a!! the genera! features of ce!!s: a nuc!eus5 cytop!asm5 !ysosomes5 mitochondria5 a ce!! mem&rane 6since this is an anima! ce!!75 and severa! other common features. Saying that the neuron is a ce!! says a great dea!. %igure AP 1.B sho s a standard ce!! and some of its parts.

Figure AP 1.4. A diagram of a "ell 5,e need a !etter image. This is from 6rays Anatomy and is not "o&yrighted !ut it is dated7.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.B

1e get so used to hat a!! is &eing said in such apparent!y simp!e statements that e sometimes forget the onder of it. It as not a! ays certain that the nervous system as !i,e the rest of the &ody and made up of ce!!s. As recent!y as the &eginning of the 38th century5 the de&ate raged even in No&e! Pri"e (ectures 6-E# A )E%E)EN$E7. But5 the fact that neurons are ce!!s does not te!! us everything. Neurons have severa! specia! characteristics that ma,e them different from other ce!!s and function in the ays needed for our nervous system. %igure AP 1.C sho s a standard diagram of a neuron. #he additiona! features of a neuron that are important to note inc!ude the dendrites $to glossary%5 soma $to glossary%5 a&on $to glossary% and terminals $to glossary%. #he dendrites receive information from other neurons. #heir function i!! descri&ed &e!o hen the synapse is discussed. #he soma is the ce!! &ody. Soma comes from the -ree, ord for &ody. #he axon is the part of the neuron that conveys the signa! from one !ocation to another. On many axons5 there are other ce!!s in the that form an insu!ating !ayer ca!!ed mye!in. #his mye!in sheath is important in speeding up communication ithin the nervous system. #he termina!s are in c!ose contact ith dendrites and somas and are the other part of a synapse. #hey may a!so contact musc!es or g!ands.

Figure AP 1.<. A standard diagram of a neuron. Function #he neuron are hard or,ing units of the &rain. #o function5 the neurons re<uire a great dea! of energy to function5 much of this energy comes from the food that e eat on a dai!y &asis. #o expend energy re<uires that the system store energy to &e a&!e to re!ease it. Dust as a f!ash!ight re<uires stored energy5 in the form of a &attery5 the neuron must store energy to &e a&!e to function. Energy that is stored5 that is5 avai!a&!e to do or,5 is ca!!ed potential energy and the energy doing the or, is ca!!ed 'inetic energy. 0inetic comes from the ancient -ree, ord for motion as energy doing or, often causes motion of some sort and this circumstance i!! not &e an exception. In the neuron5 the potentia! energy is referred to as the (esting )otential and the ,inetic energy is referred to as the *ction )otential. #he ,ey to the functioning of &oth the resting potentia! and the action potentia! !ies in the c!ever!y organi"ed ce!! mem&rane 6%igure AP 1.>7. 1ithout going into too much detai!5 the ce!! mem&rane is a phospho!ipid &i:!ayer. #he phospo!ipid mo!ecu!e5 sho as a &a!! 6the phosphate part7 and t o stic,s 6the !ipid part7 is the ,ey to the mem&rane. #he phosphate part is s!ight!y negative!y charged hi!e the !ipid part is e!ectrica!!y neutra!. #he importance of this feature of the mo!ecu!e comes from the fact that is surrounded &y ater mo!ecu!es. A ater mo!ecu!e5 hi!e overa!! e!ectrica!!y neutra!5 is s!ight!y po!ari"ed5 or charged5 in across its !ength. #he oxygen part of the ater mo!ecu!e is s!ight!y negative and the hydrogen part is s!ight!y positive. )eca!! that opposite charges attract hi!e negative charges repe! and no the formation of the ce!! mem&rane is c!ear. #he phosphate part of the phospho!ipid mo!ecu!e is dra n to ards the hydrogen part of the ater mo!ecu!e hich turns the !ipid part a ay from the ater mo!ecu!e. As a resu!t the phosphate head is referred to as hydrophi!ic or ater !oving and the !ipid tai!s are ca!!ed hydropho&ic or ater scared. #he easiest form for the phospho!ipids to group together to ma,e &oth parts of the mo!ecu!es exist in the happiest environment possi&!e is to form a sphere ith ater inside and outside and for the mo!ecu!es to &e in t o !ayers ith the phosphate heads pointed to the inside and the outside of the sphere. In other ords5 to form a mem&rane composed of a phospho!ipid &i!ayer.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.C

Figure AP 1.=. A diagram of a &ortion of the "ell mem!rane. The "ell mem!rane is made u& of two layers of &hos&holi&ids shown !y the !all 5&hos&hate &art7 and two sti" s 5li&id &art7. From Smo" 51===7. #he mo!ecu!es of the ce!! mem&rane are he!d together &y nothing more than these forces. .o ever5 this mem&rane5 unmodified5 is ho!!y impenetra&!e to the sma!! charged partic!es that are going to &e important in the resting potentia! and the action potentia!. #herefore5 proteins are produced in the ce!! that are pushed through the mem&rane5 crossing it from one end to the other. #hese proteins5 usua!!y a sma!! group of proteins5 form a pore or ho!e through hich sma!! charged partic!es5 ca!!ed ions5 can trave!. #hese proteins that a!!o for ion to cross the ce!! mem&rane are ca!!ed ion channe!s. 1ith this &ac,ground5 the resting potentia! can no &e descri&ed. 'O I 1AN# #O 'O A SE-E+EN# ON 'I%%/SION AN' E(E$#)I$A( -)A'IEN#S2 (esting )otential" 1e i!! !oo, at the axon during the resting potentia! in t o different ays. %irst5 the e!ectrica! of vo!tage of the resting potentia! i!! &e descri&ed and then the chemica! state of the neuron that supports the e!ectrica! vo!tage. #he potentia! energy for e!ectrica! energy is measured as vo!tage. *o!tage5 !i,e a!! measures of potentia! energy5 is a re!ative measure. #o measure any vo!tage you determine its vo!tage re!ative to another !ocation5 often the earth or ground. #o ma,e a vo!tage measurement of the resting potentia!5 a very fine e!ectrode5 ca!!ed a microe!ectrode5 is inserted into the axon and compared ith the reading from an e!ectrode outside the axon. In this case5 e measure the vo!tage of the inside of the neuron compared to the outside of the neuron &y convention. 'uring the resting potentia!5 the inside of the neuron is :B8 mi!!ivo!ts 6m*7 or :B8 thousandths of a vo!t 6%igure AP 1.187. %or a comparison5 a AA &attery is 1.3 *o!ts 6*7.

Figure AP 1.1>. The resting &otential has the inside of the neuron as ?4>m: when "om&ared to the outside. The measuring ele"trode is re&resented !y the !la" lines on the left side. $hemica!!y5 the resting potentia! is principa!!y supported &y three ions5 Sodium5 NaE5 Potassium5 0E5 and $h!oride5 $!:. Sodium and ch!oride are found in common ta&!e sa!t5 Na$!5 he!d together hen so!id &y their opposite charges. Potassium is a minera! e ta,e in in many foods such as &ananas. #he ma4ority of the potassium is found inside the neuron hi!e the ma4ority of the sodium and ch!oride is found outside of the neuron. #o he!p ,eep the neuron in this resting state5 there are more o&en ion channe!s for potassium than for sodium or ch!oride. As a resu!t5 it is easier for potassium ions to cross the

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.>

mem&rane than for sodium and ch!oride. #he ease for an ion to cross the mem&rane is ca!!ed the permea&i!ity of the mem&rane to that ion. #he ce!! mem&rane is said to &e semi:permea&!e ith the greatest permea&i!ity to potassium during the resting potentia!. #hese ion differences and the semi:permea&!e nature of the mem&rane is the ma4or cause for the vo!tage found during the resting potentia!. #here is one more feature of the mem&rane that generates the !ast remaining &it of the :B8 m* resting potentia! is the sodium+potassium pump. #he sodiumApotassium pump i!! attach to three sodium ions from inside the neuron and using energy shifts these sodium ions out of the ce!!. #hen t o potassium ions from outside the ce!! attach to the sodiumApotassium pump and are &rought inside the ce!! 6see Interactive I!!ustration AP 1.17. #he sodiumApotassium pump &oth generates a sma!! part of the resting potentia! and he!ps maintain the ionic concentration im&a!ance for sodium and potassium that are necessary for the functioning of the neuron. Open Interactive I!!ustration AP 1.1. 1hen the i!!ustration is opened5 you i!! see a cross section of a neuron mem&rane going across the screen. Em&edded in the mem&rane is one sodiumApotassium pump mo!ecu!e. #he mo!ecu!es of the mem&rane are dra n in orange and the &!ue. #he sodium ions i!! &e represented &y green s<uares and the potassium ions i!! &e represented &y &!ue circ!es. As sho n on the screen5 the inside of the neuron is &e!o the mem&rane and the outside of the neuron is a&ove the mem&rane. )eca!!5 that in an actua! neuron5 the ma4ority of the sodium ions are outside the neuron and the ma4ority of the potassium ions are inside the neuron. %or i!!ustration purpose5 hen you start this animation5 a!! of the sodium ions i!! &e inside the neuron and a!! of the potassium ions i!! &e outside of the neuron. Dust remem&er5 this is 4ust for this i!!ustration to sho ho the sodiumApotassium pump or,s. Before &eginning the animation5 !oo, carefu!!y at the sodiumApotassium pump. One the !eft side of the mo!ecu!e5 there are three s<uare ho!es in the side. #hese are the &inding sites for the sodium ions. On the right side5 there are t o curved ho!es and these are the &inding sites for the potassium ions. No press the Start &utton to &egin the animation and o&serve the operation of the sodiumApotassium pump. A ho!e co!!ection of sodium ions i!! start moving around inside the neuron and a ho!e co!!ection of potassium ions i!! move around the outside of the neuron. #he movement of the ions are random5 &ut the i!! &ounce off of the mem&rane. At random5 a sodium ion or more5 i!! move in the space of the sodiumApotassium pump. If the ion gets c!ose enough to the &inding site5 the ion i!! &ecome attached to the &inding site. 1hen a!! three sodium &inding sites are fi!!ed the mo!ecu!e is ready to change shape and re!ease the sodium out of the neuron. .o ever5 to change shape5 the mo!ecu!e needs energy and this energy is provided &y the energy storage mo!ecu!e of ce!!s5 adenosine triphosphate or A#P. #his energy mo!ecu!eFs ro!e is sho n &y red ATP appearing at the &ase of the mo!ecu!e. 1ith this energy5 the mo!ecu!e changes shape so that it is no c!osed to the inside of the neuron and open to the outside of the neuron. At the same time it re!eases the sodium ions hich can no move to outside the neuron. It is no possi&!e for the potassium ions to move into the mo!ecu!e. If a potassium ion gets c!ose to the potassium &inding sites5 they i!! &ind. 1hen &oth &inding sites are fi!!ed the mo!ecu!e i!! resume its origina! shape5 A#P is not re<uired for this step. #he potassium ions are re!eased and the cyc!e can &egin again. As you o&serve this animation5 the amount of sodium outside the neuron and potassium inside the neuron i!! increase. If you ish to restart the animation5 4ust press the Start &utton and it i!! restart the animation5 though the p!acement and initia! motions of the ions are a! ays random. A coup!e of additiona! points are orth ma,ing. #he movement of ions are essentia!!y random5 and as a resu!t it can sometimes &e a hi!e for the proper ions are &ound to the neuron. In our &io!ogica! systems5 it might seem a &it ris,y to re!y on such random processes. 1e!!5 there are severa! changes to the animation that can &e made to sho ho such an apparent!y random process can or, <uite re!ia&!y. In an actua!!y neuron there i!! &e more ions5 more sodiumApotassium pumps5 and the ions i!! pro&a&!y move re!ative!y faster5 given the sca!e of the animation. #he contro!s on the !eft5 Speed of Ions5 Number of Sodium Ions5 Number of Potassium Ions5 Number of Na/K Pumps5 i!! a!!o you to a!ter the animation in a ay to speed up the overa!! action of the sodiumApotassium pump and ma,e the operation appear more regu!ar. 1hen you change the num&er of sodium or potassium ions5 you i!! need to press the Start &utton again to see the changed num&er of ions. #he other t o contro!s i!! have their effect as the animation is running. #he other point is to reca!! that during the resting potentia! there are other channe!s5 particu!ar!y for potassium. #he sodiumApotassium pump restores the &a!ance of sodium and potassium and ad4ust the actua! vo!tage of the resting potentia! on!y a sma!! amount. #his pump re<uires energy and uses a !ot of the energy e eat. But ithout it5 the neuron ou!d not or,. *ction )otential" #he resting potentia! is the potentia! energy and the action potentia! is the ,inetic energy of the axon. #he action potentia! can &e descri&e in the same t o ays as the resting potentia!: in terms of the vo!tage changes on the neuron and a!so in terms of the chemica! events in the neuron and on the mem&rane. As &efore5 the discussion i!! start ith the description of the e!ectrica! changes in the neuron. #he action potentia! is &ro,en do n into t o main phases and

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.18

then some mopping up. In the first phase5 the vo!tage changes5 in a&out 1 msec5 from the :B8 m* to E98 m*5 that is5 the inside is no e!ectrica!!y more positive than the outside of the neuron 6%igure AP 1.117. #his phase is mis:termed the depo!ari"ation phase. A true depo!ari"ation ou!d &e to 8 * as 8 * means that there are no e!ectrica! differences &et een the t o sides of the neuron5 that is no po!arity. Sti!!5 the term depo!ari"ation has &y:and:!arge stuc,. Immediate!y fo!!o ing the comp!etion of depo!ari"ation5 the vo!tage moves &ac, to ard the negative vo!tage and actua!!y the neuron &ecomes more negative than the :B8 m* of the resting potentia!. #his phase is named the repo!ari"ation phase. Not as &ad of a name. #he overshoot does not !ast !ong and the vo!tage soon returns to the :B8 of the resting potentia!. Overa!!5 the action potentia! !asts 3:9 msec.

Figure AP 1.11. The a"tion &otential. The swee& of the voltage from ?4> m: to @#> m: is "alled the de&olariAation &hase. The "hange of the voltage !a" from @#> m: to even more negative than the ?4> m: of the resting &otential is "all the re&olariAation &hase. The &eriod during whi"h the voltage is more negative than ?4> m: is sometimes "alled the overshoot. #he action potentia! &ehaves in some rather odd ays. %irst5 action potentia!s are a! ays the same si"e. #he action potentia! goes from :B8 m* to E98 m* every time5 or at !east c!ose enough to not matter. Not on!y that5 &ut the action potentia! stays the same si"e as it trave!s do n the axon. Some axons are <uite !arge. #here are axons that go from the tip of your toe to the medu!!a at the &ase of your &rain. #hese t o characteristics of action potentia!s are termed the all,or, none law. #his !a is curious &ecause in passive e!ectrica! systems vo!tages are easi!y changes5 !i,e the dimmer s itch on your !ights5 and vo!tages shrin, as they trave!5 thus a!ternating current is used. #he ans er to the oddities of the action potentia! !ies in the events that ta,e p!ace on the neuron mem&rane. )eca!! that most of the resting potentia! happens &ecause of the state of the neuron mem&rane and the &a!ance of the concentrations of the different ion 6sodium5 potassium and c!oride7. #he sodiumApotassium pump provides a sma!! part of the resting potentia! vo!tage &ut most of the vo!tage comes from the e<ui!i&rium state of the ions and mem&rane. #his e<ui!i&rium ou!d ho!d the vo!tage a&out a&out :?B m* ithout the sodiumApotassium pump 6Smoc,5 1>>>7. A !arge part of that e<ui!i&rium vo!tage is determined &y the permea&i!ity of the mem&rane to the different ions. )eca!! that it is much easier for potassium to cross the mem&rane during the resting potentia! than the other ions. A!! that changed during the action potentia!. Ions need ion channe!s to cross the mem&rane. #here are more potassium channe!s open during the resting potentia!. In addition to these a! ays open channe!s there are a!so vo!tage:dependent ion channe!s that are norma!!y c!osed. In particu!ar5 there are many sodium and potassium vo!tage:dependent ion channe!s that are c!osed during the resting potentia!. #his situation i!! change during the action potentia!. Open Interactive Illustration *) -"./ The *ction )otential to he!p i!!ustrate the events at the mem&rane during the action potentia!. 1hen you open the i!!ustration you i!! see a mem&rane running across the top much !i,e Interactive I!!ustration 1.15 #he SodiumAPotassium Pump. .o ever5 instead of sodiumApotassium pump in the mem&rane5 there are no a series of vo!tage:dependent sodium 6green7 and potassium 6&!ue7 channe!s. #hey are c!osed at this point so they are dra n as so!id

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.11

&!oc,s. On the right side there is a recording e!ectrode that records the vo!tage of the neuron mem&rane at that point. #o ards the &ottom of the screen is a graph that i!! sho the vo!tage detected &y the e!ectrode. #he resting potentia! is indicated &y a hori"onta! ye!!o !ine. As in the !ast interactive i!!ustration5 sodium ions i!! &e green s<uares and potassium ions i!! &e &!ue circ!es. .o ever5 since e are i!!ustrating the action potentia!5 at the &eginning of the action potentia!5 the sodium ions i!! &e outside the neuron and the potassium ions i!! &e inside the neuron. In this i!!ustration5 the action potentia! i!! move do n the mem&rane from !eft to right. #he vo!tage at the mem&rane ith during the action potentia! is co!or coded ith green to ards positive vo!tages and red to ards negative vo!tages. #he co!or of the action potentia! graph is simi!ar!y co!or coded. 1hen the mem&rane is at the resting potentia!5 the mem&rane is dra n in orange so the ho!e mem&rane is orange at this moment. In this ay you can match up here the action potentia! is on the mem&rane ith the effects on the ion channe!s and the vo!tage &eing p!otted on the action potentia! graph. 1ith this &ac,ground5 the animation can &e p!ayed and understood. Press the Start &utton and atch the animation a first time. 1hen an action potentia! moves to ards the right5 the vo!tage starts moving more positive!y5 sho n &y the mem&rane &ecoming more green5 that triggers these sodium ion channe!s to open hich is sho n &y an opening appearing do n the !engths of the green sodium ion channe!. No 5 much more sodium can cross the mem&rane. #he permea&i!ity to sodium has increased. Sodium is positive!y charged and the inside is negative!y charged. Opposite charges attract. So the e!ectrica! difference i!! dra the sodium in. #here is much more sodium outside the neuron than inside the neuron so the concentration difference i!! a!so !ead to more sodium f!o ing inside the neuron than outside. #hese sodium ions &ring their positive charges inside ma,ing the inside of the neuron more positive exp!aining hy5 during depo!ari"ation5 the inside of the neuron &ecomes more positive. #he sodium ions &eing dra n in &y any one ion channe! are dra n in ye!!o to he!p you distinguish them from the rest of the sodium ions. #he <uestion remains as to hy the vo!tage a ays stops at E98 m*. Dust as the resting potentia! is !arge!y due to an e<ui!i&rium5 the opening of the sodium ion channe!s creates a ne e<ui!i&rium &ut ith a vo!tage of E98 m* inside the neuron. #he neuron a! ays goes to E98 &ecause that is the vo!tages ne resting state. If these ion channe!s a! ays stayed open and nothing e!se changed5 the vo!tage ou!d remain at E98 m*. So for the vo!tage inside the neuron to &ecome negative!y charged again5 severa! changes in the mem&rane must occur. #he vo!tage dependent sodium ion channe!s that opened must c!ose to reduce the permea&i!ity of the mem&rane to sodium &ac, to the !eve! it had during the resting potentia!. #his great!y reduces the trave! of sodium across the mem&rane. In the animation5 hen the vo!tage reaches its pea,5 hen the mem&rane is the &rightest green5 you i!! see the sodium ion channe! c!ose. In addition5 vo!tage dependent potassium channe!s open up so that there is more permea&i!ity to potassium than even during the resting potentia!. In the animation5 at a&out the same time5 the potassium channe! i!! open and it i!! &egin to dra potassium across the mem&rane. 1hen a potassium ion is &eing dra n out of the neuron it i!! &e dra n in cyan. It is this increase in permea&i!ity to potassium that causes the vo!tage to &ecome even more negative than during the resting potentia!. 1hen these extra potassium channe!s c!ose and the sodiumApotassium pump ,ic,s &ac, in the neuron returns to the resting potentia!. Of course in the actua! neuron there are many more ions and ion channe!s and the sodium and potassium channe!s are not attached to each other. #hey are dra n this ay to he!p in the i!!ustation. Pressing the Start &utton i!! restart the animation from the &eginning so that you can see ho 5 as the action potentia! trave!s do n the neuron5 you can atch ho each sodium ion opens as the action potentia! reaches it dra ing in sodium ions and then c!oses short!y thereafter. #hen the potassium channe!s open and then c!ose during the overshoot a!!o ing the neuron to return to the resting potentia!. After the action potentia! goes comp!ete!y off of the right side of the screen the animation stops. Notice ho there is no an increase of sodium inside the neuron and potassium outside the neuron. If the sodiumApotassium pump did not restore the ionic &a!ance to the inside and outside of the neuron you can see ho it ou!d &ecome increasing!y hard and then impossi&!e to generate an action potentia!. #o he!p you exp!ore the functioning of the neuron during the action potentia! there are severa! additiona! contro!s. #he Start &utton a! ays restarts the animation5 restoring the neuron to its &eginning state. #he Stop &utton stops the animation in its current position. #he Play/Pause &utton i!! ha!t and restart the animation from its current p!ace. So you can ha!t the animation to examine hat is going on more c!ose!y and then restart it. 1hen paused5 the Step Back (-) and Step or!ard (") &utton i!! a!!o you to move the animation of the action potentia! a short distance &ac, or for ard so you can exp!ore the opening and c!osing of ion channe!s in s!o motion and under your contro!. Gou can a!so reset the num&er of ions and the speed of the animation ith the s!iders on the !eft side of the screen. #he resu!ting picture is that the action potentia! is a ave of opening sodium ion channe!s that <uic,!y c!ose as potassium ion channe!s open. #his picture of the action potentia! indicates a !imit to ho have a neuron can generate action potentia!s. 1hi!e one action potentia! is &eing generated5 it is impossi&!e or harder for the neuron to generate another action potentia!. #his period of time hen another action potentia! is harder or impossi&!e to create is ca!!ed the refractory period. #here are t o types of refractory periods: a solute refractory period5 relative refractory period. #he a&so!ute refractory

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.13

period5 rough!y a!igned ith the period of time that the sodium ion channe!s are open5 !asts a&out 1 mi!!isecond. 'uring this time5 it is impossi&!e that the neuron cannot generate another action potentia!. 'oing a simp!e ca!cu!ation5 it is impossi&!e for a neuron to fire faster than 1888 times a second5 though in rea!ity5 the fastest neurons are o&served to fire is much !o er than that. #he re!ative refractory period pic,s up after the a&so!ute refractory period and during the re!ative refractory period it is !ess !i,e!y for a neuron to generate another action potentia!. #his !asts 3:9 mi!!iseconds. #his action potentia! is not an !i,e the ay that e!ectricity is conduction in ires at a!! and as a resu!t is much s!o er. In an unmye!inated neuron5 the action potentia! trave!s a&out 98 mAsec. #he ro!e of mye!in is to speed up neura! conduction a!ong the the axon &y reducing the resistance and having the neuron trave! under the neuron passive!y !i,e e!ectricity in the a!! a!!o ing it to trave! much faster. Passive e!ectricity5 reca!!5 !oses its vo!tage as it trave!s5 so the action potentia! is rene ed at the gap &et een the mye!in ca!! Nodes of )anvier. The Synapse: 0a'ing the Brain Fle&i le and #earn #he action potentia! is fixed in invaria&!e. 1hat changes a&out an action potentia! on a neuron is ho many occur in a given period of time &ut its characteristics5 once fired never changes. Sti!! the &rain must &e more f!exi&!e. It must !earn5 is must represent a!! the comp!ex aspects of our perceptions5 ideas5 emotions5 and experience. 1hi!e the action potentia! pro&a&!y contri&utes to this experience5 there needs to &e some feature of the &ehavior of the &rain that sho s more comp!exity than the action potentia!. #he ,ey seems to !ie in the synapse. #he synapse the p!ace here one neuron comes in contact ith another neuron5 or sometimes a musc!e or a g!and. Since the present discussion focusses on the &rain5 the synapse &et een t o neurons i!! &e discussed5 though the synapses &et een neurons and musc!es and g!ands are simi!ar. %irst5 the structure and then the activity of the synapse i!! &e discussed. Structure %igure AP 1.13 sho s a diagram of the synapse. %irst5 notice that the t o neurons do not tough. #hat is the most common arrangement. #here is a sma!! space &et een the t o ce!!s in the synapse. #his gap is ca!!ed the synaptic cleft. #his gap is extreme!y sma!!5 on the order of 1 micron 6or 1 mi!!ionth of a meter7. #he synaptic c!eft is a convenient !andmar, to use in descri&ing the synapse. Since the synapse is here t o neurons connect5 e need some !anguage to he!p ,eep straight hich of the t o neurons is &eing ta!,ed a&out. %ortunate!y5 there is a directiona! re!ationship in the synapse. #he action potentia! trave!s do n the axon to the termina!. #hen at the synapse this neuron contacts the next neuron in the chain of neura! events. So e can ta!, a&out the neuron &efore the synapse5 the presynaptic neuron5 and the neuron after the synapse5 the postsynaptic neuron.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.19

Figure AP 1.12. The stru"ture of a syna&se. 5% need a sim&ler figure. (ainly show vesi"les+ release+ and !inding. Before discussing items of the synapse. #o understand the reason for a!! the e!ements in the synapse5 notice the c!eft. #he action potentia! has an e!ectrica! component to its actions. #he c!eft !oo,s !i,e a &rea, in the !ine. #he action potentia! cannot 4ump even this sma!! &rea,. As a resu!t5 the communication &et een the t o neurons cannot &e e!ectrica!. In this case5 it i!! &e chemica!. In the presynaptic neuron5 on the top in %igure AP 1.135 there are mo!ecu!es that that serve the function of the communication &et een the t o neurons. #hese mo!ecu!es are ca!!ed neurotransmitters. #hese mo!ecu!es are stored in mem&rane sac,s ca!!ed synaptic vesicles. In addition to sodium and potassium vo!tage dependent ions5 the presynaptic termina! has vo!tage dependent ca!cium 6$aEE7 channe!s. #he a&i!ity of the termina! to a!!o ca!cium to f!o into it during an action potentia! i!! turn out to &e very important in the functioning of the synapse. On the postsynaptic neuron5 there are receptor mo!ecu!es edged in the mem&rane. #he postsynaptic neuron surface a!so is em&edded ith !arge num&er of c!osed ion channe!s that i!! &e opened &y the actions of the synapse. Function #here are many steps in the operation of the synapse. #hese steps can &e grouped into: preparation5 re!ease5 &inding5 remova!. Each of these steps i!! &e discussed in turn. )reparation" #he synapse has to &e prepared to function 4ust as the axon does. In the synapse5 the preparation re<uires the synthesis and storage of the neurotransmitters. #here are many different types of neurotransmitters that have &een discovered and depending upon the type of neurotransmitter they process of synthesis is different. Some neurotransmitters are specia!i"ed mo!ecu!es5 !i,e acety!cho!ine5 others are modified amino acids !i,e dopamine and serotonin5 others are short strings of amino acids5 the neuropeptides. /sua!!y the synthesis ta,es p!ace in the soma so the neurotransmitter must &e transmitted to the termina!. Before &eing re!eased the neurotransmitters must &e stored into the synaptic vesic!es. (elease" #he process of re!easing the neurotransmitters into the synaptic c!eft &egins as an action potentia! arrives at the termina!. HNEE' AN ANI+A#IONI #he action potentia! has a &it of a different function at the termina!. In addition to a!!o ing sodium ions to enter the axon during depo!ari"ation5 the termina! a!so has ca!cium channe!s 6$aEE7. So ca!cium enters the synapse during action potentia!s at the termina!. #his entrance of ca!cium is crucia! to the operation of the synapse. #o re!ease the neurotransmitters into the synapse5 the synaptic vesic!es must fuse ith the mem&rane of the termina!. #hat

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1;

is the mem&rane of the synaptic vesic!e must re4oin the mem&rane that is the a!! of the ce!!. #here is a &it of a pro&!em here. As mentioned a&ove hen descri&ing the mem&rane5 the outside of mem&ranes are s!ight!y negative!y charged. Both the synaptic vesic!e and the ce!! mem&rane are of the same type of mem&rane ma,ing them &oth s!ight!y negative!y charged and !i,e charges repe!. #he pro&!em is ho to get the t o mem&ranes c!ose enough that the can 4oin each other. It is to overcome the mutua! repu!sion of the mem&rane that the $aEE comes in5 inc!uding its dou&!e positive charge. #he ca!cium attaches to &inding sites on the mem&rane and the overa!! positive charge &ecause of the ca!cium ion dra s the vesic!es to the mem&rane and a!!o them to fuse5 &ecome one mem&rane again. 1hen the synaptic vesic!e mem&rane fuses ith the mem&rane of the ce!!5 the synaptic vesic!e mem&rane sp!its emptying its contents into the synaptic c!eft. #he neurotransmitters have &een re!eased 6see %igure AP 1.x for no ant an animation !ater7.

Figure AP 1.x. *iagram of the release of neurotransmitters.

Binding" Once in the synaptic c!eft5 the neurotransmitters move random!y through the c!eft. #housands of mo!ecu!es are re!eased from each synaptic vesic!e so many i!! &ump up against the mem&rane of the post synaptic neuron. It is a!so <uite !i,e!y that they i!! &ump into the receptor mo!ecu!es that are on the mem&rane. If there is a shape match &et een the three dimensiona! shape of the neurotransmitter and the &inding site on the receptor mo!ecu!e5 then the neurotransmitter i!! &ind to the receptor mo!ecu!e. But if &inding did nothing e!se5 there ou!d &e not point is riting a&out it. #he &inding of neurotransmitter to receptor mo!ecu!e !eads to events5 direct or meta&o!ic5 that cause ion channe!s to open in the postsynaptic neuron mem&rane. #he vo!tage inside the neuron i!! change. #his change in vo!tage is ca!!ed postsynaptic potentia! 6PSP7. #here are t o &asic types of postsynaptic potentia!s5 those that ma,e an action potentia! more !i,e!y or !ess !i,e!y on the postsynaptic neuron. #he postsynaptic potentia!s that ma,e an action potentia! more !i,e!y are ca!!ed e&citatory postsynaptic potentials 61)S)s7 and during EPSPs the vo!tage &ecomes !ess negative 6%igure AP 1.x7. #he postsynaptic potentia!s that ma,e a neuron !ess !i,e!y to fire are ca!!ed inhi itory postsynaptic potentials 6I)S)s7 and ma,e the vo!tage more negative 6%igurer AP 1.x7.

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1@

Figure 1.x. :oltage in a &ostsyna&ti" showing !oth an 2PSP and %PSP.

(emoval" As !ong as the neurotransmitter remains &ound to the receptor mo!ecu!e5 the ion channe!s i!! remain open and the postsynaptic potentia! i!! continue. So it is important that neurotransmitters do not stay &ound to !ong. /sua!!y the neurotransmitters re!ease after a short time. Sti!!5 as !ong as the neurotransmitter is f!oating a&out in the synaptic c!eft it cou!d &ind again and start a ne postsynaptic potentia!. #here needs to &e a mechanism to c!ear the synaptic c!eft of neurotransmitter so that the c!eft is c!ear for the next action potentia!. #he neurotransmitters are removed through t o processes: destruction and reupta'e. 'estruction occurs &ecause of en"ymes that are f!oating in the synaptic c!eft that &ind and &rea, apart the neurotransmitter mo!ecu!es. As a resu!t they no !onger have the proper shape and cannot &ind. #his method of remova! is not common for most neurotransmitters. )eupta,e occurs hen the neurotransmitters actua!!y are rea&sor&ed into the presynaptic neuron here they can &e either destroyed or used again. +ost neurotransmitters use this method of remova!. Completing the Circle: 2enerating a New *ction )otential 1ith synaptic transmission5 there remains one more step in the se<uence that e have &een considering to understand: ho does the action potentia! get started. #he discussion started ith the action potentia! a!ready going. #hen the function of the synapse as discussed. No it is time to fi!! in the missing piece of the pu""!e. Open Interactive Illustration *) -"&: Spatial and Temporal Summation hich e i!! use to he!p &ring this concept to !ife. 1hen you open the i!!ustration you i!! see in the main part of the screen i!! sho four neurons dra n schematica!!y ith circ!es for the somas and !ines for the actions and ang!es for the termina!s. #o simp!ify this discussion5 dendrites are not dra n and the synapses connect right on the soma. #hree of the neurons are input or presynaptic neurons5 and the neuron of interest is on the right and it is the post synaptic neuron5 dra n in gray at the start. #he three presynaptic neurons are !a&e!!ed A5 B 5 and # to he!p us ,eep them straight. # o of the presynaptic neurons are dra n in green 6A5 #7. #hese neurons generate EPSPs and are excitatory to the post synaptic neuron. #he other presynaptic neuron 6 B7 is dra n in red and it generates IPSPs and is inhi&itory to the postsynaptic neuron. #he graph a&ove the a&ove the post synaptic neuron records the vo!tage at the &eginning of the axon on the postsynaptic neuron during animation5 inc!uding any action potentia!s. #he resting potentia! is indicated &y the ye!!o !ine on the graph. #he green !ine on the graph represents a ne concept that is important to understand ho action potentia!s are generated. #he &eginning of the axon is ca!!ed the axon hi!!oc, and it is here that ne action potentia!s are generated. #o generate a ne action potentia! the vo!tage at the axon hi!!oc, must reach the vo!tage represented &y the green !ine hich is ca!!ed the thresho!d vo!tage. If the vo!tage reached the thresho!d vo!tage5 the action potentia! i!! fire5 if the vo!tage does not reach the thresho!d5 then the action potentia! i!! not fire. At this point5 !et us revie postsynaptic potentia!s. On the !eft side of the screen are contro!s for the si"e5 duration and starting time for the postsynaptic potentia!s to &e generated &y each of the presynaptic neurons. Each set of contro!s are !a&e!ed &y the neuron !a&e! 6A5 B or C7. On each neuron it is possi&!e to have t o action potentia!s trave! do n it and to

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1?

generate t o postsynaptic potentia!s each 6!a&e!ed irst and Second &eneath each neuron !a&e!7. On Neuron A5 ad4ust the Si$e % s!ider ti!! it e<ua!s &5 hich is indicated &e!o the s!ider. #hen press the Animate &utton. An action potentia! i!! trave! do n the A neuron and the graph i!! sho the current vo!tage on the postsynaptic neuron dra n as a cyan 6!ight &!ueAgreen7 !ine. 1hen the EPSP from neuron A reaches the axon hi!!oc, of the postsynaptic neuron5 you i!! see the vo!tage move up in a more positive direction5 &ut it does not reach the thresho!d5 so no action potentia! occurs. )eturn the Si$e % s!ider for the A neuron to 8 and ad4ust the Si$e % s!ider for the B neuron to ; and again press the Animate &utton. Neuron B generates IPSPs so hen the postsynaptic potentia! reaches the axon hi!!oc, the vo!tage &ecomes more negative. Summation It turns out that fe if any EPSPs are !arge enough to reach the postsynaptic neurons thresho!d5 so to cause an action potentia!. It ta,es t o and often more EPSP. #his is the process of summation. #here are usua!!y thousands of synapses on any one neuron. It is very possi&!e for the postsynaptic potentia!s to over!ap at the axon hi!!oc,. #he resu!ting change in vo!tage is5 !i,e the term imp!ies5 the com&ination5 !i,e addition5 of the postsynaptic potentia!s present. # o types of summation are ta!,ed a&out5 a!though in most cases5 the summation occurring is a com&ination of the t o types. Spatial Summation refers to the arriva! of postsynaptic potentia!s from different presynaptic neurons. In the interactivity5 ad4ust the Si$e % s!ider on B &ac, to 8 and put the Si$e % s!iders for &oth A and # to ? and press the Animate &utton. No hen the postsynaptic potentia!s from &oth A and # reach the axon hi!!oc,5 they are &ig enough to reach the thresho!d and an action potentia! occurs. It is a!so no possi&!e to understand hy IPSPs are tru!y inhi&itory. (eaving the rest of the settings the same5 ad4ust the Si$e % s!ider on B to 9 and press Animate. #he t o postsynaptic potentia!s from A and C used to &e &ig enough to trigger an action potentia!. No 5 ho ever5 ith the addition of the negative IPSP from B5 the summed vo!tage at the axon hi!!oc, no !onger reaches the thresho!d and no action potentia! occurs. It has &een inhi&ited. Temporal Summation occurs hen different postsynaptic potentia!s from the same presyaptic neuron arrive and add together. #empora! summation occurs &ecause it is <uite possi&!e from postsynaptic potentia!s to !ast much !onger than action potentia!s. #o i!!ustrate this concept return a!! of the Si$e % s!iders to 8 except for A. (eave that intensity at ?. No use the 'ur % s!ider on A and !engthen the first postsynaptic potentia! to 8.@5 indicated &e!o the s!ider. Ad4ust the Si$e ( for A s!ider to ? and press Animate. In this case5 t o action potentia!s i!! move do n the A neuron. 1hen the first arrives5 the postsynaptic potentia! is visi&!e at the axon hi!!oc, &ut not !arge enough to cause an action potentia!. 1hen the second action potentia! arrives5 the second postsynaptic potentia! sho s up that the axon hi!!oc, and then the vo!tage reaches the thresho!d and the postsynaptic neuron has an action potentia!. Dust a side note. In $hapter ; the terms spatia! and tempora! summation ere used to refer to the a&i!ity to reso!ve &oth detai!s and events at different times. It can &e confusing to use the same terms for different meanings &ut5 in this case5 the use of the terms in $hapter ; depend upon the use of the terms e have 4ust discussed. Spatia! summation in vision is an examp!e of the operation of spatia! summation in neurons and the same is true for tempora! summation in vision. The Corte& $overing the surface of the cere&rum is the cortex. 'epending are the part of cere&rum there are different types of cortex5 &ut most of it is Neocortex hich is the most recent!y evo!ved cortex. A!! of the sensory cortices that are discussed in the text are a!! made up of neocortex. #he neocortex is an a&out 1A;J thic, !ayer that covers a!! of the primary !o&es. #he indentations in the &rain are a!! designed to increase the surface area of the cortex. #he cortex fo!ds in and out of these indentations on the surface of the &rain. In the human &rain5 the ma4ority of the synapses of the fore&rain occur in the cortex. #he cortex is so comp!icated it ou!d &e impossi&!e to study ithout important regu!arities that a!!o use to see the organi"ation of the cortex and a!!o us a means for understanding the functioning of much of the cortex. #hree &asic organi"ationa! princip!es i!! &e discussed here. #he first is that cortex is organi"ed topographica!!y. #hat is the cortex surface can &e mapped. #hin, of a map of the earth. Ad4acent points on the or!d are a!so next to each other on the map. $anada is north of the /nited States on the Earth5 and since north is usua!!y at the top of maps5 $anada is 4ust a&ove the /nited States on a map of the or!d. #hat does not mean that maps are perfect. #he Earth is rough!y a sphere &ut maps are f!at. Inevita&!y there are distortions. In a norma! +ercator pro4ection of the earth5 areas near the t o po!es are great!y over si"ed on the map. #he north and south po!es are 4ust points on the Earth &ut turn into ide areas the same si"e as the e<uator on a map. #hese characteristics of maps are seen in the topographic organi"ation of the &rain. In the sensory and motor areas this mapping is c!earest. #he retina is mapped on the visua! cortex. Dust as images are upside do n and &ac, ards on the retina so they are on our cortex. Dust as maps have distortions so does this mapping of the retina on the cortex. #he fovea ta,es up !ess than 1K of the retina &ut a&out @8K of the cortex. Simi!ar types of mapping can &e found

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1B

for the other sensory and motor areas. #he existence of these maps are immense!y usefu! in studying the &rain. #he next princip!e is that the neocortex is organi"ed hori"onta!!y into six !ayers 6%igure AP 1.x7. Each !ayer has varies in the type of ce!!s that are found and performs a different function. (ayer 1 is at the surface of the &rain and (ayer ? is on the inside of the cortex next to the rest of the &rain. #hese !ayers are the same a!! over the neocortex and have the same function throughout. 1hat varies is the re!ative thic,nesses of the !ayers that a!!o s anatomist to identify different regions of the cortex. 1hi!e each !ayer has a different function5 the !ayer ; is the one most re!evant to the topic of this text. (ayer ; receives input u!timate!y from outside the &rain. %or examp!e5 !ayer ; of *1 is unusua!!y thin, as it received the input from the !atera! genicu!ate nuc!eus. It is the thic,ness of this !ayer5 hich is thic,er in *1 than any other region of the &rain that !eads to one of the names for this !ater5 the striate5 or stripped5 cortex. #he fina! organi"ing princip!e is that of co!umns. #he co!umns run at right ang!es to the !ayers. #he co!umns run from !ayer

Figure AP 1.x. A diagram of the layers of the "ortex.

1 to !ayer ?. #he co!umns ere first discovered &y +ountcast!e 61>@B7. +outcast!e 61>@B7 as studying the somatosensory cortex 6see $hapter 137. .e found that different areas of the cortex each responded to the different s,in senses5 for examp!e one region ou!d respond to pressure hi!e the next might respond to temperature. .u&e! and 1iese! 61>?37 refined the concept of co!umns in the visua! cortex finding that different regions of the surface respond to different orientations of the visua! stimu!us and different eyes. #he co!umns turned out to &e carefu!!y organi"ed. Sensory Coding or 3ow does the Brain a le to #et 4s 5now 6hat is 2oing on Out There7 #o conc!ude this &rief introduction to the nervous system5 it seems important to tac,!e5 &rief!y5 one very important topic. In the discussion of the functioning of the neuron5 the a!!:or:none !a as discussed. An action potentia! fires or it does not. Each action potentia! is essentia!!y the same as every other action potentia!. #his o&servation &egs the <uestion as to ho can such a monotonous5 &oring even5 response ena&!e us to experience the rich and varied range of sensory5 emotiona! and cognitive experiences that ma,e up our !ife. #his is the pro&!em of sensory coding. A secret code ta,es our norma! !anguage and trans!ates it into another pattern that u!timate!y &ehaves exact!y as the !anguage does. #he code represents the !anguage comp!ete!y in a ne form. #he search for sensory codes is trying to find out ho the &rain trans!ates the sensory

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1C

input it receives into a pattern of activity of the neurons of the &rain that in some fashion matches the &ehavior of the sensory input and ho it changes. A great dea! a&out sensory coding remains to &e discovered &ut some ays that sensory coding is accomp!ished has &e descri&ed. Some features of coding depends upon the different operations of a neuron hi!e others use the fact that the &rain is made up of many neurons to accomp!ish the coding. Actua! coding uses &oth the differences in &ehavior ithin and &et een neurons &ut for simp!icity each type of coding i!! &e discussed separate!y. Open Interactive Illustration *) -"&/ Sensory Coding to he!p i!!ustrate these different coding methods. Coding y a single cell Some information a&out the or!d experience is contained in the ce!! itse!f. In fact5 one of the the o!dest concepts in sensory coding comes from Dohannes +L!!er in the ear!y part of the 1>th century ho postu!ated that our experience depends upon hich neuron fires. #his princip!e is ,no n as the #aw of Specific Nerve 1nergies. Basica!!y5 +L!!er as trying to understand hy different regions of the &rain yie!ded such different experience hi!e the &rain anatomy !oo,ed &asica!!y the same in the different regions. .e argued that the difference as simp!y in hich neurons ere activated. #hose neurons in the visua! cortex ou!d !ead to vision and those in the auditory cortex !ead to hearing 6)E%7. Synesthesia supports this concept hich is experienced &y some peop!e. #here are those that report seeing sounds and it is this cross sensory experience that is synesthesia. In this case5 auditory stimu!ation of the visua! cortex !eads to a visua! experience of sound 6)E%7. 'reams a!so support this concept. 1hen you have a visua! experience of dreams5 your visua! cortex is active 6)E%7. (oo,ing at Interactive Illustration *) -"&/ Sensory Coding5 use the drop do n menu on the !eft to se!ect )a! of Specific Ner*e +ner,ies hich shou!d &e se!ected as it is the defau!t option. Se!ect one of the neurons 6A or B7 &y chec,ing the ire t-is neuron next to the desired neuron. #he press the Animate &utton. 1hichever neuron you have se!ected i!! fire. According to the (a of Specific Nerve Energies5 the fact that this neuron as firing is hat determines your experience. If the other neuron fired5 you ou!d get a different experience. Another means &y hich a sing!e neuron can code for more than one experience is &y its firing rate. #his is ,no n as Fre8uency Coding. %re<uency coding is found in the auditory system as one of the ays that the &rain seems to determine the fre<uency of the sound. /sing the menu on the !eft se!ect re.uency #odin,. #he arrangement of neurons changes on the screen. #here are no three neurons on the !eft 6A5 B 5 and #7 that a!! input into one neuron. #here is no a menu a&ove the somas of these three neurons that a!!o s you to Select Neuron to ire. Se!ect first A and then B and then fina!!y # in turn5 pressing the Animate &utton after se!ecting each neuron and o&serve the resu!ts. As you can see5 the num&er of action potentia!s generated differs depending upon the input neuron 6!oo, to the activity to find out the specifics7. So hat is &eing input can &e determined &y the &rain &y the firing rate. In %re<uency #heory5 the pitch of a tone in some cases seem to &e determined &y the firing rate of neurons 6see $hapter 187. Coding across many cells 1hi!e individua! neurons do seem to provide some information5 much more a&out the or!d can &e !earned hen a co!!ection neurons are used to a!!o the coding to distri&ute itse!f. #his type of coding is ,no n as distri&uted representation 6+c$!e!!and = )ume!hart5 1>C?7. (et us ta,e a simp!e examp!es of one type of distri&uted coding5 re!ative firing rate. #he cones and their coding of co!or is a type of examp!e of this type of coding. Even though they do not generate action potentia!s5 their responses fit this type of coding. Each cone c!ass has a different fre<uency that it prefers to respond to. So for each fre<uency of !ight5 the strength of responses across the three different cones differ 6see $hapter ? and Interactive I!!ustration ?.x5 #richromatic #heory7. In this ay5 three cones can he!p us distinguish thousands of different co!ors. #o i!!ustrate this means of coding5 use the menu on the !eft of Interactive Illustration *) -"&/ Sensory Coding to se!ect /elati*e irin, /ate. #here are no six input neurons 6A through 7 and three output neurons. Each input neuron is connected to each output neuron. /sing the Select Neuron to ire menu5 se!ect each input neuron in turn and then press the Animate &utton. Input neuron A i!! on!y cause output neuron % to fire. #his outcome is possi&!e &ecause neuron A has a stronger connection to output neuron % than to the other t o output neurons. It generates !arger and perhaps a !onger EPSP on output neuron % than on output neurons ( and 0. Sti!! at this point5 the outcome seems a &it !i,e the (a of Specific Nerve Energies here hich neuron fires is important. But if you !oo, at the figure c!ose!y you see that there are ? input neurons and on!y three output neurons. If the (a of Specific Nerve Energies operated5 there are too fe output neurons. No se!ect input neuron B and press the *nimate &utton. Output neuron - sti!! fires &ut it no fires three times and output neuron . fires &ut it on!y fires once and sti!! output neuron 9 does not fire. Again5 this difference in response of the three output neurons is possi&!e &ecause they each have different si"e and duration EPSPs to the one input neuron 6B7. #he

Experiencing Sensation and Perception Appendix: Basics of the Nervous System

Page Appendix 1.1>

strength of the connection &et een the input and the output neurons for * and B !eading to a different response of the output neurons. It is not as c!ear as for the (a of Specific Nerve Energies. It fact it is sort of com&ination of the (a of Specific Nerve Energies and the %re<uency $oding that ere discussed a&ove. In fact5 as you try each different input neuron you i!! find that each input neuron generates a different pattern of response on the output neurons. In addition to cones5 the response to the &asic tastse seem to use this type of coding. (et us discuss one more type of sensory coding that seems to &e used in sensory system. So far5 a!! of the codes have on!y ta,e advantage of the a&i!ity of neurons to generate excitatory connections5 EPSPs. It seems un!i,e!y that if neurons can have inhi&itory connections5 IPSPs5 that these connections do not p!ay a ro!e in ho e code our sensory systems. In fact5 throughout the &oo, severa! ays that inhi&itory connections p!ay a ro!e are discussed. In $hapter ?5 the co!or opponent theory as discussed. %or examp!e the !ong and midd!e ave!ength cones 6red and green7 input into the )edA-reen co!or opponent ce!!. If the !ong ave!ength 6red7 cone input excites the )edA-reen ce!!s then the midd!e ave!ength 6green7 cone inhi&its the same ce!!. If &oth cones respond e<ua!!y the )edA-reen ce!! does not respond at a!!. /se the menu on the !eft of the i!!ustration to se!ect Opponent Processes to see and examp!e of this type coding in action. In this examp!e there are t o input ce!!s and t o output ce!!s. #he input ce!!s connect to &oth output ce!!s &ut different!y than the !ast examp!e. Input ce!! A has an excitatory 6green7 connection to the output ce!! % and an inhi&itory connection to output ce!! (. Input ce!! B connects in the opposite fashion to the t o output ce!!s. /sing the Select Neuron to ire menu you can cause neuron A to fire5 neuron B to fire5 and Bot- input neurons to fire. Se!ect each option and press the Animate &utton and see hat happens. 1hen neuron A fires a!one so does output neuron % and hen neuron B fires a!one output neuron ( fires. But hen &oth fire5 there is no response &y the output neurons. #hey cance! each other out. #his type of coding !oo,s for differences. #he response to each neuron a!one is emphasi"ed. #he response of neither or &oth seem to &e ignored &y this type of coding. #hey !ead to the same output: no firing &y the output neurons. %or examp!e5 in out very o n co!or responses5 a hite !ight cance!s a!! co!or responses in our visua! system. 1hite is considered achromatic or non:co!ored. Summary #his chapter covered the &asics of the nervous system. %irst5 the &asic anatomy of the neurvous system as descri&e indicating the &asic distinctions in the anatomy &et een the centra! and periphera! nervous system and the &asic divisions of the &rain into fore&rain5 mid&rain and hind &rain. Next5 the anatomy and function of the neuron as covered. #he neuron is the most important functiona! neuron of the &rain. Neuron has many important parts inc!uding the dendrites and somas that &oth receive input from other neurons 6somas a!so have the genetic materia! of the ce!!75 axons hich send signa!s and termina!s hich communicate ith the next neuron in the se<uence. On actions5 neurons communicate via action potentia!s. Action potentia!s are e!ectrica! signa!s carried &y chemica! means a!ong the mem&rane of the axon. Action potentia!s fo!!o the A!!:or:none (a M they are a! ays the same si"e and do not shrin, as they trave! do n the axon. $ommunication &et een neurons occurs at the synapse. Synaptic communication is a chemica! process. #he action potentia!Fs arriva! at the termina! triggers a se<uence of events that re!eases neurotransmitters into the synaptic c!eft. #he neurotransmitters &ind to receptor mo!ecu!es on the postsynaptic neurons an change the vo!tage in the postsynaptic neuron. Some synaptic connections are excitatory hich ma,e the postsynaptic neuron more !i,e!y to fire and others are inhi&itory5 hich ma,e the postsynaptic neuron !ess !i,e!y to fire. Summation of these postsynaptic potentia! at the axon hi!!oc, is hat determines if a ne action potentia! is !i,e!y to fire. #o fire the summed postsynaptic potentia!s must reach the thresho!d vo!tage for that neuron. Next the cortex as discussed. #he cortex is the thin !ayer of ce!!s that over!ays the surface of most of the fore&rain. #hree organi"ation princip!es of the cortex ere discussed. %irst5 the cortex is topographica!!y organi"ed. One can map a sensory experience onto the cortica! surface. Second5 the cortex is made up of six !ayers. %or the purposes of this text5 !ayer ; is of particu!ar interest as that is here sensory information i!! synapse. #hird5 geing from the surface of the &rain through the !ayers of the cortex are co!umns. A!! ce!!s in a co!umn respond in some simi!ar fashion5 for examp!e a!! might respond to pain. %ina!!y5 ays that neurons can he!p represent our experience of the or!d ere discussed. #his is sensory coding. Some coding can &e done &y a sing!e neuron &ut more common!y many neurons are invo!ved. #he types of coding discussed ere (a of Specific Nerve Energies5 %re<uency $oding5 re!ative firing rate5 and Opponent Processes. 5ey Terms