Eric R. Kandel Brain and Behavior Two Alternative Views Have fiver Advaneed on ihe Relationship Between Tain and Behiinr Regions uf che Brain Are Specialized for Dillerent Functions Language and Other Cognitive Functions Arc Localized Within the Cereheal Cortex Alfective and Character Traits Are Also Anatumically Localized Mental Processes Ace Represented in the Hain hy Theie Elementary Operations AT ie gents ome amar eal ene hat al Inavior + reflection of brain tustetian, Accord, Unis tev a sew chat sve shall uy to document in this teat what we coniionty call mand fsa range of Lametions Carrie ut by ce brain, The action of the braun unslerbies phat only eelatiscly simple motor bebaviors such ay walle i, hrelung, ai siitusg, bat also elaborate allective huni cog tive behaviors such ay feeling, learn, thik To auul eomnpesitug 8 symphony. As a-corolhary, the dbs iilers of aleet dvelings) and cognition fehonght) that ee nesantie and psyehore illness ean be seen (isturbances ot ayn funetian, Tine braun is snate ap of advil units nerve cells se nevus) att hal eels, ‘Phe task of neural science 1st plant hoy tl. brary marshalls dese units tw contra behavior and fis, 1 ean, the functinning af the eonsnt hhent cells in to midividal’s beain 38 inilueneed hy that persun’s enviroment, including the behavior of other peuple th this siuapeer and the next we provide an oversth ew ol dhs task. this chapter we examne the sUItERIeS tise y the hunt beau to represent fangseage, the mont laborae cogineive behavior. We shall lacus on dhe ec bral the prt ofthe brain that has expanded must 19 higher appeets ot hunny behaviors, We illustrate hows lige groups of netirons are oxganized within the neeveats System i how even highly comptex ehaviens ean hs Tovalicelte sperdtic regions af the bean tthe next hsp rer ase shall vonsier nervous systenn Function at he ce Tatar fevel, sme simipte reflex behavior to exami hos Sensory sigials ne transformed into mote Jets.6 Mart An Overall View Two Alternative Views Have Been Advanced on the Relationship Hetwween Brain and Behavior Current views of nerve cells the fram, and behavior bave ‘emerged over the last century from the enaleseenee ot ive ‘experimental traditions: anatomy, embryology, physi ‘ogy, pltemacalogy, and psychology. The anatomical complexity of nervous tisstle way nat appreciated before the invention of the can Scope. Until the eighteenth century nervous eisse was shoughe to be glandular function, an ale that was based ‘on Galen's proposal that nerves are ducts conveying fhund svercted by the brain and spinal cord tu the periphery of the burly, Toward the end of che minetceath century, she histology of the nervous system became a more precise science, culminating in the investigations «f Camallo (Golgy and Santiago Ramon y Caial, Gubgi developed a sth ver imptegnation method that allowed mier alization of the anatomy of the whole neuron, ineluding the cell bay and its cwo major processes, the dendrites and the axon, Ramin y Cajal used his stating techni to label idividval cells, thus showing chat the nervous system 18 not a syneytium (a continous mass of fused cells sharing a commun cytoplasm) but an intricate net work of discrete eells, In the course of this work Ramon y Hal developed some of the key coneeptelal ansights and smiel of the early empirical support for the neuron aloe ne—the principle that the nervous systen 1s sale Up ol innhvidual signaling elements, the ncurons, which contact fone anther only at specialized pomnts of anteracton, called synapses Funal experimental support for the neuron deteine was provided by embryology, the seed discipline, ‘The cm lyologist Ross Harrisn devised tissue etlture methods that showed directly that the mam processes nf the nerve cel, the dendrites andl the axon, are contnnious with the cell boy and extend fro it. Harison tour turtler, 3s Ramon y Cajal had suggested, thar che cy a the axon x aon tits targets 'Newophystology, the titel sesentalic ase apfine tan reste to the growth cone, swhiel teas the advancing, mental t9 the movera analysis of neta tunetion, began at the late eighteenth century when Lugs Galvant discov ceved that maisele cells produce clectneicy. During. the rineteenth century the foundations at electiophysiology ‘were laud by Emil Dufbois-Reymond, johauiney Maller, and Hermann von Helmbolt2, who discovered that the clee trical activity of nerve cells provides 4 reas of carrying the signal that conveys mnformation frony one ent of a ell to the ather amd from vine nerve cell tu the next The ampact of the fourth discipline, pharmaco started at the end of the ninetecnth century shen Clam emmanl, Paul Elulich, and Jol Langley sdemonstraced that drugs interact with specitie recepears on cells. This dliscovery later hecame the basis of the modern stucly of ‘lemeal synaptic transmission Psychology, the fifth discipline important for under standing the brain, has the longest history. Western ideas vbowt nian! were tiast lormatlated by die classical Geek, phos tit mn the sete ee. the mis fa ers anal eccaved further det FReaie Dew antes, Paavo une, aul fli 1 Serentitic sty of behavior as the observable tanlivadial did ot box, hosvever, neil Chatkes Darwin's investigations on evolution in the second halt teonth century had paved the way for behav tw be stu ted scientitically, giving rise to expeeimental psyehology, the stusly of behavier in the laboratory, and to ethology, the study of behavior in nature he nine The mieraungof anatomy, developmental biology, phys tology, and the study of behavior hegan an a preliminary way in the nineteenth century with the phrenologists hed by the Austrian pliysie ia and neucoanatomnst Fran fo seph Gall. Gall appreciated that the functions of the mind have a biologieal hasts and, specifically, chat they are car ried out by the brain, He postulated that the brain ts not a tontary oman but a collection of at least AS domains or enters (others were wslded Tater), cach corresponding (0 a specific mental function, Gall chought that even the mast labors fe ind abstract” mental fanetinns— generosity, riother lave, sand seutetiveness.-vecur i discrete ateas of the cerebral eartex. He further believed that the center for ceaeh mental tanctinn could develup and increase in trestle ot use, much as the size ofa musele ss increased by. exererse. As each center grew, way thought to cause the overlying surfice «# progrude, Therelore the location of cramal banaps would mudicave wheel regions of the brain ine mist developed (lagune LH. By entrelatsng the person: slaty of sul ils with the bumps on their skulls, Gall Sought t develop a new objective seience for deserihin character bases ov the anatomy of the brain: anatomical less This exticrne a ncilul view was subjected exper mentat aisalvsis by Pere Flaurens at the beginning ot the rnonetcenth century, Flawens attenspeed to determine the er vis sve by retain, eaatons por iano he beau imams, Frain these experiments. Flt fut that all «sions of the fram, espectally the vetebral hemispheres of the torchsain, participate all: mental Inmiction He proposed that any part of the eezcheal hem sphere is able to poston all ot the tunetions of the hemi Sphere. Tins 6 abe area of the cerebral hemesphere Would theret-ac atleet all higher fanctons equally, Thus, ie 1824 Flot. as siete, “AH perceptions, all volitions vc ceupy the same Seat in these (cerebrall argany; the faculty ft perceiving, of cnavervang, of walle merely constitutes theiehie a haculty which # essentially one” The rapad hual tanty gene sal acceptanes nt tts belt (later called the trgeresate te che brain} sa based only panty an Hloaicny’s espoumental ork, Italso represented a phil sophieal reaction aganast the materialist basss af and the wha that smi 1s completely biological—amplied by the view that speeitie parts of che brain are dedieated to such human emotions as benevolence, hope, and selFIGURE Lot Ioan ene ot she Bunton skull This nap, taken oat a Hv The aggregate field view prevailed want the able the minereunth century, when at sas first serintsly chal lenged by the British neurslagest | ulsines tackson lackson’s clinical studies ot seterized by bray, showed that diferewt mute studies were later elabotated! systematically hy the Ger man neurologist Karl Wernc an altemauiye cew of brat tunctien called niectionivin, Aes Hulu con ding to this view, andhvdual metros are the signaling units of che brain, they are generally situated together ia tinction another i apeverse Lashiens, Wer e's we Kan parca fc meiliated by viene brain regions that are sntereumcete pathways showed! that dtterent behawes hy discrete neural The history of the sispatte herween the proponents nt the aggregate field and eetlalar connection stes6s of Carty eal function ean best be allusteates by the analy of bat guage, the hnghest and certarsly the must chaneterisee hhoman mental tanetion. Helure see eons the relevant lunical and anatonmcal studtes cancetnest th isatien of langage, fet Regions of the Brain Are Specialized for Diiferent Fun The central nervous system, whieh 9 bilateral and essen Hially sviumetiesl, consists of ty nan pants the spiel cond, the medulla ablongits, the pons (tint cerchellusn the midhrain, the diencephatan, and he cerebral heme spheres (Figures 1-2 and 18), ‘The uenleen revelations an Chapter 1 tea at Hehavier 7 2 vancty uf expoumental methods, specific hunetions have eon assigned 1 these brain aegions {Table 1A). Ax ate sul, dhe ale Oy dierent regions are specialized tor di ferent fusitinne naw accepted as ne of the eamerstones of mnadennt bate sexenee: One of the reasons chat his eo ler anvestigators far $0 many years 9 anuther viganigational prneiple of the lel processing, AS we shall see heluw, nuns sensory, mates, and ather mental func tions are subserved by more than one neural pathway wh rcvsoate partaally tor the lus, thereby wbseus ting the behavios al evidence for localization, However, the prcesvon seats rely cortaus higher fsetwons are actly clearly tram a consuderation of La Language and Other Cognitive Funetions Are Localized Within the Cerebral Cortex Hain functims rlatiog to language ace laeated primarily tiv the verebral cortex, whieh overlies the cerebral heme spheres. ln excl othe brain's ewo hemisphetes the over Iuing cortex 1 divides! ato foe anatomically distinct Iuies. jromtal qureral, aeeipital, and temporal (Figure 1a} Onginully named for overlying bones af dhe sk the lobes have specialized fnetions. The frontal lobe vs were with the planning for faeute action and ial ut qiavetent, te parietal foe wath somatic sensation ans! buy amage, the oceipital lobe with vision, and the fempural [abe with hearing as well as aspects ot Tearing, memory, and emotion, Each Tobe has several