The Structure of the Nervous System
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The Structure of the Nervous System INTRODUCTION GROSSORGANIZATION OF THE MAMMALIAN NERVOUSSYSTEM ANATOMICALREFERENCES THE CENTRALNERVOUS SYSTEM The Cerebrum The Cerebellum Ihe BroinStem The SpinolCord THE PERIPHERALNERVOUS SYSTEM Ihe SomoticPNS Ihe ViscerolPNS Afferentand EfferentAxons THECRANIAL NERVES THEMENINGES THEVENTRICULARSYSTEM r Box 7.1 Of SpecialInterest: Water on the Brain IMAGINGTHE LIVINGBRAIN ComputedTomogrophy MagneticResononce lmaging r Box 7.2 Brain Food:Magnetic Resonance Imaging FunctionolBroin lmaging r Box 73 Brain Food:Functional Imaging of Brain Activity: PETand fMRI UNDERSTANDINGCNS STRUCTURETHROUGH DEVELOPMENT FORMATIONOF THE NEURALTUBE r Box 7.4 Of SpecialInterest: Nutrition and the Neural TLrbe THREEPRIMARY BMIN VESICLES DIFFERENTIATIONOFTHE FOREBRAIN Differentiotionof theTelencepholon ond Diencephalon ForebroinStructure-F unction Relotionships DIFFERENTIATIONOFTHE MIDBRAIN MidbroinStructure-Function Relotionshrps DIFFERENTIATIONOFTHE HINDBRAIN HindbroinStructure-Function Relotionshrps DIFFERENTIATIONOFTHE SPINAL CORD SpinolCord Structure-Function Relotionshps PUTTINGTHE PIECESTOGETHER SPECIALFEATURES OF THE HUMANCNS A GUIDE TO THE CEREBRALCORTEX TYPESOF CEREBRALCORTEX AREASOF NEOCORTEX Neocortico/Evolution ond Structure-FunctionRelotionshrps r Box 7.5 Pathof Discovery:Evolution of My Brain, by Leah A. Krubitzer CONCLUDING REMARKS APPENDIX:AN ILLUSTRATEDGUIDE TO HUMAN NEUROANATOMY r68 CHAPTER 7 . THESTRUCTUREOFTHENERVOUSSYSTEM V INTRODUCTION In previous chapters,we saw how individual neurons function and com- municate. Now we are ready to assemblethem into a nervous systemthat sees,hears, feels, moves, remembers,and dreams.Just as an understand- ing of neuronal structureis necessaryfor understandingneuronal function, we must understandnervous system structure in order to understandbrain function. Neuroanatomy has challenged generations of students-and for good reason:The human brain is extremely complicated.However, our brain is merely a variation on a plan that is common to the brains of all mammals (Figure 7.1). The human brain appearscomplicated because it is distorted as a result of the selectivegrowth of some parts within the confinesof the skull. But once the basicmammalian plan is understood,these specializa- tions of the human brain becometransparent. We begin by introducing the genbral organization of the mammalian brain and the terms used to describeit. Then we take a look at how the three-dimensionalstructure of the brain arisesduring embryologicaland fetal development.Following the courseof developmentmakes it easierto understandhow the parts of the adult brain fit together.Finally, we explore the cerebralneocortex, a structure that is unique to mammals and pro- portionately the largest in humans. An Illustrated Guide to Human Neuroanatomyfollows the chapter as an appendix. The neuroanatomy presentedin this chapter providesthe canvason which we will paint the sensoryand motor systemsin Chapters8-14. Becauseyou will encounter a lot of new terms, self-quizzeswithin the chapter provide an opportunity for review. V GROSSORGANIZATION OF THE MAMMALIAN NERVOUSSYSTEM The nervous systemof all mammalshas two divisions:the central nervous system(CNS) and the peripheralnervous system(PNS). In this section,we identify some of the important componentsof the CNS and the pNS. We also discussthe membranesthat surround the brain and the fluid-filled ventricleswithin the brain. We then explore some new methods of examin- ing the structure of the living brain. But first, we need to review some anatomical terminology. Anatomical References Getting to know your way around the brain is like getting to know your way around a city. To describeyour location in the city, you would use points of referencesuch as north, south, east,and west, up and down. The same is true for the brain, except that the terms-calle d anatomicalrefer- ences-are different. Considerthe nervous systemof a rat (Figure 7.2a).We begin with the rat, becauseit is a simplified version that has all the general featuresof mammalian nervous system organization.In the head lies the brain, and the spinal cord runs down inside the backbonetoward the tail. The direc- tion, or anatomicalreference, pointing toward the rat's nose is known as anterior or rostral (from the Latin for "beak"). The direction pointing toward the rat's tail is posterior or caudal (from the Latin for "tail"). The direction pointing up is dorsal (from the Latin for "back"), and the direc- tion pointing down is ventral (from the Latin for "belly"). Thus, the rat Y GROSSORGANIZATION OFTHE MAMMALIAN NERVOUS SYSTEM I69 rIm .:! Rat Rabbit {ffi Cat dffi,':4,.\.\i#r /at '\--\\f 1 ."- Sheep Chimpanzee ChimDanzee ,d Human FIGURE7.I Mammalian brains. Despitedifferences rn complexity,the brarnsof all these specieshave many featuresin common.The brainshave been drawn to appearapproximately the same stze; their relativesizes are shown in the inseton the left, t70 CHAPTER 7 . THESTRUCTUREOFTHENERVOUSSYSTEM Spinal coro Dorsal Anterior Posterior or rostral or caudal <- :-------> (a) Ventral FIGURE7,2 Basic anatomical references in the neryous system oi a rat. (a) Sideview. spinal cord runs anterior to posterior.The top side of the spinal cord is the (b) Top view. dorsalside, and the bottom side is the ventral side. If we look down on the nervous system,we seethat it may be divided into two equal halves (Figure 7.2b). The right side of the brain and spinal cord is the mirror image of the left side. This characteristicis known as bilateralsymmetry. with just a few exceptions,most structureswithin the nervous systemcome in pairs, one on the right side and the other on the left. The invisible line running down the middle of the nervous systemis called the midline, and this gives us another way to describeanatomical references.structures closerto the midline are medial; structuresfarther away from the midline are lateral. In other words, the nose is medial to the eyes,the eyesare medial to the ears,and so on. In addition, rwo struc- tures that are on the sameside are said to be ipsilateral to each other; for example,the right ear is ipsilateralto the right eye. If the structuresare on oppositesides of the midline, they are said to be contralateral to each other; the right ear is contralateralto the left ear. To view the internal structureof the brain, it is usually necessaryto slice it up. In the languageof anatomists,a slice is called a section;to slice is /o section.Although one could imagine an infinite number of ways we might cut into the brain, the standardapproach is to make cuts parallel to one of the three anatomicalplanes of section.The plane of the section resulting from splitting the brain into equal right and left halvesis calledthe midsagittal plane (Figure 7.3a). sectionsparallel to the midsagittalplane are in the sagittal plane. The two other anatomicalplanes are perpendicularto the sagittalplane and to one another. The horizontal plane is parallel to the ground (Figure 73b]'. A single sectionin this plane could passthrough both the eyesand the ears.Thus, horizontal sectionssplit the brain into dorsal and ventral parts. The coronal plane is perpendicularto the ground and to the sagit- tal plane (Figure 7.3c). A single section in this plane could passthrough both eyesor both ears,but not through all four at the sametime. Thus, the coronal plane splitsthe brain into anterior and posteriorparts. v SELF-QUtZ Take a few moments right now and be sure you understand the meaning of theseterms: anterior ventral contralateral rostral midline midsagittalplane posterior medial sagittal plane caudal lateral horizontal plane FIGURE7.3 dorsal ipsilateral coronal plane Anatomical planes of section. V GROSSORGANIZATION OFTHE MAMMALIAN NERVOUS SYSTEM t7l The Central Nervous System The central nervous system (CNS) consistsof the parts of the nervous systemthat are encasedin bone: the brain and the spinal cord. The brain lies entirely within the skull. A sideview of the rat brain revealsthree parts that are common to all mammals:the cerebrum,the cerebellum,and the brain stem (Figure7.4a). The Cerebrum. The rostral-most and largestpart of the brain is the cerebrum. Figure 7.4b showsthe rat cerebrumas it appearswhen viewed from above. Notice that it is clearly split down the middle into two cere- bral hemispheres, separatedby the deepsagittal fissure.In general, the right cerebralhemisphere receivessensations from, and controls move- ments of, the left side of the body. Similarly, the left cerebralhemisphere is concernedwith sensationsand movements on the right side of the body. The Cerebellum. Lying behind the cerebrum is the cerebellum (the wogd is derived from the Latin for "little brain"). While the cerebellumis in fact dwarfed by the large cerebrum, it actually contains as many neurons as both cerebralhemispheres combined. The cerebellumis primarily a movement control center that has extensiveconnections with the cerebrum and the spinalcord. In contrastto the cerebralhemispheres, the left sideof the cere- bellum is concernedwith movementsof the left side of the body, and the right sideof the cerebellumis concernedwith movementsof the right side. The Brain Stem. The remaining part of the brain is the brain stem, best observedin a midsagittalview of the brain (Figure7.4c\. The brain stem forms the stalk from which the cerebralhemispheres and the cerebellum sprout. The brain stem is a complex nexus of fibers and cells that in part serves to relay information from the cerebrum to the spinal cord