Thomas A. Woolsey - · PDF fileBarrel Cortex Thomas A. Woolsey Page 3 of 15 on the cut edges of the cards, and stacking the cards (His like) I made a model (Figure 5). The region was

Barrel Cortex Thomas A. Woolsey

Page 1 of 15

Barrel Cortex

Thomas A. Woolsey

[In silico (2003); http://www.ibro.info/Pub_Main_Display.asp?Main_Id=21; 11/21/05]

Barrel CortexThomas A. Woolsey

Page 2 of 15Barrel cortex is part of an anatomically visible map of the contralateral body surface inlayer IV of the somatosensory cortex of certain mammals.

The introduction of phrenology of the early 1800’s stimulated interest in the possibilitythat different functions of the cerebral cortex were in different cortical “organs” (Gall andSpurzheim, 1810; Spurzheim, 1826; Temkin, 2002). Hitzig and Frisch first stimulatedthe exposed cerebral cortex of a dog electrically in 1870, reporting that only selectedareas produced movements and that the movements produced differed by the sitestimulated (Hitzig and Fritsch, 1870). Consistent regional patterns of histology, somematching functional findings, were described subsequently in the brains of man and manymammals (Campbell, 1905; Brodmann, 1909; Rose, 1929). By the 1930’s a significantbody of detailed information was available from direct stimulation of the brain inconscious patients related to movements, evoked sensations or perturbed functions, suchas speech (Cushing, 1919; Foerster, 1936; Penfield and Boldrey, 1937) (Figure 1).

Clinton Woolsey’s (my father), first study was of the dog motor cortex using corticalstimulation, cortical lesions and postoperative behavioral changes (Woolsey, 1933).However, when it was shown that amplified electrical recordings from the exposed brainsurface could be used to “map” responses evoked by sensory stimulation, he immediatelybegan to explore the organization of sensory cortical regions in animals and humans(Marshall et al., 1937; Woolsey et al., 1942; Woolsey et al., 1979). Clinton Woolsey andhis colleagues “mapped” somatic, visual, auditory and motor representations in the brainsof many species. In a summary, he proposed that the rat offered a general plan“prototypical” for the mammalian cortex (Woolsey, 1952) (Figure 2).

Apparently, I was witness to an evoked potential study from a bassinet, but I do not recallthe incident. Growing up, I spent time in my father’s laboratory as an observer andmenial worker. Richard Lende, a neurosurgeon, was one of many young trainees who hadworked with my father. Lende was interested in the cortical phylogeny and my chance toactually study the brain came when he asked me to spend a summer working in hislaboratory at the University of Colorado in the beautiful city Denver. It was there that Ilearned the evoked potential technique and surgical approaches to small mammals(Lende, 1970), primitive mammals (Lende, 1964) and mammals with small brains(Lende, 1963) .

I suppose someone had asked my father for a map of the mouse cortex and he suggested Irecord from mice prior to my matriculation at Johns Hopkins. The mouse’s brain issignificantly smaller than a rat’s, nevertheless I recorded regional activation of cortex bysound, light and touch. As in the rat, there was a large representation of the head and face(Figure 3). At the University of Wisconsin all experimental brains were evaluated byhistology (Welker et al., 1964). The Nissl stained brains I had mapped were available forexamination during the subsequent summer. When I started to match cytoarchitecture tophysiology the odd pattern of cells in layer IV of the first somatic area leapt out (Figure4). Jerzy Rose told me how to relate this architecture to the brain surface. By drawingoutlines of the sections on filing cards, cutting along these outlines, extending thepatterns of cell densities in layer IV to the surface with radial lines that were then inked


Page 3 of 15on the cut edges of the cards, and stacking the cards (His like) I made a model (Figure 5).The region was the same as that described by many authors, including Rose’s uncleMaximillian (Rose, 1929). The model showed a “cell dense net” and showed barrels. (Ionly appreciated the latter in retrospect.) The region obviously included the “glomérulos”described by Lorente de Nó from Golgi impregnations (Lorente de Nó, 1922). In mypaper I proposed that the “net” was related to the whiskers on the face.

Hendrik Van der Loos had read my paper and I approached him to work in his laboratory.There, I cut the brain in the plane of layer IV so as to look down on the “net”. Celloidin isa transparent embedding medium and I could see the brain surface for orientation as I cut50-100 µm sections. The very first section including layer IV showed neurons in a patternclearly resembling the whiskers on the opposite face (Figure 6). These were correlated tothicker sections cut perpendicular to the cortical surface. The neurons in layer IVevidently formed cylinders traversing the thickness of layer IV with slightly bowed sidesthat seemed to resemble a cask more than a ball of yarn as described in earliercytoarchitectonic and Golgi studies (Figure 7). Accordingly, we termed them barrels; thecortical region containing barrels is the barrel cortex. Photographs and drawings ofsections from many hemispheres indicated a remarkable constancy between individuals;in a particular region the barrels were larger, ovoid and, like the mouse’s large whiskers,patterned in five rows. The position of these larger barrels was appropriate to the locationand organization of the recorded representation of the larger whiskers (Woolsey and Vander Loos, 1970).

We hypothesized that a single barrel is related to a single whisker (Figure 8). Severaldifferent strategies were used to prove it. First, we excised selected whisker organs inearly postnatal life. The results were striking. Consistent with the functional maps, thepattern of the cortical barrels was altered and was always appropriate to the removedwhiskers (Van der Loos and Woolsey, 1973) (Figure 9). Moreover, the significantpotential of this system for studies of cortical and sensory development becamecompelling. Second, microelectrode recordings, including intracellular recording andstaining, showed that cells in a particular barrel are always activated best by deflectionsof the appropriate whisker (Welker, 1976; Simons, 1978; Simons and Woolsey, 1979).As other functional approaches were developed, the conclusion that neurons in aparticular barrel are first and best activated by and are part of a cortical column related tothe expected whisker has been amply supported (Durham and Woolsey, 1977; Greenberget al., 1979). The basis for the barrel pattern is: clustered afferents related to a particularwhisker (Lorente de Nó, 1922; Killackey, 1973; Senft and Woolsey, 1991; Agmon et al.,1993), clustered synapses from these afferents (White, 1976), concentration of dendritictargets of these synapses (Woolsey et al., 1975a; Woolsey, 1993), and displacement ofneuronal somata around these foci of information exchange (Harris and Woolsey, 1983;Feldmeyer et al., 2002).

Subsequent studies from many laboratories have expanded these findings greatly, oftenquantitatively, as follows: Comparative Neurology. Barrels are in rats, some otherrodents, lagomorphs and certain marsupials. Similar sub-nucletion has been described insomatic sensory pathways in other species, including man (Weller, 1972; Woolsey et al.,


Page 4 of 151975b; Goyal et al., 1992). Pathway. The organization of the entire pathway - brainstem,thalamus and cortex - can be determined in a single individual without special techniques(Killackey, 1973; Van der Loos, 1976; Durham and Woolsey, 1984; Ma and Woolsey,1984; Varga et al., 2002). Connections. The exact elements, their numbers, spatialrelationships, sources, targets and statistical variation have been comprehensivelycatalogued (White, 1976; Harris and Woolsey, 1983; Jacquin et al., 1984; Williams et al.,1994; Lübke et al., 2000; Petersen and Sakmann, 2000; Varga et al., 2002) . Molecules.Housekeeping, structural, transmitter, trophic, signaling, etc., molecules are spatiallysegregated within the context of the barrel map and are modulated by sustainedmodifications of activity (D'Amato et al., 1987; Glazewski et al., 1996; Maier et al.,1999; Kesterson et al., 2002). Development. Timing of neurogenesis, pattern formation,synaptogenisis and the influence of various molecules on these can be evaluated atspecific loci throughout pre- and postnatal development (Van der Loos and Woolsey,1973; Senft and Woolsey, 1991; Killackey et al., 1995; White et al., 1997; Fukuchi-Shimogori and Grove, 2001). Plasticity. Disrupting overall somatic activity and useduring development and in adults leads to long lasting and/or permanent changes instructure and function (Woolsey and Wann, 1976; Simons and Land, 1987; Knott et al.,2002). Metabolism and Blood Flow. Direct studies of cerebral blood flow, vessel growth,metabolism and signaling pathways all map precisely to the appropriate barrel (Wong-Riley and Welt, 1980; Dietrich et al., 1981; Gonzalez and Sharp, 1985; Adachi et al.,1994; Woolsey et al., 1996; Brett-Green et al., 2001). Behavior. The roles of motorcontrol, central pattern generators, active exploration, sensory processing discrimination,and integration in whisking are known (Welker, 1964; Bermejo et al., 1996; Hattox et al.,2002; Talwar et al., 2002). Genetics. Influence of different genes on brain form,development, signaling molecules and behavior have been investigated (Dun and Fraser,1958; Welker et al., 1996; Iwasato et al., 2000; Fukuchi-Shimogori and Grove, 2001).Modeling. Computational aspects of network function, neuronal structure, sensoryprocessing, exploratory activity, pattern development and plasticity have been considered(Pinto et al., 1996; Ahissar, 1998). Disease. Effects of environmental agents, geneticmodels of disease such as in mental retardation (Galvez et al., 2003), stroke (Wei et al.,1995), brain tumors (Sherburn et al., 1999), trauma (Jacobs et al., 1999) and for therapyand treatment can be interpreted in a standard context.

Thomas A. WoolseyDepartments of Neurosurgery, of Neurology, of Anatomy and Neurobiology, ofBiomedical Engineering, and of Cell Biology and Physiology4566 Scott Avenue, Campus Box 8057Washington University School of MedicineSt. Louis, Missouri 63110 USATel: 314-362-3600Fax; 314-362-8359e-mail: [email protected]


Page 5 of 15Acknowledgements

I am especially indebted to Kathryn Diekmann for her able assistance on this and manyother projects for many years. Support from the NIH, The McDonnell Center for Studiesof Higher Brain Function and the Spastic Paralysis Foundation of the Illinois-EasternIowa District of the Kiwanis International is gratefully acknowledged.


Page 6 of 15Favorite Sentences

Early thoughts on regional variations of the cerebral cortex:“At this time he [Gall] spoke of the necessity of the brain to the manifestations of mind,of the plurality of the mind’s organs, and of the possibility of discovering thedevelopment of the brain by the configuration of the head” (Spurzheim, 1826).

Rodents show a general layout of cortical function: “The general arrangement…[is]…a somewhat distorted image of the rat with its variousparts related one to another in much the same way as in the actual animal” (Woolsey,1952).

First association of cytoarchitecture and vibrissae:“The mouse then presents itself as a unique experimental animal, in which one may testthe relations here suggested, since the vibrissae can be moved individually to excite moreor less discretely the associated sensory endings and the glomeruli [barrels] can beprobed with microelectrodes to record unit cellular discharges” (Woolsey, 1967).

Functional organization of the cortex:“The data reported in these papers support the view that there is an elementary unit oforganization in the somatic cortex made up of a vertical group of cells extending throughall the cellular layers. The neurons of such a group are related to the same, or nearly thesame, peripheral receptive field upon the body surface” (Mountcastle, 1957).


Page 7 of 15 Bibliography

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Page 9 of 15Lübke J, Egger V, Sakmann B, Feldmeyer D (2000) Columnar organization of dendrites

and axons of single and synptically coupled excitatory spiny neurons in layer 4 orthe rat barrel cortex. J Neurosci 20:5300-5311.

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Page 10 of 15Wei L, Rovainen CM, Woolsey TA (1995) Ministrokes in rat barrel cortex. Stroke

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Woolsey TA, Van der Loos H (1970) The structural organization of layer IV in thesomatosensory region (SI) of mouse cerebral cortex. The description of a corticalfield composed of discrete cytoarchitectonic units. Brain Research 17:205-242.

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Page 11 of 15Woolsey TA, Dierker ML, Wann DF (1975a) Mouse SmI cortex: qualitative and

quantitative classification of Golgi-impregnated barrel neurons. Proc Natl AcadSci USA 72:2165-2169.

Woolsey TA, Welker C, Schwartz R (1975b) Comparative anatomical studies of the SmIface cortex with special reference to the occurence of "barrels" in layer IV. JComp Neurol 164:79-94.

Woolsey TA, Rovainen CM, Cox SB, Henegar MH, Liang GE, Liu D, Moskalenko YE,Sui J, Wei L (1996) Neuronal units linked to microvascular modules in cerebralcortex: response elements for imaging the brain. Cereb Cortex 6:647-660.


Page 12 of 15Figures & Legends

1. “[One of three]Diagrams illustratingthe more definitivelylocalized of thecortical centers of theexposed part of thehemisphere in relationto the main fissuresand convections; alsothe ‘word centers’(sensory and motor)involved in the specialmechanism for speech.(Receiving sensorystations in blue;discharging motorstations in red.)”(Cushing, 1919).

2. “Evolution of localization in thepostcentral tactile area as defined in aseries of mammals. The dotted areasshow the extent of the areas devoted tothe hand and foot. The figures are not allto the same scale, but the proportions ofeach are approximately correct.” Thedrawing at the upper left is of ahypothetical “limbless” mammal.Reproduced from (Woolsey, 1952)Patterns of localization in sensory andmotor areas of the cerebral cortex. In:The Biology of Mental Health andDisease, New York: Hoeber, page 204,Fig. 65 (1952), with permission fromLippincott Williams and Wilkins(www.lww.com).


Page 13 of 153. “Summary diagram of

sensory areas andanatomical field of celldense net.”Reproduced fromWoolsey TA (1967)Somatosensory,auditory and visualcortical areas of themouse. Johns HopkinsMed J 121 (2) p. 101,Fig. 5. © [CopyrightHolder], withpermission of theJohns HopkinsUniversity Press.

4. “Photomicrograph, coronalsection, left hemisphere …showing striking cell columnsof head face subdivision of SI …” Lines mark thethickness and plane of thetangential section in Fig. 6through layer IV. Reproducedfrom Woolsey TA (1967)Somatosensory, auditory andvisual cortical areas of themouse. Johns Hopkins Med J121 (2) p. 104, Fig 8. ©[Copyright Holder] withpermission of the JohnsHopkins University Press.


Page 14 of 15

5. Above. Filing cards stacked after cuttingalong section outlines drawn with acamera lucida attachment and marking“cell densities” on the cut edge andbound with rubber bands to show thesurface projection of the cell densitiesillustrated in Fig. 4. Below. Cardsmatched to mouse brain. The barrelpattern related to activation from thelarge whiskers is present in thereconstruction but was not appreciateduntil later.

6. The first thick (100 µm)section cut in 1968 in theplane of and throughlayer IV showedgrouping of cells in ringsorganized into 5 rows.The cytoarchitecturematched the pattern ofwhiskers that activatedthis cortex.

7. “Model” of the arrangement of cells in layer IV of the mouse cortexhypothesized to constitute part of a cortical column related to a whisker.Reproduced from Fig. 7 in Woolsey TA, Van der Loos H (1970) Thestructural organization of layer IV in the somatosensory region (SI) of mousecerebral cortex. The description of a cortical field composed of discretecytoarchitectonic units. Brain Research 17:205-242, with permission fromElsevier.


Page 15 of 15

8. Direct correspondence of the whiskers on the face and the large barrels in cortex that whiskerstimulation activates in the opposite cerebral hemisphere (Woolsey and Van der Loos, 1970).Reproduced from Fig. 15 in Woolsey TA, Van der Loos H (1970) The structural organization of layerIV in the somatosensory region (SI) of mouse cerebral cortex. The description of a cortical fieldcomposed of discrete cytoarchitectonic units. Brain Research 17:205-242, with permission fromElsevier.

9. The middle row of barrels was altered in the right cortex (drawing- above, tangential section through layer IV - middle) of a mousewhen the middle row of whiskers on the left face (bottom panels)was cauterized shortly after birth (Van der Loos and Woolsey,1973).

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Thomas A. Woolsey - · PDF fileBarrel Cortex Thomas A. Woolsey Page 3 of 15 on the cut edges of the cards, and stacking the cards (His like) I made a model (Figure 5). The region was