LEE WILLERMAN ROBERT SCHULTZ University of Texas at Austin J. NEAL RUTLEDGE University of Texas at Austin Austin Radiological Association ER1N D. BIGLER University of Texas at Austin Austin Neurological Clinic It is widely believed that human brain size and intelligence are only weakly related to each other. Using magnetic resonance imaging, we show that larger brain size (corrected for body size) is associated with higher IQ in 40 college students equally divided by high versus average IQ, and by sex. These results suggest that differences in human brain size are relevant to explaining differences in intelligence test performance. St udi es have shown pos i t i ve but modes t r el at i ons bet ween ext ernal head si ze and ps ychomet r i c i nt el l i gence ( e . g. , Van Val en, 1974, f or a revi ew of ear l i er re- search; Sus anne, 1979). An as s umpt i on in much of t hat wor k was that head and br ai n si ze wer e f ungi bl e, and i f head si ze wer e not power f ul l y pr edi ct i ve of i nt el l i gence, br ai n si ze woul d not do much bet t er ( Goul d, 1981). Lar ge bet ween- per son var i at i on in aver age skul l t hi ckness compr omi s es ext r apol at i ons of ext er- nal cr ani al meas ur ement s t o pr edi ct cr ani al capaci t y, however ( Roger s, 1984). The evol ut i on of br ai n si ze, sex di f f er ences, and i nt el l i gence have been cl assi c quest i ons ( Jer i son, 1973). Pos t mor t em evi dence br ought to bear in t he 19th and ear l y 20t h cent ur i es was def i ci ent in s ampl i ng and met hodol ogy: Sex, age, body si ze, br ai n- f i xat i on pr ocedur es , wei ghi ng bef or e or aft er f i xat i on, i ncl usi on or excl usi on of meni nges , and cause o f deat h wer e oft en i nadequat el y cont r ol l ed. El der l y per s onages wi t h smal l br ai ns wer e sai d t o i l l ust rat e t he i r r el evance of br ai n si ze in hi gh achi evement , and l ar ge- br ai ned per sonages wer e t hought to Erin D. Bigler is now at the Department of Psychology, Brigham Young University. We thank N. Gopal and P. Watson of the University of Texas Advanced Graphics Laboratory for computer programing, and J.C. Loehlin, T. Schallert, and W. Wilczynski for advice and encouragement. Correspondence and requests for reprints should be sent to Lee Willerman, Department of Psychology, University of Texas at Austin, Austin, TX 78712. 223 224 WILLERMAN, SCHULTZ, RUTLEDGE, AND BIGLER champi on the opposi ng view. However, brain size at the height of intellectual power would have been a better index than aged brain size postmortem. Relating intelligence to intracranial volume rather than to post mort em brain size might have finessed some of these probl ems, but no such study has been done. The advent of in vivo brain imaging would now seem to have rendered studies of head size and intelligence obsolete. Brain size in living younger subjects can be measured reliably, and statistical controls for body size and socioeconomic back- ground can be applied. Gi ven the limitations of previous work, this study was designed specifically to examine brain morphometric correlates of intelligence in an ostensibly healthy college-age sample using magnetic resonance imaging (MRI). METHOD Subjects The final sample consisted of 40 right-handed Anglo introductory psychol ogy students (M = 18.9 years, SD = 0.6) who had indicated no history of alco- holism, unconsciousness, brain damage, epilepsy, or heart disease. These sub- jects were drawn from a larger pool of introductory psychol ogy students with total Scholastic Aptitude Test scores of --> 1350 or <-- 940 who had agreed to satisfy a course requirement by allowing the administration of four subtests (Vocabulary, Similarities, Block Design, and Picture Completion) of the Wechsler (1981) Adult Intelligence Scale-Revised. With prior approval of the Uni versi t y' s research review board, students selected for MRI were required to obtain prorated full-scale IQs of >- 130 (M = 136.4, SD = 3.95) or --< 103 (M = 90. 5, SD = 8.12), and were equally divided by sex and IQ classification. Instruments and Analyses Eligible volunteers made appointments for MRI at a local facility. Using the lowest margin of the cerebellum in a midsagittal view to align the first axial (horizontal) MR slice, 18 mi xed-wei ght ed images (spin-echo pulse sequence with a TR of 2000 msec and a TE of 30 msec) were obtained from a Signa MRI unit with a field strength of 1.5 Tesla. All images were 5 mm thick and separated by 2.5 mm. Each image was 256 256 pixels in area with 256 levels of gray. The MRI tape was read into a VAX comput er and the images analyzed after removi ng identifying information. Analyses were done blindly with respect to IQ and sex. For each slice, a Roberts gradient traced the boundary of the scalp by outlining large-intensity differences between adjacent pixels. All gray scale intensity values of <96 within this boundary were converted to zero. This deleted skull, most of the meninges, and interhemispheric fissure; other brain coverings were deleted man- ually with a cursor. Interrater agreement for the number of pixels contained BRAIN SIZE AND INTELLIGENCE 225 within a slice was r = . 99 (n = 10). The comput er then count ed all pixels with nonzer o gr ay scale values for brain size in each slice, their summed value serving as the index of overall brain size. We report here onl y on total brain size; a future report will focus on cortex versus subcort ex vol umes. RES ULTS Based on scores on a iaterality quest i onnai re (Oldfield, 1971), the hi gh- l Q group was mor e ri ght -l at eral i zed (p < .05), but this difference did not interact with any ot her results. Aver age- I Q men were taller than hi gh- I Q men (p < .05). Mid- parent years of compl et ed educat i on f or the t wo I Q groups did not differ signifi- cant l y: hi gh I Q = 16. 95 years, S D = 1.79 and average I Q = 16.0 years, S D = 1. 95, p > . 10. Cont rol l i ng f or mi dparent educat i on had no effect on the results to be report ed. Brain tissue was present on onl y 17 of 18 slices for 12 of the subjects (9 women). Aft er cont rol l i ng f or sex, correl at i ons of height and wei ght with brain size were onl y r = .09 and r = . 10. Body size, nevertheless, was partialled out because studies using larger sampl es have shown moderat e br ai n- si ze- body- si ze correl at i ons ( Dekaban & Sadowsky, 1978; Ho, Roessman, St raumfj ord, & Munr oe, 1980; Hol l oway, 1980), and brain size adjusted for body size is believed t o approxi mat e mor e cl osel y the proport i on of brain devot ed to intellectual func- t i on (Jeri son, 1973). Anal ysi s of covar i ance ( ANCOVA) cont rast ed the t wo di chot omous variables: hi gh versus average 1Q and mal e versus femal e, cont rol l i ng for body size (height and weight). Results showed that the hi gh- l Q gr oup had greater brain size, F( 1, 34) = 6. 6, rph = . 40, p > . 05, as did the men, F( 1, 34) = 4. 7, r~, h = . 35, p < . 05, wi t h no i nt eract i on bet ween I Q and sex. An analysis adjusting for height and wei ght separat el y by sex yi el ded essentially the same results. An analysis that excl uded a single outlier wi t h l arger body size did not affect the br ai n- s i ze- I Q relation, but reduced the sex difference to margi nal significance (p = .08); thus, the sex di fference in adjusted brain size must be consi dered tentative. A stepwise multiple regressi on with the di chot omous I Q classification entered first and the actual I Q scores entered second, tested whet her IQ scores cont ri b- ut ed t o the prediction of adjusted brain size within the t wo I Q subgroups. Results showed a significant cont ri but i on of residualized I Q scores (partial r = . 41, p < .05) after cont rol l i ng f or IQ cl assi fi cat i on, suggest i ng generalizability t o a con- t i nuous I Q distribution. Correl at i ons of brain size with I Q scores were hi gher among men t han women, al t hough not si gni fi cant l y so. Among men, 1Q scores as a cont i nuous variable correl at ed with brain size, before and after adjusting f or body size, r = .51 (p < .05) and r = .65 (p < .01). Correspondi ng correlations f or women were r = .33 and . 35, bot h n. s. Wi t h sexes pool ed, the IQ-adj ust ed brai n-si ze correl at i on was r = .51 (p < .01). This correl at i on is hi gher than 2 2 6 WI L L E RMAN, SCHULTZ, RUTLEDGE, AND BI GLER expect ed f or the general popul at i on because of sel ect i on of ext reme I Q groups. Appl yi ng a statistical correct i on (Gui l ford & Fruchter, 1973) predi ct ed a correl a- tion of r = .35 f or a mor e represent at i ve sampl e. Not all brai n levels cont ri but ed equal l y t o the br ai n- s i ze- I Q correl at i on (Fig- ure 1); size di fferences were greatest f or ventricular-level slices in men. These levels i ncl ude l anguage circuits, associ at i on fibers, and associ at i on cortex (Dea- con, 1988), but mor e di rect anat omi c compar i son in speci fi c regi ons of interest is requi red f or substantiation. The axial i mage approxi mat i ng t he gl abel l a-opi st hocrani on plane was used to est i mat e head perimeter. Head- per i met er - I Q correl at i ons in men (r = . 17) and women (r = .31) were not si gni fi cant , nor was the unadj ust ed br ai n- si ze- head- peri met er correl at i on in men (r = .24). However , the cor r espondi ng correl at i on was si gni fi cant in women (r = . 68, p < .01). The pe r i me t e r - l Q correl at i ons are consi st ent with previ ous reports of such relationships ( e. g. , Van Valen, 1974), and mor e variability in skull t hi ckness in men coul d account for the sex dif- ference in the head- per i met er - br ai n- si ze correl at i on ( Roger s, 1984). 1.0 n ~ 0 .) 0 1 0.5 Z o.o n N - 0 . 5 Z m - I . 0 - I 0 I I I I I I I I - 8 - 6 - 4 - 2 0 2 4 6 SLICE LEVEL F I G. 1. Br ai n ar ea i n s t andar d scor es (M = 0 +- SD = l ) by hi gh ver s us aver age I Q and sex, adj us t ed f or hei ght and wei ght . Squar es r ef er t o me n and ci r cl es t o wome n; e mpt y s ymbol s r ef er t o hi gh I Q, f i l l ed s ymbol s t o aver age IQ. Sl i ce 0 i s at t he mi dve nt r i c l e l evel as s hown i n t he i nsert ; 5 mm t hi ck MR1 i ma ge s are separ at ed by 2. 5 r am. Lar ge b r a i n - a r e a - I Q di f f er ences i n men near vent r i cl es ( sl i ces - 3 be l ow t o + 1 a bove 0) i ncl ude neur al s ubs t r at es of l a ngua ge and as s oci at i on. A r epeat ed- meas ur es ANOVA on vent r i cl e s i ze acr oss s l i ces - 3 t o + 1 r eveal s no ef f ect of sex, I Q, or t hei r i nt er act i on (al l p s > . 20) s o ve nt r i c l e s ar e i ncl uded i n ar ea me a s ur e me nt s . Ti s s ue above or be l ow gr aphed l evel s ar e e xc l ude d be c a us e al l s ubj ect s c oul d not be r epr es ent ed. BRAIN SIZE AND INTELLIGENCE 227 DI SCUSSI ON A previous study of brain size and IQ (Yeo, Turkheimer, Raz, & Bigler, 1987) obtained a correlation of r = .07 with sexes pooled, using eight or nine con- tiguous computerized t omography (CT) slices maxi mal l y encompassing about 53% of the brain. The patients had medically unconfirmable neurologic symp- t oms and many had elevated psychopat hol ogy scores. A correlation of r = .57 (p < .01) obtained for relative hemisphere size and superiority of verbal or nonver- bal IQ within subjects is difficult to reconcile with the low between-subjects correlation for brain size and IQ. A recent CT study found an eta correlation of .35 (p < .05) between area of a single ventricular-level slice and occupational class in normal adults, but the sampl e' s racial heterogeneity makes interpretation ambi guous (Pearlson et al., 1989). Corresponding results were obtained for Frontal Lobe Size Educational Achievement (r = .31) in an MR1 study (Andreason et al., 1990). Neither of the last two studies controlled for sex, and it is, thus, not feasible to examine possible sex differences in the correlations. Although the sex difference in the adjusted br ai n- si ze- I Q correlation was not significant here, its absolute magnitude (r = .35 vs. r = .65) warrants mention of possible sex differences in brain organization. Aphasia in women is com- paratively less frequent following focal left hemisphere damage, except in one region where it is more likely (Kimura, 1987). There is also some indication that men and women have the same number of cortical neurons despite differences in overall brain size (Haug, 1987). That the sexes may package similar processing capacities differently implies alternative routes to achieving similar adaptive goals. It is possible that unknown environmental factors common to both brain size and IQ could be responsible for their correlation. One candidate is nutritional variation, but that seems improbable because the middle-class background of our subjects makes prenatal or postnatal undernutrition unlikely. Future research, however, should be directed to establishing whether the br ai n- si ze- I Q effect occurs within, as well as between, families (Jensen & Sinha, in press). Brain size is correlated with cortical surface area (Haug, 1987) so that larger size might reflect more cortical columns available for analyzing high-noise or low-redundancy signals, thus enabling more efficient information processing pertinent to IQ test performance (Raz, Willerman, & Yama, 1987). Embryologic factors may be important because the panoply of stem cells giving rise to cortical neurons is fully present by the 40th day after fertilization, and the full comple- ment of cortical neurons is present before birth (Rakic, 1988; Williams & Her- rup, 1988). Glial cells contribute to postnatal brain growth, but their influence on intelligence is unknown. Attractive hypotheses to explain larger cortices are a greater number of stem cells, an increased number of mitotic divisions producing more descendant neurons, or different rates of neuronal death. 228 WILLERMAN, SCHULTZ, RUTLEDGE, AND BIGLER R E F E R E N C E S Andreasen, N. C. , Ehrhardt, J. C. , Swayze, V.W., II, Alliger, R. 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