Muhammad Khairul Hevrizal 24110754 3KB05

Chapter 36 Statistical Outliers in Neuron Population and Myelinated Fiber Development in Human Neocortex
David ! Cooper" #ames $! %entle and #ames ! Olds &bstract Between 1939 and 1967, JL !nel "er#!rmed detailed hi$t!l!%i&al e'aminati!n$ !# 54 human $u()e&t$ at ei%ht !($ervati!n$ a%e$ ran%in% #r!m new(!rn$ t! 6 *ear$ !ld+ He made detailed !($ervati!n$ !# neur!n den$itie$, ,ei%ert and -!l%i. !' $tained #i(er$, #i(er diameter$, and &ell $ize$ (* ne!&!rti&al la*er #!r u" t! 43 ne!&!rti&al area$ #!r ea&h a%e+ Here we u$e thi$ data t! &!n$tru&t $tandard $am"le &!lumn$ t! &al&ulate &han%e$ (etween !($ervati!n a%e$, and $h!w that it i$ well mat&hed (* a %amma di$tri(uti!n+ ,e then $h!w that a devel!"mental di$tan&e metri& /00M1 tran$#!rmati!n #a&ilitate$ a $tati$ti&al e'"l!rati!n !# inter.area di##eren&e$ durin% devel!"ment+ 'ntroduction JL !nel2$ 314 #!ur milli!n !($ervati!n$ !# neur!n den$it*, la*er thi&5ne$$, $!mal $ize, -!l%i. !' and ,ei%ert.$tained #i(er den$itie$ #r!m human $u()e&t$ thr!u%h 6 *ear$ !# a%e /N 6 541, "r!vide$ a uni7ue $!ur&e !# in#!rmati!n (el!w &urrent ima%in% re$!luti!n+ 8han5le et al+ 324 u$ed a multivariate !rdinati!n te&hni7ue &alled &!rre$"!nden&e anal*$i$ t! redu&e !nel2$ data int! 1727 #eature "r!#ile$ #!r the 43 re%i!n$ !# the human ne!&!rte' that !nel e'amined+ 9he "r!#ile$ reveal di$tin&t time."ha$ed &han%e$: #!r e'am"le, neur!n "a&5in% den$it* de&rea$e$ with a%e, while #i(er m*elinizati!n in&rea$e$+ 9hi$ anal*$i$ "r!vided a &!m"ellin% ar%ument #!r a (r!ad devel!"mental "attern !# &han%e$ a&r!$$ individual$+ H!wever, &!rre$"!nden&e anal*$i$ d!e$ n!t "re$erve in#!rmati!n !n the di$tri(uti!n$ !# &han%e #!r the individual mea$urement$, and thu$ d!e$ n!t hi%hli%ht di##eren&e$ (etween ne!&!rti&al area$+ 9h!$e di##eren&e$, i# the* e'i$ted, mi%ht well (e !# #un&ti!nal im"!rtan&e+ Here we a""l* a n!vel tran$#!rmati!n meth!d that"re$erve$ that in#!rmati!n !n di$tri(uti!n$, while e'"andin% in#!rmati!n !n the tem"!ral detail$ !# devel!"ment+ ,e $h!w that the mea$urement !# neur!n$ and m*elinated #i(er$ in the human ne!&!rte' durin% devel!"ment &!n#!rm$ t! %amma di$tri(uti!n$+ Methods ,e em"l!*ed tw! tran$#!rmati!n$ t! !nel2$ neur!n den$it* and ,ei%ert.$tained /m*elinated1 #i(er den$it*+ 9he #ir$t tran$#!rmati!n wa$ t! &!nvert den$it* in#!rmati!n (* &!rti&al area and la*er int! $tandard $am"le &!lumn$ /88 $1+ ;n the &a$e !# neur!n "!"ulati!n$, !nel made hi$ mea$urement$ #!r $tandard v!lume$ !# 100<m3+ =!r #i(er "!"ulati!n$, he &!unted $tained #i(er$ in a $am"le v!lume !# 50<m > 50<m > 25<m+ 88 $ were &!n$tru&ted u$in% !nel2$ la*er thi&5ne$$ mea$urement$ #!r the$e unit v!lume$+ ,hen !nel re"!rted the den$itie$ #!r $u(la*er$ rather than &!m"lete la*er$, we u$ed the mean den$it* #!r the la*er+ Here, we "r!du&ed 88 $ &!rre$"!ndin% t! 19 $am"le Br!dmann ?rea$ related t! audi! and vi$ual "r!&e$$in%+ 9he $e&!nd tran$#!rmati!n wa$ t! &reate 00M$ u$in% the &han%e$ (etween a%e$ within ea&h 88 t*"e /neur!n !r #i(er1+ ,e treated !nel2$

mea$urement$ at the ei%ht !($ervati!n a%e$ a$ ei%ht e'"eriment$ !n a ne!&!rti&al ma&r!arra*, &!m"ara(le t! mi&r!arra* anal*$i$ !# %ene !r "r!tein e'"re$$i!n /#!r e'am"le, 3341+ 8imilar t! 8trand et al+, we #ir$t t!!5 the l!% "r!"!rti!n !# neur!n !r #i(er "!"ulati!n$ within ea&h !($ervati!n a%e 3@7+ /36+114+ Ae't, rather than the z.$&!re meth!d u$ed in 8trand et al+, we n!rmed &han%e$ t! the l!% "r!"!rti!n$ a&r!$$ !($ervati!n a%e$ (* u$in% the "!!led varian&e 3@7+ /36+214+ =inall*, a$ in 8trand et al+, we e'amined #!r &lu$terin% !# the $&!re$ (* ta5in% the @u&lidean di$tan&e$ (etween the n!rmed $&!re$ area.(*.area+ !nel made hi$ !($ervati!n$ at ei%ht di##erent a%e$ #r!m new(!rn t! 6 *ear$ !# a%e, $! the 00M$ "r!vide &umulative d!u(le.n!rmed mea$ure$ #!r $even $et$ !# &han%e$+ LP/?reai 1 6 l!%/ !unti /<j !untj 1 /36+11 D 6 /1 B 212//2 1 C 2 2 1 /36+21 9! e'amine the &han%e data, we #ir$t mea$ured unad)u$ted 88 &han%e data #!r ea&h la*er, and then e'amined the di$tri(uti!n$ &reated (* d!u(le.n!rmed 00M$+ (esults Dverall neur!n &han%e data #!r the 54 (rain$ $h!w$ a ne%ative mean and ne%ative $5ewne$$ /=i%+ 36+1a1+ Dn avera%e, a la*er l!$t 36+9 neur!n$ within a 100<m2 &r!$$$e&ti!n (etween an* tw! !($ervati!n a%e$+ 85ewne$$ #!r an entire 88 wa$ B1.74+ 9he m*elinated #i(er &han%e$ have "!$itive $5ewne$$ /2+36 #!r the 88 1 and a 36 8tati$ti&al Dutlier$ in Aeur!n E!"ulati!n and M*elinated =i(er 0evel!"ment 205 Fi)! 36!* Aeur!n and m*elinated #i(er di$tri(uti!n$+ Aeur!n hi$t!%ram$ u$e $!lid line$+ =i(er hi$t!%ram$ u$e da$hed line$+ /a1 &han%e$ t! neur!n and #i(er &!unt$ in a la*er #!r 19 ne!&!rti&al area$+ /(1 &!m"ari$!n !# &han%e data #!r neur!n l!$$e$ and #i(er in&rea$e$ (* la*er+ /&1 00M neur!n and #i(er di$tri(uti!n$ "!$itive mean /23+2 #!r a 50<m2 &r!$$.$e&ti!n #!r ea&h la*er1+ 9! te$t #!r a %amma #it, we &!m"ared neur!n l!$$e$ t! m*elinated #i(er %ain$ /=i%+ 36+1(1+ ,e re#le&ted the neur!n &han%e data a(!ut an a'i$ e7ual t! the ma'imum value, and tran$lated #i(er data t!ward the !ri%in, e'&ludin% the $i' di$&!nne&ted ne%ative !utlier$ in =i%+ 36+1a+ 9he di$tri(uti!n$ then (e&ame "!$itivel* $5ewed !ver a n!n.ne%ative ran%e+ ,e &!m"ared the$e em"iri&al di$tri(uti!n$ t! (!th a %amma di$tri(uti!n and a l!%n!rmal di$tri(uti!n with &!rre$"!ndin% value$ (a$ed !n ma'imum li5elih!!d e$timate$+ ,e then a""lied a K!lm!%!r!v.8mirn!v %!!dne$$.!#.#it te$t #!r the$e di$tri(uti!n$ a%ain$t a &umulative den$it* #un&ti!n e7ual t! the mean !# a 10,000. iterati!n rand!m #un&ti!n with the &!rre$"!ndin% e$timated "arameter$+ P.value$ a""ear in 9a(le 36+1+ 9hi$ reveal$ that a %amma di$tri(uti!n m!del$ the em"iri&al data well+ =!r an 88 , ma'imum li5elih!!d e$timate$ #!r %amma "arameter$ #!r neur!n$ are < 6 5.4764 /95 "&t &!n#iden&e interval 6 34.97F7, 6.023941, and < 6 0.0745 /95 "&t&!n#iden&e interval 6 30.0675, 0.0F2441+ 9he P.value #r!m a K!lm!%!r!v. 8mirn!v %!!dne$$.!#.#it te$t i$ 0+3116+ =!r m*elinated #i(er$ < 6 4.3359 /95 "&t &!n#iden&e interval 6 33.F409, 4.F94641, and < 6 24.FF31 /95 "&t &!n#iden&e interval 6 321.FF24, 2F.295241+ 9he a$$!&iated P.value i$ 0+3334+ ?lth!u%h d!u(le.n!rmed 00M$ are %amma.li5e /the K!lm!%!r!v.8mirn!v P.value i$ 0+1546 #!r 4F and 72 m!nth$ #!r neur!n$, and hi%her #!r m*elinated +able 36!* P.value$ a$$!&iated with K!lm!%!r!v.8mirn!v %!!dne$$.!#.#it te$t$ #!r %amma and l!%n!rmal di$tri(uti!n$ &!m"ared t! the mean &umulative den$it* #un&ti!n #r!m 10,000 iterati!n$ u$in% e$timated "arameter$ #!r neur!n "!"ulati!n and m*elinated #i(er data (* la*er P.value Aeur!n %amma Aeur!n l!%n!rmal =i(er %amma =i(er l!%n!rmal La*er ; 0+3759 0+3755 0+4219 0+4169

La*er ;; 0+35F2 0+342F 0+403F 0+4026 La*er ;;; 0+2435 0+1405 0+3553 0+3391 La*er ;G 0+3F99 0+3F97 0+2671 0+3045 La*er G 0+3F40 0+3649 0+3732 0+361F La*er G; 0+3F39 0+3605 0+2961 0+15FF #i(er$1, their hi$t!%ram$ a""ear t! (e multim!dal /=i%+ 36+1&1+ ? 50.(in hi$t!%ram !# the &umulative di$tan&e$ at 72 m!nth$, #!r e'am"le, ha$ tw! l!&al ma'ima t! the le#t !# the !verall ma'imum, and ha$ $everal l!&al ma'ima at the ri%ht tail $e"arated #r!m the re$t !# the di$tri(uti!n (* em"t* (in$+ -enerall*, the $h!rte$t d!u(le.n!rmed neur!n di$tan&e$ &!rre$"!nd t! La*er ;G /the thalami& in"ut la*er1 #!r m!$t !# the 19 area$, wherea$ the l!n%e$t d!u(le.n!rmed di$tan&e$ "ertain t! "rimar* vi$ual &!rte' /B? 171, the anteri!r &in%ulate &!rte' /B? 241, and the d!r$!lateral "re#r!ntal &!rte' /B? 91+ @'treme di$tan&e$ #!r #i(er$ &!rre$"!nd t! "rimar* m!t!r &!rte' /B? 41+ Discussion 00M$ a""ear t! Hun#!ldI the $tate.$"a&e a$$!&iated with neur!n l!$$e$ and m*elinated #i(er %ain$ durin% devel!"ment #r!m (irth thr!u%h 6.*ear$ !# a%e+ 9he di$tan&e metri& em(ed$ the &han%e data in a hi%her.dimen$i!nal mani#!ld and ena(le$ the $e"arati!n !# &han%e #a&t!r$ that are initiall* nearl* ad)a&ent in $tate $"a&e 344+ 9he "re$en&e !# la*er !r area.$"e&i#i& a$$!&iati!n$ in the !utlier area$ indi&ate$ that thi$ i$ a "r!mi$in% avenue t! "ur$ue in #uture re$ear&h+ (e,erences 1+ !nel JL /1939J19671 E!$tnatal devel!"ment !# the human &ere(ral &!rte', G!l$ 1JF+ Harvard Kniver$it* Ere$$+ 2+ 8han5le ,L, L!mne* ?K, Landin% BH, Hara J /199F1 0evel!"mental "attern$ in the &*t!ar&hite&ture !# the human &ere(ral &!rte' #r!m (irth t! 6 *ear$ e'amined (* &!rre$"!nden&e anal*$i$+ Er!& Aat ?&ad 8&i K8? 95M4023J402F+ 3+ 8trand ?0, ?ra%a5i ?K, Ba7uet N , H!d%e$ ?, unnin%ham E, H!lman$ E, J!ne$ KL, J!ne$ L, K!!"er(er% , Dl$!n JM /20071 !n$ervati!n !# re%i!nal %ene e'"re$$i!n in m!u$e and human (rain+ EL!8 -enet 3/41M e59 d!iM10+1371O)!urnal+"%en+0030059+ 4+ -ra$$(er%er E, Er!&a&&ia ; /19F31 Mea$urin% the $tran%ene$$ !# $tran%e attra&t!r$+ Eh*$i&a 0 9M1F9J20F+