Pisareva 1965 - translation.pdf

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    BIOELtCTnIC RF.AC':IOIiS or TIlt :UDBRAItl TECTUH OFCHICK EMBRYOS ro LICl:T S ' l ' I ~ L I

    ZhW'Dal EyollUtalonnoy lUokh1ll11 i H. L. P l l ~ v a . , !"hiol0611. Vol. , 1, lIo. 2, pp 176-1e2,1965. Laboratory of Compa.rative Onto~ e n e & i s o f the Nervous System,Brain Inst i tute ot the A c a d ~ or Medical Dciences of the UGSR,Molcow.

    ABSTRACT. I lec t r iea l relponsea to , taulat1on with single,rhythmical or paired flashes vere recorded t'r0lll the dorsallur taee or the Illidbrain teet\llll. Potentials evoked by singlef l uhe . were shown to occur in the contralateral midbrainteet... a r ~ .. the 13th dl or incubation. If. ::lont. of thecues , relponsea ..lUlled a tn-phasic pattern (pas! t1 ve-neRaUve-posit ive) , the negathe eoaponent b e l ~ of larger UIJll1tude. In l ~ d - . v emb!'Joa, U1P11 tude ot the nep,atl "Ie CClIImOnentvas tOWld to vary vithin to 00 \.IV, in 19-1lay Hlbryo!l - ", i t l : in 50-100 being over 100 In 20-day e M b ~ o s . Both amnl1~ u d e and shape ot the eYOked potentials d 1 a p l ~ e u considerableindividua1 variations, they alftO tended to verI in the StlllM! embryoduring an experiaent. The r e . ~ n . e to rhythmical flashes shoved180rhytbmicity a t frequencies uJ) to 5 t ' luhes per sec 1n 1 ~ 1 9 en'bJ7oa, and l1Jl to 10 nuh e s per sec In 2o-da,y embryos. On . ~ r : a -ulat ion \I1t.h pairod f luhea , the ablolute retract ive period YQtound to lut 60 to 200 lUec In 18-2o-da,y eIlIbr:--oa. It haa thusbeen shown,that in th e chick eabr,ro tunctional develOJlllent ortbe v':.ual analyzer attains a fair ly high level in the l a s t rrenatal d ~ 8 .

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    ~ ~ c e n t . l y th e attention of' nhY3iolor;i3ts has bep.n ,lravn to a stu \ J . D . : : ' r_"le.l:.::cr uron !l.:lequate atir.lulat1on ".:.1.(> r ~ " ; i lle. "T nurinp. pl{'c

    t r l c ~ exci tati(;>n of tile optic nerve. ha.ve fI.1.wnys seemec1 pr01"klsinc for t:Xplar.ntior. of t ~ , e 1:E'culiarities or

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    lap o f the paaowat1mulator vu CIlI I lVIlY ~ r ( l m th

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    d A ' t "V, 11-" secB I~ - - + - _ - - - I ~ - - ~ - - -

    , $

    IOn no. III 0 ' ,_ , 11 100 to . , 1 f I

    0'' ' ' ' ' , pp "

    , "" ' , . ' 4ft . ." 1'$ , t I _d t ' . . . . .,"

    au to b-- .. .t.,tIlt

    F l ~ e 1. P.loelectr ic react ions of the of 1 ~ ~ 8 : ' old chick eMbr,yo to photost1Jllulation F r e o u e n ~ of stimulr.tion(numher of f l lU lh( '1 I ~ r second): A and L - ' 0.2 indirect continuation of A);C - 1; - 2; - 3; F - r, - 10; l: - ! l a ind f lesheswith interval of 150, 300and 100 microseconds. 1-

    r e c o r d i n ~ of electr ical nct iv i ty ot ' i , ~ . 1 - s t i m u l u . ~ [The upward movement or thepen c o r r e 3 ~ o n d a to n e ~ a t i vity ot actlTe e l ~ c t r o d e ] .

    responses began to appear again, e o n s 1 s t i ~ 1!ULinly of nep:ative eomr-()nentsof EP (Figure 1 . A-F).In res!'onse to tbe use or 8. fl iekef ' lng 111")lt vi th a frequency or Inflaz:.es a second or more. a certain exaltat10n or compression of bacltr;roun,l

    act iv i ty vas observed which vas not aeeom!,af.ied by 13orh:rthM.1c res!,onsec.Orten on-type and. less rare ly . orr-type relponses arORe in resnonse to nwitchIna on the st1aula't1oa.In 1 9 - d ~ r old elllbr;yoa the LP were d i s t 1 ~ s h c d by a l a r ~ ~ 8Illplitu(]eand a remanent mani fe s t a tion Indi vi dual va.riab i l i ty . a l thoup,h observed. vaanot as 3ignU'icant &8 1n lE-cia)- olto cIIlbry08. The a v e r a ~ e ampli tude of thenegative c o m ~ n e n t vas 50-100 aierovolts (up 20n ~ i e r o v o l t s in isolated

    t e l t ~ ) ; duration - about 50 Ill1croseconds. The shape of the Er and the ro lar l tyof i t s cex:lponenta did not dif fer trom thOle in the If.-/le.y old ~ b 1 ' ) ' o s ( ~ i C ' ; u r e 2. A).

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    , .a t -+ + 4 4 -A ,1.:, H e~ ~ ~ .. , .1 .. ~ ~ .." I _.s , 1 , I I 00

    v~ _ J M W J M ,I II

    F i g u r ~ 2. Dioelectrlc react ion of ttBT of 19-da.y oldchick mcryo to !>hotostilLulat ion.Frequency of stimulation(number of nashes nersecond): A 0.2i P. - 1.C - 2. 0 - 3; E 5. r -10 . n - paired flashes vithintervals of 200 and 100microseconds. Other I ~ bol l sane as 1n ~ i ~ r e 1.

    ,,"w'.bI.t 1'1 . . . . . 14 ~.. ----'l

    o , :.....-----+------.....-Continuous stimulation o t th e eye by l ight f l a s h ~ s vith a frequeneyor 0.1 pe r second. continued ~ C " r 20-30 Ildnutel. did not evoke a noticeable re4uctlon of EP eapl1tude. Hovever. EP 8IIlP11tude did not renain str ict lyc:onltant duriD!J th e ent i re period ot stiJllU1ation: i t increased and decreasedperiodic:al1. altbough to a very inaip;n1ticant Cr second vas not reproduced ( r i : ~ e 2. r-). Incertain cues . the rhenomenon of rh:Ithm t.ransformat-ion oec:urred a t t h l ~ S tlJllwation frequency. A rhythm or 5 nashes ~ r seeond vas \Wed :s nstead of 10 .

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    C ~ '_ _ _A ~ F - 't I-I t.::., I te

    B . ..... . ~ . ~ . ~ l ..... . _ ~ ...........- . " - I ~ ' ' ' ' ' - - - ::-i"',;:'(' '. . . .:Ol.lcC'tri:: '{',.act,ion of ?o-d,v 01 r1 ('\,1 etir,!":rc to nhot03t1Jnulnt1vn._ ~ - - ~ , . J , . t l

    r r e q u c n c ~ or c t l ~ u l n t i o nc , " " . ~ I I-+'-+I-+ l _ ( n u r . ~ e r or r l n 8 h ~ s ~ r s e ~ o n d ) : A - 0.2; - 1;C - 1; D - ?; r. _ v _ f ---IJ,JJ,J;J),J,J,lJ,J,I,J).ItJ)J,J,J,J,IJ)J,J)AJ, lr.. - Other ~ J ~ t c l sI) sane in F i ~ r e 1., _---lllo6l.&.I... l l o l l ~ l ... ~ I I o 4 ....... +,,11104' . . .....---:-... l+I+' " ~ I ~ l " ' I " " .... I... 1+,.- ' ~ . b . ' . m " " P I I I I . ~ I I I L ' I i I I__ "'''IllI' _

    , . 1 ' l ' I " " I I l l I " l l I l 1 l l l W l l I l l l l ~ l l I I I M I l I I l ' I I I lllll'

    t ~ " W M l " " . I I I ' l I l ' " ' _ - ' ' ' ""',J".I. ..W' l_....... . . .p , __ -4_. ' ' ' '._''- f - - - ~ - - - - - " - - -G 1__ -:-- -..----

    Figure 3 shows iili'r' respon to l 1 ~ h t st imuli or var,,-ing treQuenc:. lr ,ar. embryo ..t te r 20 ~ . ot incubation. A characterist. ic of F.P eTOke4 b7 asingle Tlhoto.t1mulus IS .. re.1d\&&l cha.rge, vbieh consists of a R1"OUl'l or Slowvaves that tollow EP (ngu.re 3. A). In I lIOBt cues t.he az:splitude ot thenegative EP cCDJ'lOnent exceeded 100 microvolts. In stimulation or the eye bya tlickering 118ht vtth a frequency or 3 flashes per second the :mT reactionuau&ll:r eoaa1sted of tY1'lIc:&l PI. a t each f l u h . in contrast to the '.tnT re-action or 10- and 19-day emb17oB, which v u exnressed prll'l.!U"lly by slov \ I n . v e ~ (Figure 3. D). In response to photostimulatlon vi tl l a freet-eney of 10 flas:J.s per lecond a rbTthla or 10 per aecond vas recorded in the :.mT. but i: : themaJorl t;y ot instanc the index or . . . .iJ!li1ation vas no h i ~ h e r than O. St1z:1ulaUon at a frequency ot 15 per second evoked the on-reaction phenanenon(Ie.. otten. the off-reaction) and did not lead to any ch&nfl':e of back4round

    r ~ h I I l (figure 3, G).

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    In etlaulatlOil by pa.1re4 l1ght f luhes . aeJ)a1"&ted troIIl one another'by in tenals ot 3 ueollc!s to 50 aicl'Oleeonds. the relponse to th e aeeoadf l uh eU.appeared a t a certala cr lt i ea l interval between st.imuli. v h o ~ e v a l ~ e cbanged vi thin l ~ t . o t 60 to 200 aicroseconds d e ~ n d i n ~ on a numberor ul t te rent eoacl1t101l8 ( r 1 ~ 1. Hi 2. 0) . Anal7sh of the data o b ~ i n e d . ab0ve4 that. lD41'f'1clual Yariat10u of this f t lue are IIIOre a1gn1t1eant in coapartlon to IP t1uetuat iou ot tbe re tractive !'eriod.

    DISCUSSIONStudT or the bioelectr ic reactionl of 41fterent aeetlonl ot th eYiaual ~ . r to 11lbt s tb lu l i 1n chick BbryM v u conducted b7 Peters

    e t ale (11). Howftr . In th is york he U8ed an 1aproper lead (ld4bra1n - cerebellum) th e e.bryoa " . . . DOt l . -obl l ized. the electrodes ve1"e lneened toa 4epth ot 3 _. which cornsponded to different MBT layera in embryo. atYV!oua per10da ot 4eYeJ.oJ-At.. Th e r e t o r e . " attempted lI D InveaUatlonvb1eh vould Dot. 1llclu4e th..e M'thodologleal lrwlequaeiea. In calP&rtng thedata traa our experiment. v i th those of Pe ten 1t turned out tha t . alongwith certalD cUfferenee there V& l a c:orrelJlODdence ot the b u ic resul ts .POl' l na tuee . deteJ"ll1na'-1011 or the t1JDe of deft lopeent or the _bryo . whenMB'1' relpanae. to lipt . t1aul i appeared (18 dq a o t 1neubation). TarlabUi tyof VP llADit.. ~ t l O D . preT&leDCe ot tbe n e ~ t 1 f t coeponent and eatablish!lent ot u upper l1a1t for repl"Odueed frequenciea of l igh t flickering (10/.econd) coatlna Pe ten t flncUllS.

    Ulltortunate17. e x c ~ t tor Peters ' work. we did not find any I tudiesot the proee.. or the tuDetlOD&1. maturation ot the visual analyzer in towl.Hovever. a certaln congruence 1n th e atructure ot fowl and the neocortex or' ermlls (12). ~ 1 1 tn e pre.ence of aix sharply pronounced 1. ,er l . gives UIa b u i s to r dravlna certain puoallela to reault . of ont.ogenetic It.udies ofthe r t a u l area ot the cortex In MIII'.al.s. A n ~ s i s ot the l i terature showsthat in ua1llaU 'bom blin4. ( rabbit . cat) the Ell in the v h u a l zone of th ecortex appears duriDS the t i r s t 4Al.Ys ot postnatal 11te. Thus. Harty [5] 1eacr1bea th . appearance or the EP In relponse to a1l1Rle l 1 ~ t n u h e a on the2-5th da.Y o t poetDatal deftlOlll8Dt in kit tens uad a t the en d of the tintft_ 1n rabbit . . ID th e work of faneI ' [8] the EP ill reaponae to 11Sht t l u h ea v.. reeol'4ed In kit tena an d rabbita CD the 3-5th" _ ar ter bi r tb . Ell1D1SOD an d ~ i l c o t t obaenell EP appearance 1n relpoue to single pbotoatlmuli Inkit tena trca the eighth dIlf ot po8tnatal l i te . In nevbol"ll children the r;preaponae to l ight f l u b e . la recorded during the f .irst houri or Ur e . Ina4ditiOil. EP can be recorded nen In J'l't!IIl&ture children v e l ~ i n g 1300 gr8r. lBon the second _ ar ter b i r th [10]. Thi. tact c:oatinlUl th e hi,,h d e ~ r e e oftuDctional aatW'l ty of the r t . ua l analyzer 1n newborn children in eClllJ)&l'l.on with &Dial . bam blind.

    The frequency r-ange vlthln vhich i80rbYthmic r e l l ! l ~ n l l e s or th e visualcenters are recorded i s .000000t videI' In chick .wryOI than in the represent . a t i f t an1a&la bieh were .tucUed. The capaclt7 to reT'roduce a rhythm of

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    l ight tlickeJ"1n8 of ~ per second appeus 1n cata and rabbits onlr on th e22-30th ~ a of postnatal deft low-ent [13], while in newborn children th eupper l imit 18 a tnqUIDCJ of 3-6 per second [9 ] or 3-5 per second [10].Our data confirm the e v ~ c l e T e l o ~ D t ot the visual t u n c t ~ o n inchick embl708, u d are 1n direct correlation vith llI01"!lhologie&l . ~ _ l .. will have sbovD a quite high degree ot ..tur i ty ot the studied sections of th evisual 8Ilalyzer in chick eaabry08 in th e t inal d.,ys of prenatal development[llt-18] Blocb-.1cal studies by Linc1eman [19] shov that 'by the 19tb d ~ ofincubaUDD the acetylcholine COI1te11t .. vell .. th e cholinesterase and a l-

    ka l i. . " . . . . .~ . a c U . i ~ r the ret lna Incre..e sbur ly . Thus, 1n chick_bl'7Os, b7 the IS-19th ~ of incubation, a definite cr i t ical period in def t l ~ n t begina, a period c h a l " & c ~ r 1 z e d by abrupt biocheJ:11cal, structural .uel tlmctloaal cbarges, vh1ch sbow an 1ncreulag degree ot _tun ty of th ensua l analyzer.Certain researchen [5 , 6-10] noticed a rapid adyance or fatigue ofspee1!1c reaponses of th e "f1sual an d auditory are.. of the cortex during acClBpvatively Slow rbtbm ot st iaulat lon (0.3-1 p .r second) 1n newborn children u well as in rabbi ta and eats in the tint dqa ot postnatal l i re .The results of our tests are s ~ t contradictory v i th respect to th e ci teddata. The Yisual centers in the chick eI!lbryo are capable or responding toeach l ight f l uh at a st1aulaUon frequency or 0.1 to 1 per second withcontinuous .t1lllulation in the cour ot several lo-a1nute J'l4!rioda withoutsignificant reduction of EP .-p11tude.The considerable individual Yariation in the size and shape ot th eEF, noticed b7 u.s 1n tb e lut dIt. of embl')"Onic deTelopment, is in a,o;reenentv i th tb e data obtained by Farber [9], Hrbek [10) u d others. However, onth e backp'ound or tb i . Yarlabll1t7 one can detect a tendency ot EP 1IIIIl'11-t.ude to increase in the cOUl' le or dnelopDeDt. Further, in 'j)asl1ng troe th e18th to 19th dtti1 at de-felOJMnt there i s alia an expansion or the rang\! orpaired frequencies. 'rhea. dat.a al .o I n d 1 c : a ~ t ha t b7 th e 19th ~ o t embl'Jocic: d 4 m ! l ~ t the Yin&! anal7ser a t th e chick "ryo 11 well-developedtuDet10na1l.We could find only oae work (Farber [8]) vbich t reated the cycle ors c i tabll1 ty or response. of the r i .ua l area ot th e cortex 1n eat . and rabbits . In 7 - U - _ old ulJ11&18 the absolute retract-lYe ~ r 1 o d v u 120-200

    micro.econds. In our tests th e value or the absolute refractive reriod fluctuated trca 60 to 200 IIl1crosecoada, depending on various caDdi tiona. Theseare entirely c

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    C

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