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~ESKOSLOVENSKA AKADEMIE V$D
ACADEMIA SCIENTIARUM BOHEMOSLOVENICA
FOLIA
BIOLOGICA
TOMUS II
FASCICULUS 1
-M~LLIl D~Illf iYf
(c~A~~~
Fol. biol. (Praha) Tom. 2 - Fasc. 1 Praha 20. 3. 1956
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FOLIA BIOLOGICA (PRAHA)
ll~l~~asra~~rapoa~rne raaaa.r~ue alci~~~~n.2nrs Ceskoslovenskcc biologde w C'esb;oslove~~,Icri
7ni1?robzologie
h e ;(a IS i~ l[ O lI ]I a fI IC O JI :I C P L[ ft
Ai~:I;~entn~ti lI. Ala~u~~~ (real{tu,Iii pe,SaleTOp), B. Bpnrlaclci2il, M. I'aniclt, 4aL-Itopl~. T1C:~I[
(U, hP.I19113L~ alSa~[e MLIK (]. IIIIOIt(', L~~ IO. 1~Ia1[yha~ 7KaJ~0 MIiIS ~.. IIj)aT~ Ii. I'O(711 ~ISllll ~CC IiII, I1C~[.
ho~l~erl-IU.), JI. rIeplu,Ii~, fl. IIITepI~.llb.
1[epei;o~~a Ita hyccl.IIGI srzr~n;: ;[oiS. i[-p IIIIIpoI;a, IYa anrsr~I`iclelaK xei,Ile: f[-p Pla;secoi;a, IrI ue-
~cllxuil xaiah: a-p ~ailrzb
Iia;~aeTCSt IUIIO~IOrxvechu~I I,IIICTI-ITyroM rlexoc~rol3al~I~oilAltauemnla IIay~I~ x Ila,~aTe~iLr.Ti+e'1CAIf.
13iato~ZwT 6 IIa.; ~i roiS. IIo;41IicIrIH Ihxa xa 1 ro,~ K'u' GU.-, I~cIIa o;~uoro IroMeha I{~u, 1n.-.
~l;[I CPC he)~.1 KI~[4IL' Ii710.~01'Ii'IP,CItH ~i If IICTnTyT tI (;AII, IIa l~IS1I9111I PP11 2~ II})al'a ~1I~. ~U Ka:SLI:
11~,1~nsr, CMe~ncx 3u, llpara II, `Ie~oc~IOi;a~~nsl.
FOLIA BIOLOGICA (PRAHA)
In.icrn~r.tional I~;~diti-on of tjae Journals G~eskoslovenslca biologde a~~~l ~'eskosloverasl,?ii
mikrobiologie
Aca.domiciwn I, M~lek(Ckiief P:ditor), I,. horny, M. Hc~sek, Curresponding Member of the Czechoslov~rli
Academy of 5
with turkey (Hayek, Hraba,
Majer, not published)
I
Benesova, Hlavackova 1955)
Hen - intra-embryonal injec-
--
tion of duck blood (Hraba,
j
Duck -embryonal parabiosis
Ha~kova, Lengerova, Vojti~-
i
with hen (Ha?ek, Hraba, Be-
kova 1956)
I
nesova, Hlava~kova 1955,
Hen - intra-embryonal injec-
Brent 1955)
tion of guinea fowl blood
Duck - intra-embryonal injec-
(Hraba, Ha~kova, Lengerova,
tion of goose blood
Vojtiskova 1956)
__ _
Hen - intra-embryonal injec-
~
Turkey - intra-embryonal in-
tion of turkey blood (Hraba,
jectionofhenblood(Simonsen
Haskova, Lengerova, Vojti~-
1955)
kova 1956)
~
Pheasant -embryonal pars-
Duck - intra-embryonal in-
'~
biosis with hen (Hayek 1954)
jection of hen blood (Ha~kova,
Pokorna, not published)
Goose-intra-embryonal in-
i
jection of duck blood
Guinea fowl - intra-embryon-
al injection of hen blood
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which complete suppression of antibody formation or taking of a skin transplant
may be found. Even this approximation, however, has its q~xantitative aspect. In
inter-special relationships, complete suppression of the immunity response is not
found in most cases, either with transplants or with immunisation. Only in hens,
embryonal parabionts with turkey, however, a marked decrease in reactivity to
turkey blood cells was found (Hraba 1956). Partial approximation is indicated by
the survival of hen skin heterotransplants in a pheasant (21 days, Hasek 1954) and
in a duck (40 days, Brent 1955) in which approximation had been produced and also
by the finding in some cases of a decrease in antibody formation, which are likewise
given in tab. 4. Finally, in a series of further combinations, no approximation effect
was found.
2. Special differences exist in the effectiveness of the production of immunological
approximation. For example, in ducks, embryonal parabionts with hens, an obvious
decrease in antibody formation was found, but in hen embryonal parabionts with
ducks this was not found (Hayek, Hraba, Bene~ova, HlavaL~kova 1955). In the same
way, in the present work, a marked decrease in antibody formation was found in
ducks after intra-embryonal injections of goose blood, but in geese injected with
duck blood there was no decrease.
3. From the aspect of producing approximation, embryonal parabionis is far more
effective than a single intra-embryonal injection. For example, there is a significant
difference between the degree of approximation in hen parabionts with turkey
(Hayek, Hraba, Bene~ova, Hlava~kov~ 1955, Hraba 1956) and hens injected intra-
embryonally with turkey blood (Simonsen 1955, Hraba, Ha~kova, Lengerova,
Vojtf~kova 1956).
4. It is probable that in a number of cases approximation was not found because
adaptation to the foreign antigen was so weak that it could no longer be demon-
strated in post-embryogenesis. Skin transplants are used for testing 14-30-day-old
birds, while immunisation is carried out in birds which are at least six weeks old and
frequently even older. Finally, differences in the immunisation method may also be
of importance.
Summary
1. A decrease in the formation of immune agglutinins against goose blood cells
was found in ducks in which approximation had been produced by the intra-embryonal
injection of the same antigen. Precipitins were not found either in the experimental
birds or in the controls.
2. In geese which had been injected intea-embryonally with duck blood and inguinea-'
fowl injected intra-embryonally with hen blood, no decrease was found in the forma-
tion of immune agglutinins.
B i 1 1 i n g h a m, R. E., Brent, L., Medawar, P. B.: Actively Acquired Tolerance
of Foreign Cells. Nature 172:602, 1953.
Brent, L.: Some Consequences of Parabiosis in Chick Embryos. Summaries of Int. Embryol.
Conference, Brussels 1955.
H as a k, M.: Vegetativnf hybridisace zivoLichu spojenfm ~krevnfch obLhu v embryonalnim
v$~voji. Ls. biologie 2:265, 1953.
H a a e k, M : Projevy vegetativnfho sblizovanf v adspptaci vyssiho zivoLicha na cizoroda
antigeny. ~s, biologie 3:327, 1954. T a ui a x, M.: IIpoRBnaxHR BereTaTHSHOro c6~HxceHHR
B aAaHTauHH B~Cmxx xtHBOTHhIX Ha Yy3KeppO~[HSie aHTNreHEd, tICJI. BHOnorxx 3:344, 1954.
Hasek,M.,Hraba,T.,Benesoya,H.aHlavaLkova,H.: Imunologick~vztahy
u embryonalnfch parabiont>~ mezi kachnou a slepici II. Gs, biologie 4:135, 7.955.
51
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H r a b a, T.: Imunologicke chovani embryonalnich parabiontu mezi krutou a slepici. I. L`s.
biologie, 5:89, 1956.
Hraba, T., Haskova, V., I~engerova, A. and Vojtiskova, M.: The Influ-
ence of Intra-embryonal Injections of Foreign Blood on the Formation of Antibodies
I. Observation of Reactivity in Hens. Fol, biol. (Praha) 2:00, 1956.
Simonsen, M.: Artificial Production of Immunological Tolerance. Nature 175:763, 1955.
BJIIIFIHYIe BHyTpYI3c`1pOAbILTIOBbIX BH~JbICI{I~IBaHIII~I LIy?ISepOJ~HOYI IfpOBI~I Hcl
OrOpa80BaHHe aHTHTBJi
II. I~Icc~IeRoBaxxH peaxTxsxocTx y yTOx, ryceH H uecapol~
M. I'AIIIEK
Pearo.,~ce
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y yTOx, ryceH H >~ecapolc. MbI snpblcxIIBa~IH BxyTpxsexxo rycxxylo xpoBb yTxxblM
3apOJjbIIIIaM, yTHHyIO xpOBb ryCIIHbIM 3apOj(bIIIIaM YI xypHxylo xpoBb aapoAbllIIaM
uecapxx. B BoapacTe 2-3 MecHt~eB MbI onpe?~e~Ixalg y peuHHIIexTOB o6paaosaxIIe
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aM6pxoxa~Ibxo. MbI ycTaxosxalx c~axT cxIIxcexHH cnoco6xocrx x o6paaoBaHIIIo
IIMMyHHbIX arrJIIOTHHIIHOB IIpOTHB PyCHHbIx 3pHTpOI~IITOB y yTOx, xoTOpble c6~IH-
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6bIJIII y IIepBOI~ cepxl3 c6~Igxcasmgxcx c rycHMi3 yTOx: 1, 2, - y xoxTpo~Ibxblx: 16,
128, 32; Bo BTOpoI~ cepIIII y c6~IIIx~aBnTHxcsl yTOIC: 4, 4, 8, 4, - y xoxTpo~Ibxblx: 128,
64, 64. IIoc~Ie peIIMnlyxIIaaijIIII THTpbI y c6~III~xaBHlHxcx yTOx 6bIJix 8, 8, y xoxTpo~Ib-
xblX: 2048, 1024. IIpeL[HIIHTYIHbI IIe rObI7II3 HaLIJ~eHbI xII y C~JIHSxaBIIIHXCH, HH y KOH-
Tp07IbxbIX yTOx. y ryCOII, ICOTOpbIM B 3apOIjbIIIIeBOM COCTOHHIIII BnpbICxIIBaJIaCb
yTgxaH xpoBb, II y izecapox, IIo~IyxaBHlHx xypllxylo xpoBb, xe ~bIJIO oTMexexo
cHIIxcexge o6paaollaxxx IIMMyxxblx arr~IIOTHHIIxoB.
OxaablBaeTCH, uTO x y ApyrIIx BIIAos, xpoMe xyp, uacTO xenoaMO;xxo 6blsaeT
oTMeTIITb IIMNryxo~IOrII~ecxoe c6~Ixx;eHl3e c x~IeTxaMH Roxopa Apyroro BIIAa IIoc~Ie
BxyzpgBexxblx BIIpbICxHBaHIIx aTIIx axTIIrexos aapoRblHly. BecbMa BblcoxoH oxaabl-
BaeTCR cTexexb c6alIIxceHxsl y yTOx, xoTOpbIM Bnpblcxxsa~Iacb rycIIxaH xpoBb.
B Me?xBIIj[OBbIX OTHOIIIeHIIxX, - B OT7IH=IHe OT I330CepOJIOPI3=IeCxIIX HCCJIej(OB8HIII3
xMMyxo~IOrIIgecxoro co~IIIxcexHH y xyp, - B 6o~IbIIIIIxcTBe calyuaeB xa6~IloAaeTCx
xanIIxHe ecTecTBexxblx reTepoarralloTIIxxxoB. B xarIIeK cepHll oIIbITOB xII y oAxoro
rycx IIJIH >~ecapxII He 6bI7I0 oTMeuexo IIpgcyTCTBge ecTecTBexxblx reTepoarr~IloTII-
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]ICeHIIFI. Ha OCHOBBHIIH IIpexcHHX HaIIIIIX H IIpHBOjjIIMbIX 3]Zecb peGyJIbTaTOB OIIbITOB
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po~IOrHxxoro Roxopa (Ta6. 4):
1. Taxcoxo.ntiu~recKOe eu8oeoe po8cmao aorcopa u peilunuewma. I~IMMyxo~IOrguecxoe
c6~Igxcexge c IIxRHBgAyanbxo Ag~ic~epeHTpbIMII xaeTxaMH Toro xce caMOro sII~a
HbI3bIBaeT cocTOHHIIe, IIpH xoTOpoM MoxcHO Ha6~IIOUaTb Ho~IHOe yrxeTexlle o6paao-
saHHH aHTHTe~I H IIpIIxcHB~IeHge xoxcHOro roMOTpaxcII~IaxzaTa. ORHaxo xo~Ig~eczsex-
Hafi cTOpoxa aTOro c6~IHxceHgH He6espaa~IgxHa. - B McHCBPIJ1;oBbix oTHOmeHnxx
B 6o~IbiIIHHCTSe c~Iy~aeB HH HpA TpaxcHnaHTai;Hg, xH Hpg xMMyxgaai~HH He xa6nlo-
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RaezcR Honaoe yr$eTeHHe gMMy$HOg peaxugH. Tonbxo y xyp-aM6pgoaanbHbix
Hapa6noxTOB c gHRioHixoi~ 6bino oTMeueao aaa~HTenbxoe cxHxceHHe peaxTgBHOCTH
HO OTHOHIeHgio K HpOBHHraIM Te7IbI~aM HHRioHiKg (I'pa6a 1956). 0 ~acTHLIHOM c6nx-
xceHgg CBI3jjeTeJIbCTByeT IIepexcHBaHHe KypgHbIX KOH{Hbix reTepOTpaHCII7IaHTaTOB
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Brent 1955), a Taxxce HexoTOpbte AaHHbie oTHOCgTenbao cHHxceHxR o6paaosaHHR
axTHTen, HpxBOAgMbte B Tab. 4. Haxogeu, B pHRe AanbHei~Hixx KoM6gHaugA ac~c~eKT
c6nxxceHHH He Ha6nioRaxcR.
2. I~I.,~ceromcs~ euaoebre paar~uuuR a sg5y6exmuatcocmu ucrcyccmeerctco ebasbreae.~ozo
u.~.roc~teorroau~recxoao c6~ruarcexuR. Hanp., y yzox-3M6pHOHa7IbxbIY Hapa6ROHTOB
c KypaMg Ha6nioAanocb saMeTHOe cHgaxeHHe cnoco6xocTH K o6paaosaHHi0 aHTI3Ten,
Ho y Kyp-aM6pIIOaanbxbix Hapa6goHTOB c yTxaMH oxo HaHAeao He 6bino (I'a~ex,
I'pa6a, Beae~osa, I'nasauxosa 1955). PaBHbIM o6paaoM B sTOi~ pa6oTe 6bino oTMe-
xeao aHaKgTenbxoe cHgxceHHe o6paaosaHxH aHTxTen y yTOx B peayxbTaTe BHyTpH-
aapoRbimeBbix BHpbrcKxBaHHK rycHHOH xpoBH, Ho oxo He 6bino oTMexeao y rycei~
HOJj ReHCTBHOM yT$HOI3 KpOBg.
3. C TOLIKH 3peHnH C6ngxteHgR rOpa3J;0 607iee 3(~(~eKTgBHbIM, YeM OJ~HOKpaTHaA
BHyTpyI3apOj[bIIIIeBaFi I3H'bexIZI3H, IIpeJZCT8B7IFIeTCH 3M6pI3OHa7IbHbII3 IIapa6gO3. Cy-
II~eCTByeT, HaIIp., 3aM0THaH pa3HgI[a MexcRy CTeIIeabIO CCJIHHfeHI3H y Kyp-IIapa6HOH-
ToB c gHRioHixoir (I'a~ex, I'pa6a, Beae~osa, I'nasa*~xosa 1955) g y Kyp, xoTOpbiM
xposb gHRioHixH BBORHaacb HHTpaaM6pgoHanbHO (Simonsen 1955, I'pa6a, I'a~xosa,
JIeareposa, BOI3THHIHOBa 1956).
4. B pRAe cnyuaes, BeposTHO, c6ngxceHHe He 6bino aaMe~eao, Tax xax aAaHTa~HH
Ha uyxcepoAHbii~ aaTHreH 6bina HacTOnbxo cna6og, PTO B HoczaM6pgoreHese ee yxte
H0BO3MOH{HO 6bIJI0 OTM0TI3Tb: TOCTbi TpaHCHJIaHTaI;HH HpOH3BOJjHTCH B BO3paCTe
14-30 Z[HeI3, a HMMyHx3auHR - B BO3paCTe 6 HeRenb H C)OJIbIIIe. HaKOHeR, 3j[eCb
MOryT HrpaTb pOJIb H pa3JIHiIHH HpgeMOB HMMyHH3aI~gH.
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FOLIA BIOLOGICA
Suppression of the Formation of Immune Agglutinins in Chicks in which
Approximation was Produced on the First Day after Hatching
by NTeans of Injections of Erythrocytes and Leucocytes
M. HA~`EK, T. HRABA and M. ESSLOV[~
Institute of Biology, Czechoslovak Academy of Science,
.Department of Experimental Biology and Genetics, Praha
It has been shown by us that approximation to a skin transplant cannot be
produced by the injection of a suspension of washed blood cells in which there are no
white cells present (Hayek, Lengerova, Maternova 1955). Immunological approxima-
tion was, however, produced experimentally by the temporary joining of the circula-
tions of chick embryos (embryonal parabiosis), which resulted in inability in post-
foetal life to form immune agglutinins against the blood cells of the partner
(Hayek 1953) and also by injections of blood administered to the embryo, in which
resultant tolerance to skin transplants was found (Billingham, Brent, Medawar 1953,
1955). Tolerance to skin transplants was also found in chick embryonal parabionts
(Hayek 1954). These results show the effectiveness of leucocytes in the development
of immunological approximation as manifested in the production of tolerance to
skin transplants.
The problem under consideration here involves the decision as to whether erythro-
cytes freed from leucocytes can lead to inability to form immune agglutinins in
post-foetal life. The fact that erythrocytes alone are not capable of producing
tolerance to skin transplants may be due to the fact that only leucocytes are capable
of producing immunological approximation or that only leucocytes have completely
corresponding, common antigens with the cells of the skin. Since immunological
tolerance can be produced in a large percentage of newly hatched chicks by the
injection of foreign cells (Hayek 1955), this form of inoculation was also used in the
present experiments.
The Production o/ Immunological Approximation
Newly hatched chicks (none of them older than 20 hours) were injected either intravenously (35 chicks)
or subcutaneously (40 chicks) with a suspension of blood cells in amounts of 0.5-1.0 ml. The intravenous
injections were made into the wing vein, the subcutaneous injections through the thigh musculature,
below the skin in the area of the abdomen. Aa donors, three cocks were used (Nos. 71, 85 and 89). The
suspension of erythrocytes from all cocks was prepared in the same way while the preparation of leuco-
cytes was modified in every cock.
Preparation of Erythrocytes
The citrated blood was centrifuged in small glass tubes, which had been sealed by melting, for
15 minutes at 2,000 revolutions/minute. After centrifuging the tubes were cut off about half a centi-
meter from the level of the sediment and in this way the leucocyte layer was removed together with
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part of the erythrocytes. The remaining erythrocytes were pipetted into centrifuge tubes, diluted with
the same volume of Tyrode solution and re-centrifuged for 15 minutes at 2,000 revs.~min. This process
was repeated once again. The washed erythrocytes were then suspended in Tyrode solution in the
ratio of 1 : 1.
Prepar?txon of Leucocytea
20 ml. of blood were collected from cocks Nos. 89 and 71 (4 ml. sodium citrate to 10 ml. Mood). The
leucocytes were isolated from the blood of cock No. 89 by centrifuging. The citrated blood was centrifuged
in sealed glass tubes for five minutes at 1,b00 revs.~min. After centrifuging, the leucocytes, which
formed a white layer above the erythrocytes, were carefully pipetted off and re-suspended in Tyrode
:solution inaconcentrationof37,960,000/ml. Erythrocytes were presentintheproportionof4,200,000/ml.
In the case of cock No. 71, the leucocytes were isolated by the agglutination method (Lee and Osgood
1949).
Extract of phytohaemagglutinin was added to the citrated blood (0.2 ml. of phytohaemagglutinin
solution~l ml. blood, titre 1:10). The supernatant fluid was pipetted into glass tubes which were sealed
and centrifuged for 2-5 minutes at 1,000-1,500 revs.~min. After centrifuging, the supernatant fluid
containing the leucocytes was pipetted off from the sedimented residue of erythrocytes into new glass
Tab. 1. Reactivity in First Series of Birds in which Approximation was Produced.
The first immunisation was carried out at the age of eight weeks in doses of 0.5 ml. (the last two
doses were reduced to 0.2 - 0.4 ml. because of insufficient donor blood) administered on alternate
days. Re-immunisation was carried out at 15 weeks with three doses of 1 ml. administered on alter-
nate days.
Production of Approximation by Leueocytea
No. of birds and reactions
1
Donor
Method
No. of birds
After 1st immure.
After re-immure.
Positive
Negative
Positive
Negative
89
71
i. v.
s. c.
i. v.
s. c.
5
5
6
6
5
5*
5
5
0
0
1**
I 1
1
0
Total
22
20
2
1
0
I
89
i. v.
I
5
2
I
3
I
3
0
s. c.
5
5
0
71
i. v.
3
3
0
s. c.
3
2
1
0
1***
Total
16
12
4
3
1
N. B.: * Zn one case positivity was not absolutely clear. After re-immunisation the reaction was
strongly positive.
** On remimmunisation, this bird was immunised with the blood of a different bird from the
original donor and formed agglutinins.
*** This bird did not form agglutinins even after the administration of a provocative dose at
18 weeks (0.5 ml. ).
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Tab. 2. Reactivity in Second Series of Birds in which Approximation was Produced
The donor in this series was cock No. 85. The first immunisation was carried out at eight weeks,
with five doses of 0.4 ml, on alternate days. Re-immunisation was carried out at 11 weeks with three
doses of 2 ml. on alternate days.
No. of birds and reactions
- - -
No. of
birds
After lat immun.
After re-immun.
Positive
Negative
Positive Negative
Approximation prod. with injection
~
of erythrocytes
12
11*
1 1 ~ 0
Approximation prod. with injection
of leucocytes
3
2
1 ~ 1 ~ 0
~ I
Controls 9
8
0 1**
0
* In one case positivity was not absolutely clear (after re-immunisation strong positivity).
** This control bird was immunised only at the time of re-immunisation.
tubes, diluted with the same of Tyrode solution and again centrifuged for five minutes at 1,500 revs./min.
The sedimented leucocytes were pipetted off and re-suspended in Tyrode solution in a concentration
36,900,000~ml.; erythrocytes were present in the proportion of 3,800,000~m1.
A mixture of washed erythrocytes and leucocytes, in which the leucocytes were concentrated in the
proportion of approx. 1 : 5, was injected from the third donor, cock No. 85.
Testing of Approximation
Approximation was tested by immunising or re-immunising the chicks under observation, between
eight and 18 weeks, with citrated blood (citrate 1:10) from the original donor. The immunisation
methods and doses are given in the tables. The collection of blood from the immunised birds was carried
out five to seven days after the last immunisation dose. Agglutination was carried out by the test-tube
method, one drop of aonce-washed suspension of blood cells in physiological saline being added to two
drops of serum or plasma. The results were read off after 15 and 60 minutes. The test-tubes were left at
room temperature and were shaken before being read off. In the experimental series given in tab, 3
the titre was ascertained. The absence of agglutination was controlled microscopically.
The results are given in tabs. i-3. The individuals which acquired approximation
and did not form agglutinins constituted 15%. Out of 30 birds which were given an
injection of leucocytes on the first day, five did not form agglutinins. Of 36
birds which were given an injection of pure erythrocytes on the first day, six did
not form agglutinins. Previous results were also confirmed in that there was no
difference in the production of approximation after the intravenous and subcutaneous
injection of the antigen in one-day-old chicks. Neither do the results show any
striking relationship to the amount of injected material within the limits used. The
re-immunisation results show that non-reactivity might disappear more rapidly after
injection of the antigens.
It follows from the results that pure erythrocytes are also capable of producing
immunological approximation. The effectiveness of leucocytes in producing tolerance
to a skin transplant is therefore evidently due to the fact that, in contradistinction
to erythrocytes, the leucocytes contain completely corresponding, common antigens
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with the skin cells, which are responsible for the adaptation leading to the formation
of immunological tolerance.
This explanation is closely connected with the observation (Medawar 1946) that
transplantation immunity to skin can be induced in rabbits by the intradermal
injection of foreign, homologous leucocytes, but cannot be produced by the injection
of erythrocytes. Our contribution to this question is the observation that leucocytea
contain all the antigens which are decisive for transplantation immunity in relation
Tab. 3. Reactivity of Third Series of Birds in which Approximation was Produced
The first immunisation was carried out at 13 weeks with three doses of one ml. on alternate days.
A provocative dose of 0.5 ml. was administered at 18 weeks.
a) Approximation produced with erythmcytea:
Donor
Eaper, bird
Titre after lat
immunisation
Titre after
lat immunisation
Titre after
provoc. dose
552/5
i. v. 1.0
8
542/5
0.8
4
562/5
0.8
-
gg
b58/5
0.5
8
559/5
0.5
2
8
563/5
0.5
84
71
565/5
0.8
84
89
b46/5
I s. c. O.b
128
32
2
b47/5
0.5
2
2
548/5
0.5
2
564/5
0.3
4
71
568/5
0.5
8
571/5
0.5
4
b) Approximation produced with leucocytea:
89
519/b
i. v. 0.5
-
-'
71
534/5
0.5
8
89
502/b
s. c. 0.5
4
b07/5
0.5
64
2
612/5
520/5
0.5
0.5
2
2
16
71
540/6
0.6
-
-
541/b
0.6
32
I
1
576
585 I
2 ~
582
84
8
60b
32
816
32
818
16
621
128
819
64
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to epidermal cells. Medawar's observations did not make it clear that leucocytes
contain all the .decisive antigens of epidermal cells. For producing immunity to
a skin graft, only partial correspondence to these antigens would be sufficient. We
further extend Medawar's observations by the clear demonstration that antigen
correspondence is actually to be found between the leucocytes and the epidermal
cells and not perhaps only between the leucocytes and the cells of the subcutaneous
connective tissue of mesenchymal origin.
The number of birds which acquired approximation by injection on the first day
after hatching, which in earlier work were tested by skin transplants at 14 days,
amounted to 40% (Hayek et al. 1955). Iri these experiments immunological tolerance
was ascertained by immunising the chicks under observation at the age of eight to
18 weeks. The number of experimental birds not forming agglutinins is far lower. In
our view this might be connected with the quantitative character of the approxima-
tion. Immunological approximation obtained by the injection of blood is not nearly
so strong as after the joining of the circulations in embryonal parabiosis and the
low degree of approximation probably causes a more rapid decrease in the percentage
of non-reactive birds in relation to their ages.
Summar.~
A suspension of erythrocytes or of leucocytes was injected intravenously or
subcutaneously into newly hatched chicks. After immunisation with the blood of the
donors at eight to 18 weeks, five out of a total of 30 chicks injected with leucocytes
and six out of 36 injected with erythrocytes did not form agglutinins. It follows from
the results that pure erythrocytes are capable of producing immunological adaptation
and six out of 36 injected with erythrocytes did not form agglutinis. It follows
from the results that pure erythrocytes are capable of producing immunological
adaptation.
Literature
B i 1 1 i n g h a m, R. F,., Brent, L., Medawar, P. B.: Actively Acquired Tolerance
of Foreign Cells. Nature 172:602, 1953.
B i 1 1 i n g h a m, R. E., Brent, L., Medawar, P. B.: Acquired Tolerance of Skin
Homografts. Ann. 1~Tew York Acad. Sci. 59:3, 277, 1955.
H as e k, M.: Parabiosa ptaku v embryonalnim vyvoji. ~s. biologie 2:25, 1953. I' a m e x, M.:
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H as a k, M.: Vegetativni hybridisace zivo~ichu spojenim krevnich ob~hu v embryonalnim
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H as e k, iVL: Projevy vegetativniho sblizovani v adaptaci vys""siho zivoSicha na cizorode
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H as e k, M., L e n g e r o v a, A., M a t e r n o v a, E.: Analysa uLasti krvinek pii po-
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MoTpaxcx~axTaTOB y Tex.xoxpoxxblx. Fol. biol. (Praha) 1:319, 1955.
Hass"ek, M., Lengerova, A., Majer, J., A4aternova, E.: Ik uloze bunek ve
sbliiovacim procesu. ~s, biologie 4:627, 1955. I' a Iu e H, M., JI e x r e p o s a, A., M a ~ e p,
A, x M a T e p x o s a, 8.: K Boxpocy po.nH H~IeTOx B xpouecce c6.>rxHtexHH. Fol, biol.
(Praha), 2,1956.
Lee, J. G., Osgood, E. E.: A Method for Rapid Sepparation of Leucocytes and Nucleated
Erythrocytes from Blood or Marrow with the Aid of Phytohaemagglutinins from Red Beans
(Phaseolus vulgaris). Blood 4:670, 1949.
? Medawar, P. B.: Immunity to Homologous Grafted Skin. I I. The Relationship between
the Antigens of Blood and Skin. Brit. J. Exp. Pathol. 27:15, 1946.
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IIOJ~aBJIeHI~Ie OCpa30BaHIIFi IIMMyHHbIX aPPJIIOTI3HYIHOB y I(bIIiJIfiT~
I{OTOpbIM Ha IlepBbII3 J~eHb IIOCJIe BbII{JIeBbIBaHI3FI BIIpbICKIIBaJII3Cb
uyxcepoAHbie apHTpouHTbt H ,~e~xouHTbt
I~1MMyxo~IOrggecxoe c6nHx{eH.He, IIoAaB~ISIIOItzee y c6~IHxcaeMblx opraHH3MOB cuo-
C06HOCTb K gMMyHIITHOII peaKUI3II IIpH TpaHCIITIaHTauIIH, 6bIJI0 yCIIeIIIHO OCyII;O-
cTBneao B oIIblTax BBeReHgsl xyx{epoRHOg xpoBH 3apoJ.[bIIIIy (I'amex 1953, Billingham,
Brent, Medawar 1953, I'amex 1954, Billingham, Brent, 11Redawar 1955). OAHaHO MbI
IIOHa3aTIII, ~ITO CCJIHHfeHHH IIO OTHOmeHHIO K KOH{HOMy TpaHCIIJIaHTaTy HOB03M0?KHO
j[OCYITbCH II}~TOM IIHyTpH3apOJ~bIIIIOBIiIX BIIpbICKI3BaHIII3 B3BOCH IIpOMbITbIX 3pHTp0-
uIITOB, - 6e3 ~IeHKOIIxTOB (I'aIItex, JIeareposa, MaTepxosa 1955). OTCIORa o*IeBHAHo
3HaLIeHIIe 7IOI3HOUHTOB J[JIFI C03J~aHEH HMMyHOJIOrH~teCKOTO COTII3H{eHIIH H COBMOCTH-
MOCTH K05ICHOI'O rOMOTpaHCII7IaHTaTa. - B HaCTOHII~eI3 pa60Te pOIIIaeTCH BOIIpOC,
MOryT IIII BIIpbICKIIBaHHH 3pIITpOuHTOB 6e3 JIeHKOuHTOB BbI3BaTb HOCIIOCOGHOCTb
H 06pa30BaHIIIO IIMMyHHTiIX arr7IIOTHHHHOB B TeLIOHIIe IIOCT3M6pIIOHaTIbHOTO IIepxOAa.
r1pI3TIIIHOI3 HOB03M0?KHOCTII C IIOMOHjbIO OJ~HBX TO7IbH0 3pHTpOuIITOB C03T,jaTb COBMO-
CTHMOCTb I{OHfHOPO TpaHCIIJIaHTaTa MOHCOT GbITb HJII3 TO, TITO TO7IbH0 7I0HKOIjRTbI
CII0006HbI BT,I3B8Tb HMMyHOJIOPI3LIeCKOe C67III?HOHIIe, HJIH x{e TO, iITO TO7IbK0 7IOHK0-
uHTSI COj[ep1KaT IIO7IHOCTbIO COOTBOTCTByI0IuI3e, 06IIZHe C KJIOTHBMH KO1KH aHTHreHbT.
`raK KaK C IIOMOIIjbIO BIIpbICKIIB3HIIT3 LIyH{epORHbIX K7IOTOK MOH{HO ;jOOIITbCH HMMyHO-
JIOTIIileCKOTO CC7IIIIKOHI3H y 3HagATOJIbHOrO IIpOuOHTa CBOSKOBbIJIynHBmHXCH ubIII7IfiT
{I'ameK 1955), MbI IIOJIb30BaJII3Cb 3TI3M MOTOJ~OM II B HaczoxlueI3 pa6oTe. BHyTpH-
Beaxble BIIpbICxI3BaHi~H 6bIJII3 cRenaHbI 35, a IIoRxox{Hble 40 csexteBbI7IyIIHSmI3MCH
ubIII7IHT8M. rlOJIOBIIHe ubIII7IHT 6bI7Ia BBeueHa B3BOCb LIHCTTiIX 3pgTpOuHTOB, a BTOpOH
IIO7IOBHHO - B3BOCb JIeHKOuHTOB. e~pIITpOuHTbI IIpArOTOB7IHJIHCb IIyTOM ueHTpHC~yrH-
posaHgsl, IIoc~Ie *Iero oTpe3anacb H oT6pacblBa~Iacb BepxHHH *IacTb ocaAxa B czex-
aIHHHOi~ Tpy6o~Ixe BMecTe co cIIOeM ~Iei3xoui3TOS. IIoc:Ie IIOBTOpaoro ueHTpt3c~yrH-
posaHxll IIpoMblTble 3pIITpouHTbl pa3BORm~IHCb 1 : 1 pacTBOpoM `I'yrode. JleuxoTjHTbi
IIpHroTOB~IRnr3cb H7iI3 IIyTeM oTZ[eJIeHHII sepxxeir =lacTx ocaAxa IIoc~re ueHTpP[(~TyrH-
posaHIIa uIITpaTHOK KpoBII, IIIIII c IIOMOII[bI0 (~TI3TOreMarr~IIOTHHgHa, IIIIg x{e, Ha-
KOHOu, IIpIIMOHRJIaCb CMOCb IIpOMbITHTiIX 3pATpOuHTOB H 7IeI3KOuI3TOB TpeTberO j[O-
Hopa B COOTHOIIIOHI3~I J : 1. IIOpBble j[Ba i33 3THX IIpeIIapaTOB JIOHKOuHTOB 6bIBaJIH
Bcerua 3arpH3HOHbI 3pI3TpOuHTaMII. l~ICIIbITaHIITi CCJIII9KOHHfi IIpOPI3BOT(H7IIICb B B03-
paCTO 8-1g HOJjeJIb IIyTOM IIMMyHII3auIIH (PI7ITd peIIMMyHH3auIII3) C67IH1HaBIIIHXCA
ubIIIJIHT uHTpaTHOH HpOBbIO OT TOI'O ?KO j~OHOpa. AI'rTIIOTIIHaIjIITi IIpOH3BOJ[IIJIaCb IIO
MOTO;[y II1J06IIpOK. OTpIIuaTe7IbHbIH p03yJIbTaT peaxuIIII arr~IloTt~xauIIg xoHTpo~IH-
posa~ICH MIIKpOCKOIIH~IOCKII. Pe3yJIbTaTbl IIpxBOJ~TITCH B Tab. 1-3. Iloc7le BIIpblcxH-
BaHIIH JIeIIKOuHTOB I3MMyHHble arr7IIOTHHIIHbI He OrOpa30BaJII3Cb y 5 H3 30 ubIII7IHT,
IIOC7Ie BIIpbICKIIBBHYII3 tIIICTbIX 3pI3TpOuIiTOB - y 6 ubIIIJIHT g3 36. P03yJIbTaTbl OIIbITOB
peHMMyHI33auIIH IIOKa3bIBaIOT, ~ITO apOaKTHBHOCTb MOr7Ia 6bI HC~Ie3aTb 6blcTpee
IIoc~Ie Harpy3KII aaTxrexaMH. IIoRTSepl~i~~IHCb Taxx{e IIo~IyueHHble HaMx paxee
j(aHHble OTHOCI3TeJIbHO TOI'O, iITO y OJ~HOJ~HOBHbIx ubIIIJIHT He 6bIBaeT pa3HHuTiI
B pe3yJIbTBTaX C6JII3HCeHHA IIyTOM BHyTpgBOHHbIX HJIg IIOJjKO}HHbIX BIIpbICKHBaHHI3.
113 JjaHHbIX OIITiITOB BTiITOKaOT, iITO H iIHCTbTO 3pHTpOugTbI CIIOCOCHIaI BbI3bIBSTb
HMMyHOdIOrII90CKOe C6JIIIIKOHI3e. r103TOMy 3(~(~OKTgBHOCTb BIIpbICKIIBaHI3I3 7IeHKOuH-
TOB J~7IH C03~aHHTi COBMOCTHMOCTg HO?KHbIx TpaHCIIJIaHTaTOB o6yc~IOBnHBaeTCH, ?
IIOBHj[T3MOMy, TOM, ~ITO OHI3 - B OTTIH~IHe OT 3pgTpOuHTOB - COueplKaT IIOJI-
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HOCTbIO COOTBOTCTByIOLT~I3e, O6x~YIe C IC7IeTHaMYI KO1IiI3 aHTI3reHbI. B xarIIeH IIpe?IC-
xer~ pa6oTe (I'amex c corp. 1955) xozinuecTBO i~bixzixT, c6~igxcaBrIIxxcil c AoxopoM
IIpg IIOMOH~YI YIH'beKIjYII3 Ha xepBbIx xce J~eHb IIOCTIe BbIxJIeBaxHx I~3 xI3Lja, y ICOTOpbIX
MbI xa 14-bIx ~eHb }ICII3HII IIpOYI3BOJ~HJIH I3CHbITaHI3ii C IIOMOII[bI0 HOSISHbIX TpaHCxJIaH-
TBTOB, COCTaB7iFIJIO ~}~% BCOX C67IH}IfaBIIIgXCx OCO6eyI. IIpII HaCTOSIruIIX OIIbITaX
CTeHOHb YiMMyHOJIOPYILIeCKOI'O CCJIH?ICeHYIH YICC7I0JZyeMbIX i~bIxJIFIT OxpeJ~e7Ix7IaCb C x0-
MOIL~bIO YIMMyHI33aI~YIYi B BO3paCTe g-18 xej[e7ib. KO7IYILIeCTBO CGJIxHCCHHbIX OCO~eI3,
y KOTOpbIX xMMyHHble arrJiiOTIIHI~HbI He O6pa3OBaJII3Cb, ~bIJIO rOpa3J(O HIISI{e. MbI
IIOJIaraeM, iITO 3T0 MOrTIO HaXOj[YITbCFi B CBii3g C KO7II3LIeCTBBHHOII CTOpOHOH C6JIiI-
xceHliH: BepOxTHO, I3MMyHO7IOPHLIeCKOe c6nxaxexIIe, iloJIy9alou~eecH B peay~IbTaTe
BIIpbICKHB8HI3H KpOBx, Ra~Iexo He TaIC CYI7IbH0, ICax C6Jix5ICeHxe IIOJ[ BJIiIHHYIeM
O61i;HOCTg HpOBOO~paILjOHIIH npx 3MOpHOxaJIbHOM IIapa61iO3e, a C BO3paCTOM ~OJIee
Hg3xax CTeIIeHb C6~IIIxcexr3H CTaHOBI3TCx, BepOSITHO, IIpIi9I3HOx 6o~Iee 6bicTporo
CHx1xeHHH IIpOI[eHTa apeaKTYIBHbIX xcxBOTHbIX.
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FOLIA BIOLOGICA
Tom. II. (1956) - Fasc. 1
1'OMOTpaHC1IJIaHTaI~I~IH I{O}xYI y OJ~HOAHBBHbIX L~bIIIJIFIT~ yTSiT I3 I3HJ~IOIiIaT
T. I'PAFA x M. I"AiIIEI{
TIIOnorH*~ecxxft xxcTxTyT LICAH, axcnepxMexmanbIIax 6xonorxII x rexeTxxa, IIpara
llocmynuno a pe8ak~yuw 11 X 1955
Yxce Danforth (1929) ycIIeIIlHO IIepecaxcHSan i~blIInxTaM oT I(bIII7IHT Apyrgx
IIOpOi~ KO~Ify, Ha KOTOPOH BbIpaCTaJIg IIepbH paCIjBOTKH J~OHOpa. rOMOTpaHCII7IaHTaTbi
y xypHHblx aapoRblIIleK coxpaHllloTCR B TexeHge 3-5 MecHizeB IIocTaM6pgoxanbaog
xcgaHH (Eastlick 1941). Cannon H Longmire (1952) ycTaxoBIIIIH, xTO y op[xoRHesxblx
I~bIIIJIHT IIpI3HHMaeTCH g coxpaasleTCS B Te*IeHge 6onee, ueM 8 Heuenb 16% xoxcablx
rOMOTpaHCIIJIaHTaTOB, y 3-J~HOBHbix I[bIIIJIHT 13%, y 4-AxeBHbtx yxce To7lbxo 1%,
a y 2-HeAenbHbix OHH He yualoTCH BOBCe. e~Tg aBTOpbI IIpgBOAIIT, ~ITO y IjbIIIJIHT
Mo7IO?xe 3 AHei~ ycTOHxgBOe IIpgxcHBneHHe HOH(HbIx rOMOTpaHCIIJIaHTaTOB Ha67II0-
Aanocb IIHOrAa Aaxce B 10% cIIy*IaeB. 8TO IIa6nIOReHHe Tecxo cBSlsaHO c AaHHbIMH
HMMy$onorg*Iecxoro c6ngxcexHx. IIprlxcgBneHIIe TpaxcII~IaHTaTa y Mo~IOZ[bIX xcR-
BOTHbIX o6yc7loBnexo, IIOBIIjjHMOMy, TOM, LITO OHH H0 CII0006HbI x aRaIITa>~gH IIO
OTHOIIIeHHIO x xyxcepOAHbIM aHT$reHaM, - B J~aHHOM CIIygae x x0}xHbIM TpaHCIIJIaH-
TBTaM. ~pfiMUe Z[OKa3aTeJIbCTBO TOI'O, LITO y CBexceBblJIyIIIIBIIIIIXCFI I~bIIIJIFiT B 3Ha*IH-
TeJIbHOM IIpOI~eHTe CnyxaeB MO}xeT ~bITb J;OCTgrHyTO IIMMyHOJIOrHileCKOe CCJII35xeHge,
6bIJI0 IIOJIyLIeHO IIyTOM BIIpbICxgBaHgI3 xpOBH OJ~BOJjHOBHbIM I;bIIIJIfITflM C IIOC7Ie-
RyloIIjei~ ycneIIlHOH IIepecaRxog xoxcII oT Toro xce AoHOpa (I'aIIrex 1955).
B HaczoHII~eI~ pa6oTe MbI HccneAyeM Boupoc, Tax xce 7IH orpaHH~Ieaa cTaAIiH
B03M0?xHOCTII IIMMygo~IOrHUecxoro c6n13xceHHH H y Apyrgx BHRoB. RxR IIposepxH
B03M0]xHOCTg gMMyH07IOrgLIeCHOPO C6JIgH{eHHH MbI IIO7Ib30Ba7IIICb H03xHbiMH TpaHC-
IIJIaHT8TaMH y OJ;HUJ{HOBHbIX I[bIIIJIFiT, yTHT g HH;~IOIIIaT.
MemoBuxa
yIcnoIIraosa~iacs Texxxxa nepecaAox no Cannon-y H Longmire (1952). IIyx IIa cnxxxe uba-
IInexx~ yAaaIHSCH, cIIYIxx~ IIepeR oIIepagge# cMaab[sanacb pacTBOpoM xo~i~oRxsi, a~iacTxYIIOCTb
HOTOpOrO IIOBbiIIIaJIaCb IIyTeu IIpg6asneHxx xaMC~Op~ x HaCTOpOBOPO MaC7Ia (20 r xaM~)Opr~,
30 r xaeu;eaxxsoro Mama, 750 Mn ac~xpa, 250 MJI aTgIIOBOro cuxpTa H 40 r IIxpoxcxnxxa).CMaaax-
xaR xonnoRHeM xoxta IIe paa~acb H se cTpaPfana npx IIepecaAxe. I~saApaTxxx xo}xn oxono 1 cM'
nepecaxcxsanxcb MextRy ubtIInxTaMH saaxMIIO. B cny~as~, xorAa xoxte~ o6MexHBanxcb IIopoubt
c paaIIx~tIIblM onepeIIHeM, TpaHCIInaHTaT pacnonaranca IIop~a~Ibao; ec~in sxe B oxpacxe IIepbeB
Aoxopa x peuxnxeIITa He 6r~no abipaaHTenaao~t pasxxi;bl, TpaxcIInanTaT IIosopasxBancx Ha 180?.
JIocxyTxH xoxcH xe IIpxmxsanxeb, a TI~aTenbxo yxataAa~Ba~cxeb x IIoxpblsanxeb AnR c~xxcaRxx
~OJIbIIIxM xoJIx~tecrosoM xo7t7loJ{HH. j{onno7{x# IIa*~gIIan omsanHBaTacn IIa 7-o~-9-t~~ AeIIb IIocne
pIIepauHx. KOHTpOJIbHIaie aBTOTpaHCIIJIaHTaTEi B aTO~ Cepxx OmdTOB HQ AettaIIxCb, IIO C IIOMOII~bIo
sumeonIICaaHO~ TegIIxxH y 1-j{IIesar~i% D;IaIIIJTAT TpaECIIJIaxTaTH IIpxxcxsanH B 90-100% cnyYaes.
O~exxa IIpHxcxsneHIIR TpaxcIInaHTaTa IIpoHasoAgnacb IIa ocsosaaxH ero onepeaHx.
Peaynbmamat
Sbino IIpoHaBeReHO B o6II~eM 184 roMOTpaHCIInaHTa>;IIII xoxcg y oAHOZjHeBHbIx
i[bIIITIHT. npHHiHBJIeHHO H OIIepeHHe TpaHCIIJIaHTaTa Ha6JIIOj[a7IOCb y 6 I;bIII7IHT (3,3% .
~3 18 yTHT OIIepeHHe xOH{HOI'O TpaHCIIJIaHTaTa ~bIJIO IIO7Iy~IeHO y 11 (61 %). I~I3
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10 oAHOAHeBHbIx IIxAIOIIIaT TpaxcII~IaxTaT ollepIIIICx To~Ibxo y 1. Ha Tab. 1 IIoxaaaxo
IIepexcgBaHIIe oxepIIBIIlIIxcu xoxcHblx TpaxcII~IaxTazoB. 4 TpaxcII~IaHTaTa paapy-
IIIIIJIgCb MexcRy 5O-M H 8O-M JZHOM xOC7Ie onepauIIx, OJjIIH COXpaHx7ICFI BIIJIOTb Jj0
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ulomaTaMx 6bI7I OTMeLIeH rOpa3J;0 6oIIee BbICOICIIII IIpOueHT oco6ei3, crloco6Hblx enge
K aj[aIITaIIxx.
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Rexl3x cylljecTByeT : BBeRexIIe =IyxcepoRxbIx axTgrexoB He BblabI-
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g y upyrIIx TeIInoxpoBHblx xcIIBOTxbIx.
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(I'aIIlex, Ppa6a g 3ccnoBa 1956, I'aIIlex, JIexrepoBa II MaTepxosa 1955).
Hpx ocyIIxecTB~Iexxx xoxcxb3x roMOTpaxcH~IaxTauIIK xa IIepBbIK Rexb IIoc~Ie
BblxnesblBaxgx MbI y6eRx~Ixcb, uzo Ro~IroBpeMexxoe HepexcIIBaxxe II Raxce ycTOH-
ugsoe IIpIIxcIIB~IeHIIe y yTHT Ha6TIIOj~aeTCH ropaaRo uaII~e (oxo~IO 65%), ueM y i;blII-
7IHT (3,3%) x y I3xj[IOIIIaT (1 cIIyuaH xa 10).
JIHTepaTypa
I' a HI e x, M., JI e H r e p o B a, A., M a T e p H o B a, 3.: Axanxa yuacTHH xpolsxxb3x TeIIeI;
IIpI3 aKCIIepI3M0HTaJIbHOM IIpeOJ~OJIeHI3I3 HQCOBMCCTHMOCTH HO}ILHbIX POMOTpaHCHJIaHTaTOB
y TeI3aloxpol3xb3x. Fol. biol. (Praha), 1:319, 1955.
B i l l i n g h a m, R. E., Brent, L., M e d a W a r, P. B.: Acquired Tolerance of Skin
Homografts. Ann. New York Acad. Sci., 59:277, 1955.
Cannon, J., L o n g m i r e, W.: Studies of Successful Skin Homografts in the Chicken.
Ann. SI3rg., 135:60, 1952.
Danforth, C. The Effect of Foreign Skin on Feather Pattern in the Common Fowl (Gallus
domesticus). W. Roux' Archie fur lJntwicklungsmech, der Organismen, 116, 1929.
Approved For Release 2008/04/10 :CIA-RDP80T00246A002900500016-8
 Approved For Release 2008/04/10 :CIA-RDP80T00246A002900500016-8
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