INDIVIDUAL BLOOD DIFFERENCES IN MEXICAN INDIANS, WITH SPECIAL REFERENCE TO THE Rh BLOOD TYPES AND Hr...

8/13/2019 INDIVIDUAL BLOOD DIFFERENCES IN MEXICAN INDIANS, WITH SPECIAL REFERENCE TO THE Rh BLOOD TYPES AND

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I N D I V I D U A L B L O O D D I F F E R E N C E S I N M E X I C A N I N D I A N S , W I T HS P E C I A L R E F E R E N C E T O T H E R h B L O O D T Y P E S A N D

H r FA C T O R *

Bx ALEXAN DER S. WIEN ER, M .D., J. PRECIAD O ZEPEDA, M.D., EVE B. SONN,H. R. POLIVKA

(From the Serological Laboratory of the O ice of the Chief Medical Examiner of the Cityof New York, and the University of Guadalajara, Mexico)

(Received for publication, March 30, 1945)

Recen t inves t iga t ions have shown tha t the re a re s t r ik ing d i ffe rences in thed i s t r ibu t ion o f the Rh b lood types in d i ffe ren t r aces (1) . To da te , s tud ieshave been carr ied out on white individuals (2 , 3) , Negroes (4) , Chinese (5) ,

Japanese (6, 7 ) , and As ia t i c Ind ians (8 ). Because the resu l t s a l r eady ob ta inedhave p r oved to be o f va lue fo r the s tudy o f rac ia l r el a t ionsh ips , i t s eems wor thwhi le to ca r ry such s tud ies fu r the r.

In th i s paper, we p ropose to p resen t the resu l t s o f Rh typ ings on a se r i e s o f98 Mexican Ind ians and to compare the d i s t r ibu t ion o f the Rh b lood types inMexican Ind ia ns wi th tha t o f o the r r aces . The b lood samples o f these MexicanInd ians hav e a l so been t e s t ed fo r the A-B-O groups , the subgroups o f A ,the M -N types , the agg lu t inogen P, and the Hr fac to r. Whi le a num ber o fs tud ies have a l ready been ca r r i ed ou t on the b lood g roups , subgroups o f A andM- N types in Amer ican Ind ian s (9 -11) , and whi le the re has a lso been one s tud yon the Rh fac to r (12) , thus f a r no da ta have been co l l ec ted fo r th i s r ace con-ce rn ing the Rh b lood types , o r the p roper t i e s P and Hr.

Materials and MethodsThe Mexican Indians studied by us came from Tuxpan, this locality being selected because

of the special asdmtancO available there an d because the Indians there kept themselves apartfrom the Mexicans and whites.

In prehistoric times, Tuxpan is said to have been inhabited by nom ad tribes of unknownorigin. Th e existence of these tribes is inferred from the primitive ar t work the y left behind,in particular very crude pottery and rudimentary stone carving. The Indians themselves aredescendants o f the family of Toltecas, which migrated from a certain locality known as Tollan(also called Huehuetlapallan) in the United States. In their trek from the United Statesto the state of Jalisco, the Toltecas Indians passed throu gh Sinaloa, stopped near the city ofTepic for a short time, and then moved on to Tuxpan which they reached in the year 542A.D., according to Ft. Antonio DeTello. The bulk of the tribe then moved on to the valleyof Mexico, but a nucleus of the tribe remained behind in Tuxpan. These may have mixedwith the nomad tribes and were the ancestors of the Indians tested b y us. During the Spanish

* Aided by a grant fro m the United Hospital F und o f the City of New York.I We wish to acknowledge our indebtedness to J. M. Ruvalcab a for his valuable assistance

in obtaining the cooperation of the Indians.559


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560 INDIVIDUAL BLOOD DIF]~ERENCES IN M~XICAN INDIANS

domina t ion of Mexico , the on ly whi te people who came in con tac t wi th the Ind ian s a t T uxpanwcre the government employees .bu t they d id no t in te rmar ry wi th the Ind ians . Whi te peoplecame to Tuxp an abo ut 1812. According to the stat is t ics available, 90 per cent of the presentpopula t ion of T t lxpan a re fu l l-b looded Ind ians . Tuxpan i s loca ted a t 19 22 ' nor th l a t i tudeand 4 west long itude from the m eridian of Mexico City. I t is 1,137 meters abov e sea leveland i t sc l imate i s t empera te . I t i s a fe r ti l e va l ley sur rounded bym ount a ins and the reare twovolcanoes in the locali ty.

The blo od samples were collected by venip unctu re by one of us (J . P. Z.) from 100 Indiansa t Tuxpan , Mexico , and sh ipped a t o~ce by a i r express to New York Ci ty where they a r r ivedwithin 7 days. The blood samples were received on a Sunday mornin g and no tests were mad eunti l the fol lowing day. I t too k 3 days for al l the tests to be completed. At the beginning ofeach day's work blood suspensions were prepared by shaking up the clots in sal ine solution;coarse part icles were al lowed to set t le and the supernatant smooth suspensions transferredto a series of numb ered tubes. The suspensions were washed once by centrifuging the m atlow speed and resuspcnding the sediment in fresh sal ine solution, to form a 2 per cent sus-pension ( in terms of blood sediment) . At the end of each day' s experiments al l the bloodcell suspensions were discarded and fresh suspensions were prepare d the fol lowing mornin g

f rom the b lood c lo ts which were kep t in the re f r igera tor dur ing the in te r im.As in our previous studies, the anti-A and anti-B sera used were of high f i ter and avidity,

hav ing b een prepared f rom the se ra o f donors who ha d been in jec ted in t ravenous ly wi thp lasma (Sharpe and Dohme) , o r in t ramuscula r ly wi th so lu tions of A and B group subs tances .2The absorbed B se rum was prepared f rom the b lood of a g roup B ind iv idua l whose se rumconta ined an unusua l ly la rge amoun t o f a l agg in t in in . The an t i -M and an t i -N tes t ing f lu idswere p repared in the usua l m anner by absorp t ion of our s tock rabb i t : immun e sera .

For P-,h typing, sera of specifici ties anti-Rh0 (standard , 85 p er cen t posi t ive for bloo ds ofwhite indiv iduals) , anti-R.h ' (70 per cent posi t ive) , and anti- Rh p (30 per cen t posi t ive) arerequ i red . The an t i -Rh0 se rum used in the p resen t s tudy came f rom a g roup B, type Rh 'mother o f an e ry throb las to t ic in fan t o f type Rhl . The se rum was of h igh t i t e r (128) and forthe tests was used in a di lut ion of 1:8, by mixing 1 part of serum, 1 part of autoclaved groupA saliva solution (or group substance), and 6 parts of sal ine. Two different reage nts ofspeci f ic i ty an t i -R h ' were used : one f rom an Rh-negaf ive mother o f an e ry throb las to f ic in fan t ,the second f rom a type Rh2 mo ther o f an e ry throb las to t ic in fan t . The se rum f rom th e fo rmer

individual was of specif ici ty anti-Rh~ (87 per cent posi t ive) with weak anti-Rh0 agglutinins.The an t i - Rh ' reagen t was p repared f rom th i s se rum by b lock ing the ac t ion of the an ti -R .h0agglu t in in by us ing the fo llowing formula : 1 par t o f the s tock se rum was mixed wi th 1 par teach of po ten t b lock ing se rum and pooled A and B sa l iva so lu t ion and fu r ther d i lu ted wi th 2parts of sal ine solution. The second anfi- Rh' serum, from the type Rh2 moth er, besidesbe ing of h igh t i t e r, had the ad vantage th a t i t con ta ined the an t i -Rh ' agg lu t in in in pure fo rmwith out any anti-Rh 0 agglutinin. For this reagent the fol lowing formula gave excellentresu l ts : 1 par t s tock se rum was mixed wi th 1 par t poo led A and B sa l iva so lu t ion and 3 p ar t sof sal ine solution. Two an ti-R h reagents were also used, one prepared with the aid of block-ing se rum f rom an an t i -Rho se rum and the second f rom the se rum of a type Rhl mother o f anery throb las to t ic in fan t . The two an t i -Rh / reagen ts were p repared f rom these two s tock se rain a m ann er similar to that used for the anti-R_h' reagents.

The an t i -Hr se rum was excep t iona l ly po ten t and was made ava i lab le to us th rough thecourtesy of Dr. Peter Vogel . I t came from a group O, type Rh~ mot her of an erythro blastot icin fan t . I t s excep t iona l po tency may b e ev iden t f rom the fac t tha t a l l Hr-pos i t ive b loods gaveunifo rm strong reactions. The single gene and double gene dose effect f irst noticed by Race

2 For this material we are in debted to ~Dr. J . A. Leig hty of the Eli Lil ly Comp any.


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V ~ IE N ER , PP ~ CI A I) O Z E P E D A ~ S O N N ~ A N D P O L I V K A 561

and Taylor (13), and also observed by Wiener, Davidsohn, and Pot ter (14) was not evidentwith this serum because this effect is only apparent with weaker anti-Hr sera. Despite itsexceptionally high titer, the anti-Hr serum was used undilut ed for testing the samples of allthe group O Indians. For testing Indians belonging to groups A and B, it was first mixedwith an equal volume of pooled A and B saliva.

The anti-P serum came from a group A individual who had become isoimmunized to theagglutinogeu P as a result of repeated blood transfusions (15). This serum was diluted withan equal volume of saline solution for the tests, and was not used for testing the few group Bblood samples in the series.

With regard to the saliva solution used for the inhibition of the anti-A and anti-B aggiutin-ins in the anti-Rh sera, we have found the following to be the simplest and most satisfactorymethod of preparing it. Equa l volumes of group A and B saliva from secretors were pooledand autoclaved at 15 pounds pressure for 15 minutes. The coagulated mucus and othersuspended material were then readily removed by centrifugation. The supernatant, opales-cent fluid was diluted with 2 volumes of saline solution and distributed among a number ofvials of convenient size . After a second autoclaving for 15 minutes at 15 pounds pressure,the vials were capped with sterile rubber stoppers and stored in the refrigerator until used.

If the saliva could not be processed immediately after collection, it was placed in a boilingwater bath for 15 minutes to destroy the blood group enzymes and then stored in the refrig-erator unt il processing could be carried out.

The A-B and M-N tests were carried out on Boerner well slides at room temperature asdescribed in our previous papers. All group A bloods were further tested with absorbed Bserum. The Rh typings and Hr tests were carried out in small test tubes in a water bath a t38 C. as already described (2, 14 ). The tests for agglutinogen P were also carried out intest tubes bu t at refrigerator temperature (1S).

Of the 100 blood samples received, two were hemolyzed and therefore nottested because it was felt the results would be unreliable. The findings on theremai ning 98 blood samples are presente d in Table I.

Blood Groups, Subgroups, and M- N Type s

In Table I I, we have su mmarize d the results of the tests for the blood groups,subgroups of A, and M-N types on the 98 Mexican Indians . For the purposeof comparison, we have also listed in this table the groups, subgroups of A, andM-N types of a different group of Indians and of white individuals in NewYork Ci ty.

The Indi ans in both studies differ from whites in the high incidence of groupO. At one time it was held that all full-blooded Indi ans belonged to group O(16), but Matson and Schrader (17) encountered a number of Indian tribeswith a high frequency of group A (as high as 76.7 per cent). The two In dia ntribes in Table II show somewhat different distributions of the blood groupsand t hey bot h con tain significan~ numbe rs of group A individuals. All of thegroup A individua ls in the In dia n tribes listed in Table I I be long to subgroupA1, in contrast to the situ ation in white individuals where as man y as one-

fourth or one-fifth of all A or AB indivi duals belong to subgroup A~. Theseresults are in agreement with those of Levine, Matson, a nd Schrader (18) who


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562 INDIVIDUAL BLOOD DIF~ 'ERENC ES IN MEXICAN INDIANS

report that subgroup A2 is virtually absent even in Indian tribes with a high

freq uency of group A. In this connection, it should be mention ed that sub-TABLE I

Cl~sification of Blood Samples from 98 Mexican Indians

1. OMNR hfflr +P +2. OMRhzRhffIr+P +3. OM'NRh~Hr+P +4. O M R h , R h ~ I r + P +8. OMR h,Hr-P +6.* OMNRh~Rh~-Ir+P +7. OMNRhxHr-8. OMRh,RhffIr +P +9. OMRhzRh~Ir-P-

10. OMRh~Rhff Ir+P +11. OMRh,Rhff IrP +12. A,MNRhxRhffIrP 13. OMRh,Hr-P-14. OMRh~Rh~Hr-P +15. BMNRh ~Hr +16. OMNRh,Hr -P +17. OMNRhl Hr-p +18. A x N R h , H r - P +1 9. O M R h , H r - P -

2 0. A , M R h , H r - P +

2 1. A , M N R h , P -

2 2. O M N R h ~ R h ~ X - I r + P

23. OMR h,Hr+P 24. OMRh~Rh~25. OMRhxHr-P-

26, B M N P . h , R h ~27. O M R h ~ H r + P +28. O N R h l H r - P +29.* O M N R h , H r - P +3 0. O M N R h ~ I r + P +

3 1. O M R h * H r - P +

32. OMRhxHr-P +33. OMRh*Hr-P +34. OMRh~I-Ir+P +35. OMNRh*Rh~Hr+P *36. OMNRh,Hr -P +37. OMRh tHr- P 38. OMRhaHr- P +39. OMNRh, RhffIr+ P +40. OMR hiHr-P +41. OMNRh~ RhffIr +P +42. OMNRh,Rh~Hr+P -43. OMNRh ,Hr+P +44. A,M Rh~RhffIr +P +45. OMRhxHr+P -46. OMRh,Hr- P +47.* OM RhoHr+P +48. OMRh,Hr-P +49. OMRh,Rh2Hr+P 50. OMRh,Hr -P +

51. OMRh,Rh ffIr+P 52. OMRhaRh~Hr+P +53. OMRhIHr-P-54. OMRhxHr+P +55. OMRh iRh:Hr +P +56. OMRhxRhzHrP 57. ONRh,Hr- P +58. OMRhlHr -P +59. OMRh, Rh~Hr+P +60. OMR h,RhffIr+ P +61 OMRhIHr-P-62. OMRhxHr+P +63. OMRh,RhffIr-P +64. OMNRh,Hr- P +65. OMR h~Hr-P 66. OMNR h,RhR-I r+P +67. OMRh ,Rhg'Ir+ P +68. OMRh,Rh~-*'Ir+P +69. OM_NRhffIr+P70. OMNRhlHr-P-71.* B MNRh, RhffI r +72. OMRh~ Hr-P +73. OMNRh, RhzHr+P -74. OMNRhlRh~ Hr+P +75. OMRhxHr-P-

76. OMRhIHr-P-77. OMR hIRh~Hr+P +78. OMNRhlRh~Hr+P +79. OMNR h1Hr-P +8O. OMRhffIr+P81. OMNRh,R hffIr+ P 82. O M N R h , H r - P +83. OMNRh,Hr-P-84. O %fNRhxHr-P +85. OMRh,Rh~ 'tr+P +86. OMgh,RhffIrP +87. A,MR h,Hr+P +88. OMRhffIr +89. OM Rh,Hr+P +90. OMRhlRh2Hr+P91. OMRh IHr- P +92. OMRh,Hr- P +93. OMNRh *RhzHrP +94. OMRh*RhzHr+P -95. OMRha Rhd~r+P -96. OMRh,Hr- P +97. OMNRh, Hr-P +98. OMNRh~RhH-Ir+P

* Indians known t o b e mixed.

TABLE IIDistribution of the Agglutinogens A -A~-B and M-N among 98 Mexican Indians As Compared

witk U. S. Indians and Whites

Populat ion

Mexican IndiansFull-blooded Indi-

ans (varioustribes) FortLewis, Wash.

Whites (U. S.)

Invest igators

Present studyLandsteiner,

Wiener, andMatson (12)

Wiener (10)

~o. ofindi-'iduals~ested

98120

1,077

Distr ibuti on of groups andsubgroups

_ _ 0_0 _ ~ Ip e r p e rc e n l c e n l

90.8[ 6.173.3]25.8

141.7129.0

per p e r

0 [3.1

0 10.8

8.9113.9

p e r [ p e rc e n t [ c e n l

05.2 1.4

Distr ibut ionof types

her I z 0 e r IPer. . [ c ~ I . .

2 1 . 2 1 3 . 11 3 s . 7

29.2121.3149.5

group A2 is lacking or extrem ely rare in Chinese (5) and Japa nese (7), an obser-vation which supports the theory that American Indians are of Mongolianorigin.

It is not cer tain whether the group B individuals among the Mexican Indianswe tested derived their B gene from their Mongolian ancestors or from mixture


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V C- IE NE R P R E C I A D O Z E P E D A ~ S ON Iq A N D P O L I - V K A 563

w i t h o t h e r r a c e s. W h i l e S n y d e r 1 6 ) h a s d e m o n s t r a t e d t h a t t h e p r e s e n c e o f

i n d i v i d u a l s o f g r o u p s A , B , a n d A B i n m a n y I n d i a n t r ib e s i s d u e t o m i x t u r e w i t hwhi tes or o the r races , i t would be imposs ib le to expla in on th i s bas i s f requenc ieso f g r o u p A a s h i g h a s 75 p e r c e n t , a s h a s b e e n re p o r t e d b y M a t s o n a n d S c h r a d e rf o r B l a c k f e e t I n d i a n s . M o r e o v e r, B o y d 1 9 ) h a s p r o v e d s t a t i s ti c a l l y t h a te v e n i n I n d i a n t r i b e s w i t h l o w fr e q u e n c i e s o f g r o u p s A a n d B , t h e d i s t r i b u t i o n sa r e u s u a l l y i n c o n s i s t e n t w i t h t h e t h e o r y o f r a c i a l a d m i x t u r e . A s im p l e w a y o ft e s t i n g f o r a d m i x t u r e i s t o s u b g r o u p t h e g r o u p A i n d i v i d u a ls , a s w a s d o n e i l l t h ep r e s e n t s t u d y ; t h e p r e s e n c e o f i n d i v id u a l s o f s u b g r o u p A 2 w o u l d i n d i c a t e t h ep r e s e n c e o f r a c i a l a d m i x t u r e . A s s h o w n in Ta b l e I , w i t h o n l y f o u r e x c e p ti o n s ,a l l t h e I n d i a n s t e s t e d i n t h e p r e s e n t s t u d y a r e b e l i e v e d t o b e f u l l- b l o o d e d .

I t w i ll b e s e e n fr o m Ta b l e I I t h a t t h e d i s t r i b u t i o n o f th e M - N t y p e s i n t h et w o I n d i a n t r i b e s i s v e r y s i m i la r, a n d s t r i k i n g l y d i f fe r e n t f r o m t h e d i s t r i b u t i o ni n w h i te s . I t i s o f i n t e r e s t to m e n t i o n t h a t a l l t h e I n d i a n t r i b e s t e s t e d t o d a t e

TA B L E I I IDistribution o f gglutinogen P among M exica n Ind ian s s Compared with Other Racvs

Population

Mexican Ind ians . . . . . . . . .Whi tes N. Y. C. ) . . . . . . . .Negroes N. Y. C.) . . . . . . .

Investigators

Presen t s tudyWiene r and Unger 15 )Wiene r and Unger 15 )

~ O. 0indi-idua::ester

95328

73

requencieso f t y p e s

P__5_~er eeut{per een78 .9 I 21 .173 .2 [ 26 .897 .7 I 2 .3

requenciesof g n s

per~nt

54 .1 I 45 .948 .2 [ 51 .884 .9 I 5 .1

1 0 ) , i n c l u d i n g t h e o r i g i n a l s t u d y b y L a n d s t e i n e r a n d L e v i n e 11 ) , a ls o e x h i b i t

v e r y s i m i la r d i s t r i b u t i o n s f o r t h e M - N t y p e s . T h e q u e s t i o n m a y a r i s e h o wi t i s p o s s ib l e f o r t r i b e s o f A m e r i c a n I n d i a n s w i t h s u c h w i d e l y d i f f e r e n t d i s t r i b u -t i o n s o f t h e b l o o d g r o u p s t o r e s e m b l e o n e a n o t h e r s o c l o s e ly w i t h r e s p e c t t o t h eM - N t y p e s . A p o ss i b le e x p l a n a t i o n i s t h a t t h e b l o o d g r o u p s a r e d e t e r m i n e db y a s e r i es o f th r e e a l l e li c ge n e s, w h i l e th e M - N t y p e s a r e d e t e r m i n e d b y a p a i rof a l le l ic genes , so there i s g rea te r l eew ay for var ia t ion in the fo rme r case.

The gglutinogen P

T h e o b s e r v a t i o n s o n a g g l u t i n o g e n P m a y b e o f i n t e r e s t b e c a u s e s o fe w s tu d i e sh a v e b e e n c a r r ie d o u t t h u s f a r o n t h i s p r o p e r t y. I n t h e o ri g in a l s t u d y o f L a n d -s t e i n e r a n d L e v i n e 2 0 ) , t h e s e i n v e s t ig a t o r s p o i n t e d o u t t h e s t r ik i n g d i ff e r e n c ei n t h e d i s t ri b u t i o n s o f a g g l u t i n o g e n P a m o n g w h i t e s a n d N e g r o e s . A s i d e f r o mt h e s t u d ie s o f D a h r a n d h i s c o l l a b o r a t o r s 2 1 ) o n w h i t e in d i v i d u a ls in G e r m a n y,

a n d t h e s t u d ie s b y F u r u h a t a a n d h i s c o l l a b o r a to r s 2 2) o n t h e r e l a t e d p r o p e r t yQ i n J a p a n e s e , t h e o n l y o t h e r s t u d ie s o n t h e a g g l u t i n o g e n P a r e t h o s e l i s te d i nTa b l e I I I . A l t h o u g h t h e n u m b e r o f M e x i c a n In d i a n s t e s t e d is s m a l l, it w il l


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5 6 4 I N D I V I D U A L B L O O D D I T ] ~ E R E N C E S I N M E X I C A N I N D I A N S

be seen tha t the distr ibution of property P is significantly different from that

in Negroes, there being a substantial number of P-negative individuals, justas in whites.

The Rk Blood Types

The results of the tests for the R h blood types on the 98 Mexican Indiansare summarized in Table IV. Inasm uch as the Rh blood types were onlyrecent ly discovered, it seemed of interest, for purposes of comparison, to sum-marize in the table al l the available data on the d istr ibution of these types indifferent races.

T A B L E IVDisvbu___ ono_ f he R h Type s among Mexica n India ns As Compar~_~ witk Otker Ra~es

P o p u l a t i o n I n v e s t i g a t o r s I~ .o f ] D i s t r i b u t i o n o f R h b l o o d t y p e s[ G e n ef requenciesi

f i- . . . . . .

t e d -~ 2 2 , -~ ~ ,~. . ,~ ~ -~

per p er per per p er per per per per per per lper Per lPer

Present study 98] 0 148 .0 ] 9 .2141 .8J 1 .0 ] 0 I0 I 0 I0 16S .1130 .0 ] .9 0 ] 0W iener, U ng er , 468113 .3154 .4113 .7115 .01.41 1 .010 .2 0 r6 .5143 .3114 .513 .2 1 .31 0 .3

W iener,andonnBelkin,(3) 2231 8.112 0.212 2.415 . 4 /4 1 . 2 1 2 . 7 ] 0 0 1 2 8 , 4 11 . 7 11 4 . 4 1 4 2 . 1 2 . 7 1 0

W iener,andSnn4)Sonn, 156] 7 .1170.51 5.1112.8 1.91 2.6 10 0 126 .656.216 . 0 1 3 . 4 4 . 4 1 0

] g u c h i(7)

M e x i c a n I n d i a n sW h i t e s ( N . Y. C . )

N e g r o e s ( N . Y.C . )

A s i a t i c I n d i a n s

Chinese

J a p a n e s e

J a p a n e s e

As shown in the table, the distr ibution of the Rh blood types among theMexican Indians differs str ikingly from the distr ibution in whites, Negroes,and Asiatic Indians. The absence of Rh-negative individuals conforms withthe p revious report of Landsteiner, Wiener, and Mat son (12), 3 who tested aseries of No rth American Indians usinganti-rhesus guinea pig serum. As inthe Chinese and Japanese, th e M exican Indians belong almost exclusively totypes P,hb Rh~, and RhlRh~ , and no bloods giving intermediate reactions (23)were encountered. These findings, therefore, further su pport the the ory ofMongolian derivation of the American Indians.

The calculation of the gene frequencies is simplified if one ma kes use of the

a A t th e t i m e t h is s t u d y w a s m a d e t h e e i g h t R h b l o o d t y p e s w e r e n o t k n o w n . H o w e v e r ,s o m e o f t h e I n d i a n s i n t h e s t u d y w e r e t e s t e d w i t h a n t i - R h t s e r a .


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W l EN E R , P R E C I A D O Z E P E D A , S O N N , A N D P O L I V K A 6

fac t th a t the e igh t Rh b lood types fa l l in to four na tu ra l pa i r s , g iv ing r i se to thefour classes as fol lows:

W ---- Rh0 + Rh negativ e (1)

U = Rh I .31- Rh' (2)

V = Rh2 + Rh (3)

UV --- RhlRh2 + Rh 'Rh (4)

As was po in ted ou t in p rev ious papers , these four c lasses a re t r ansmi t t edby t r ip le a l l e li c genes jus t l ike the four b lood groups . Accord ing ly, the f re -quenc ies o f the th ree genes can be ca lcu la ted wi th the a id o f the fo l lowingf o r m u l a e :

= ~ / ~ ( s )

. = ~ / ~ + u - ~ / ~ ( 6)

, -- ~ + v - ~ (7 )

I f one wishes to com pute the f requenc ies o f the s ix Rh genes , the fo l lowingformulae a re f i r s t used :

= V ~ n e g . ( s )

R// = V/R_h + gh neg. -- V / ~ neg. 9 )

R h - - ~ / R h + R h n e g . - - V / ~ n e g . 1 0 )

The f requenc ies o f the o ther genes a re then ob ta ined by sub t rac t ion , us ingthe fo l lowing re la t ionsh ips :

w = R h o + r h (11)

u = g h + R h ' (12)

v - - R h 2 + R h (13)

Whi le these fo rmu lae a re no t 100 per cen t e ffi c ien t, they a re cons i s ten t andwere sa t i s fac to ry fo r ca lcu la t ing the gene f requenc ies fo r the popula t ions o fwhi tes , Negroes , and As ia t i c Ind ian s l i st ed in Tab le IV. The fo rmulae becomehigh ly ine ff ic ient , however, whenever the f re quen cy of any o f the th ree c lassesW, U, V i s ve r y low. The fo rmulae were the re fore unsa t i s fac to r y fo r theMexican Ind ians , Chinese , and Japane se because o f the low f requen cy of c lassW. Accord ing ly, the fo llowing fo rmulae were used ins tead fo r these races.

= 1 - ~ (14 )

w = 1- - (u + v) (16)


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566 I N D M D U A L B L O O D D I F F E R E N C E S I N M E X IC A N IN D I A N S

where ( Rh'-) r epresents the percentage of individuals whose blood gave nega-

tive reactions with the anti-Rh' serum, and (Rh"-) represents the percentageof negative reactions with the anti-Rh" serum.Inasmuch as there were no Mexican Indians of types Rh ' , Rh", Rh 'R h",

or Rh negative, the frequencies of genes R h ' , R h , and rh were all zero or verysmall. Therefore, for this race:

R/t1 = u (17)

R/t~ = v (18)

Rk0 ~- w (19)

It is of interest to apply the chi square test to our sample of Mexican Indians.This is done by recalculating the four classes with the aid of the formulae:

W ~- ua (20)

U ~ u ~ + 2 u w ( 21 )

V = ~ + 2 w v ( 22 )

UV =- 2uv (23)

Accordingly, the expected frequencies of the four classes are as follows:

w -- Rh0 = (0.019) 2 = 0.04 per cent, or 0.04 individuals (24)

U = Rhl = (0.681) 2 -4- 2(0.681) (0.019) = 49.0 per cent, or 48 individuals. (25)

V = Rh~ = (0.30) ~ -4- 2(0.30) (0.019) = 10.1 per cent, or 9.9 individuals. (26)

UV -- RhlRh~ = 2(0.681) (0.30) -- 40.9 per cent, or 40.1 individuals. (27)

The observed frequencies of the Rh types were:

Rh0 - 1Rhl -- 47Rh~ ~ 9RhlRh2 = 41

x ~ -_ :~ x - - x o ) ~ _ ( 1 - - 0. 04 ) ~ + ( 4 7 - - 4 8 ) 2 + ( 9 - - 9 . 9 ) s + ( 4 1 --40.1) 2x0 0.04 48 9.9 40.1

-- 23.0 + 0.2 + 0.08 + 0.02 = 23.1

Since n = 1, P is much less tha n 0.01.These findings, therefore, do not conform with the expectations under the

theory. The difficulty obviously centers about the single indivi dual encoun-tered of type Rh0. This might be attrib uted to chance because only a single

individ ual is involved, except tha t the same difficulty has arise n in the studieson Chinese and Japanese. This suggests some complication in the genetic

Therefore,


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WIENER, PRECIADO ZEPEDA~ SONN , AND POLIVKA 567

mecha nism. The na ture of th is compl ica t ion wi l l be d iscussed in connect ion

wi th t he Hr f ac to r.The Hr Factor

In T able V are l i s ted the resul t s of the H r tes ts on 95 of the Mex ican Indians ,and, for compar ison , a l so a series of whi te indiv idua ls and a ser ies of Negroes .I t wi l l be seen tha t the inc idence of Hr-negat iv e indiv iduals i s h ighes t amo ngthe M ex ican Ind i ans and l owes t among the Negroes.

Acco rd ing to t he gene t i c t heo ry o f Raceet al. 24), the Hr fac tor i s presen tin the a gglu t inogen s de termined by genes Rha,Rh , Rho,a n d rh b u t n o t i n t h eagg lu t i nogens de t e rmined by genes Rh l andRh ' . The f r equency o f t he Hr-nega t ive t yp e shou ld t he re fo re be equa l t o t he squa re o f t he sum o f t he f re -quencies of genes Rhl andRh r. As shown in Table V, the observed f requencies

TA B L E V

Dis t r ibu t ion of the Hr Fac tor among Mexica n Ind ian s s Compared wi th Other Races

Population

M e x i c a n I n d i a n s . . .W h i te s . . . . . . . . . . .N e gr oe s . . . . . . . . . .

Investigators

Present s tudyWiener, Davidsohn, and Pot te r 14)Wiener, Davidsohn, and Pot te r 14)

qo. ofindi-idualsested

9523949

Frequencies oftypes

Hr___~ Hr_____~_

~er cen t pe r cen t

55.8 I 44.272.0 ] 28.098.0 I 2.0

(Rhl +Rh )

per cen t

46.419.92.1

ag ree c lo se ly w i th t he expec t ed va lues fo r t he Mex ican Ind i ans and Negroes,bu t t he f r equency o f t he Hr-nega t ive t yp e among w h i t e i nd iv idua l s i s too h igh ,sugges t ing some compl ica t ion in the genet ic mechanism.

Amo ng the M ex ican Ind i ans t e s t ed by u s , p r e sumab ly on ly t h r ee o f the s i xa l lel ic Rh genes are present , nam ely, Rhl , Rh2, and R/ t0(cf. Tab le V) . Unde rthese condi t ions , the Rh types which are poss ib le , the corresponding genotyp es ,the H r reac t ions to be exp ected , and th e ca lcula ted f requencies of the pheno-types are as fo l lows: - -

Rh types

Rhl

Rh2

RhlRh2

Genotypes Hr reactions FrequenciesRh0Rh0 P osi tive w ~RhlRhl Negative u*

RklRhs Positive 2 u w~Rh~Rh2~Rh2RhoJ Positive v~ q- 2vwRhaRh2 Positive 2uv

I t wi l l be seen tha t Hr-n egat iv e indiv iduals a re theore t ica l ly poss ib le on lyin t ype R h l . Moreove r, u sing t he ca l cu la t ed f r equenc i es g iven in equa t ion


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568 INDIVIDUAL BLOOD DIFFERENCES IN MEXICAN INDIANS

U22 5 ) t h e e x p e c t e d p r o p o r t i o n o f H r- n e g a t i v e i n d i v i d u a l s in t y p e R h l i s u 2 + 2 u w

0.464- - 0 . 49 0 o r 9 5 p e r c e n t . T h a t i s, o n l y 1 o u t o f 2 0 M e x i c a n I n d i a n s b e l o n g i n g

t o t y p e R h l w o u l d b e e x p e c te d t o b e H r - n e g a t i v e , a n d t h e o b s e r v e d n u m b e r7 ) o f s u c h i n d i v i d u a l s i n o u r s e r i e s o f 9 8 I n d i a n s d o e s n o t d i f fe r s i g n if i c a n t l y

f r o m t h e e x p e c t a t i o n s(cf. Ta b l e V I ) .

TABLE VI

Rela t ion of the Hr Fac tor to the Rk Blood T ypes among 91 Mex ican Ind ians

H r r e a c t i o n

H r + . . . . . . . . . . . . . .

H r - . . . . . . . . . . . . .Tota ls . . . . . . . . . .

Rh

No. of individuals in types

R h t R h ~

7 9

39 0

46 9

R h l R h t

3 6

3

3 9

To t a l s

53

42

95

TABLE VI I

Rela t ion of the Rare Genes Rhu an d Rh , to the S ix S tandard Rh Genes

Genes

rhRhoR h 'R h tR hRh2

Rg (Rk~)Rk l ~ (Rh,)

R e a c t i o n s i t h h a n t i se r a

Rh

N e g

N e g

P o s

P o s

N e g

Neg.

P o s

Pos.

P, h R h 0

N e g N e g

Neg. Pos.N e g N e g

N e g P o s

P o s N e g

Pos. Pos.

Pos. Neg.P o s P o s

Reaction with H rantiserum

Pos.Pos.Neg.Neg.P o s

P o s

Neg.Neg.

I n T a b l e V I i s g i v e n a c o m p a r i s o n o f th e H r r e a c t i o n s a n d t h e R h b l o o dt y p e s i n t h e 95 M e x i c a n I n d i a n s . I t w i l l b e s e e n t h a t , c o n t r a r y t o e x p e c t a t i o n s ,a s m a n y a s 3 o u t o f t h e 3 9 M e x i c a n I n d i a n s b e l o ng i n g to t y p e R h t R h ~ w e r eH r - n e g a t i v e . T h i s o b s e r v a t i o n s e e m e d so st a r t li n g t o us t h a t t h e t e s t s w e r er e p e a t e d a n u m b e r o f t i m e s , b u t a l w a y s w i t h t h e s a m e r e s ul t s. R a c e e tal.2 4) h a v e a l r e a d y e n c o u n t e r e d a f e w H r - n e g a t i v e i n d i v id u a l s o f t y p e R h t R h 2

a m o n g C a u c a s i a n s a n d o n t h i s b a s i s p o s t u l a t e d t h e e x i s t e n c e o f o n e o r p o s s i b lyt w o r a r e g e n e s(Rh~ a n d Rhz) in a d d i t i o n t o t h e s i x s t a n d a r d R h g e n e s 2 5 ) .T h e r e a c t io n s d e t e r m i n e d b y t h e r a r e g e n es R h u a n d R h , i n c o m p a r i s o n w i t h t h es i x s t a n d a r d g e n e s a r e g i v e n i n Ta b l e V I I . A s i n d i c a t e d i n t h e t a b l e , th e s e t w o


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B q E N E R ~ P R E C I A D O Z E P E D A , S O N N , A N D P O L I V K A 569

genes are perhaps more appropriately named Rh 'p and~{h 1 2, respectively,

in keeping with the designations for the other Rh genes (26, 27). Accordingto some unpublished observations of Wiener, Sonn, and Unger, the genesRh r and R hl 2, appear to be q uite rare among white individuals, because onlyone Hr-negative individual was found among 132 individuals of type RhlRh2in contras t to the relat ively high incidence of this phenotype am ong MexicanIndians.

Since the effects of genes Rh' and R hl 2 would be practically indist inguish-able among Mexican Indians because they would occur in combination withRhl, or Rh2 or Rh0, in the succeeding discussion, for the sak e of simplicity, weshall consider only gene Rht ~. If we assign to this gene the frequency x, theRh types, their theoretically corresponding genotypes, the Hr reactions to beexpected, and the theoretically expected frequencies of the six possible pheno-types would be as follows:--

R h types GenotypcsR h o RhoRho

f RhtRhtPOa ~ Rh Rho

J R h 2 l ~P.h2 ~Rh2RhoJ

[ RhlRh2| Rhl Rh

RhlRh2 JP,hx 2Rho| Rht 2Rhl( Rhl ~Rht

Observed frequenciesTheore t ica l Abso- Per

H r reactions frequencies lute centP o s i t i v e w 2 1 1 . 1

N e g a t i v e u s 3 9 4 1 . 1

P o s i t i v e 2 u w 7 7 . 4

P o s i t i v e v ~ + 2 v w 9 9 . 5

P o s i t i v e 2 u v + 2 ~ x + 2 w x 3 6 3 7 . 9

N e g a t i v e 2ux + x s 3 3.1

So that ,

u = 0%/-6AH.411= 0 . 6 4 1 0 r 6 4 . 1 p e r c e n t

2uw = 0 . 0 7 4

2 ( 0 . 6 4 1 ) w = 0 . 0 7 4

w = 0 . 0 5 8 o r 5 . 8 p e r c e n t

Rh0 + R h2 = ze + 2vw + v2 = 0 . 1 0 6w + v = ~ = 0 . 3 2 6 o r 3 2 . 6 p e r c e n t

Moreover,

Then,

The theory can be teste d by compu ting the values of the four gene frequenciesw, u, v, and x, and using these values to recalculate the frequencies of the sixphenotypes in order to apply the chi square test .

Since,

u 2 = 0 . 4 11


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570 I N D I V I D U L B L O O D D I F F E R E N C E S I N M E X I C N I N D I N S

Therefore,

-- 26,8 per centand x -- 1 -- (u + v + w) = 1 - 0.967 -- 0.033 or 3.3 per cent

Using these values for the four gene frequencies, the expected absolutefrequencies of the six phen otyp es are as follows: Rh0 = 0.3; Rh~ ( Hr -) =39.0; Rhl (Hr +) = 7.0; Rh2 = 9.8; RhlRh2 (Hr +) = 34.7; and RhlRh2 ( Hr- )= 4 . 1 .

Therefore,

1 - 0.3)* (9 - 9.8)* (36 - 34.7)* (3 - 4.1) 2x * - - - + 0 + 0 + - - + +

0.3 9.8 34.7 4.1

= 2.04, for 1 of freedom

So that , P = 0.16Accordingly, the agreement between the observed frequencies and those

expected under the genetic theory is satisfactory.In conclusion, it should be mentioned that in view of the similarity in dis-

tribution of the Rh blood types among Chinese, Japanese, and American In-dians, it would be reasonable to expect a similar incidence of the genes Rh ppand/or Rhl 2 in these peoples.

S U M M R Y

98 Mexican I ndians were tested for the blood properties A-B-O, A1-A2, M-N,P, Rh~-RhPt-Rh0-rh, and Hr . Of the 98 Indians , 90.8 per ce nt belonged togroup O, 6.1 per cent belonge d to Ax, and 3.1 per cent to group B. There were61.2 per cent of type M, 3.1 per cent of type N, a nd 35.7 per cent of type MN.

Of the 95 Mexican Indians tested with anti-P serum, 21.1 per cent were foundto lack the P aggintinogen.

In tests for the Rh blood types, 48.0 per cent of the Indians were found tobelong to type Rh~, 9.2 per cent to type Rh2, 41.8 per cent to type RhlRh2,and 1 per cent to type Rh0. There were no bloods giving intermediate re-actions. Of the 95 Indians tested for the H r factor 44.2 per cent were foundto lack this property.

The reactions for the Rh blood types and Hr factor were correlated witheach other and the results supported the conclusion of Race e t a l that in addi-tion to the six standard allelic genes and the so called intermediate genes,there is one or possibly two genes having the pr oper ty of determining agglutino-gens which react with anti- Rh ~ and anti- Rh p~sera, but not with an ti-Hr serum.This gene (or genes) appears to be relatively common among Mexican Indians(approximately 3.3 per cent) in contra st to its rareness in white individuals.


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W I E N E R , P R E C I A D O Z E P ED A ~ S O NN , A N D P O L I V K A 571

BIBLIOGRAPHY

1. Wiener, A. S., Sonn, E. B., and Bel~n, R. B.,Proc. Soc. Exp . Biol. and Med.,1943, 54, 238.2. Wiener, A. S., Sonn, E. B., and Belkin,R. B. , J . E xp. Med.,1944, 79, 235.3. Wiener, A. S., Unger, L. J., and Sonn, E. B., Proc. Sac. Exp. Biol. alut Med.,

1945, 58, 89.4. Wiener, A. S., Belkin, R. B., and Sonn, E. B.,Am. J. Physic. Anthropol.,1944,

2 , N . S . , 187.5. Wiener, A. S., Sonn, E. B., and Yi, C. L.,Am. J. Physic. Anthropol.,1944, 2,

~. s., 267.6. Waller, R. K ., and Levine, P., Science, 1944, 100, 453.7. Miller, E. B., and Taguchi, T., data to be published.8. Wiener, A. S., Sonn, E. B., and Belkin,R. B. , J . Immunol. ,in press.9. Boy d, W. C., The blood groups,Tab. Biol., 1939, 17, pt. 2, 148.

10. Wiener, A. S., Blood groups and transfusion, Springfield, Illinois, Charles C.

Thomas, 3rd edition, 1943, chapt er 18.11. Landsteiner, K., and Levine,P., J . Immunol. , 1929, 16, 123.12. Landsteiner, K ., Wiener, A. S., and M atson,G. A., J . Exp . Med. ,1942, 76, 73.13. Race, R. R., and Taylor, G. L.,Nature, 1943, 152, 300.14. Wiener, A. S., Davidsohn, I., an d Pot ter,E. L. , J . Exp. Med. ,1945, 81, 63.15. Wiener, A. S., and Unger, L. J.,Am . J . Clin. Path.,1944, 14, 616.16. Snyder, L. H ., Blood grouping in relation to ]egal and clinical medicine, Baltimore,

The Williams & Wilkins Co., 1929.17. Mat son, G . A., and Schrader,H. F. , J . Immunol. ,1933, 25, 155.18. Levine, P., Matson, G. A., and Schrader, H. F.,Proc. Soc. Exp. Biol. and Med.,

1936, 35, 46.19. Boy d, W. C., Am. J. Physic. Anthropol.,1939, 25, 215.20. Landsteiner, K., and Levine,P., J . Immunol. , 1929, 16, 123.21. Dahr, P., and Zehner, G.,Deutsch. reed. Woeh.,1941, 67, 71.

22. Furuhata, T., and Imamu ra, I.,Japan. J. Genetics,1936, 12, 50.23. Wiener, A. S., Science, 1944, 100, 695.24. Race, R. R., Taylo r, G. L., Cappell, D. F., and McFarlane, M. N .,Nature, 1944,

1 5 3 52.25. M urray, J., Race, R. R., and Taylor, G . L.,Nature, 1945, 1Ni, 112.26. Wiener, A. S., Science, 1944, 99, 532.27. Wiener, A. S. , J . Am. Med. Assn. ,1945, 127 , 294.

INDIVIDUAL BLOOD DIFFERENCES IN MEXICAN INDIANS, WITH SPECIAL REFERENCE TO THE Rh BLOOD TYPES AND Hr...

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