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HUMAN

BLOOD

GROUPS:

THEIR

INHERITANCE

ANDRACIAL SIGNIFICANCE*

DR.LAURENCE

H.

SNYDER

Genetics Laboratory, Dept.

of

ZoOlogy

and

Entomology,

N.

C.

State College

TABLE

OF

CONTENTS

I.

INTRODUCTION

II.

THE INHERITANCE OF THEGROUPS

A.

Mathematical

considerations

of

the

mass

data.B. Individual

family

inheritance

III.

THE DISTRIBUTION

OF

THE GROUPS

A.

The

effect of crossing

on group proportions.B.

The

classification of races.

IV.

DISCUSSION

V.

TECHNIQUE

VI.

SUMMARY

VII.

LITERATURE

INTRODUCTION

This paper

is a

presentation

of some racial

considerations evolving

from

the

heredity

and

distribution

of

the

blood groups among

human

beings.

In

previous papers(Snyder

1924a, b, c; 1925)

the

general

question

of

the

blood

groups

has

been

taken

up.

It

was

pointed

out

that

they

occur

rarely, if

at

all,

in

the

lower animals.

Recently

evidence

has beenbroughtfonvard

that

theymay

occur

in

the

anthropoidapes(Land

steiner

and

Miller

1925, Hirszfeld,

L.

1926).

In

the hmnan

race the

~ S ;

S ' - " ~

~ O >

blood

groups

occur

as

fixed bio-chemical conditions,

subject

to the

laws

. ~

I

of

heredity.

As

such

they

provide

a

method

of

studying

racial origins

'"

~ . , ;

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oci

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....

01-01-01

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01-01

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--..--

---.--

.--.

and

relationships.Blood

groups

were first discovered

by

Landsteiner

(1900)

and Shat

tock (1900) working

independently.

They

found

that

there

were definite

substances

in

the

serum

of some bloods

that

would

agglutinate,or

clump,

the

cells of

certain other

bloods.

On

the

basis of

the

agglutina

1

i

ting reactions, blood

can be

classified

into four

groups.

Two

classifications.

the

Moss

andthe

Jansky,

are

in

current

use.

The

Jansky

classifi

8

~

E

8 8

cation is recognized

ashaving

priority

rights,

and

is

used

here.

The

.E

~

.E

~

O'

~

iO'

>.

"

>.

scheme of

this

classification is given

in

table

1.

<

8 ~

«:

" ' ~

'"

"

'

<<

~

.

"

'.

§

~

The

specific

substances in theserum, causing

the

clumping

of

the

cells,

Il

.r:

O

~

'"

.r:

O

"

1i

"

Eo

8

. ~

Eo

1;;

"

I'l

.r:

O

.<l

'"

1i

are

known as

agglutinins.

The

equally

specific

receptors

in the red

cells

'"

E'"

Eo Eo

E

"

.

0

1 < 1 ~

0

1<1

~

'"

0

~

0

~

1<1

•

gj

oJ

!:lObO

b/:

.

•

00

~

·Published

with

the

approval

of

the

Director

of

Research, as

paper

no. 7

of

the

>>>

!&

!&!&

""

>

<<

«

<

~ ~

~

<<<

<:

"

Journal

Series.

!t33

AM.

].

PHYS.

ANTHROP

••

1926, Vol.

IX, No.2.

I

.J

~ 3 4

LAWRENCE

H.

SNYDER

TABLE

l.

SCHEME

OFTHE

JANSKY

CLASSIFICATIONOFBLOOD

GROUPS.

Group

Effect ofserumCapacity ofcells

I

Agglutinates

cells of

II,

III,

IV.

Notagglutinatedby any

serum.

II

Agglutinates

cells of

III

and

IV. Agglutinated

by

serum

of I

and

III.

III

Agglutinates

cells of

II

and

IV. Agglutinated

by

serum

of I

and

II.

IV

Does

not

agglutinate

any

cells.

Agglutinated

by

serum

of I,

II

and III.

are

designated

agglutinogens. Agglutinogen A is

the

one found ingroup

II.

Agglutinogen B is

the

one

found

in group

III.

Group

IV

contains

both,

while

group

I

contains

neither.

The

four blood groups,

thought

at

first

to

bepresent

only

in

diseasedpersons, were soon found

to

be

of general occurrence.

Within

each race

there

occur fairly definite

and

characteristicproportions

of

the

fourgroups. As few

unit

characters

areyet

known

in

man,

the

blood groupsform a welcome

addition

to the

list, since

they can

b€

accurately

studied

on a bio-chemical basis,

and,

depending

as

they

apparently

do, on two

dominant mutations

at

the

same

genetic locus,

they

afford a

means

of

investigating

racial origins

and

relationships

by

correlating

the

frequencies of

the

hereditary

factors concerned.

Observations

on

the

proportions

of

the

four

groups

among

peoples

have

been

made by

physicians

in

all

parts

of

the

world.

The

occurrenceof

groups

having

from

its very nature presented

itself first

as

a medicalproblem,

it

is

natural

that

the practical

applications

have

been chieflyin blood transfusion

and

medico-legal cases, so

that

until very

recentlyscarcely

more

than

a

paperor two

on

the

subjecthasbeen

found outside

the

medical

literature.Thus

far

observations

have

been reported

individually,

mainly

in

the

form of figures,

and in only

a few cases has

any

attempt

been

made

to

correlate

the

observations

withother

reports.

It

is of

interest

to

compare the

results found

on

the

basis of

the

bloodgroups

with

the

known

racial relationships

based

on anthropologicalcriteria.

Such

a procedure, however, as I

have already pointed

out,

(Snyder

1925)

must be undertaken

with extreme

caution.

The

rarity

of

pure

races,

the

fact

that

racial

characters are

in general blending in

inheritance

and

not sharply

defined

in

succeeding

generations

following

race

mixture,

while

the

blood

group

factors

are

unit

factors

with

com

plete

dominance,

retaining theiridentity

from

generation

to

generation,

the

great

racial divergences

that

have

obviously occurred since the

blood group

factors

became integral

parts

of

the

hereditary

complex,

and the

possibility of linkage

of

the

blood

group

factors

with

suscepti

bility

to

specific disease, all

combine

to make the

problem

a difficult one.Although

most

of

the

early investigators

of

the

blood

groups

reported

the

frequency

of

the

groups in

the

peoples

among

which

their

observa

tions

were made,

the

first work

undertaken

for

the

express

purpose

of

INHERITANCE

AND SIGNIFICANCE OF

BLOOD

GROUPS

~ S 5

studying

the

group

frequency in

various

parts

of

the

world

was

that

ofHirszfeld

and

Hirszfeld (1919).

These authors took advantage

of

the

concentration of

nationalities

in

the

Macedonian

battlefront

during

the

war,

andstudied

five

hundred

members

of each of sixteen differentpeoples.

Theyreported

the

frequency of occurrence of each of

the

four

groups in each of

the

nationalities.

They

found

that

the

sixteen peoplescould

be

roughly

divided

into three

types, as follows.

One

type

contained a high

percentage

of agglutinogen

A,

and

a correspondingly lowpercentage of

B.

This

type

comprised

nearly

all

the

European

countries,

and

was

therefore

named

the

European type.Anothertype contained

ahigh frequency of B

and

a correspondingly low

percentage

of

A.

The

Mongolian

and Ethiopians

fell

into this

category,

and

it

was

accordingly

named

the

Asio-African

type.

The third

type,

which

the

authors

called

the

Intermediate type, contained approximately equal

proportions

of A

and

B,

and

comprised Russians,

Turks,

Arabs

and

Jews.

The

Hirszfelds suggested a "biochemical index" for

reporting

ongroup

distribution.

This

was

to

bearrived

at

by

dividing

the

percentage

of

A

in

a given

race

by the

percentage

of

B; inother

words,

by

dividing

the percentage

of

groups

II

and IV

(which

contain

A)

by

the

percentage

of

groups

III

and

IV

(which

contain

B).

It

will

thusbe

seen

that

a

race

having twice

as

much

A as B would

have

a bio-chemical index of

2,

while a

race having

only

half

as

much

A

as

B would

have

an

index of0.5.

Of

the

Hirszfelds'

three

types,

the

European

type

had

an

index of

2.5-

4.5.

the

Asio-African

type

showed

an

index of 0.5-1.09,

and the Inter-

mediate

type varied

from1.3

to

1.8.

Later reports

from

other

investigators

have

given indices of

intergrading magnitude,

so

that

the

three

types

are

no

longer

sharply

defined.

Furthermore the

index is

notbased

on

sound mathematical

reasoning,

and

must

be discarded entirely,

as has

already

been

pointed

out

(Snyder

1925,

Bernstein

1925).

In

its

placemust

beused

the

frequency of each of

the

factors concerned.

I

The

work of

the

Hirszfelds

stimulated

a

large

number

of workers invarious

parts

of

the

world

tostudyother

peoples,

and

b i o ~ c h e m i c a l

indices of

various

nationalities

have been determined

everywhere.

In

l

general

authors have been content

to

report

the

frequencies of

the

groups,

the

bio-chemical index,

and

thetechnique

used,

without

at-

tempting

to

draw

conclusions.I use in

this

paper

the

terms "nationalities" or

"peoples,"

where

other

authors

have

used

the

term"races." This

distinction

mustbe

carefully

drawn

from

the

anthropological

standpoint.

The study

of

the

American

Indians

reported

in

this

paper

is

the

nearest approach

to

a

true

"racial"study

of blood groups.

~ 3 6

LAURENCE H.

SNYDER

Schutz

and

Wolisch (1924), from a

study

of

the

blood

glOUpS

of Ger

mans,

note variations in the

percentages of

the

four

groups

in different

parts

of

Germany.They

conclude

that

the

point

is

not

proven

that

the

group

frequencies

are

characteristic

of

the

race.

For

the

various

parts

of

Germany they have

only small

numbers,

however,

andthe

averageof all

their

figures is

in

close

agreementwith

those

of

other

investigatorsin

Germany.

Moreover,

the

German nationality

is

not

a racial unit.

In

general, where sufficiently large

numbers are

used,

the

results

of different

investigators

in

the

samecountry are in

close agreement,

and

Isee

no

necessity of considering seriously

the

objection

of

Schutz and

Wolisch.

It

has

been

amply

established

that

the

agglutinogens

are

present

at

birth,

but that

the

agglutinins

maybe

delayed

in

their

appearancefor several

months

(von

Dungern

and

Hirszfeld 1910, Hirszfeld

and

Hirszfeld 1919,

Jones

1921,

Robertson,

Brown and

Simpson 1921, Unger1921, de Biasi 1923,

Kirihara

1924).

Tebbuttand

McConnel

(1922),

noting

this, suggest

that

the

agglutinogens

probably

preceded likewise inevolution.

This

assumption,

however, is

not warranted

by

the

fact

that

prim

itive or

isolated peoples show a

very

high

percent

of

group

1.

As will

appear

later,

group I reaches

more

than

90%

among

full-blooded Amer

ican Indians.

The

peoples which show

more

than

50%

of

group

I are

the

American

Indians,

the

Filipinos,

the

Melanesians of

New

Guinea,

the

South

African

natives,

the

Australian

aborigines,

and the

Icelanders: all island folk or otherwise isolated.

In

order

that the

distribution

of

the

groups

may be used in

studies

of

racial relationships,

it

must be

shown

that

they are hereditary,

and

the

mode

of

their inheritancemust bedemonstrated.

Only

then

can thefrequency

of each

factor

be

computed,

and

a classification of races bemade.

THE INHERITANCE

OF

THE

GROUPS

The

heredity

of

the

groups

has

long

been assumed

to

be

on

the

basis

of

two

independent

pairs

of factors,

the

agglutinogens (A

and

B) being

dominant to their

respective iso-agglutinins (a

and

b).

This

hypothesis,proposed fifteen

years

ago

by

von Dungern

and

Hirszfeld (1910), was

based

on

inheritance

studies

of 72 families, comprising 348 individuals.

The

hypothesis was

accepted

and

confirmed

by

all

later

investigatorswho

made inheritance

studies

on

the

blood

groups

of families (Lear

month

1920, Weszeczky 1920,

Ottenberg

1921,1922,1923,

Keynes

1922,

Tebbutt and

McConnel

1922, Jervell 1923,

Dyke and Budge

1923,

INHERITANCE

AND

SIGNIFICANCE

OF BLOOD GROUPS

~ 3 7

Kirihara

1924,

Pluss

1924,

Mino

1924, Hirszfeld, Hirszfeld

and Brokman

1924)

with

the

exception of

Buchanan

(1923) whose

lack

of

grasp

of

the

whole

situation has already

been

referred

to

(Snyder 1924b).

More

than

six

hundred

families

have

been

studied altogether

by

these various

investigators.Recently, however,

Bernstein

(1925)

has

denied

the

theory

of

two

in-

dependent

pairs

of factors,

and has

proposed

in

its

place

the

hypothesisthat the

blood

groups

are inherited

as a series of

threemultiple

allelomorphs.

He

bases

thisnot

on

any

study

of family inheritance,

but

entirely on

mathematical

considerations of

the

grouppercentages

invarious peoples as

reported

by

other

invesitgators.

He

assumes

that

the

agglutinogens (A

and

B)

are both dominant

to

the

same

recessive, which

he

calls

R.

The

heterozygote

AB forms

group

IV.

This theory takes

the

agglutinins

out

of

hereditary

control, leavingonly

the

agglutinogens

to

be

inherited.

It

requires a difierent

immuno-

logical

explanation

(see

Bernstein

1925, pp. 246-248), which, however,could

probably be assumed

if

the

facts

of

the inheritance

of

the

groups

really

support

the

hypothesis.

The

genetic formulae of

the

four groups

for

both

hypotheses

are

given

intableII.Our

American

usage would

require

A,

a'and

a

as

the

symbols for

the

series of allelomorphs,

but

the

agglutinogens

have

been

known

as

A

and

B

for.

so long

that

it

seemsadvisable

to

retain

these

symbols, using R

to

represent the

recessive.

TABLE

II.

GENETIC FOItMULAE

OF

THE FOUR

BLOOD

GROUPSTwo

ThreemultipleThree

multiple

Group

independent

allelomorphs Group Two

independent

pairs

of

factors

allelomorph::;

pairs

of

factors

I

aabb

RR

III

aaBB, aaBb

BB,

BR

II

AAbb,

Aabb

AA,

ARIV

AABB,AaBB,AABb,

AaBb

AB

The

facts

upon

which

the

hypothesis

of

three multiple

allelomorphs isbased

are

these. Since A

and

B

areseparatemutations,

group

IV

(AB)must

originate

by

unions

of A

and

B.

Knowing

the

proportions

ofgroup

II

(A)

and group

III

(B)

in any

race,

the

proportion

of

groupIV

(AB)

can be

calculated.

On

the

basis of

two

independent

pairs

offactors,

this

would

be

done as

follows.

Let

p

equal

the

frequency

of

A,

q equal

the

frequency

of a, r

equal

the

frequency

of B,

and

s

equal

the

frequency

of

b.

Then

p

+

q

=

1,

r

+

s

=

1.

The

possible

combina-

tions of

these

factors

are

expressed

by the

equation

(p

+

qF

.(r

+

sF

=

1.

Fromthis

the

following

combinations

may

be

formed.

AA

2Aa

aa 2Bb

2AABb 4AaBb

2aaBb

p2

2pq

q'

2rs

2p2

rs

4pqrs

2

q

2

rs

BB

AABB2AaBB

aaBB bb

AAbb 2Aabb

aabb

r'

p'r'

2pqr'

q'r' s'

p's'

2pqs'

q2

s'

J

Category:	School Work
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Upload Date:	11/04/2010
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