H E R E 66-6

OBSERVATIONS ON THE BLOOD

GROUP RECEPTOR PI AND ITS
DEVELOPMENT I N CHILDREN
By AAGE H E I K E N
STATE INSTITUTE FOR BLOOD GROUP SEROLOGY, STOCKHO1.M GO, SWEDEN

(Received May 13th, 1966)

I. INTRODUCTION

B ASED on the fundamental observations of LANDSTEINER and LEVINE

(1927),LEVINEet al. (1951),SANGER(1955), MATSON et al. (1959)
and their own extensive investigations, KORTEKANGASet af. (1965) have
recently advanced a tentative model for the genetical pathways of the P
blood group system.
According to this model the hypothetical precursor substance, p, is
acted on successively by the genes Z, Y and P,. Z is a dominant and very
frequent gene, which converts p to Pk substance. Y is also frequent and
dominant in effect but not necessarily linked to Z; it changes Pk to P
substance. In the absence of Z, i.e., if the genotype zz occurs, the pre-
cursor substance is not converted; and in the absence of Y, i.e., if the
genotype yy occurs, the Pk substance is not converted. The P and Pk
substances possess antigenic activity while the precursor substance is
lacking such activity. P, is capable of acting on both P and Pksubstance
but not on p substance. If P substance is available, the genotypes P,P,
and P,P, produce a P, antigen but leave some P substance unchanged
(phenotype PJ, the genotype P,P, is unable to change the P substance
and the phenotype P, possess therefore only the P antigen. If Pk sub-
stance is available, the genotypes P,P, avd P,P, produce a P, antigen but
leave some Pk substance unchanged (phenotype P?), the genotype P,P,
is unable to change the Pk substance and the phenotype P/ possess
therefore only the Pk antigen. The remaining phenotype characterized
by lacking antigens is designated by p. Y and P, are not closely linked.
This basic scheme corresponds to the occurrence of three antigens, P,,
P and Pk,defined by three antibodies anti-P,, anti-P and anti-Pk. Anti-P,
is present in the serum of some Pa individuals and in the serum of cer-
84 AAGE HEIKEN

tain species of animal. Anti-P and anti-Pk may be isolated from the
serum of p individuals by means of absorption techniques. - It should
be noted that Pkand p phenotypes are extremely rare, and it has not yet
even been possible to estimate their frequencies.
A characteristic feature of the P, antigen is its variation in strength;
the practical difficulties of this fact is pronounced by the gradual devel-
opment in strength of the P, antigen in children, and the scarcity of
specific potent antisera. Already in their first works on the P types,
LANDSTEINER and LEVINE(1927, 1930, 1931) attempted a subdivision
according to the grades of reaction. Subsequently, several research
workers have studied this problem and tried to show that the receptor
strength is inherited. On the basis of a comprehensive family material,
HENNINGSEN (1949 a, b; 1952) stated that the P, strength is determined
by several closely related allelic genes, all of which dominate over the
gene conditioning to the absence of the P, antigen. A thorough analysis
of HENNINGSEN’S data was performed by FISHER (1953), but he attribut-
ed the variations in antigenic reactivity mainly to dosage effects.
Although many extensive family studies have been carried out (cf.
Table 14),the frequency of the P, antigen is so high that up to the pres-
ent only 164 P,-XP,- matings with 429 children have been tested.
Owing to the relationship between the amount of material for which the
frequency of exceptions from the theory of inheritance is exactly known
and the biological reliability with which paternal exclusions can be
made, there exists obviously a grelat need of further studies. One of the
purposes of the present paper is to report the results of an additional
family material.
It has often been quoted in the literature that the P, antigen is con-
siderably weaker in infants than in adult persons; but in spite of the
importance of this question in paternity tests and other blood group
genetical investigations, little is known about the development of the P,
antigen after birth. The family material mentioned above, and a large
sample of mother and child pairs P,-tested in connection with medico-
legal investigations, afforded an opportunity to scrutinize this problem
more closely.

11. MATERIAL AND METHODS

1. Material
The material consisted of 101 unrelated families with 231 children,
who voluntarily contacted this laboratory in order to participate in the
 BLOOD GROUP RECEPTOR P, 8.5

TABLE 1. Titration experiment with the human anti-P, serum used in

the investigation.
The technique applied is described in the text.

Cells 1 Undiluted 1:2 1:4 1:s 1:16

Pl +
(S. B.) ~ 3.5 3.0 2.0 0.5 -

Pl +
(R. 8815*) ~ 3.5 3.0 2.0 -3 -

p1-
(I. H.) - - - -

* Established to possess a strong P, receptor on the basis of experiments with a very weak
anti-P, serum of human origin.
The strength of agglutination was indicated as follows:

++++ L . 5 1
+++($1 f strong agglutination
+++ '3.0)

++
++(+) 2'5} medium agglutination
2.0

+ weak agglutination
(+) 0.5
- 0 no agglutination

investigation, and a series of 3609 mother-child pairs, which were ran-

domly selected from cases of disputed paternity. Further details regard-
ing the composition of the samples will be given when describing the
results obtained.

2. Serological techniques
The serological investigations were carried out during the years 1961
-1965. They included two agglutination tests and one absorption ex-
periment. Two different anti-PI sera were employed: one of human
origin (obtained from a A,B P,- test donor), and one of animal origin
(obtained from a non-immunized horse). These two sera were used
throughout the investigation. The test donor was bled every third
month. The horse serum originated from a single batch. The inactivat-
ed, unabsorbed sera were stored frozen at -20' C .
The equine anti-P, serum was prepared by absorption. Equal volumes
of washed, packed erythrocytes from A,B P,- and OP,- test donors
86 AAGE HEIKEN

TABLE 2. Titration experiment with the horse anti?, serum used in

the investigation.
The technique applied is described in the text,

Time of
Cells reading, Undiluted" 1:2 1:4 1:s 1:16 1:32 1:64
minutes

P1+ 10 3.0 2.5 2.0 1.0 - - -

(S. B.) 20 3.5 3.5 3.5 3.0 1.0 - -
35 3.5 3.5 3.5 3.0 1.5 -? -

Pl +
(R. 8815*)
10
20
3.5 3.5 3.0 2.0 0.5 -
-? -
-
3.5 3.5 3.5 3.0 1.5
35 3.5 3.5 8.5 3.5 2.5 0.5 -
P1- 10 -? - - - - - -
(I. H.) 20 0.5 - - - - _ -
35 0.5 - - - - - -

* Established to possess a strong P, receptor on the basis of experiments with a very weak
anti-P, serum of human origin.
* * The serum has been absorbed, but not yet finally adjusted for use.

were mixed, and this mixture was added to an equal volume of horse
serum and allowed to stand two hours at 5' C. The procedure was re-
peated three times, but the final absorption was carried out with only
half the volume of packed cells. Before use, the absorbed serum was
standardized by titration against known P,+ and P,- test cells and, if
necessary, carefully adjusted through addition of saline. An example of
the titre of the undiluted serum is given in Table 2. For comparison, a n
example of the titre of the human anti-P, serum used in the investiga-
tion is presented in Table 1.
The two agglutination tests were carried out at a controlled tempera-
ture of 17' C. The tests with the human anti-P, serum were performed
according to the test tube technique, while the tests with the horse anti-
P, serum were performed according to the slide method. The absorption
experiments were carried out with the horse anti-PI serum, the readings
were made after a period of incubation of two hours at 5' C. All read-
ings were made microscopically-when using the slide method after 10,
20 and 35 minutes.
The difficulties met with in P typing have been emphasized by many
authors ( c f . , e.g. MOHR, 1963), and, during this study, it has been neces-
sary from time to time to carry out much more profound investigations
 BLOOD GROUP RECEPTOR P, 87

TABLE 3. The P, blood group frequencies obtained in the mother-child

com binations.
PI + PI -
Age of
children
No.
tested
Mothers I Children Mothers 1 Children

abs. I % 1 abs. 1 % abs. I % I abs. I %

0-3 months 451 349 77.38 323 71.62 102 22.62 128 28.38
4-7 9, 1371 1039 75.78 971 70.82 332 24.22 400 29.18
8-11 ,, 695 547 78.71 514 73.96 148 21.29 181 26.04
1-2 years 734 551 75.07 530 72.21 183 24.93 204 27.79
3-14 ,, 358 282 78.77 284 79.33 76 21.23 74 20.67
Total
On the basis of the blood group frequencies obtained in mothers, the following
gene frequencies were estimated: P,=0.5173; P,=0.4827.

than those described above in order to arrive at a final conclusion

whether a weak P, antigen did occur or not. In such cases different
haemagglutinins and varying methods have been employed, and, owing
to the importance of using fresh corpuscles, several blood samples from
the same person have often been tested. However, while the grouping
results given in Table 3 are based on the total amount of information
obtained, the tables dealing with the antigen strength are solely derived
from the agglutination results obtained with the two antisera mentioned
above.

111. THE DEVELOPMENT O F THE P, RECEPTOR

In the course of testing blood specimens from 403 foetuses between
the age of twelve and twenty-eight weeks, IKINet al. (1961) observed
that the incidence of the P, antigen was considerably below the adult
level, but that it (rather unexpectedly) occurred more frequently in the
younger than the older foetal samples. A tendepcy for the strength of
the P, antigen to decrease with increase in foetal age was also observed.
Among the older foetuses (with a crown-rump length of more than 10
cm) 78.86 per cent were found to be PI-.
HENNINCSEN (1949 a) tested fourty-one newborn children and observ-
ed that 41.4 per cent were PI-.In a group of 333 infants between birth
and the age of twelve months the corresponding percentage was 23.4,
which is somewhat above the adult level in the Danish population.
Moreover, in the former group 14.7 per cent and in the latter group 1.8
per cent of the infants could not be determined with certainty.
TABLE 4. Agglutination strengths in the mother-child combincctions obtuined on the basis of tests with the
horse anti-PI serum.
1 Total 0 - 1.5 2.0 - 2.5 3.0 -3.5
__
Age of children Moth- Mothers 1 Children 1 Mothers ! Children Mothers 1 Children
Child-
ren abs. % abs.
I
: I 1 % %

0- 3 months 349 323 20 5.73 121 37.46 18 5.16 74 22.91 311 89.11 128 39.63
4-7 ,, 1039 971 32 3.08 327 33.68 80 7.70 254 26.16 927 89.22 390 40.16
8-11 ), 547 514 20 3.66 143 27.82 35 6.40 124 24.12 492 89.95 247 48.05
1- 2 years 551 530 13 2.36 98 18.49 45 8.17 127 23.96 493 89.47 305 57.55
3-14 ,, 282 284 7 2.48 25 8.80 21 7.45 46 16.20 254 90.07 213 75.00
-- - -
2622 92 7.19 2477 48.93

TABLE 5. Agglutination strengths in the mother-child combincctions obtained on the basis of tests with the
human anti-PI serum.
I 1 Total 1 0-1.5 I 2.0 - 2.5 3.0 - 3.3
Children 1 Mothers Children Mothers 1 Children
-
% % abs. %
-
0- 3 months 349 323 110 31.52 208 64.40 63 18.05 52 16.10 176 50.43 63 19.50
4- 7 ,, 1039 971 359 34.55 685 70.55 185 17.81 124 12.77 495 47.64 162 16.68
8-11 ), 547 514 186 34.00 335 65.18 97 17.73 78 15.18 264 48.26 101 19.65
1- 2 years 551 530 187 33.94 296 55.85 91 16.52 88 16.60 273 49.55 146 27.55
3-11 ,, 282 284 100 35.46 137 48.24 54 19.15 47 16.55 128 45.39 100 35.21
34.03 17.70 389 I 14.84 1336 21.82
 BLOOD GROUP RECEPTOR P, 89

1. Investigation of mother-child combinations

In the present investigation the mother and child pairs were divided
into five classes according to the age of the children ( c f . Tables 3-5).
Subdivision of the fifth class (3-14 years) would have been very de-
sirable, but the number of children tested was considered to be too small
to justify this. The grouping results are summarized in Table 3.
The differences between the five classes of mothers are not signifi-
cant, but the difference between the mothers and their children are so
in the classes I (~'=3.941; d.f.=l; P=O.O5-0.025), I1 (f=8.617; d.f.=
= 1 ; P=O.O05-0.001) and I11 ( ~ ' ~ 4 . 2 4 6d.f.=l;
; P=0.05-0.025), i.e.,
in all classes which include children under a year old. When the dif-
ference between children and mothers is analysed by testing each class
of children against the total group of mothers, class I11 does not differ
significantly (~*=1.990;d.f.=l; P=O.20-0.10), while class IV does
(f=6.665; d.f.=l; P=0.01-0.005). In the total material of mothers
23.30 per cent were found to be PI-. Previous studies on the P blood
group frequencies in the Swedish population have been reported by
JONSSON(1959), who found 23.90 per cent to be PI-, and by BECKMAN
(1959), who obtained a somewhat lower frequency (21.19 per cent).
While the frequencies achieved in the present study closely agree with
those reported by JONSSON( l . c . ) ,they deviate significantly from those
given by BECKMAN (lx.).
The quantitative results of the investigation are shown in Tables 4
and 5. The heterogeneity-XPanalyses of the results were performed by
separating the macroscopic agglutination reactions from the microscopic
agglutination reactions. No significant heterogeneity could be found be-
tween the classes of mothers either using horse anti-P, serum or human
anti-P, serum; but the differences between the mothers and their chil-
dren were highly significant in all five classes. However, as appears
from the tables, the tendency of giving weak reactions was much more
marked in the younger than in the older children.
The differences in the strength of reaction observed between mothers
and children are possibly accentuated by the fact that the blood samples
from the children have been taken from the capillaries, while the moth-
ers have been bled by venipuncture. However, most children aged two
years or more are handled in the same way as adults by the Swedish
physicians. It seems therefore most unlikely that this source of error
should have played any role for the results obtained in the fifth class of
children.
90 AAGE HEIKEN

2. Investigation of families
In the family material, the following distribution was obtained with
the horse anti-P, serum:
Strength of agglutination
n weak medium strong
Mothers 80 2.50 % 2.50 % 95.00 %
Fathers 78 2.57 70 1.28 % 96.15 %
Children 182 3.85 70 10.44 70 85.71 %

The following distribution was obtained with the human anti-]?,

serum :
Strength of agglutination
n weak medium strong
Mothers 80 15.00 % 15.00 % 70.00 %
Fathers 78 15.39 % 20.51 % 64.10 %
Children 182 21.08 % 10.78 % 58.24 "/o

As is seen, there is a clear tendency for the children to give weaker

reactions than their parents. However, since the children constitute a
rather heterogeneous group with regard to age, fifty-nine families were
selected from the material in which all the children were at least seven
years old at the time of the investigation. The distributions within this
group did not vary much from those given above. When tested with the
horse anti-PI serum 2.08 per cent of the mothers, 2.17 per cent of the
fathers and 4.63 per cent of the children were found to give weak reac-
tions. When the human anti-PI serum was used the corresponding per-
centages were as follows: 12.50, 13.04 and 20.37. - It ought to be noted
that the family sample is rather small for quantitative studies and too
far-reaching conclusions cannot be drawn from these results; none of
the differences are statistically significant.
If the strengths of agglutination observed in the family sample are
compared with those presented in Table 4 and 5, it is recognized that
the reactions observed in the mother-child sample are somewhat
weaker. The deviation can certainly be explained by the fact that while,
in the mother-child material, about 75 per cent of all the blood speci-
mens have been submitted to the Institute from all parts of the country
and often mailed under less favourable conditions, most of the blood
specimens from the families were taken at the Institute and tested im-
mediately.
 BLOOD GROUP RECEPTOR PI 91

50
5
s
Z
Lu
kL!u
P
40

30

20

10

0- 0,6- 1,l- 1.6- - 3J-

0,5 1,O 13 2 ,O 35
STRENGTH OF AGGLUTINATION

Fig. 1 . The variation in strength of agglutination in 3609 mothers based on the

average values of two anti-PI sera.
92 A A G E HEIKEN

3. Conclusions
Considering the results given in Tables 3-5, it appears justified to
conclude that P, determination cannot be made with certainty in chil-
dren younger than one year. Even if present the PI antigen is still in-
completely expressed in many children aged three years. The results
obtained at the family investigations indicate that the age for full devel-
opment of the P, antigen might be as much as seven years or more.
However, the establishment of an exact age for the complete develop-
ment of the antigen requires a more comprehensive study of children
older than three years.

IV. THE STRENGTH OF THE P, RECEPTOR IN ADULTS

The variation in strength of agglutination within the group of mothers
in the mother-child series, based on the average values of both the two
antisera used in the investigation, is graphically presented in Fig. 1. On
the basis of titration experiments, HENNINCSEN (1949 a) ascertained
that, in adult individuals, the P, activity shows approximately a curve
of normal distribution. The results obtained in this study do not con-
tradict this statement. It seems quite possible that titrations would have
disclosed heterogeneity in at least the higher score groups.

V. THE INHERITANCE OF THE P, RECEPTOR

The phenotypes Pkand p are so infrequent that they can be ignored in
statistical studies of the inheritance of the P, antigen. In the present in-
vestigation, the family material was analysed according to the method
devised by SMITH(1956). The results are given in Tables 6-13.
The gene frequencies used for the analyses shown in Table 8 for
P,+ XP,+ matings and in Table 11 for P,+ XP,- matings are based
on the blood group frequencies obtained among mothers in the inde-
pendent mother-child sample (see Table 3). Since, however, BECKMAN
(h) has reported a significantly lower PI- frequency than that found
in the sample of mothers, the goodness of fit was checked by applying
the gene frequencies given by him; but no significant difference could
be disclosed by application of these frequencies either.
As a final check all the xB values for the comparisons presented in
Tables 7 , 8 , 10 and 11 have been summarized in Table 13 to give a single
xp; the corresponding degrees of freedom have also been added. As ap-
 BLOOD GROUP RECEPTOR P, 93

TABLE 6. Observed distribution of families o f mating type P,+XP,+.

No. of children in family tested with anti-P,
No. of children in
Total
family of group P,- 2 3 4

I 0 I 8 29 13 1 1 51

'
1
2

Total no. of families 8 32

3 5
1

19
1

2
1 : 61
I I
Total no. of families
with at least one P, -
child 0 3 6 1

LNo. of children
No. of P, - children
in fam,ly
No. of families
Obs. 1 Exp. I Variance

1 0 0 0.000 0.000
2 3 3 3.429 0.366
3 6 7 7.782 1.578
4 1 1 1.463 0.420

No. children Total no. of 1 No. of families with a t least one P,- child
in family families 1 Obs. I Exp. I Variance

1 8 0 0.848 0.758
2 32 3 5.941 4.839
3 19 6 4.655 3.515
4 2 1 0.580 0.412

x2 = 2.024a/9.524 = 0.430; d.f. = 1; P = 0.7 - 0.5

94 AAGE HEIKEN

TABLE 9. Observed distribution of families of mating type PI+ X Pl-.

1 No. of children in family tested with anti-P,
No. of children in
Total
family of group P, - I' 1 2 3 4 5

0 I 1 10 4 1 1 1
I
17

1
2
3 8
4
2
2 I
Total no. of families 1 4 22 8 1 1 i 36

Total no. of families

with a t least one PI-
child 3 12 4 0 0 19

No. of families
in Iamily

1 3 3 3.000 0.000
2 12 16 15.996 2.664
3 4 6 6.856 1.960

x2 = 0.852'/4.G24 = 0.157; d.f. = 1; P = 0.7-0.5

TABLE 11. Mating type Pl+ XPl-. Analysis of families with and
without children of group PI-.
No. of families with at least one P,- child
No. of children Total no. of
in family families Obs. I Exp. 1 Variance

4 3 1.302 0.878
22 12 10.743 5.500
8 4 4.558 1.960
1 0 0.611 0.238
1 0 0.631 0.233
Total 1 36 I 19 I 17.845 I 8.809

x2 = 1.155'/8.809 = 0.151; d.f. = 1; P = 0.7 - 0.5

 BLOOD GROUP RECEPTOR PI 95

TABLE 12. Distribution o f families o f mating type PI- X PI-.

Total no. of children

of group
in family lested
PI+ 1 p,-

1 0 0 0
2 1 0 2
3 1 0 3
4 2 0 8

I Total I 4 1 0 1 1 3

No. of P,- children, given m, PI+ x P,- 0.157 1 0.7-0.5

No. of Pl- children, given M , P1+ x P1+ 1.185 1 0.3-0.2
m,, given n, PI+ x PI- 0.151 1 0.7-0.5
M c , given N c p1+ x p1+ 0.430 1 0.7-0.5

pears from the table, also for the total material, the figures observed
agree closely with those expected; - and no exceptions to the theory of
inheritance were encountered (cf. Table 12).

VI. DISCUSSION
The application of a blood group system for exclusion of paternity
depends in the first hand on its exclusion capacity and the biological
and technical reliability with which the exclusions can be made. For
practical reasons, however, it is also of importance that the blood group
antigens should be developed at birth or soon thereafter.
Regarding the P blood group system, the theoretical exclusion rate
can be estimated to 2.81 per cent on the basis of the gene frequencies
obtained in this investigation. As compared with the other blood and
06 AAGE HEIREN

NO. of Pi- x
N O . Of
' Cliildren of pi-
pi- matings
x
Author
-

families Pi - matings
PI- I PI+

563 50 173 4 DAHR(1942)

40 2 7 0 JUNCMICHEL (cit. HENNIXGSEN,
1952)
12 2 7 MOHARRAM (1942)
50 1 2 BERGER (cit. HACKER, 1964)
12 0 0 APPELHOFF (cit. HACKER, 1964)
45 9 13 KORBEL(cit. HACKER, 1964)
85 2 6 SANGER el al. (1949)
89 7 13 BRENDEMOEN (1952)
32 1 3 GROSJEAN(1950, 1952)
304 17 38 HENNINCSEN (1952)
306 9 17 RACEet aI. (cit. RACEand SANCER
1962)
104 6 11 0 WIENER(cit. RACEand SANCER,
1962)
50 0 0 OERTEL(cit. HACKER, 1964)
4 0 0 PETTENKOFER (cit. HACKER, 1964)
176 12 20 LINNET-JEPSEN et a!. (1958)
96 7 13 GALATIUS-JENSEN (1960)
134 12 24 KERDE(cit. HACKER, 1064)
195 14 41 HACKER (1964)
18 1 1 LINNET-JEPSEN (1965)
125 8 27 MOHR(1966)
101 4 13 Present investigation
2541 1 164 429 5*

* Four of these children have been stated to be extra-marital.

serum group systems used in genetico-legal investigations in Sweden,

this exclusion rate must be designated as low. However, due to the fact
that about 68 per cent of the blood samples from children involved in
cases of disputed paternity are submitted to the Institute before the
children have aged one year, the practical utility of the system is even
more limited than could be expected from the theoretical exclusion rate
(cf. Table 3). In considering the use of the P system in paternity cases,
it must be realised that paternal exclusions can be established only on
the basis of absolutely clear-cut'results.
 BLOOD GROUP RECEPTOR P, 97

Table 14 summarizes P,- XP,- matings of all those family studies

on the P blood group system, which, as far as is known to the author,
have been published. Four of the five children which represent excep-
tions from the mode of heritance have been stated definitely to be extra-
marital. Thus, the genetical security of the system can be estimated to
99.4 per cent, given that only one child of each mating is considered to
be informative. If all children of each mating are considered to be
equally informative, the genetical security of the system can be estimat-
ed to 99.8 per cent. Attempts to estimate the security rate more exactly
have not been made, as the number of children of each mating is not
quoted by all authors. In the first case, the upper 95 per cent confidence
limit is 0.029, in the second case 0.011, if the formula given by HALD
(1949) is used in the calculations.

SUMMARY
The purpose of the present investigation was to study the develop-
ment of the P, receptor in children and to examine the mode of inheri-
tance postulated for the P blood group system.
The development of the PI receptor was studied on 101 families with
231 children and on a series of 3609 mother and child pairs. On the
basis of the results obtained in the mother-child series, it is concluded
that P, determination cannot be made with certainty in children younger
than one year. Although present the P, receptor was observed to be in-
completely developed even in children older than three years. Thus, in
a group of children between the age of three and fourteen years the re-
ceptor strength was significantly below the adult level. The results of
the family investigations indicate that the age for the complete develop-
ment of the PI receptor might be as high as seven years or higher.
The theory of inheritance was examined on the basis of the family
material mentioned above. The segregation ratios observed were in satis-
factory agreement with those expected, and no exceptions to the genetic
theory of the system were encountered.
Finally, the results were discussed with special reference to the ap-
plication of the P blood group system in genetico-legal investigations.

Literature cited
BECKMAN, L. 1959. A contribution to the physical anthropology and population genet-
ics of Sweden. - Hereditas 4 5 : 1-189.
BRENDEMOEN, 0. J. 1952. P blood group in 89 families. - Acta Path. Microbiol.
Scand. 31: 71-72.
7 - Ileredilas 58
98 AAGE HEIKEN

DAHR,P. 1942. uber die bisher im Kolner Hygienischen Institut gewonnen Unter-
suchungsergebnissen iiber das Blutmerkmal P. - Z. Immunitatsforsch. 101: 346
-355.
FISHER,R. A. 1953. The variation in strength of the human blood group P. - Hered-
ity 7: 81-49.
GALATIUS-JENSEN, F. 1960. The Haptoglobins. A Genetical Study. - Dansk Vidensk.
Forlag, Kebenhavn.
GROSJEAN,R. 1950. Sur I'hCrCditC de l'agglutinoghe P. - C.-R. SOC.Biol. 144: 1011.
- 1952. Nouvelles recherches sur l'agglutinoghe P. - Sang 23: 490-507.
HACKER, G. 1964. Zur Vererbung des Blutfaktors P. - Blut 10: 161-172.
HALD,A. 1948. Statistiske Metoder. - Det private Ingeniarfond, Kebenhavn.
HENNINGSEN, K. 1949 a. Investigations on the blood factor P. - Acta Path. Microbiol.
Scand. 26: 639-654.
- 1949 b. On the heredity of the blood factor P. - Ibid. 26: 769--785.
- 1952. Om blodtypesystemet P. - Dansk Vidensk. Forlag, Kabenhavn.
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