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Anemia Falciforme - Tulio Arends - 1965
1. Brit.J. Huetrmt., 1965, 11, 350.
Haemoglobin Types and Blood Serum Factors in British
Guiana Indians
TULIOARENDSAND M. L. GALLANGO
Iristittito Verzezolarzo de Investigaciories Cient$cus,
Laborutorio de Heiiiatologia ExperinieiztuI,
Caracas, Veilex11ekz
THEstudy of population genetics of South American Indians, since they live in relatively
isolated communities,may furnish valuable information on the genetic factors of t h s racial
group, which participated in such an important manner in the formation of the population
of SouthAmerica. It may also indicate whether the existing populations are mixed with other
racial groups.
This report refers to the search for abnormal haemoglobins and the determination of
haptoglobin, transferrin and Gm groups in Wapishana, Macushi and Acawai Indians of
British Guiana. Additional observationson their plasma proteins were also made.
MATERIAL
Three hundred and twenty-five blood sampleswere obtained from the following tribes:
Wupisharm One hundred and twenty samples were collected in Aishalton, Southern
British Guiana Savannas (lat. 2" 30' N, long. 59" 16' W) (Fig. I). The term Wapishana dis-
tinguishesa tribelinguisticallyand culturallyrelated to the Arawak Indians,whose population
was calculated in 1918 to be only 1200 persons (Steggerda,195oa) although in the lastcentury
it occupied the territory extending from the Essequibo river to the Branco river, between
lat. 2" and 3" N (BastosD'Avila, 1937).
Mucushi. One hundred and eighteen samples were obtained in Lethem, situated in the
Savannasbetween the Kanuku mountains and the Takatu river (lat. 3" 7' N, long. 59" 46' W).
This tribe belongs to the so-called Tropical Forest culture (Lowie, 1948). They are specialized
in the preparation of manioc0 and curare poison and, linguistically,are Caribs.
Acaruai. Eighty-seven sampleswere obtainedin Kamarang (lat. 5" 58' N, long. 60" 40' W),
located in the depression made by the Merume and Parakaima mountainsat theconfluenceof
the Kamarang and Mayaruni rivers. Culturally and linguistically they are Caribs ( G i h ,
Blood, drawn by venous puncture, was mked with Alsever solution. The samples were
obtained from adults apparently in good health and approximately in the same proportion
from both sexes. Persons directly related were excluded. In regard to the Guianas it is quite a
complex problem to decide whether these samples came from anthropologically unmixed
groups or not. According to Steggerda(IgSob) 'the population of the Guianas is more varied
than that of any other country in the world. Almost every race is represented, and mixed
breedsarepresentin allpossiblecombinations.It appearsthat the peopleliving alongthe rivers
arelargely mixed-African, Indian, andEuropeanelements; Negro blood, however, seemsto
be the most evident.' However, with regard to the Indiansof thepresent study,it is understood
that dueto the remoteness oftheir habitat they could be consideredtohaveremainedunmixed.
According to their physical features (Layrisse, personal communication) only the Acawai
might have negro admixture.
1948).
3so
2. Haeriioglobitz Types in British Gtiiatia Itzdians 3-51
METHODS
.Haemoglobiizs
The haemoglobin solution was prepared according to the method of Houchin and Robi-
nette (1959) and kept frozen at -20' C. until the time of its study. Paper electrophoresiswas
performed in a home-made tank where two paper strips (46.5x21.0 cm. each one) were
suspended between two crystal rods at an inclined plane of 45O, with the anode towards the
higher end (Arends, 1960). The size of the sheets allowed a auick visual comDarison of theI I
d
.KAMARANG
DUTCH
GUIANA
i-
"3UNTAIN
APOTLRI
8
AWARUWAUNAWA
q Wopshono
FIG. I. British Guiana. Approximate location of the Indian tribes studied.
relative migration of seven samples in each one. Whatman No. 3MM paper was used with
veronal buffer pH 8.6 and ionic strength 0.03. The migration lasted 1 6 2 0 hours at room
temperature. In all cases in which it was suspected that the Hb-AZ might be augmented,
starch-block electrophoresiswas performed following the technique of Kunkel and Wallen-
Haptoglobins
To type the haptoglobins, starch-gel zone electrophoresis was performed according to
Smithies (1955) horizontal technique with borate buffer, using a few minor modifications
ius (1955).
3. 352 Tulio Arends and M.L. Gallango
(Arends and Gallango, 1960). Haemoglobin was added to the plasma in sufficient quantity
to saturate the haptoglobin. At the end of the electrophoreticrun the starchplate was divided
into approximately equal halves. One was stained with benzidine for the haptoglobins and
the other with amido black IOB for the protein fractions. A sample of Hp 2-1 was included
as a control in eachplate of six samples.
Trarzsferrins
The transferrins were studied by means of the same electrophoretic method, including in
each plate a sample of Tf B I - ~ Cas control, and using 59Fe in the proportion of 5 vc. per ml.
of plasma, according to the technique of Giblett, Hickman and Smithies (1959). Autoradio-
graphs were made using Ansco, non-screen, X-ray safety filmfor 24 hours.
Gm Groups
The technique of Harboe and Lundevall(I959) was employed for the classification of the
y-globulin Gm groups using as anti-Gm(a) rheumatoid arthritis sera No. 61258 (Grubb) and
No. 1604 (Henningsen), as anti-Gm(x) Rolfserum (Henningsen), as anti-D human serum
No. 367507 (Henningsen),and Rh-positive red cells of probable genotype CDe/cDE.
Plasma Proteins
The amido black IOBstaining used in one half of the starch-gel plate allowed observations
of the plasma proteins fractions.In every casewhere any qualitativeor quantitativedeparture
from normal was observed,paper electrophoresiswas performed using a Durrum cell type.
Photographic Record
A photographic record was kept of each starch gel plate, which allowed the detailed study
of each experiment at any time. Agfa Agepe 35 mm. film was used, with tungsten lighting
without filter and the developingwas done with Agfa Final.
RESULTS
Haemoglobins
All the 325 samples examinedwith paper electrophoresisshowed only Hb-A. Fifty samples
were examined on starch block and visual comparison showed that the quantity of Hb-A2
was normal. The mean value of Hb-Azobtainedin 29 sampleswas 2.46k0.45 per cent of the
total haemoglobin. This is in agreement with the values obtained in our laboratory in 76
normal adult Venezuelan natives, in whom Hb-A2 was found to be 2.46A0.48 per cent of
the total haemoglobin (Arends, 1962).
Haptoglobins
There was no difficulty in classi+ing the haptoglobin type since the majority of the
population studied showed the three common types of haptoglobin, Hp 1-1, Hp 2-1 and
Hp 2-2 (Table I). However, in some samples a low haptoglobin level was observed, which
could not be quantitatively determined due to the slight haemolysis present in the samples
and the Alsever solution used. No haptoglobin was found in six Wapishana, 18 Macushi, and
two Acawai samples. These samples were not included in the statistical calculations for
reasons to be explained in the discussion. One sample (Acawai, No. 460) showed a highly
abnormal quantity of Hp (Fig. 2). No cases of Hp 2-1 (mod), Hp-Johnson or Hp(Ca) were
observed.
5. Tirlio Areiids arid M. L. Gahrigo
FIG.4. Starch-gel electrophoretic analpis of blood scruni: conditions the sanic as for Fig. 2. The abnormal
post-albumin spot corresponding to Wapishana sample No. .+j is indicated by the arrow (Cl).
6. Hminoglobiri Types in British Gzriarza Indians
globin
detected*
353
P
(%I (%I (%) (%)2-1 I1-1 2-1 2-2 (mod) S.E.t Hpl xz
Transferrins
The three populations studied showed d o r m i t y in regard to the total absence of trans-
ferrins different from the common Tf C. The autoradiography consistently confirmed the
results obtainedwith amidoblack IOB.
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.7
1.1
TABLEI
DISTRIBLITION OF IiAPTOGLOBN PHENOTYPES IN VARIOUS SOUTH A M W C A N INDIAN POPULATIONS
50.035 0.559
=0.031 0.733
b0.043 '' 0.714
~ 0 . 0 2 9 0.739
50.055 0.750
t o . 0 ~ 0 0.720
+0.020 0.780
=0.019 0.749
~ 0 . 0 4 3 0.455
Huptoglobiii plieriotypes
I I 1120-INo.
testedPopulation
VENEZUELA
Makiritarel
Makkitare2
Paraujanol
Irapa (Yupa)l
Macoita (Yupa)l
Pariri (Yupa)z 2370 }Shaparu
Guaharibol
Panare2
Pernonz
Guahiboz
Piaroaz
Yaruro2
WaiCaZ
COLOMBIA
Pam3
PERU
Aymara4
Quechua4
CHILE
Araucanianj
Mapuche (Araucanian)G
Pehuenche (Araucanian)G
BRAZIL
Caingang7
Xavantes
BRITISHGWA
Wapishanag
Macushi9
Acawaig
1 ~ ~ 3
59
85
0
6
2
0
0
0
0
3
12
2
I
2
3
30.5 23.7
38.0 46.8
49.6 21.8
39.5 59.7
54.1 29.7
33-3 5.4
21.1 0.0
55.2 37.9
55.9 24.3
39.1 13.1
33.7 6.1
45.5 12.9
35.3 2.9
5 0.045
5 0.038
AO.025
=0.041
$0.030
LO.050
5 0.062
;0.025
-10.031
=0.030
=0.024
i0.033
=0.034
0.610
0.342
0.530
0.205
0.432
0.780
0.895
0.478
0.674
0.345
0.771
0.644
0.795
7.601
1.918
0.114
5.762
0.756
0.114
0.265
1.413
2.865
1.417
0.005
0.220
0.858
0.001
0.2-0.1
0.8-0.7
0.02-0.0I
0.5-0.3
0.8-0.7
0.7-0.5
0.3-0.2
o.1-o.og
0.3-0.2
0.95-0.9
0.5-0.3
0.7-0.5
45.8
15.2
28.6
0.8
16.2
61.3
6.9
47.8
19.8
60.2
41.6
61.8
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
5
0
28.4
52.4
55.0 16.5
41.8 5.8
1.493
0.477
0.3-0.2
0.5-0.3
0.8-0.7
0.95-0.9
0
0
50.0
54.7
42.9 7-1
38.5 6.8
0.140
0.007
34
113
116
0
0
0
58.8
56.0
62.8
32.4 8.8
33.6 10.4
34.5 2.7
0.645
1.290
2.645
0.5-0.3
0.2-0.1
0.5-0.3
56.2
16.2
326
78
0
I0
0.017
1.855
o.g-o.8
0.5-0.3
29.8 50.9 19.3 0.0 1 =0.046 0.553 0.103 0.8-0.7
19.8 53.5 26.7 0.0 ~ 0 . 0 4 9 0.465 0.546 0.5-0.3
44.7 49.4 5.9 0.0 =0.049 0.694 2.295 0.2-0.1
6
I8
2
I20
I18
87
*The samples of this column were not included in the statistical calculations.
t Standard error was calculated according to the standard formula, u = (p4//T)*where p and q are observed fiequencies for
1 Arends and Gallango (1962a).
2 Arends and Gallango (1962b).
3 Gallango and Arends (In preparation).
4 Giblett and Best (1961).
5 Parker and Beam (1961).
6 Nagel and Etchevemy (1963).
7 SaLano and Sutton (1963).
8 Neel, Salzano,Junqueira, Keiter and MayburyLewis (1964).
9 This report.
the two alleles and T represents the number of iiideperiderit genes analysed (Parker and Beam, 1961).
7. 354
Gni Groups
showed a variable frequency(Table11).
Tdio Atwds and M. L. Gallaiigo
The Gm(a)factor was positive in 100per cent of the population studied. The Gm(x)factor
Plasnia Proteiiis
The plasma proteins, as observed after electrophoretic migration in starch gel and with
amido black IOBstaining, showed the following patterns: (i) decreasedy-globulin was found
in one out of 120Wapishana samples studied. (ii)Hyper-y-globulinaemia was found in one
Wapishana,two Macushi,and two Acawai samples.Thesepatterns were confirmed by paper
electrophoresis. (iii) h two Wapishana samples, an unusual fraction, that deserves detailed
description was observed:
Sample No. 112.Hb-A, Tf C and Gm(a+x+) phenotype. With benzidine staining, this
sample showed no haptoglobin. The only visible spot corresponds to free haemoglobin.
With amido black staining an unusual spot with a mobility of post-albumin was observed
(Fig. 3). This fraction does not bind haemoglobin or radioactiveiron.
TABLEI1
FREQUENCY (PER CENT) OF Gm FACTORS AND PHENOTYPES IN VARIOUS BRITISH GUIANA INDIAN
POPULATIONS
Grtifartors G ~ Ipheriotypes
a+x+ I n+x- I a-x- I a-x+
0.0
Macushi 0.0
Akawai
Sample No. 45. Hb-A, Hp 1-1, Tf C and Gm(a$x-) phenotype. With amido black this
sample showed a heavy spot in the post albumin region, which barely separated itself from
the albumin. This globuh, as well as the one of the previous samples, does not bind haemo-
globin or iron. No abnormality was observed in paper electrophoresis(Fig. 4).
DISCUSSION
Studies on the distribution of haenioglobins and serum groups in the Guiana Indian
populations are very scarce (Collier and Parra, 1952; Liachowitz, Elderking, Guichirit,
Brown and Ranney, 1958;Jonxis, 1959; Steinberg, Stauffer and Fudenberg, 1960). Data for
comparison and evaluation of our own results are, therefore, limited.
Haenloglobins
Liachowitz et al. (1958) studied the haemoglobins,blood groups, malaria infection and the
anaemia incidence in 343 Djukas, who descend directly from slaves brought to Surinam
from West Africa in the seventeenth and eighteenth centuries. These authors found an incid-
ence of 15 per cent of Hb-S and 3 per cent of Hb-C, 20 per cent malaria infection and 74 per
cent of anaemia.Jonxis (1959) studied 966 bush negroes, fmding in the population of Kabel,
Moengo and Stoelman Island (Dutch Guiana)Hb-S percentagesof 17,21and 12respectively,
and of Hb-C 6, 6 and 3 respectively. These results are possibly also valid for the British
8. Haeinoglobiiz Types in British Giriana Iizdiaizs 355
Guiananegro population. The high percentage of abnormal haemoglobin observed, indicates
that, if the Indian populations studied by us had mixed with these groups, in all likelihood
abnormal haemoglobins would have been present; thus, since only Hb-A was found
apparentlyno admixture has occurred.
The absence of abnormal haemoglobins in the Indian of the American continent is an
interesting finding. When abnormal haemoglobin has been found in them it is generally
of the African types (Hb-S, Hb-D and Hb-C) and a clear picture can be traced of their
admixture with negroes and Mestizos (Rucknagel, 1957; Pollitzer, Chernoff, Horton and
Froelich, 1959; Scott, Griffith, Hosking and Schneider, 1959; Blumberg, Allison and Garry,
1959; Sutton, Mattson, Robinson and Koucky, 1960; Tondo and Salzano, 1960; Eisen and
Abildgaard, 1960; Arends, 1960, 1961). An exception to this statementis the finding of a fast
haemoglobin (Hb-Mtxico) in heterozygous form in two brothers out of a group of 172
Nahoa Indians (Lisker,Loria, Gonzalez, Guttman and Ruiz, I962). Very recently, Schneider,
Haggard, McNutt, Johnson, Bowman and Barnett (1964) found a new type of Hb-G in an
Alabama Coushatta Indian famdp.
TABLE111
TEST OF HOMOGENEITY OF THE BRIITSH GLXANA INDIAN POPULATIONS
STUDLED
I 600 I tX= 338 I ... I t p X = 195.280
x2 = 20.268; d.f. = 2 ; P < 0.001.
Haptoglobiizs
The frequencies obtained for the Hpl gene in the three populations studied show certain
variations, although they possibly were subjected to sirmlar environmental factors. The x 2
test for goodness of fit (TableI) indicatesthat the variations found between the observed and
the expected values are not statistically significant. Applying the x2 test for homogeneity
(Table111) highly significant evidence was obtained (Pco.001) that they are a heterogeneous
population, or at least that they arc not similarin relation to the frequency of the Hpl gene.
The x 2 test for comparison of the haptoglobin phenotypes of the three tribes (Tables IV, V
and VI), revealed that the difference between the Wapishana and Macushi is not significant
(o.Io>P>o.o~),but the difference between these two populations and the Acawai is highly
significant(P = O.OOI),although the latter is Lnguistically related to the Macushi.
If the figures obtained are compared with those found in other South American Indian
populations (TableI) a marked variationin the Hp1 gene frequencyis observed. Thisvariation
does not allow a characterization of Indian populations by this gene, instead it agrees with
anthropological observations on the physical, linguistic, cultural and serological differences
among the Indians (Boyd, 1950; Mourant, 1954; Layrisse and Arends, 1958; Steward and
Faron, 1959;Rivet, 1960; Cruxent and Rousse, 1961).
No haptoglobin different from the ones described up to now was observed. The fact that
no haptoglobin of the 2-1 (mod)phenotype was found coilfirms the conclusionalready made
9. 356 Tulio Arena's and M. L. Gallango
Observed Expecfed 1 x2
~
H p 1-1
Hp 2-1
Hp 2-2
1 Observed
28.63 25.37
59.39 52.61
25.98 23.02
I 14.00 101.00
Expected X2
2(?= 3.52; d.f. = 2; 0 . 2 0 ~ P ~ 0 . 1 0 .
Wupishaiia Akniuni Total
34 38 72
58 42 I00
5 2722
114 8s I99
Wapishnrin Akniuni
41.25 30.75
57.28 42.72
15.47 11.53
114.00 8j.00
Hp 1-1
Hp 2-1
Hp 2-2
Total
Wnpisliaria
I .27
2.76
0.01
Akniuni
1.71
0.01
3.70
Hp 1-1
Hp 2-1
Hp 2-2
Total
7cO-E,"= 9.46; d.f. = 2; P<O.OOI.
Observed Expected
Macirslii Akniuni Total
20 38 58
54 42 96
27 5 32 17.38 14.62
I01 85 I 86 101.00 85.00
TABLEVI
COMPUTATION OF X2 FOR A COMPARISON OF THE HAPTOGLOBIN PHLYOlTF'ES FOUND AMONG MACUSHI AND
AKAWAI INDIANS
4.19
0.07
5.32
4.98
0.08
6.33
X'
iMnciislii I Akaivni
Allison, Blumberg and Rees (1958) in 3 2 per cent of Nigerian negroes and 2.8 per cent of the
London white population. This has been confirmed by Giblett (1959), Sutton, Neel, Living-
stone, Binson, Kunstadter and Trombley (1959) and Smithies (1959)in United Statesnegroes.
10. Haeiiioglobin Types in British Guiatia Indians 357
If the cases without haptoglobin found among the British Guiana Indians are examples of
congenitalahaptoglobinaemia,it could be assumed that this populationhas had a high degree
of mixture with negro populations. This has not been confirmed by the haemoglobin study.
Studiespreviouslycarried out in I 16 ParaujanoIndians from Venezuela(ArendsandGallango,
1960) revealed two cases without haptoglobin. In one of them a haemolytic anaemia, a
positive direct anti-globulin (Coombs)test was found, whdein the other no additionalstudies
could be made. Further studies (Arends and Gallango, 1962a) performed in 281 Venezuelan
Indians (Irapa, Macoita, Makiritare and Guaharibo) showed no case without haptoglobin.
Therefore,it is assumed that this phenotype is not frequent in the Venezuelan Indian popula-
tion and if it is present in the British Guiana Indians it possibly represents the effect of an
environmental factor. In this connection, it is important to remember that the ecological
conditions where these Indians dwell greatly favour the existence of malaria infection and
this could easily explain an acquired ahaptoglobinaemia as a consequence of a haemolytic
process.
Transferrins
No transferrinsdaerent from Tf C were found in the British Guiana Indians. This absence
of polymorphism has already been described in other South American Indians. Giblett and
Best (1961) found only bne case of B2C transferrin in 173 Peruvian Indians, who had a
certainamount of white andnegro mixture. However, previousstudies(Arendsand Gallango,
1962) among two Yupa Indian tribes (Irapa and Macoita) and in Paraujano Indians revealed
the presence of a slow moving transferrin classified as Tf DI. Recent studies(Arendsand Ga-
Ilango, 1964) showed that a slow moving variant commonly found in another two Yupa
tribes(Paririand Shaparu)has a different mobilityfrom standardTfD1, but is electrophoretic-
ally indistinguishablefrom Tf DChi,a transferrinhitherto only found in Chinese living in the
United States. The finding of only transferrin C in British Guiana Indians, suggests either
that these populations are not subjected to the selectiveagents that in other Indian tribes have
produced an appreciable incidence of aberrant transferrins or that the population sample
studied from British Guiana was too small.
G m Groups
The finding of the Gm(a) factor in IOO per cent of the three populations examined is in
accordancewith that found by Steinberg, Stauffer, Blumberg and Fudenberg (1961) in other
Indians of the American continent. With regard to the frequency of the Gm(x) factor, the
results obtainedrepresent the highest incidencefound, to date, of this factor. Finally, it must
be emphasizedthat phenotype Gm(a-x-), whch is only observed in Caucasoidpopulations
or in Mestizos with Caucasoidadmixture (Gallango and Arends, 1963) was not found in any
of the subjectsstudied. This could be another proof against the possibdity of admixture with
other racial groups.
Plasma Proteins
The most remarkable fact in relation to the plasma proteins is the presence of the globulin
with a migration in starch-gel electrophoresis s d a r to that of post-albumins whch was
observed in two Wapishana Indians. The only reference in the literature to a similar protein
componentis that reported by Frazer, Harris and Robson (1959) who studied a British family
in which several members showed a genetically determined ‘unusualprotein’. This ‘unusual
protein’ on paper electrophoresis had the same mobility as albumin, while in horizontal
11. 358 Tulio Arends and M. L. Gnllango
starch gel electrophoresis,using borate buffer, a mobility slightly faster than F,z (fast alphaz)
was found. According to Harris, Robson and Siniscalco (1959) this protein does not bind
radioactive iron but binds haemoglobin. The globulin found in the two Wapishana Indians
had an electrophoreticmobility very sirmlar to this ‘unusualprotein’, but with the difference
that it did not bind either haemoglobin or iron. Therefore it probably is a protein with
characteristics not reported to date, although the possibility remains that it could be related
to the Gc serum groups, since Parker, Cleve and Bearn (1963) demonstrated a post-albumin
mobility for the Gc antigens.
SUMMARY
A study was made on the haemoglobins, haptoglobins, transferrins, Gm groups and
plasmaproteinsof the Wapishana,Macushiand Acawai Indiansof British Guiana.
In 325 haemoglobin samples, examined by paper electrophoresis,only Hb-A was found.
Hb-AZ was comparativelynormal in 50 samplesstudied by starch-block electrophoresis.The
quantitative determination of this minor fraction made in 29 samples, gave a value of 2.46&
0.45 per cent of the total pigment.
Haptoglobin studies, using starch-gel electrophoresis, gave the following results: in 26
samplesno haptoglobin was found and the Hpl gene was 0.553, 0.465 and 0.694 for Wapi-
shana, Macushi and Acawai, respectively. The phenotype frequency showed a statistically
significant difference between the Acawai and the other two populations.
All sampleshad only transferrin C. The Gm groups gave a 100 per cent positivity for the
Gm(a) factor and for the Gm(x) a frequency of 61.2, 45.7 and 77.4, respectively,was found.
Thestudy of plasma proteinsrevealedtwo sampleswith a globulintypehtherto unreported.
Few samplesshowed‘hyper-or hypo-y-globulinaemia.
ACKNOWLEDGMENTS
Ths study was made with the technical assistance of Miss Rosario Suinaga, Mr. Gilbert0 Garh
and Mrs. Trina Arends. The blood sampleswere generouslyprovided by Dr. Miguel Layrisse, who
was helped by Mr. E. Banfor, Commissionerof the Interior, and Dr. B. B. G. Nehaul, Ministry of
Health, both from Georgetown, British Guiana. The Photographc Department of the hstituto
Venezolanode InvestigacionesCientificascooperatedin maintaining a photographicrecord.
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