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An anthropobiological study in Basse Kotto (Central Africa). I. Erythrocyte and sero-genetic markers An analysis of the genetic differentiation

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 6039-47 11983)
An Anthropobiological Study in Basse Kotto (Central Africa). I.
Erythrocyte and Sero-genetic Markers: An Analysis of the
Genetic Differentiation
GABRIELLA SPEDINI, HUBERT WALTER, ENRICO CAPUCCI, MARIA
FUCIARELLI, OLGA RICKARDS, MARIA LUISA AEBISCHER, AND
NICOLETTA CROSTI
Institute ofAnthropology, Uniuersity of Rome, Rome, Italy (G.S., E. C., M . R ,
0.R.1; Department of BiologpHuman Biology/Physical Anthropology, University o f Bremen, Bremen, Federal Republic o f Germany (HW); National
Centre ofBlood Transfusion of the Italian Red Cross, Rome, Italy (M.L.A.);
and Institute of Human Genetics, Catholic Uniuersity, School of Medicine,
Rome, ftaly IN C.)
KEY WORDS
Central Africa, Mbugu, Yakpa, Sango, Erythrocyte
polymorphisms, Serum polymorphisms, Genetic heterogeneity, Genetic
distances
ABSTRACT
Phenotype and allele frequencies for hemoglobin types (Hbp),
acid phosphatase (AcP), phosphoglucomutase (PGM1 and PGMz), esterase D
(EsD), 6-phosphogluconate dehydrogenase (6-PGD), glyoxalase I (GLO), superoxide dismutase (SOD A), and adenylate kinase (AK) as well as for haptoglobins (Hp), group-specific component (Gc),transferrin (M,Gm, and Inv groups
and albumin, are reported in the Mbugu, Sango, Yakpa, and Baya Mandja
ethnic groups in the Basse Kotto district of the Central African Republic. The
total sample size amounts to 133 males and 128 females aged from 16 to 60,
unrelated and healthy. A new albumin variant (albumin Mbugu) is described
and discussed. The average heterozygosity is high in each group because of a
high degree of exogamy. The FSTaverage standardized value among the four
groups indicates that the genetic differentiation in Basse Kotto is at level of
about 2 4 . This indicates that the four examined groups might be considered
genetically homogeneous, in spite of their different ethnic origins. The genetic
distances among the four groups show that only the Baya Mandja are less
closely related to the other three groups because of their foreign origin.
The district of Basse Kotto, Central African Republic, where the present survey was
carried out, is included in the southern area
of Central Ubangui, which consists of a n undulating plateau at 600 m above sea level
from 4" to 6" latitude N and from 20" to 22"
longitude E. To the east, the Lower Kotto
River fixes the boundary with the district of
Mboumu, and southwards the plateau slopes
down to the right bank of the Middle Ubangui rather roughly. For most of its length the
river marks the boundary between the Central African Republic and Zaire (Fig. 1).
The plateau is crossed by a track that links
Bangui, the capital of Central Africa, with
Bangassou. This track winds for a total of
650 km along the shrubby savanna, alternating wide strips of gallery forests along the
water courses with coffee and tobacco plantations and with cotton, sesame, and peanut
0002-948318316001-0039$03.00C2 1983 ALAN R. LISS, INC.
plantations particularly in Basse Kotto. During the rainy season (from March to October)
this track is impassable for' long distances,
causing connections between the western and
eastern areas of the country to become very
difficult, and trade activities, particularly retail, to be greatly curtailed, just because of
the poor state of the roads.
The main economic source for a family is
represented by the work which the active
male population carries out either directly in
the plantations or in the factories connected
to them (i.e., in the important oil mill at
Alindao). Women attend to the cultivation of
vegetables.
On the plateau, along the track, hamlets of
typical Sudanic houses (circular huts with
Received January 4,1982; accepted April 15,1982
40
G. SPEDINI ET AL.
-~
THE
CENTRAL AFRICAN
*m. 30
D
80
_ - - ,
m
Do
15
120
REP
Irn
rra
V
16.
011
1 I
2 0.
Lp--2
2 2.
1=
NZAKARA
Fig. 1. Geographic location of the four considered ethnic groups.
conical roofs) follow one another close to the
savanna. However, these villages are not
permanent. One often sees the carbonized
ruins of complete villages along the track,
because the villagers set fire to the savanna
during the dry season in order to flush small
game.
The energy intake is generally sufficient;
in fact, it covers 98% of the total needs, especially carbohydrates (manioc and maize).
In contrast, lipid intake is low (40 g per capitdday) and proteins amount to less than 50
g per capitdday (O.M.S., 1977). Children
often show clear signs of protein malnutrition, with frequent avitaminosis (riboflavin,
thiamine, vitamin A) (Cresta, 1965).
The nutrition situation is made more precarious by various dietary taboos that are
particularly in force in the most vulnerable
population: children and pregnant women
(e.g., among the Banda, eggs and fish are
forbidden to them because “these foods produce skin spots and children’s caries”).
The hygienic and sanitary situation is also
very poor; the child mortality rate is close to
19%(demographic inquiry in 1959).The more
frequent causes of death are the anemias
associated with malnutrition and parasites,
primarily malaria and the genetic hemoglobinopathies; 26%of deaths in the first year
of life and 13% from 1 to 4 years are due to
malnutrition (compared to 11%found at the
Civil Hospital of the capital city, Bangui).
Goiter is also frequent; Cresta (1965) found
that about 19% of 450 Banda adults were
suffering from adenomatous goiter and about
7% from visible goiter.
In central Ubangui, the district of Basse
Kotto is inhabited by two main ethnic groups:
the Banda, settled more in the interior, and
the Ngbandi-Yakoma, along the sandbanks
of the upper and middIe Ubangui River. The
GENETIC DIFFERENTIATION IN CENTRAL AFRICA
Banda belong to the equatorial branch of the
eastern Nigritic peoples (Murdock, 1959).
They reached Central Ubangui in about the
19th century, when some Sudanic groups
were forced to cross the Bahr el Ghaza to
escape Nubian and Arab slave raids, separating into various small groups and settling in
a large area including from 4" latitude N
and from 16" to 25" longitude E. They did
not reach the Ubangui River banks, however, because the Ngbandi-Yakoma populations, already settled, prevented them.
A t present the Banda represent 53%of the
Central Ubangui population. The social
structure is based on clans, each clan consisting of several families with a polygamous
chief. Because of their settlement in different
geographic areas, the Banda cannot be considered an homogeneous unit. Within the
subdivided groups endogamy is not observed.
About 77% of marriageable females (45%
marry at age 14-20 years) are chosen outside
of the husband's village of residence, but inside the Banda area. Exogamy is more common among the Ngbandi-Yakoma, more than
78% of the females coming from Yakoma villages other than the husband's. In addition,
no less than 54% of males over 15 years come
not only from villages on the right (Central
African Republic), but also on the left bank
(Zaire) of the Ubangui River. All of that
shows the great mobility inside the group.
However, the environmental conditions connected with the great river and the language
tend to preserve the groups' cultural homogeneity. A t present they amount to about
20,000 individuals; they border eastward on
the N'Zakara and the Azande.
The Banda tribes, settled in Basse Kotto,
are the Langba, the Dakpa, the Yakpa, and
some smaller ones. Girard (1901) and Daigre
(1947) count the Mbugu among the Banda.
According to Murdock (1959) and Biasutti
(1967), however, they do not belong to the
same stock, being a residual group of a tribe
that belongs to the southern branch of the
Cushitic subfamily.
The Ngbandi-Yakoma tribes of the same
area include the Sango and Banziri.
Owing to the uncertain position of the Mbugu, we have compared them with the Yakpa (Banda) and Sango (Ngbandi-Yakoma) either as to their morphometric traits or as to
the distribution of several genetic markers.
MATERIALS AND METHODS
The Basse Kotto survey was carried out in
January 1979 (Spedini) in the villages of
41
Kongbo (4"33' latitude N, 660 km east of
Bangui) and of Mobaye (4'22' latitude N, 60
km south of Kongbo), an important center
built on the banks of the Ubangui River
(Laurentin-Retel, 19791, and in November of
the same year (Spedini and Capucci) in Mobaye. The sample reported consists of 133
males and 128 females 16 to 60 years, unrelated and healthy, and a group of 77 infants
less than 2 years.
Body measurements, finger and palmar
dermatoglyphics, and blood were obtained in
the Dispensary of the Swiss Evangelic Mission at Elime (4 km from Kongbo) and in the
hospital a t Mobaye. Venous blood was collected in ACD and stored a t about 0°C both
during transportation to Bangui and shipment by air to Rome.
A portion of the whole-blood sample was
utilized, within 2 weeks, for typing blood
groups as ABO, MNSs, Rh (-C, -c, -D, -E, -e),
and Duffy in the National Centre of Blood
Transfusion (NCBT)of the Italian Red Cross,
Rome. Hemolysate and plasma samples,
divided into 1-2 cc aliquots, were stored
a t -20°C until used. In the same center, triiodothyroxine (T3), thyroxine (T&
and thyrotropin (TSH) hormone levels and
analysis for HbsAg typing by radioimmunological techniques were carried out, and
Treponema pallidurn was shown by VDRL
test.
In the Laboratory of the Institute of Anthropology, University of Rome, Hb types, serum proteins, and serum lipids were detected
by electrophoresis, using cellogel strips, according to Helena Laboratories (Beaumont,
Texas), The total levels of serum proteins,
serum lipids, cholesterol, and hemoglobin
were determined using Boehringer kits and
a Bausch and Lomb spectrophotometer. The
electrophoretic support for the determination
of the erythrocyte enzyme polymorphisms
acid phosphatase (AcP),phosphoglucomutase
(PGMI and PGMZ), adenylate kinase (AK), 6phosphogluconate dehydrogenase (6-PGD),
and esterase D (EsD) was cellogel RS and the
staining was carried out by the method of
Chemetron (Milan). The electrophoretic support for determination of haptoglobin types
(Hp) was cyanogum (Hoppe et al., 1972; modified). In a few cases, for further verification,
electrophoresis on starch gel, according to
Karp and Sutton (1967), for the RA and RB
phenotypes was utilized.
In the Institute of Human Genetics (Sacred
Heart University, Rome) glyoxalase I (GLO)
and superoxide dismutase (SOD A) were de-
42
G.SPEDINI ET AL.
tected using as support cellogel RS (MeeraKhan and Doppert, 1976; Chemetron, Milan,
respectively), and in a few cases, for further
verification, according to Crosti et al. (1976);
the support for transferrin types (TO was
cyanogum (Matson et al., 1965).
Finally, Gm and Inv (Km) groups and Gc
types were determined in the Laboratory of
Human Biology, Department of Biology, University of Bremen (West Germany).
Data on blood group polymorphisms (ABO,
MNSs, Rh, and Duffy), hemoglobin types, and
HBsAg have been reported (Spedini et al.,
1981; Capucci et al. 1980-81b) as those regarding some anthropometric variables
Capucci et al., 1980-81a), a partial sample
for the AcP system (Spedini et al., 19801, and
GLO and SOD A polymorphisms (Spedini et
al., 1982). The dermatoglyphics analysis is in
progress.
In this paper we report AcP, PGMl, AK, 6PDG, Hp, Gc, Tf, Gm, and Inv frequencies.
We have been able to analyze blood samples
of 139 unrelated Mbugu, 34 Yakpa, 73 Sango,
and 15 Baya Mandja. This last sample has
been included in spite of its small number
because until now no data have been available. The Baya Mandja are aliens to the territory of Basse Kotto; they represent about
15% of the entire population of Central Africa, and occupy the region to the west from
north to south, and the comparison with the
local groups of Basse Kotto could give some
interesting indications about the eventual
contribution of the actual genetic structure
of that territory.
RESULTS AND DISCUSSION
Phenotype and gene frequencies are shown
in Table 1for the red cell enzyme polymorphisms AcP, PGMI, EsD, 6-PGD, and GLO as
well as for Hbp variability and for the serum
polymorphisms Hp and Inv. No variation was
found in any group in the PGMz locus.
For the AK locus five Mbugu showed AK
2-1 phenotype (AK1 = 0.9810) and for SOD A
locus one Mbugu showed SOD A 2-1 phenotype (SOD A' = 0.9950.) (Another subject,
SOD A 2-1, sister of this heterozygote individual, was found).
With respect to serum polymorphisms the
following results were obtained: Gc locusone Mbugu in a total of 43 subjects and four
Sango in a total of 48 subjects showed Gc 2-1
phenotype (Gel = 0.9880 and 0.9580, respectively); Tf locus-three Mbugu out of 44 subjects and one Sango out of 40 subjects showed
Tf CD phenotype (Tfc = 0.9660 and 0.9880,
respectively). Only one Mbugu out of 45 subjects showed Gm 5, 13 allotype, and another
one out of 139 individuals showed an albumin variant.
Every locus indicated Hardy-Weinberg
equilibrium (for verification we utilized Levene's formula; see Li, 1955).The only exception is that of the Mbugu for the 6-PGD locus
(P < 0.0025, 1d.f.1.
On the whole, the red cell isoenzymes and
hemoglobin allele frequencies observed in
Basse Kotto are within the range of those
observed in sub-Saharan areas (Spedini et
al., 1978). The Hp', Gc', and Tfc allele fre-
TABLE 1. Erythrocyte and serum phenotypes and gene frequencies in the Basse Kotto
Locus
HbR"'
Phenotype
A
AS
S
Total
$1
Mbugu
Yakpa
Sango
Baya
116
14
1
29
5
0
57
16
0
11
0
0
131
0.715
34
0.169
73
1.030
11
0.939
0.061
0.926
0.074
0.890
0.110
I
-
Gene frequency
8"
PS
10
80
37
5
n
1
RB
Total
$3,
139
6.100
17
10
2
2
5
39
22
1
6
34
1.920
73
1.222
3
1.000
-
1
6
5
0
0
12
0.015
(Table 1 continued on next page)
43
GENETIC DIFFERENTIATION IN CENTRAL AFRICA
Locus
TABLE 1. Erythrocyte and serum phenotypes and gene frequencies in the Basse Kotto (continued)
Phenotype
Mbugu
Yakpa
Sango
Baya
Gene frequency
P
0.223
0.734
0.043
Pb
P’
PGMl
1-1
2-1
2-2
Total
x:1 I
0.265
0.676
0.059
0.226
0.726
0.048
0.292
0.708
-
8
3
1
113
20
0
24
8
2
53
16
4
133
0.832
34
1.501
73
3.266
12
1.087
0.925
0.075
0.824
0.176
0.836
0.164
0.792
0.208
Gene frequency
PGM:
PGM?
EsD
1-1
2-1
2-2
Total
x 6I
105
27
1
25
8
1
60
13
0
11
1
0
133
0.231
34
0.215
73
0.640
12
0.891
0.109
0.853
0.147
0.911
0.089
-
Genefre uency
EsDsl
ESD~
6-PGD
A
AC
C
Total
XflI
0.958
0.042
70
3
0
12
0
0
-
73
0.022
12
0.985
0.015
0.979
0.021
125
7
1
33
133
5.845
34
1
0
-
Gene frequency
PGD*
PGD~
~ ~ 0 ‘ 3 )
0.966
0.034
1-1
2-1
2-2
Total
X;l I
3
8
1.000
-
3
5
7
10
30
52
10
3
27
40
92
2.854
21
0.430
70
0.350
15
1.191
0.333
0.667
0.236
0.764
0.367
0.633
Gene frequency
GLO’
GLO~
HP
0.272
0.728
45
50
12
26
1-1
2-1
2-2
0
Total
X;l I
4
6
1
1
10
11
5
28
30
3
12
133
0.085
34
0.518
73
1.858
12
0.350
0.654
0.346
0.596
0.404
0.705
0.295
0.636
0.364
16
31
1
8
23
23
2
6
47
9
46
8
0.188
0.057
a
Gene frequency
~
~
“
4
)
HP2
Inv
Inv
1
Inv - 1
Total
Gene frequency
Inv
~
~~
“”his figure does not include the sample of 151 boys of Mobaye reported in Spedini et al., 1981.
“This figure includes the sample of 90 Mbugu and 50 Sango reported in Spedini et al., 1980.
‘‘’This is the sample of Spedini et al., 1982.
‘‘‘Individuals with Hp 0.0 phenotype were excluded from gene counting.
0.293
0.134
44
G. SPEDINI ET AL
quencies are also in good agreement with
hitherto published data from the negroids of
sub-Saharan Africa. Our average values
come to Hpl = 0.65; Tfc = 0.97; Gcl = 0.99
(Mourant et al., 1976: Hpl = 0.28-0.87;
Tfc = 0.86-1.00; Gc' = 0.82-0.97).
In Basse Kotto, temperature and humidity
create excellent conditions for the diffusion
of Plasmodium Falciparum. Most likely due
to this, HbP5 frequency is 8%, and that of the
P" is rather low (about 4%).This figure would
confirm the hypothesis suggesting the correlation between a low incidence of this allele
and such climatic conditions as are found in
sub-Saharan areas. The reason for this correlation may be a selective disadvantage of
the P' allele in malaria areas (Spedini et al.,
1980).
The HbPC allele is absent in Basse Kotto.
This is a function of the large geographic
distance from the Alto Volta area.
Of particular anthropological interest is the
presence of a n SOD A 2-1 individual among
the Mbugu. This is the first documentation
of SOD A2 allele presence in Africa. As gene
flow from Caucasoids, in which the frequencies of this allele come to 2% (Welch and
Mears, 1972; Beckman and Pakarinen, 1973;
Eriksson, 1973; Carter et al., 19761, can be,
excluded, the presence of SOD A2 allele in
the Mbugu seems most likely to be caused by
"an independent mutational event, which occurred in more or less recent times" (Spedini
et al., 1982).
Of great anthropological interest also is the
appearance of a slow-moving albumin variant of the Mexico type (Fig. 2). Unfoi-tunately, it has not been possible to perform a
family study to ascertain the genetic nature
of this variant, and it has not been possible
to get additional samples to confirm this variant by electrophoresis on starch gel. The
excellent condition of the sample in question
and the fact that the propositus was in good
health led us to consider this variant to be a
genetic one, which tentatively has been
called albumin Mbugu. It seems to be a new
variant different from albumin Cayemite, a
slow-moving variant, the only one hitherto
found in American Negroes (Weitkamp et al.,
1969). It should be emphasized that this is
the first report on the existence of an albumin variant among African Negroes.
Fig. 2. Albumin variant observed in a Mbugu individual. Photograph of cellulose acetate electrophoresis (pH
8.6) (top) and scanned profile (bottom).
45
GENETIC DIFFERENTIATION IN CENTRAL AFRICA
Since genetic frequencies reflect exactly the
genetic composition of populations, they are
the most suitable for population comparisons
on a genetics scale. However, before intergroup comparisons, intragroup should be carried out. The simplest and most direct
approach for this is the estimation of heterozygosity, considering a s many loci as possible.
Since the average heterozygosity (heterogeneity index) of a population is defined not
only by interlocus variance (owing to drift),
but also by intralocus variance (given that in
the multiallelic locus every allele may cause
a variation), we have calculated the average
heterozygosity for every locus, adopting Nei
and Roychoudhury’s (1974a) formula. For
each of the groups of Basse Kotto the results
obtained are reported in Table 2. Considering the presence of loci at very low heterozygosity or those for which a gene is fixed in
the population, as for AK, Gc, Tf loci, it was
concluded that the heterozygosity is high in
each group. Nevertheless, the small number
of individuals in the Yakpa and Baya Mandja
groups (less than 50) could affect the sampling variance of heterozygosity (Nei, 1978).
In Negroids, Nei and Roychoudhury (1974b)
have estimated a n average heterozygosity of
0.162 i 0.035 for 34 blood group loci but only
0.092 k 0.019 considering the enzymatic and
nonenzymatic proteins too, for a total of 64
loci. In Basse Kotto, however, considerable
heterozygosity within each group exists:
about 30% in the Yakpa and Sango and 24%
in the Baya Mandja. These results appear
congruent with the exogamy practiced in a
high degree by the Mbugu and also by the
Yakpa of Kongbo, and to a higher degree by
the Sango (78%).
To estimate the intergroup genetic differentiation we have calculated FST.This was
possible by hypothesizing that the high average heterozygosity found in every subpopulation was the result of exogamy, not only
among members of a single ethnic group but
among members of different groups, so that
the effect of the division in ethnic subpopulations would be annulled. In this case too,
we have considered that in a multiallelic locus, each allele can produce a different FST
and so we have estimated the mean and
standard deviations among FSTcoefficients
for each locus (Chakraborty et al., 1977;
Chakraborty and Ghosh, 1981).
The corrected values per allele are given in
Table 3. The alleles that contribute most to
the total variance are IB,Fya, PGM:, HbPA
(about 3-5%), and cDe allotype (about 4%).
Thoma (1974) concluded that the IA allele
frequency has not reached stability in subSaharan Africa. On the contrary, in Basse
Kotto region this allele has reached a considerable stability. This could suggest the presence in the region of a specific selective factor,
stabilizing the I* allele. For the Duffy system the greatest variablity of the Fy” allele,
in comparison with Fyb and Fy, confirms all
TABLE 2. Average heterozygosity per locus
Mbugu
Sango
Yakpa
Baya
h*
vh**
h
Vh
h
Vh
h
Vh
0.005
0.010
0.004
0.0001
0.001
0.002
0.00001
0.00006
0.002
Inv
0.307
0.322
0.710
0.252
0.251
0.000
0.413
0.330
0.080
0.401
0.000
0.463
0.000
0.000
0.232
0.0007
0.017
0.0001
0.0001
0.066
0.461
0.647
0.449
0.334
0.195
0.419
0.275
0.162
0.360
0.000
0.416
0.080
0.025
0.414
0.002
0.009
0.002
0.0001
0.0006
0.001
0.0002
0.001
0.002
Tf
0.003
0.013
0.003
0.0002
0.002
0.001
0.001
0.0004
0.001
0.004
0.002
0.004
0.003
0.0001
0.550
Gc
0.463
0.697
0.472
0.211
0.115
0.410
0.139
0.194
0.395
0.043
0.452
0.023
Locus
ABO
MNSs
Rh
Duffy
Hbp
AcP
PGM,
EsD
GLO
AK
HP
a = 0.285 f 0.003
0.660
0.511
0.239
0.136
0.469
0.291
0.251
0.444
0.000
0.481
0.000
0.000
0.108
0.003
-
0.002
= 0.296 & 0.002
-
0.001
0.003
0.005
0.001
= 0.303 f 0.002
-
H
-
0.003
0.0008
0.001
0.002
-
0.004
-
0.0005
= 0.245 f 0.001
Note: The average heterozygosity for ABO, MNSs, Rh, and Duffy loci was calculated using the data reported in Spedinl et a]., 1981.
46
G. SPEDINI ET AL.
TABLE 3. Heterogeneity o f gene frequencies among four
ethnic groups of Basse Kotto lFs7 coefficientl
Allele or
aplotype
I*
P
2x
10-3
FST
0.1815
0.1231
0.6954
0.17
4.56
5.01
0.001
0.042
0.024
Ms
NS
Ns
0.1783
0.3710
0.0760
0.3747
1.06
3.99
5.72
0.50
0.007
0.017
0.008
0.002
CDe
cDE
cDe
cde
0.0373
0.0584
0.7000
0.2043
0.02
0.83
7.12
1.29
0.001
0.015
0.039
0.008
FY”
FY
Fy
0.0698
0.0714
0.8590
1.88
1.15
0.92
0.029
0.017
0.004
Hb BA
0.9249
1.76
0.0%
P“
0.2325
0.7229
0.0446
0.36
0.36
0.12
0.002
0.002
0.003
0.8790
0.8948
0.7272
0.6610
0.2173
3.67
1.51
1.55
1.69
- 3.56
0.034
0.016
0.008
0.007
0.021
0.003
IB
i
MS
Pb
P‘
PGM
ESD’
GLO~
HD’
The estimated genetic distances (Nei, 1972),
on the other hand, show that the Mbugu are
closely related to the Sango and Yakpa but
much less to the Baya Mandja (Table 4).
This result agrees with the earlier statement that the Baya Mandja represent an
ethnic minority of Basse Kotto who recently
emigrated from the western regions, their
place of origin.
ACKNOWLEDGMENTS
We wish to thank the staff of the Swiss
Evangelic Mission at Elime, and Mr. Mario
Ferrari and the missionary sisters of Mobaye
for handling the logistics of getting the sample to the Civil Hospital a t Mobaye. We
should also like to express our gratitude to
the local authorities in making easier the
survey and to the people of Mobaye and
Kongbo for their cooperation. Finally, we
thank Dr. M. E. Danubio and Dr. S. Romagnoli for typing Tf system.
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