Distribución de los linajes de Mycobact=
erium
tuberculosis en Sudamérica
Leslie
Cabezas Vinueza. [1]. & Patricia
Jiménez Arias. [2]
Recibido:
05-05-2021 / Revisado: 16-05-2021 /Aceptado: 02-06-2021/ Publicado: 05-07-2=
021
|
=
Abstract. Molecu=
lar
genotyping of Mycobacterium tuber=
culosis
allows for the identification of circulating lineages and sublineages
in the population and their relationship with migratory movements. The purpose of this review is to describe the |
|
Resumen. La genotipificación
molecular de Mycobacterium tuberculosis permite identifica=
r los
linajes y sublinajes circulantes en la
población y su relación con los movimientos migratorios. Este artículo de
revisión tiene por objetivo describir la filogeografía de Mycobacterium tuberculosis reportada por los =
países
de Sudamérica mediante el uso de técnicas de genotipificación, analizar l=
os
puntos críticos de Tuberculosis para la región y determinar el impacto de=
la
pandemia de COVID-19 en el programa de control de la Tuberculosis. El lin=
aje Latin American Mediterranean
(LAM) perteneciente al linaje Euro-Americano (linaje 4) presenta la mayor
prevalencia en Sudamérica
y le sigue el sublinaje Beijing, pertenecient=
e al
linaje Asia oriental (linaje 2). El sublinaje
Beijing considerado de interés mundial por su asociación con la Tuberculo=
sis
multirresistente (MDR-TB), se encuentra distribuido casi en su totalidad =
en
Sudamérica, siendo Perú el país con mayor prevalencia de este sublinaje. Por otro lado, se han reportado los sublinajes pertenecientes a: Indo-Oceánico (Linaje =
1),
India-Asia Oriental (Linaje 3) y África Occidental 2 (Linaje 6) con menor
prevalencia en Sudamérica. En la actualidad, Brasil y Perú son los puntos=
calientes
de la Tuberculosis y la TB-MDR en Sudamérica, donde el control de la Tube=
rculosis
se vio totalmente afectado por la pandemia de COVID-19. Por lo tanto, ha
habido impactos significativos en los programas de vigilancia y contenció=
n de
la Tuberculosis, dando como resultado diferentes escenarios post-pandémicos, de tal manera que las contribucion=
es
científicas deberán ser evaluadas e implementadas con nuevas estrategias =
de
prevención, diagnóstico, tratamiento y control de la Tuberculosis. Palabras claves: molecular, genotipificación;
Tuberculosis; enfermedades infecciosas;
micobacterias. |
Introduction.
Tuberculosis (TB) is a preventable, transmissible
bacterial disease caused by Mycobacterium tuberculosis (Koch bacillu=
s)
and is one of the top 10 causes of death worldwide from an infectious agent.
This bacterial infection is cataloged as a health problem in both the social
and health fields, is associated with poor living conditions and causes the
death of 1.3 million people worldwide per year, which is 300,000 more than =
the
human immunodeficiency virus (HIV) (World Health
Organization, 2020). The World Health Organization (WHO) has
developed a global strategy to end the TB epidemic and has set targets to
reduce TB deaths and incidence of the disease by 2035 by combining efforts =
to
provide timely diagnosis and treatment and to promote research (García & Astudillo, 2019; PAHO, 2020)<=
!--[if supportFields]>=
.
In 2018, in the American region, it was
estimated that almost 50,000 people with TB were unaware of their disease <=
/span>(PAHO, 2018). This region has 2.9% of the global TB
burden (10 million cases) and an incidence rate of 10 per 100,000 inhabitan=
ts (World Health
Organization, 2020). It was estimated that 87%=
of
TB cases are concentrated in Brazil, Peru, Mexico, Haiti, Colombia, Venezue=
la,
Argentina, Bolivia, the United States, El Salvador, Panama and Guyana (PAHO, 2020).
There are 7 Mycobacterium tuberculo=
sis
(MTB) lineages that are
distributed worldwide, and they divided into two ancestral and modern group=
s,
with the ancestral lineages being Lineage 1 (Indo-Oceanic), Lineage 5 (West
African 1) and Lineage 6 (West African 2), while the modern lineages are
Lineage 2 (East Asia), Lineage 3 (India-East Asia), Lineage 4 (Euro-America=
n)
and Lineage 7 (Ethiopia). Lineage 7 represents an intermediate phylogenetic
branch between the ancestral and modern lineages. Lineages 5 and 6 correspo=
nd
to strains traditionally known as Mycobacterium afr=
icanum, and lineages 2 and 4 a=
re
more virulent and prevalent worldwide. There are 16 su=
blineages,
and the most important ones are as follows: lineage 1, MANU and EAI; lineag=
e 2,
Beijing; lineage 3, Central Asian (CAS) and Delhi; lineage 4, Haarlem (H),
Latin American Mediterranean (LAM), T, X, S, Ghana, URAL, TUR, Uganda and
H37Rv; and lineage 6, AFRI and West African (Augusto et al., 2018; Salvato et al., 2=
019).
This review describes the phylogeography=
of
MTB sublineages reported in South American coun=
tries
that was analyzed using genotyping tools, analyzes the TB hotspots in South
America, and determines the impact of COVID-19 on TB control strategies.
The molecular epidemiology
techniques =
The molecular epidemiology of MTB aims=
, by
a comparative analysis of nucleic acid sequences of strains to determine the
relationship when the stains descend from a common ancestor and where the l=
evel
of closeness and similarity between isolates varies. Molecular markers allow
for the determination of the level of genetic relationship between strains
according to variations in the DNA sequence due to single nucleotide
polymorphisms (SNPs), long sequence polymorphisms (LSPs), or repeated seque=
nce
polymorphisms; this method allows for the characterization of the lineages =
of
strains as phylogenetic contributions to control the disease and disrupt the
transmission chain <=
!--[if supportFields]>
The techniques used for genotyping are typing by complementary
oligonucleotide spacer sequences (spoligotyping=
), restriction-hybridization patterns (IS6110-RFLP,
PGRS-RFLP), mycobacterial interleaved repeat units-variable number tandem
repeats (MIRU-VNTR), and whole genome sequencing (WGS). The most advanced
technique in the analysis of MTB from clinical samples is WGS, which allows=
the
study of genome microevolution and the preparation of genomic libraries to
identify epidemiological events of interest, with greater discrimination wh=
en
detecting outbreaks, virulence, pathogenesis, transmission chains and mutat=
ions
associated with resistance to first- and second- line antituberculosis drug=
s (Guthrie et al., 2019; =
Meehan
et al., 2019). MIRU-VNTR is also a highly
stable, fast, reproducible and highly discriminatory technique, which is wh=
y it
is considered for epidemiological studies, population genetics and phylogen=
etic
studies of pathogens belonging to the Mycobacterium
tuberculosis complex (MTBC) (Babai Kochkaksaraei et =
;al.,
2019; Jiménez et al., 2017).
=
Description
of the circulating lineages in South America
The molecular epidemiological analysis=
is
stratified by country through the molecular characterization of MTB and all=
ows
us to know the dynamics of disease transmission and to identify high-risk
groups in order to provide an early response to outbreaks. In this way, sev=
eral
studies in South America report the circulating sublin=
eages
in each country and their percentage of prevalence is of interest for the
control of TB in South America (Hill et al., 2020).
Brazil
It is the country with the highest number of TB ca=
ses
and the second highest concentration of rifampicin- and multidrug-resistant
tuberculosis (RR/MDR-TB) in the American region. In 2019, 76,000 new cases =
of
TB were reported, and nearly 4,500 deaths occurred. Most of the cases repor=
ted
in this country were concentrated in the southeast, where the state of São
Paulo (SP) represented 20% of the disease incidence in the country. As one =
of
the countries with the largest geographical extension in South America and
having several borders, Brazil has a great variety of circulating lineages =
of
MTB. There are foreign MTB lineages due to the movement of the population f=
or
tourism and commercial events. Currently, TB transmission routes are presen=
t in
both rural and urban populations, so special attention has been given to st=
udy
the disease through molecular epidemiology techniques, early diagnosis, and
monitored treatment. The HIV-infected population and people deprived of the=
ir
liberty are vulnerable groups, so plans for diagnosis and observed treatment
have been implemented to disrupt the chain of transmission (Esteves et al., 2018; PAHO, 2020)
The circulating sublineages
in Brazil with the highest percentages of prevalence are LAM (48.87%), RDRio
(22.58%), Haarlem (13.7%), T (16.5%), X (3.6%), and S (2.05%), belon=
ging
to lineage 4, which prevails in the American region, due to the stability of
its genotype and transmissibility. The lineages with the lowest prevalence =
are:
Beijing (0.65%), Uganda (0.65%), MANU (0.58%), EAI (0.48%), CAS (0.45%), We=
st
African (0.26%), H37Rv (0.24%), Ural (0.13%), Cameroon (0.065%), BOV (0.02%)
and Ghana (0.02%) (Table 1). The lineages 3, 4 and 6 have been incorporated
into a variety of MTB lineages found in Brazil by the migration of people <=
/span>(Esteves et al., 2018; Salvato et al., 2=
019). Brazil has a low incidence of
unidentified strains (10.5%), which is due to the implementation of molecul=
ar
techniques such as spoligotyping, MIRU-VNTR (24 and 12 loci) and WGS to determine the transmissi=
on
routes of TB (Cardoso
Oelemann et al., 2011; Dalla Costa et al., 2013; Gomes et al=
.,
2012; Luiz et al., 2013; Machado et al., 2014; Martins et al=
.,
2013; Medeiros et al., 2018; Nogueira et al., 2016; Noguti
et al., 2010; Soares et al., 2013; Vasconcellos et al., 2014=
).
In
2007, MTB isolates from Rio de Janeiro were analyzed for genomic deletions.=
The
RDRio sublineag=
e
was characterized as having a new deletion of a single long-sequence
polymorphism (> 26.3 kb) that included 10 genes. This sublineage
was derived from a common ancestor that belongs only to the LAM sublineage, and is thus a predominant clade that caus=
es TB
in Rio de Janeiro and the most important source of recent transmission.
Peru
After Brazil, Peru is the country with the second
highest concentration of TB cases and the first in terms of RR/MDR-TB cases=
in
the American region. In 2018, 31,668 TB cases were reported of which 1,457
corresponded to RR/MDR-TB and 121 to extremely resistant TB (XDR-TB).
Metropolitan Lima and Callao account for 64% of TB cases in this country. S=
ome
indicators show progress in disease control, such as decreases of 2% and 3%=
per
year for the reported incidence and total morbidity rates; however, the num=
bers
of XDR-TB cases have increased (Centro
de Epidemiología, Prevención y Control de & enfermedades, 2019; PAHO, 2=
020;
Soto Cabezas et al., 2020). Peru,
because of its
political and geographical history, has had visitation of different
nationalities that have increased the variety of MTB lineages in this regio=
n.
Among the visitors are persons from Asia, a continent with a high incidence=
of
TB worldwide and with the presence of lineages with high virulence and
pathogenicity, such as Beijing (Farhat et al., 2019).
The
most prevalent sublineages in Peru are LAM (35%=
),
Haarlem (31%), and Beijing (10.27%); these findings are
consistent with the data for the incidence of MDR/XDR-TB cases, since the B=
eijing
sublineage is associated with cases of resistan=
ce to
both first- and second-line anti-tuberculosis
drugs. In several studies, the Haarlem sublineage had
the highest incidence rates due to the stability of this genotype in the
population and the high rate of recent transmission. The sublineages
with the lowest prevalence in Peru are X (3.1%), T (7.4%), S (0.32%), U
(0.32%), MANU (0.01%), Ghana (0.01%), and EAI (0.01%); the presence of thes=
e sublineages correlates with the movement of people fr=
om the
Asian region (Table 1). The percentage of unidentified strains (12.2%) is
relatively low due to the implementation of molecular techniques such as
Colombia
In
the American region, Colombia has the fifth-highest concentration of TB cas=
es
after Mexico and Haiti and the sixth-highest concentration of RR/MDR-TB cas=
es (PAHO, 2020).
In 2019, there were 14,684 cases of TB. The territorial entities with the
highest rate are Amazonas, Guainía, Risaralda, Barranquilla, Meta, Arauca, and Casanare. TB is
concentrated in the most vulnerable segments of the Colombian population, s=
uch
as HIV patients (11%), indigenous people (5%), street dwellers (4%), and
prisoners (6%) (Instituto
Nacional de Salud de Colombia, 2020).
The most prevalent sublineage=
s
in Colombia are LAM (39.14%), Haarlem (27.48%), T (10.41%), U (4.2%), S (1.=
9%),
X (1.7%), and Beijing (1.63%); the lineage with the highest prevalence is L=
AM
due to the stability of this genotype and transmissibility. The Beijing
Venezuela
It is the country with the sixth highest number of=
TB
cases and with the eighth highest concentration of RR/MDR-TB in the American region (PAHO, 2020). In 2019, a TB infection rate of 47.8 per 100,000
people was reported (World Health
Organization, 2020). In 2017, 10,952 cases of TB were reported (PAHO, 2018).
The most prevalent lineages in Venezuela are LAM
(54.06%), T (11.11%), and Haarlem (4.72%), belonging to lineage 4, and those
lineages with the lowest prevalence are X (1.25%), Beijing (0.41%), EAI (0.=
35%)
and MANU (0.07%); there is evidence of lineages that are typical of the Asi=
an
region, which shows population movement. The RDRi=
o
lineage (55%), which is part of the LAM lineage, has a relatively high
prevalence (Table
1). The unidentified strains (0.83%) show a percentage close to zero=
due
to the application of highly discriminatory molecular techniques such as
MIRU-VNTR 24-loci (Abadía et al., 2009; Maes et al., 2008)=
.
Argentina
It is the country with the seventh highest number =
of
TB cases and with the seventh with the highest concentration of RR/MDR-TB <=
/span>in
the American region (PAHO, 2020). In 2018, 11,670 cases of TB were
reported, and 720 deaths occurred. Buenos Aires and Greater Buenos Aires had
45% of the cases at the country level, the incidence rate was 26.2 per 100,=
000
inhabitants, of which 6 out of 10 were men (Ministerio de
Salud Argentina, 2020).
The sublineages with t=
he
highest prevalence are those belonging to lineage 4: T (31.55%), LAM (30.9%=
),
and Haarlem (18.4%), with the T sublineage bein=
g the
one with the highest concentration both in Buenos Aires and in Greater Buen=
os
Aires due to its adaptability in the urban population. The sublineages
with the lowest prevalence are S (2.19%), X (1.02%), and Beijing (0.23%),
belonging to lineages 2 and 4. The RDRio sublineage (11.3%), part of the LAM sublineage,
has been reported as a foreign sublineage imple=
mented
by population movement between Argentina and Brazil (Table 1). Unidentified
strains (14.6%) present a relatively low percentage due to the use of molec=
ular
techniques such as spoligotyping, RFLP, and MIRU-VNTR 24-loci. The number of migrants from Europe and the Middle East=
has
contributed to the diverse
Bolivia
It is the country with the eighth highest number o=
f TB
cases and with the ninth highest concentration of RR/MDR-TB in the American
region (PAHO, 2020). In 2018, 7,762 cases of TB were reported, where =
the
Department of Santa Cruz had the highest incidence of TB in the country (42=
%),
with 3,240 cases and 103 deaths (PAHO/WHO, 201=
9a).
The most
prevalent sublineages are Haarlem (39.4%), LAM
(26.3%), and T (22%), belonging to lineage 4. The Haarlem sublineage
presents high transmissibility and stability in rural geographic areas, which justifies its high
prevalence. The identified sublineages with low=
er
prevalence are S (2%) and X (1%), belonging to lineage 4 (Table 1). =
This
study used spoligotyping and MIRU-VNTR (Monteserin et al., 201=
3).
Guyana
Guyana has the twelfth highest concentrations of TB in the American region, after the Dominican
Republic, El Salvador and Panama (PAHO, 2020). In 2017, an
incidence of 109 cases per 100,000 people and 35.1% of patients with TB and=
HIV
were reported. TB control still has some challenges in this nation, and one=
of
the most critical being the HIV epidemic in the population. This epidemic h=
as
been identified as a major concern and underlying cause of the increase in =
both
TB morbidity and mortality (PAHO/WHO, 2019b). In 2018, Guyana was =
reported
to be the country with the highest number of TB deaths in the American regi=
on (PAHO, 2020).
The
sublineages with the highest prevalence are T
(72.62%), EAI (10.8%), and Beijing (1.09%), with the T sublineage
being the one found at higher concentrations due to its adaptability in the
urban population (Table 1). Unidentified strains (16.2%) were reported in the studies,=
and
the molecular techniques of spolygotipyng and
15-loci-MIRU-VNTR were used (Millet et al., 2014; Streit et al., 201=
4).
Ecuador
In 2018, 6,094 cases of TB were reported, including
252 cases with RR/MDR-TB, 805 cases of TB in people with HIV, 637 cases of T=
B in
people deprived of their liberty, and 154 cases of TB in children under 15.=
Men
infected with TB are predominant, with a ratio of 2.40 men to women, and the
most affected age group is 25-34 years old. The urban coast region has the
highest concentration of TB (Ministerio de Salud Pública et al., 2019).
The most prevalent circulating sublineages
are LAM (42.9%), Haarlem (26%), S (11.53%), Ghana (7.12%) and X (3.35%). The
Beijing sublineage is of phylogenetic interest =
due to
its high virulence and pathogenicity has been found in this country with a
prevalence of 1.69%, which shows that the movement of people from frontier
countries such as Peru has been able to migrate foreign lineages, increasing
the transmission of TB. The sublineages with the
lowest prevalence are Cameroon (0.83%) and Delhi (0.4%), which are sublineages belonging to lineages 3 and 4 (Table 1).
The percentage of unidentified isolates (5.87%) is a relatively low value d=
ue
to the use of more sensitive and specific techniques such as MIRU-VNTR 24-l=
oci
and WGS (Garzon-Chavez
et al., 2019, 2020; Jiménez et al., 2017; Zurita et al., 201=
9).
Paraguay
In 2018, a rate of 43 per 100 000 individuals was
reported for TB in Paraguay, with Asunción, Central, and Alto Paraná being =
the
regions that comprised 54% of the country's cases. The disease had a greater
incidence among the indigenous population (16.4%) and persons deprived of t=
heir
liberty (14.4%) (Ministerio de Salud Pública y Bienestar Social, 2=
019).
The most prevalent lineages are LAM (46.06%), Haar=
lem
(17.27%), RDRio (11.3%), T
(10.9%), and S (9.7%), belonging to lineage 4. The RDRio
lineage shows the migration of people between Brazil and Paraguay. The sublineages with the lowest prevalence are X (1.51%) =
and
Beijing (0.3%) (Table 1). There has been only one MTB genotyping study in th=
is
country; the study reported a case of a foreign lineage belonging to lineag=
e 2
(Beijing sublineage), which was isolated from a=
South
Korean immigrant residing in Asunción. This study used spoligotyping, and to confirm the results obtained, the molecular techniques of RFLP and MIRU-VNTR we=
re
applied (Candia et al., 2007).
Chile
Very low rates of TB =
in
Chile have been achieved after decades of sustained economic
development-related decline and a robust and monitored National TB Program.=
In
2017, 2,740 cases of TB were reported. In 2018, the incidence rate was 14.7
cases per 100,000 people (Ministerio de
Salud, 2018).
The most prevalent sublineage=
s
are LAM (39.4%), T (33.77%), Haarlem (13.6%) and X (3.64%), belonging to
lineage 4. The LAM lineage is more prevalent due to the stability and
transmissibility of this genotype. The sublineages
with the lowest prevalence are Beijing (0.99%), S (0.3%), Cameroon (0.16%),=
and
AFRI (0.16%), belonging to lineages 2 and 4 (Table 1). The Beijing sublin=
eage
is of interest because of its high pathogenicity and virulence present in t=
he
urban population of Chile due to migration from Peru and Venezuela <=
!--[if supportFields]>
Suriname
In 2018, an incidence of TB of 38 cases per 100 000 people and 4.7% o=
f patients
with TB and HIV was reported; in 2017, an incidence of 29 cases per 100 000
people was reported, suggesting an increase in TB transmission in the count=
ry (Commiesie et al., 2019).
The
sublineages with the highest prevalence are T
(26.25%), EAI (25%), Beijing (4.3%), and Haarlem
(10.8%), with the Beijing sublineage
having
the highest percentage among Guianas (Table 1).
Unidentified strains (31.2%) were reported in the studies using the molecul=
ar
techniques of spolygotipyng and
15-loci-MIRU-VNTR (Millet et al., 2014; Streit et al., 201=
4).
Uruguay
In 2018, an incidence of 33 cases per 100 000 peop=
le
was reported (PAHO, 2020). This result=
is
currently all the information that is available since no MTB molecular
genotyping studies have been performed in this country.
French Guyana
French
Guyana has the highest burden of TB among all French territories with a sha=
rp
increase in the incidence of TB in recent years to 9.5 cases per 100,000
people. TB is the fourth most frequent opportunistic infection among HIV
patients in this country (Nacher et al., 2020).
The
sublineages with the highest prevalence are T
(32.6%), LAM (23.3%) and Haarlem (22.6%), with
the T sublineage having the
highest concentration due to its adaptability in the urban population and t=
he
LAM sublineage being the
main circulating sublineage in South America (T=
able
1). The prevalence of more sublineages is unkno=
wn due
to a single genotyping study carried out in the French region, and
the molecular technique used was spoligotyping =
(Guernier et al., 2008).
Table
1: References of shown
data in this table are cited in the text of each country. Country Number of Samples LAM Haarlem X Cameroon T S Ugand TUR Ghana RDRio H37Rv Beijing MANU EAI Delhi CAS West African AFRI Unidentified Brazil 4581 2239 48.87% 628 13.70% 165 3.6% 3 0.065% 756 16.5% 94 2.05% 30 0.65% 1 0.02% 1 0.02% 89 22.6% 11 0.24% 30 0.65% 27 0.58% 22 0.48% - - 12 0.26% - 481 10.5% Peru 8863 3157 35% 2745 31% 277 3.1% - 652 7.4% 28 0.32% - - 1 0.01% - - 910 10.3% 1 0.01% 1 0.01% - - - - 1085 12.2% Colombia 1834 718 39.14% 504 27.48% 31 1.7% 1 0.05% 191 10.41% 36 1.9% - - - - 1 0.05% 30 1.63% 4 0.2% 2 0.1% - 1 0.05% 1 0.05% - 136 7.4% Venezuela 1=
439 7=
78 5=
4.06% 6=
8 4=
.72% 1=
8 1=
.25% - 1=
60 1=
1.11% 2=
5 1=
.73% - 4=
7 5=
5% 4=
7 5=
5% - 6=
0=
.41% 1=
0=
.07% 5=
0=
.35% -=
- - - 1=
2 0=
.83% Argentina 1=
274 3=
94 3=
0.9% 2=
34 1=
8.4% 1=
3 1=
.02% - 4=
02 3=
1.55% 2=
8 2=
.19% - - 6=
1=
1.3% 6=
1=
1.3% - 3=
0=
.23% - - -=
- - - 1=
86 1=
4.6% Bolivia 9=
9 2=
6 2=
6.3% 3=
9 3=
9.4% 1=
1=
% - 2=
2 2=
2.2% 2=
2=
% - - - - - - - - - - - - 9=
9=
.1% Guyana 7=
4 -=
-=
-=
- 5=
4 7=
2.62% -<=
o:p> - - - - - 1 1.09% -=
8 10.8% - - - - 1=
1 1=
6.2% Ecuador 885 2=
05 4=
2.9% 1=
24 2=
6% 1=
6 3=
.35% 4=
0=
.83% - 5=
5 1=
1.53% - - 3=
4 7=
.12% - - 1=
5 1=
.69% - - 2=
0=
.4% - - - 2=
8 5=
.87% Paraguay 3=
30 1=
52 4=
6.06% 5=
7 1=
7.27% 5=
1=
.51% - 3=
6 1=
0.9% 3=
2 9=
.7% - - - 6=
1=
0% - 1=
0=
.3% - 1 1% - - - - 3=
3 1=
0% Chile 6=
04 2=
38 3=
9.4% 8=
2 1=
3.6% 2=
2 3=
.64% 1=
0=
.16% 2=
04 3=
3.77% 2=
0=
.3% -=
-=
- - - 6=
0=
.99% - - - - - 1=
0=
.16% 4=
8 7=
.4% Suriname 8=
0 -=
9=
1=
0.8% -=
-=
2=
1 2=
6.25% -=
-=
-=
-=
-=
-=
1=
1=
.25% -=
20 25% - - - -=
2=
5 3=
1.2% French Guyana 2=
73 6=
4 2=
3.3% 6=
2 2=
2.6% 1=
6 5=
.8% -=
8=
9 3=
2.6% -=
-=
-=
- - - -=
- - - - - -=
4=
2 1=
5.38% Table
1: References of shown
data in this table are cited in the text of each country. Source:
Own elaboration
Molecular epidemiology studies in South
America used to determine the lineages and sublineages=
circulating in each country, which the most prevalent are represented
graphically according to their distribution and incidence rate of TB (Figur=
e 1).
Distribution of the
most prevalent MTB sublineages in South America=
.
Figure 1: Data are included in =
Table
1 with de percentage of
prevalence per country. Each country has been shaded according to the
estimated TB incidence rate in concordance with the “Tuberculosis in the
Americas 2019 regional report”, by Pan American Health Organization (20=
20) [2]. Country codes
(http://www.worldatlas.com/aatlas/ctycodes.htm).
Source: =
Own elaboration.
The dynamics of lineage
distribution in South America
Susceptibility
to MTB varies in every individual and this heterogeneity influences the rou=
tes
and frequencies of transmission in a population. By understanding the magni=
tude
and distribution of these differences, it is possible to predict the dynami=
cs
of the disease to control and reduce TB transmission (Rodrigues et al., 2017). The imminent
interactions between host, environment and bacterial factors makes the
epidemiological study of MTB difficult due to the independence of strains in
phenotypic versus genotypic manifestation, explicitly in terms of virulence,
where morbidity and mortality are the measures of proportion, since the
virulence of MTB is directly related to transmission (Zhang et al., 2019).
Lineages 2, 3 an=
d 4 are
more transmissible than other lineages, and their subl=
ineages
are predisposed to adaptability to specific host populations; thus, their
evolution has allowed them to be distributed worldwide (Zhang et al., 2019)=
. In South Ameri=
ca,
these lineages are widely dispersed in both rural and urban areas, where
factors such as air pollution, smoking, malnutrition, population density,
crowded living conditions, HIV incidence and mobility among migrants increa=
se
TB transmission and distribution (Esteves
et al., 2018). This dispersion is how the =
LAM sublineage appears in cities with high population
concentrations, promoting a more severe disease due to its high mutation ra=
te.
In turn, a new mutation from this sublineage ha=
s been
reported, giving rise to a new RDRio
strain that appears in the population of Rio de Janeiro with a clinical pic=
ture
that has with high bacillary loads (Mor=
aes
et al., 2017); lineage 4 sublineages =
such
as Haarlem and T are directly related to their stability and transmissibili=
ty
in urban areas and the S sublineage in rural ar=
eas,
especially in indigenous communities (Díaz
Acosta et al., 2019; Garzon-Chavez et al., 2020). The Beijing
Brazil and Peru are hotspo=
ts
for tuberculosis in South America
Brazil is the country
with the highest incidence of TB in South America, is an economic and touri=
sm
power, and has the greatest phylogenetic variety of MTB (lineag=
es 1, 2, 3, 4 and 6). T=
his
diversity of lineages arises from the Portuguese impact on the country when=
it
was a colony and the European and Asian impac=
ts in the economic
development of the country; thus, the most prevalent lineage is 4, from whi=
ch
comes the RDRio sublineage
that is currently found not only in the
population of Rio de Janeiro but also in Paraguay, Argentina and Venezuela =
(Díaz
Acosta et al., 2019; Esteves et al., 2018; Moraes et al., 20=
17;
PAHO, 2020). The geographic extens=
ion
of Brazil provides border proximity to almost all countries in South Americ=
a, and thus, political and econom=
ic
relations increase the movement of people and consequently the distribution=
of
MTB. This phylogenetic diversity that Brazil brings to South America has led =
to
complex outbreaks and new routes of infection to countries that do not have=
the
molecular tools or control strategies to counteract the transmission of TB<=
/span> (Esteves et al., 2018; Medeiros et al.,
2018; Rodrigues et al., 2017)(Esteves et al., 2018). Peru is the country w=
ith
the second highest incidence of TB but the first in terms of RR/MDR-TB, whi=
ch
is related to the level of prevalence of the Beijing s=
ublineage.
Prevalence of the Beijing sublineage in Peru is=
the
highest in South America (10.3%) due to population migration from China
associated with labor, commercial and economic factors. Chin=
a is one of the hotspots =
with
the highest level of TB prevalence in the world (Grandjean et al., 2017; Huang et al., 2=
020;
PAHO, 2020).
XDR-TB reports also have a higher incidence in this nation than in South
America as a whole. The highest density of cases are concentrated in the
districts of Lima and Callao and in more than half of the country's departm=
ents
with high poverty rates (Grandjean et al., 2017; Soto Cabezas
et al., 2020). Soto Cabezas et al., 2020 reported that almost half of the XDR-TB cases analyzed (48=
%)
had no previous anti-TB treatment, which highlights the need to evaluate TB
control programs. For bordering
countries, both Ecuador and Chile have reported an increase in Beijing sublineage strains, making Ecuador and Colombia addit=
ional
areas identified as critical or hotspots for the prevalence of lineage 2 (Garzon-Chavez et al., 2020; Lagos et al=
.,
2016). Impact
of COVID-19 on TB disease It is likely that the high prevalence of TB and increasing
COVID-19 allow a temporal association that has a synergistic effect both
economically and socially, where comorbidity would increase the rate of dea=
ths
in both vulnerable individuals and the general population (Homolka et al., 2020). In 2020, there has been a significant decrease in
presumptive, confirmed cases and notifications of TB deaths compared to 201=
9, which is associated with restricted access to the
diagnostic testing and treatment centers during confinement and associated =
with
the use of resources such as laboratories, people and supplies that are foc=
used
on emergency planning and containment of the COVID-19 pandemic (Buonsenso et al., 2021; Homolka et al.,
2020). One of the possible post-pandemic effects is the
increase in morbidity and mortality from TB and RR/MDR/XDR-TB due to
overcrowded conditions, treatment abandonment and lack of an active search =
for
cases during confinement and outbreaks of COVID-19, allowing the appearance=
of
new outbreaks and chains of transmission, especially in vulnerable populati=
ons;
thus, measures should be taken and TB control programs should be evaluated =
and
implemented (Buonsenso et al., 2021; Comella-del-Barrio
et al., 2020).
TB control requires surveillance interventions, clinical assessment, diagno=
stic
testing, contact tracing, and confirmation of diagnosis with supervised treatment regimens; as all
these important actions have been limited, all the recent advances in the E=
nd
TB Strategy have been affected (Homolka et al., 2020; McQuaid et al., 2=
020;
Motta et al., 2020). It is necessary to respond to the COVID-19 pande=
mic
without neglecting an epidemic such as TB, which is the fifth leading cause=
of
death worldwide. This statement is even more true in South America where
countries that are low development regions cannot cope with a coinfection of
two infectious diseases or coinfection of up to three diseases if HIV is
included. Conclusions. ·
The
genotyping of MTB in the American region, mainly in South America, has prov=
ided
results of epidemiological interest by identifying the circulating lineages
that each country has and how migratory movements have contributed to the
dissemination of TB and the increase in MTB
genotypes in each country. ·
In this c=
ase, Brazil and Peru are the countries with the major
incidences of TB and increased phylogenetic diversity of MTB in South Ameri=
ca.
According to the scientific reports analyzed in this review, a high prevale=
nce
of sublineages belonging to lineage 4 can be ob=
served
as LAM, Haarlem, Cameroon X, T, S, and Ghana. Additionally, the new sublineage RDRio has a
more severe clinical profile and is currently found not only in Brazil, the
country of origin, but also in Argentina, Venezuela and Paraguay. Another ·
The ·
Finally, the TB control directions
are from the WHO,
and each country implements these directions according to the country’s
resources (laboratories, supplies and people), which has been considered an
impediment to harmonize strategies aimed at the control and elimination of =
the
disease. Therefore, the response to the COVID-19 pandemic has a possible
post-pandemic effect of a significant resurgence of TB cases; thus, as a
scientific community, we must propose projects or seek diagnostic resources=
to
actively search for cases, supervise and complete treatment, and
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