Polymorphism in the flanking regions of the PbGP43 gene from the human pathogen Paracoccidioides brasiliensis: search for protein binding sequences and poly(A) cleavage sites
BMC Microbiology
BioMed Central
Research article
Open Access
Polymorphism in the flanking regions of the PbGP43 gene from the
human pathogen Paracoccidioides brasiliensis: search for protein
binding sequences and poly(A) cleavage sites
Antonio A Rocha1, Flávia V Morais2 and Rosana Puccia*1
Address: 1Department of Microbiology, Immunology and Parasitology, Federal University of São Paulo (UNIFESP), 04023-062 São Paulo, SP,
Brazil and 2Laboratory of Molecular Genetics and Genomics, University of Vale do Paraíba (UNIVAP), SP, Brazil
Email: Antonio A Rocha - ; Flávia V Morais - ; Rosana Puccia* -
* Corresponding author
Published: 30 December 2009
BMC Microbiology 2009, 9:277
doi:10.1186/1471-2180-9-277
Received: 23 June 2009
Accepted: 30 December 2009
This article is available from: http://www.biomedcentral.com/1471-2180/9/277
© 2009 Rocha et al; licensee BioMed Central Ltd.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Abstract
Background: Paracoccidioides brasiliensis is a thermo-dimorphic fungus that causes
paracoccidiodomycosis (PCM). Glycoprotein gp43 is the fungal main diagnostic antigen, which can
also protect against murine PCM and interact with extracellular matrix proteins. It is structurally
related to glucanases, however not active, and whose expression varies considerably. We have
presently studied polymorphisms in the PbGP43 flanking regions to help understand such variations.
Results: we tested the protein-binding capacity of oligonucleotides covering the PbGP43 proximal
5' flanking region, including overlap and mutated probes. We used electrophoretic mobility shift
assays and found DNA binding regions between positions -134 to -103 and -255 to -215. Only
mutation at -230, characteristic of P. brasiliensis phylogenetic species PS2, altered binding affinity.
Next, we cloned and sequenced the 5' intergenic region up to position -2,047 from P. brasiliensis
Pb339 and observed that it is composed of three tandem repetitive regions of about 500 bp
preceded upstream by 442 bp. Correspondent PCR fragments of about 2,000 bp were found in
eight out of fourteen isolates; in PS2 samples they were 1,500-bp long due to the absence of one
repetitive region, as detected in Pb3. We also compared fifty-six PbGP43 3' UTR sequences from
ten isolates and have not observed polymorphisms; however we detected two main poly(A)
clusters (1,420 to 1,441 and 1,451 to 1,457) of multiple cleavage sites. In a single isolate we found
one to seven sites.
Conclusions: We observed that the amount of PbGP43 transcripts accumulated in P. brasiliensis
Pb339 grown in defined medium was about 1,000-fold higher than in Pb18 and 120-fold higher than
in Pb3. We have described a series of features in the gene flanking regions and differences among
isolates, including DNA-binding sequences, which might impact gene regulation. Little is known
about regulatory sequences in thermo-dimorphic fungi. The peculiar structure of tandem repetitive
fragments in the 5' intergenic region of PbGP43, their characteristic sequences, besides the
presence of multiple poly(A) cleavage sites in the 3' UTR will certainly guide future studies.
Page 1 of 13
(page number not for citation purposes)
BMC Microbiology 2009, 9:277
Background
Paracoccidioides brasiliensis is a thermo-dimorphic pathogenic fungus. It causes paracoccidiodomycosis (PCM) in
man, which is an endemic mycosis in Latin America that
affects mostly the lungs, but can disseminate to other
organs [1]. P. brasiliensis is multinucleated in both pathogenic yeast and infectious mycelial phases. Genetic transformation in the species has recently been optimized [2],
however genetic manipulation is still in its infancy. It is
now recognized that most P. brasiliensis isolates diversified
into an S1 main species, which is genetically close to the
PS3 group of Colombian isolates, while PS2 is composed
of a few isolates that constitute a phylogenetically cryptic
species [3].
Gp43 is the main diagnostic and prognostic antigen so far
characterized in P. brasiliensis [4,5]. It is a secretory glycoprotein whose peptide structure bears antigenic properties
that are peculiar to the species [6]. Therefore, it confers
high levels of sensitivity and specificity for PCM patients'
sera when used as antigen in diagnostic tests such as
immunodiffusion and capture ELISA, as well as by antigen
detection in biological fluids [7]. Antibody titers are
directly proportional to the severity of active PCM; they
are probably not protective in advanced stages of the disease, but experimental protocols in mice point to the
immunotherapeutic potential of anti-gp43 monoclonal
antibodies [8]. On the other hand, gp43 contains T cell
epitopes that are protective to vaccinated mice [5]. The
best studied T-cell epitope is 15 aminoacid-long P-10,
which showed additive effect in the treatment of murine
PCM when administered with anti-fungal agents [9]. In
addition, gp43 has adhesive properties to extracellular
matrix proteins that may help fungal dissemination
[10,11].
The complete PbGP43 ORF has originally been found in a
cloned 3,800-bp EcoRI genomic region from the Pb339
(B-339) isolate. It comprises 1,329 bp that contain a
unique 78-bp intron [12]. The EcoRI genomic fragment
includes 326 bp from the PbGP43 5' intergenic proximal
region and about 500 bp of the 3' intergenic sequence,
which is shared by a neighboring RanBP homologue. This
gene encodes a nuclear Ran-binding protein in Schizosaccharomyces pombe, or importin 11 in Aspergillus fumigatus,
that transports ribosomal proteins to the nucleus [13].
PbGP43 and PbRanBP are linked in twelve P. brasiliensis
isolates, as observed by Feitosa et al. [14].
Our group has carried out original and detailed studies on
sequence polymorphism in the PbGP43 ORF [15] and 5'
intergenic proximal region [16], which defined at least
five genotypes [17]. When compared to a consensus
sequence, the most polymorphic A genotype carries three
substitutions in the 5' intergenic proximal region and up
to fifteen informative sites in the ORF, mostly concen-
http://www.biomedcentral.com/1471-2180/9/277
trated in exon 2. So far, the A genotype has been detected
in all six PS2 isolates [3]. It is of note that PbGP43 was the
most polymorphic gene in the multilocus analysis performed by Matute et al. [3] in P. brasiliensis. Isolates Pb2,
Pb3 and Pb4, which belong in PS2 group [3], evoked
milder experimental PCM in B10. A mice than representative isolates from the main species S1, including Pb18
[16]. This isolate has been long used in experimental PCM
due to its high virulence.
P. brasiliensis Pb339 has traditionally been employed in
antigen preparation [18]. It secretes high amounts of
gp43, however that is not a rule among isolates [19]. (...truncated)