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PubMed2003

Identification and characterization of HtsA, a second heme-binding protein made by Streptococcus pyogenes.

Group A streptococci (GAS) can use heme and hemoproteins as sources of iron. However, the machinery for heme acquisition in GAS has not been firmly revealed. Recently, we identified a novel heme-associated cell surface protein (Shp) made by GAS. The shp gene is cotranscribed with eight downstream genes, including spy1795, spy1794, and spy1793 encoding a putative ABC transporter (designated HtsABC). In this study, spy1795 (designated htsA) was cloned from a serotype M1 strain, and recombinant HtsA was overexpressed in Escherichia coli and purified to homogeneity. HtsA binds 1 heme molecule per molecule of protein. HtsA was produced in vitro and localized to the bacterial cell surface. GAS up-regulated transcription of htsA in human blood compared with that in Todd-Hewitt broth supplemented with 0.2% yeast extract. The level of the htsA transcript dramatically increased under metal cation-restricted conditions compared with that under metal cation-replete conditions. The cation content, cell surface location, and gene transcription of HtsA were also compared with those of MtsA and Spy0385, the lipoprotein components of two other putative iron acquisition ABC transporters of GAS. Our results suggest that HtsABC is an ABC transporter that may participate in heme acquisition in GAS.

ATP-Binding Cassette Transporters↗

PubMed2005

Role of the N-terminal region and of beta-sheet residue Thr29 on the activity of the omega2 global regulator from the broad-host range Streptococcus pyogenes plasmid pSM19035.

The dimeric regulatory protein wild-type omega (wt omega2) binds to arrays of 7-bp sequences (heptads) present in the operator DNA region of copy control and partition functions of plasmid pSM19035. Each omega2 protein probably binds with an antiparallel beta-sheet structure in the major groove of the 7-bp subsite of the operator DNA. Exchange of threonine at position 29 to alanine (T29A) drastically affects the activity of variant protein omega2T29A both in vivo and in vitro, and reduces the thermodynamic stability deltaG(o)u, but does not change the conformation. Likewise, the binding affinity to DNA is reduced and the association of the two monomeric subunits of the omega2T29A dimer is weakened, as manifested by an increase in the dissociation constant from 3.2 microM for wt omega2 to 6.3 microM for omega2T29A. Denatured dimers are formed upon thermal unfolding of wt omega2 and omega2T29A at ca. 45 microM (D(n)<-->D(u)). Removal of 8 (omega2deltaN8), or even 18 (omega2deltaN18) N-terminal amino acids has no obvious effect either on the core structure or on the activity in comparison to wt omega2. The stability of variants omega2deltaN8 and omega2deltaN18 is similar to that of wt omega2, and their binding to operator DNA is not impaired.

Bacterial Proteins↗

PubMed1983

Type 49 Streptococcus pyogenes: phage subtypes as epidemiological markers in isolates from skin sepsis and acute glomerulonephritis.

Studies of group A, M type 49 streptococci from England, Trinidad and Alaska indicate that isolates of this serotype often differ with respect to phage subtype from one geographical area to another, but are generally homogeneous in one place at one time. The findings support the conclusion that acute glomerulonephritis can be associated with a variety of phage subtypes of M type 49 streptococci. In outbreaks of skin sepsis without nephritis in England, the phage subtypes of M type 49 streptococci isolated from skin lesions of meat handlers were the same as those recovered from skin lesions of non-meat handlers in the same community. The findings on the Trinidad isolates suggest that M type 49 streptococci of one phage subtype may persist in a population for 9 years and may result in a second outbreak of acute glomerulonephritis. In an Alaska Eskimo population in whom acute glomerulonephritis was occurring, most of the M type 49 isolates available for testing were of a single phage subtype. Equally prevalent in this population were group A streptococci that exhibited the same T antigen as the type 49 isolates but differed in their serum opacity reaction and phage subtype. This apparently related strain was not typable with available M antisera but showed functional evidence of M protein and is probably a new M type.

Alaska↗

PubMed1998

Molecular co-operation between protein PAM and streptokinase for plasmin acquisition by Streptococcus pyogenes.

Bacterial surface-associated plasmin formation is believed to contribute to invasion, although the underlying molecular mechanisms are poorly understood. To define the components necessary for plasmin generation on group A streptococci we used strain AP53 which exposes an M-like protein ("PAM") that contains a plasminogen-binding sequence with two 13-amino acid residues long tandem repeats (a1 and a2). Utilizing an Escherichia coli-streptococcal shuttle vector, we replaced a 29-residue long sequence segment of Arp4, an M-like protein that does not bind plasminogen, with a single (a1) or the combined a1a2 repeats of PAM. When expressed in E. coli, the purified chimeric Arp/PAM proteins both bound plasminogen, as well as plasmin, and when used to transform group A streptococcal strains lacking the plasminogen-binding ability, transformants with the Arp/PAM constructs efficiently bound plasminogen. Moreover, when grown in the presence of plasminogen, both Arp/PAM- and PAM-expressing streptococci acquired surface-bound plasmin. In contrast, plasminogen activation failed to occur on PAM- and Arp/PAM-expressing streptococci carrying an inactivated streptokinase gene: this block was overcome by exogenous streptokinase. Together, these results provide evidence for an unusual co-operation between a surface-bound protein, PAM, and a secreted protein, streptokinase, resulting in bacterial acquisition of a host protease that is likely to spur parasite invasion of host tissues.

Antigens, Bacterial↗

PubMed2003

Susceptibility of skin and soft-tissue isolates of Staphylococcus aureus and Streptococcus pyogenes to topical antibiotics: indications of clonal spread of fusidic acid-resistant Staphylococcus aureus.

Staphylococcus aureus (SA) isolates (n = 255) from outpatients with skin and soft-tissue infections were collected in 3 different areas in Norway. Group A streptococci (GAS, n = 68) were isolated from skin or pharyngotonsillar specimens from outpatients. Minimum inhibitory concentrations (MIC) of bacitracin, fusidic acid and mupirocin were tested using the E-test. Pulsed field gel electrophoresis (PFGE) patterns of fusidic acid-sensitive (FusS) and -resistant (FusR) SA were compared. All GAS isolates showed MIC of bacitracin of < or = 1.0 mg/l, of mupirocin of < or = 0.125 mg/l and of fusidic acid 1.0-4.0 mg/l. All the SA showed MIC of mupirocin < or = 0.5 mg/l and of bacitracin of > or = 2.0 mg/l, 91% with MIC > or = 16 mg/l. FusR was shown by 32.5% of the SA strains with similar prevalence rates in 3 different geographical areas of Norway. One particular PFGE pattern (type 1) was shown by 76% of the FusR SA. SA of type 1 belonged to phage group II and produced exfoliative toxins. Thus, the results demonstrated a high prevalence of FusR among SA causing skin infections and that this was mainly due to dissemination of clonally related FusR SA.

Administration, Topical↗

PubMed2004

Molecular characterization of Streptococcus pyogenes isolates collected during periods of increased acute rheumatic fever activity in Utah.

BACKGROUND: Salt Lake City, Utah has seen a continuing resurgence of rheumatic fever (RF) since 1985. METHODS: emm genotyping and multilocus sequence typing of streptococcal isolates from periods of increased RF activity were performed. RESULTS: Multiple genotypes were present during 1985 and 1998, two peak years of RF activity, and in 1992, a year with reduced RF activity. emm3 and emm18.1 were present in 1985 and 1998, but not in 1992. Two other emm types, 12 and L28, were significantly elevated in 1998 (a peak RF year) over 1992 (a non-peak RF year). Allelic profiles for the emm3 and emm18.1 isolates exhibited clonality. CONCLUSIONS: During years of increased RF activity multiple emm types, including emm18.1 and emm3, were circulating in the community. During a year of decreased RF activity, emm3 and emm18.1 genotypes were absent. The clonality of the emm3 and emm18.1 types suggests that specific clones of both types are important in the resurgence of RF during these peak years. Two other genotypes, emm12 and emmL28, may also be associated with the persistence of RF in the Salt Lake City, UT area.

Bacterial Outer Membrane Proteins↗

PubMed2003

[Molecular epidemiology of a streptococcus pyogenes related nosocomial outbreak in a burn unit].

BACKGROUND: Group A Streptococcal (GAS) infections have increased in frequency and severity worldwide. During April 1996, a nosocomial outbreak associated to GAS infections affected seven patients admitted to a pediatric burn unit. The causative organism was likely disseminated from the source patient to another child in the emergency room before he was transferred to the burn unit. Patients developed burn infections or invasive disease. One of them died due to a toxic shock syndrome and 3 other lost their skin grafts. Perineal and nasal microbiological surveillance of 42 related health care workers identified only one of them as carrier of S pyogenes. AIM: To report a molecular analysis of an apparently clonal outbreak. MATERIAL AND METHODS: The available isolates were analyzed by molecular methods including random amplified polymorphic DNA analysis (RAPD) with 4 different primers, Sma-I pulsed field gel electrophoresis (PFGE) analysis, and speA, speB and speC detection by polymerase chain reaction (PCR). RESULTS: Two phylogenetically distant and sequentially isolated bacterial groups were identified either by RAPD analysis with selected primers or by Smal-PFGE analysis. The first group involved isolates identified in two patients that included the lethal case. The second bacterial group comprised 5 clinical isolates and the perineal and nasal isolates obtained from a health care worker. Only strains belonging to the first group harbored the speA gene and were associated with invasive disease. The second group could be split further in two subgroups according to their speB profile. CONCLUSIONS: RAPD analysis with selected primers can reproduce the PFGE-discriminating ability on the epidemiological analysis of GAS infections.

Burn Units↗

PubMed1984

Streptococcus pyogenes streptolysin O as a cause of false-positive CAMP reactions.

The synergistic hemolysis of sheep erythrocytes in the CAMP reaction by the sequential action of staphylococcal beta-lysin and the CAMP factor of group B streptococci is the only known function of this extracellular product of group B streptococci. The reaction forms the basis of the CAMP test used to identify group B streptococci because the CAMP factor is believed to be restricted to this group of organisms. However, on occasion other streptococci, notably group A streptococci, may produce a similar synergistic lysis of sheep erythrocytes. The nature of the synergistic lytic factor of group A streptococci responsible for this sequential hemolysis was investigated in a tube CAMP reaction system. The properties of this synergistic lytic factor were found to correspond to those of streptolysin O of group A streptococci. The synergistic lytic factor, like streptolysin O, was produced during the logarithmic phase of growth; the activity was increased by reducing agents and greatly decreased or abolished by heat, trypsin, cholesterol, and anti-streptolysin O, and it was immunogenic in rabbits. This would suggest that the synergistic hemolysis seen in the CAMP reaction system with group A streptococci is due to the action of those small amounts of streptolysin O which remain unoxidized and thus have a capacity to lyse the fragile beta-lysin-treated sheep erythrocytes.

Animals↗

PubMed1988

Molecular evolution of streptococcal M protein: cloning and nucleotide sequence of the type 24 M protein gene and relation to other genes of Streptococcus pyogenes.

The structural gene for the type 24 M protein of group A streptococci has been cloned and expressed in Escherichia coli. The complete nucleotide sequence of the gene and the 3' and 5' flanking regions was determined. The sequence includes an open reading frame of 1,617 base pairs encoding a pre-M24 protein of 539 amino acids and a predicted Mr of 58,738. The structural gene contains two distinct tandemly reiterated elements. The first repeated element consists of 5.3 units, and the second contains 2.7 units. Each element shows little variation of the basic 35-amino-acid unit. Comparison of the sequence of the M24 protein with the sequence of the M6 protein (S. K. Hollingshead, V. A. Fischetti, and J. R. Scott, J. Biol. Chem. 261:1677-1686, 1986) indicates that these molecules have are conserved except in the regions coding for the antigenic (type specific) determinant and they have three regions of homology within the structural genes: 38 of 42 amino acids within the amino terminal signal sequence, the second repeated element of the M24 protein is found in the M6 molecule at the same position in the protein, and the carboxy terminal 164 amino acids, including a membrane anchor sequence, are conserved in both proteins. In addition, the sequences flanking the two genes are strongly conserved.

Amino Acid Sequence↗

PubMed1995

Increased incidence and severity of Streptococcus pyogenes bacteremia in young children.

An increase in the incidence and severity of bacteremia caused by group A streptococci was noted in 1993 and 1994 in the Hadassah University Medical Center, Jerusalem. During the 6-year period 1987 to 1992, 12 children with group A streptococcal bacteremia were hospitalized, whereas in 1993 and 1994 there were 17 patients, 5 of them with 1 each of the following severe clinical manifestations: meningitis and septic shock; streptococcal toxic shock syndrome; septic shock; pleural empyema; and fatal outcome. Our 29 patients with group A streptococcal bacteremia were younger than those reported in the literature: 10 (35%) were < 3 months of age; 17 (59%) were < 1 year old. Most children were previously healthy and only 3 had an underlying immunodeficiency predisposing to infection (1 case each): leukemia; Di George syndrome; and congenital nephrotic syndrome. Two children were recovering from varicella. The skin was the most common site of primary infection (16 of 29). The average white blood cell (WBC) count was 18 150 cells/mm3 (range, 2200 to 34,200). The cases were not related epidemiologically and were caused by a variety of M-protein types. Polymerase chain reaction amplification of the genes encoding exotoxins A (speA) and C (speC) was done on 19 isolates and disclosed 2 strains positive for speA and 5 positive for speC. One of the speA-positive isolates was from the single patient with toxic shock syndrome.

Age Distribution↗

PubMedDate not supplied

Attachment of Streptococcus pyogenes to mammalian cells.

Accumulated evidence indicates that lipoteichoic acid (LTA) is centrally involved in the attachment of group A streptococci to epithelial cells of the host. The binding of LTA to a variety of host cells is mediated by the glycolipid end of the LTA molecule, which can form ionic complexes with streptococcal surface proteins, permitting the reorientation of the LTA to expose some of its lipid ends toward the surface of the organism. The ability of albumin to block the adherence of streptococci to epithelial cells and to bind to LTA-M protein but not deacylated LTA-M protein complexes supports the idea that the lipid ends remain free to interact with cell membrane receptors. The cell membrane receptors appear to consist of a lipid-binding region(s) on fibronectin molecules on the surfaces of oropharyngeal epithelial cells. Although streptococci exhibit LTA-sensitive binding to phagocytic cells in a serum-free system, in the presence of serum, they do not; rather, complement bound to the streptococcal cell surface is required for recognition by phagocytes. The binding of fibrinogen to the M protein on the surface of M-rich streptococci specifically blocks the recognition of the organisms by opsonic complement components. The attachment of M-rich streptococci to phagocytic cells requires the development of antibodies directed specifically toward regions of M protein not blocked by fibrinogen.

Adhesiveness↗

PubMed1991

Molecular analysis of pyrogenic exotoxins from Streptococcus pyogenes isolates associated with toxic shock-like syndrome.

Toxic shock-like syndrome (TSLS) is characterized by hypotension or shock, fever, multiorgan system involvement, and a concurrent group A streptococcal infection. We analyzed 34 streptococcal strains isolated from patients with clinically well-documented TSLS for their pyrogenic toxin profiles and M-protein types. Although strains of nine different M types were represented in the sample, 74% of the isolates were of either M type 1 or 3. It was determined that 53% produced streptococcal pyrogenic exotoxin type A under in vitro growth conditions and that 85% contained the gene encoding this toxin. These values are in contrast to the published value of 15% for the incidence of this gene in a sample of general group A streptococcal isolates. As has been found with all group A streptococci examined to date, regardless of disease association, 100% of TSLS-associated isolates contained the gene encoding pyrogenic exotoxin type B. This toxin was detectably produced by 59% of isolates. The gene encoding pyrogenic toxin type C was found in only 21% of isolates. We conclude that the pyrogenic exotoxin type A gene is associated with group A streptococcal strains isolated from patients with TSLS and may play a causative role in this illness. However, other factors are also likely to be important, since not all strains from patients with TSLS contained the A toxin gene.

Antigens, Bacterial↗

PubMed2002

Processing, stability, and kinetic parameters of C5a peptidase from Streptococcus pyogenes.

A recombinant streptococcal C5a peptidase was expressed in Escherichia coli and its catalytic properties and thermal stability were subjected to examination. It was shown that the NH2-terminal region of C5a peptidase (Asn32-Asp79/Lys90) forms the pro-sequence segment. Upon maturation the propeptide is hydrolyzed either via an autocatalytic intramolecular cleavage or by exogenous protease streptopain. At pH 7.4 the enzyme exhibited maximum activity in the narrow range of temperatures between 40 and 43 degrees C. The process of heat denaturation of C5a peptidase investigated by fluorescence and circular dichroism spectroscopy revealed that the protein undergoes biphasic unfolding transition with Tm of 50 and 70 degrees C suggesting melting of different parts of the molecule with different stability. Unfolding of the less stable structures was accompanied by the loss of proteolytic activity. Using synthetic peptides corresponding to the COOH-terminus of human complement C5a we demonstrated that in vitro peptidase catalyzes hydrolysis of two His67-Lys68 and Ala58-Ser59 peptide bonds. The high catalytic efficiency obtained for the SQLRANISHKDMQLGR extended peptide compared to the poor hydrolysis of its derivative Ac-SQLRANISH-pNA that lacks residues at P2'-P7' positions, suggest the importance of C5a peptidase interactions with the P' side of the substrate.

Adhesins, Bacterial↗

PubMed1993

PAM, a novel plasminogen-binding protein from Streptococcus pyogenes.

The ability of group A streptococci to bind human plasminogen and plasmin has attracted interest, because it could provide the bacteria with a mechanism for invasion. M or M-like proteins account for the binding of several plasma proteins to group A streptococci. To investigate whether M or M-like proteins were responsible for the binding of plasminogen to group A streptococci, acid-extracted material from a type M53 streptococcal isolate was tested for its ability to bind plasminogen. Indeed, a 42-kDa plasminogen-binding protein was solubilized. Two oligonucleotides homologous with conserved sequences in known M protein genes were used as primers in the polymerase chain reaction, with chromosomal DNA from the M53 isolate. When cloned and expressed in Escherichia coli, a resulting fragment encoded a 43-kDa plasminogen-binding protein. Nucleotide sequence determination of the gene fragment revealed an open reading frame encoding a polypeptide of 43,580 Da, which matched the amino-terminal amino acid sequence of the plasminogen-binding protein extracted from M53 streptococci. The DNA sequence data also proved the relationship of the encoded protein, named PAM, to the M proteins. The plasminogen-binding domain was mapped to the amino-terminal third of PAM. Plasminogen absorbed by M53 streptococci or by immobilized PAM could be activated by streptokinase. The results provide further evidence of the diversity of the M protein family and suggest a new mechanism whereby these proteins contribute to the virulence of group A streptococci.

Amino Acid Sequence↗

PubMed2002

Comparison of two rapid Streptococcus pyogenes diagnostic tests with a rigorous culture standard.

BACKGROUND: Comparisons of rapid antigen tests for the diagnosis of pharyngitis are often made between published studies but may not be reliable given differences in study design, sampling and reference standard definitions. Tests are rarely compared against each other in a single study. METHODS: The sensitivity and specificity of two rapid diagnostic tests were compared against a multiplate culture standard defined as the identification of on any of four culture plates. Paired swabs were tested for antigen using Genzyme's OSOM Ultra Strep A Test and compared with Biostar's Strep A OIA Max Test. RESULTS: Ninety-four (31.1%) of 302 matched samples were identified with The sensitivity of Genzyme's OSOM Ultra Strep A Test against the multiplate culture standard was 92.6%, significantly higher ( P= 0.001) than that (75.5%) of Biostar's Strep A OIA Max Test. Specificities were 92.8 and 97.1%, respectively. Data analysis of culture results and statistical modeling showed that cultures of two or more samples are necessary for a sensitivity of 95% or greater for a comparison standard compared with true disease status. CONCLUSIONS: When comparing the performance of rapid antigen tests for pharyngitis, a rigorous culture standard should be used consisting of at least two separate samples (swabs and/or pledgets), ultimately plated on selective agar. Genzyme's OSOM Ultra Strep A Test was significantly more sensitive than Biostar's Strep A OIA Max Test in comparison with a multiplate culture standard and a same swab, single plate culture standard.

Antigens, Bacterial↗

PubMed1996

[Serotype distribution and antimicrobial susceptibility of group A streptococci (Streptococcus pyogenes) isolated in Taiwan].

T-protein serotypes and antimicrobial susceptibility of a total of 139 group A streptococci (GAS) strains isolated in Taiwan area in 1993 and during the outbreak of scarlet fever in 1994 were analyzed. All strains were T-typable, and T12 (42.46%) and T4 (38.85%) were the dominant T types. According to the results of analysis of antimicrobial susceptibility, all GAS strains were divided into 9 resistotypes, A (all susceptible), B (resistant to tetracycline), C (resistant to erythromycin and tetracycline), D (resistant to chloramphenicol and tetracycline), E (resistant to chloramphenicol and clindamycin), F (resistant to chloramphenicol, clindamycin and tetracycline), G (resistant to clindamycin, erythromycin and tetracycline), H (resistant to chloramphenicol, clindamycin, erythromycin and tetracycline), and I (resistant to chloramphenicol, clindamycin, erythromycin, tetracycline and vancomycin). Type B (37.42%) was the dominant type. Type A (25.91%), and type H (26.63%) also appered with high incidence. Most of strains isolated from Mid-Taiwan were type H. Only one strain, that was isolated in I-lan, was resistant to vancomycin, in addition to resistant to chloramphenicol, clindamycin, erythromycin, and tetracycline. All strains were susceptible to penicillin G, ampicillin, and ceftriaxone. Some strains were resistant to chloramphenicol (32.38%), clindamycin (30.22%), erythromycin (31.66%), tetracycline (73.39%), and vancomycin (0.70%). During the outbreak of scarlet fever in 1994, the dominant T types of strains isolated in North-Taiwan and Mid-Taiwan were T4 and T12, respectively, and the major resistotypes of those strains were B and H types, respectively. These clues suggested that the outbreaks occurring in North-Taiwan and Mid-Taiwan may have no epidemiological linkage between each other.

Disease Outbreaks↗

PubMed1993

Staphylococcus aureus toxic shock syndrome toxin 1 and Streptococcus pyogenes erythrogenic toxin A modulate inflammatory mediator release from human neutrophils.

We studied the influence of staphylococcal toxic shock syndrome toxin 1 and streptococcal erythrogenic (pyrogenic) toxin A (ETA) on intact and digitonin-permeabilized human polymorphonuclear granulocytes (PMNs). As was shown by reversed-phase high-performance liquid chromatography analysis, toxic shock syndrome toxin 1 or ETA alone, in the absence of any additional stimulus, did not induce the generation of the chemoattractant leukotriene B4 (LTB4) from PMNs in a wide range of concentrations. In addition, pretreatment of intact PMNs with either toxin potentiated formyl-methionyl-leucyl-phenylalanine (fMLP)- and washed Staphylococcus aureus cell-induced generation of LTB4 in a time- and dose-dependent manner. This increase included LTB4 as well as its inactive omega-oxidated compounds. Further studies revealed evidence that toxin exposure was accompanied by enhanced cellular receptor expression for fMLP as well as for LTB4. The intrinsic GTPase activity of membrane fractions was modulated by both toxins. Short-term incubation with ETA increased the GTPase activity of PMNs up to 141%. Inhibitory effects were obtained when GTP-binding protein functions were stimulated with sodium fluoride (NaF). In addition, specific binding of Gpp(NH)p to GTP-binding protein was inhibited by both toxins during the first 10 min of incubation and was restored at later times of incubation. Our data therefore suggest that both toxins significantly affect the signal transduction pathways of human PMNs, which results in immunomodulatory functions.

Bacterial Proteins↗

PubMed2005

M protein conserved region antibodies opsonise multiple strains of Streptococcus pyogenes with sequence variations in C-repeats.

The development of a group A streptococcal (GAS) vaccine has focused on the M protein, a major virulence factor. Antibodies against the amino terminal domain of the M protein are generally protective but only provide type-specific immunity. J14, a 29-mer peptide sequence which contains a conserved epitope from the C-repeat region of the M protein, offers the possibility of a vaccine which will elicit protective opsonic antibodies against multiple GAS strains. In this study we have shown that antibodies raised against J14 are capable of opsonising 37 GAS isolates representing different emm types derived from a region in which GAS infection is endemic. We also demonstrate that J14 antisera is capable of opsonising GAS isolates containing J14 homologues but not J14-specific sequences, further increasing the strain coverage of this vaccine candidate. Isolates with three C-repeats were opsonised more efficiently than isolates with two repeats. Opsonisation of a strain with only a single C-repeat was dramatically lower than other strains tested. The number of C-repeats present in the M protein of individual isolates therefore appears to be the critical factor in determining bactericidal capacity of J14 antisera. The reduced opsonic capacity of sera against this strain was shown to correlate with a reduced capacity to bind J14 antisera, as demonstrated by immunofluorescence microscopy and FACS analysis. In vivo challenge experiments also confirmed the protective efficacy of immunisation with J14 peptide.

Adolescent↗