Actinobacillus suis: Respiratory and Septicemic Disease in Pigs – Diagnosis and Control
Etiology and Taxonomy
Actinobacillus suis is a Gram-negative, facultatively anaerobic, pleomorphic coccobacillus belonging to the family Pasteurellaceae. The organism is phylogenetically related to Actinobacillus pleuropneumoniae and Haemophilus parasuis (now Glaesserella parasuis) [1]. A. suis requires nicotinamide adenine dinucleotide (NAD; V factor) for growth but does not require hemin (X factor), distinguishing it from certain other Pasteurellaceae. Colonies on chocolate or supplemented blood agar are small, smooth, and grayish, with a characteristic sweetish odor.
The bacterium possesses a range of virulence factors, including capsular polysaccharide, lipopolysaccharide (LPS), outer membrane proteins, and RTX (repeats in toxin) toxins. Genomic analyses have revealed that A. suis isolates harbor variable numbers of toxin gene copies (apxI, apxII) which correlate with virulence potential [2]. Serotyping is not routinely performed, but most clinical isolates belong to serovars 1 and 2, with serovar 1 associated more frequently with septicemia.
Epidemiology
A. suis is a commensal inhabitant of the upper respiratory tract and tonsils of clinically healthy pigs, including both domestic swine and feral populations [3, 4]. The primary reservoir is the carrier pig, and transmission occurs horizontally via direct contact, aerosolized respiratory secretions, and contaminated fomites. The tonsil serves as a major colonization site; A. suis adheres to extracellular matrix components of the tonsillar epithelium, facilitating persistent colonization [5].
Disease is most commonly observed in nursery and early grow-finish pigs, typically between 3 and 12 weeks of age, when maternal antibody wanes and stressors such as weaning, mixing, transport, and concurrent viral infections predispose to clinical disease [1]. Outbreaks can occur in herds naïve to the pathogen or in situations where virulent strains are introduced. The prevalence of A. suis in respiratory disease complex cases varies; in diagnostic surveillance from a US veterinary laboratory (2017-2022), A. suis was detected in approximately 5.2% of submissions [1], while cross-sectional studies in China reported detection in 7.3% of porcine respiratory disease complex (PRDC) cases [6, 7]. Infection dynamics are influenced by co-infections with primary viral pathogens such as porcine reproductive and respiratory syndrome virus (PRRSV) and swine influenza virus [8, 9].
Clinical Signs and Pathogenesis
Two major clinical presentations are recognized: acute septicemia and subacute/chronic respiratory disease. In peracute septicemia, pigs present with fever (40.5-42.0°C), lethargy, anorexia, cyanosis of the extremities, and sudden death. Cutaneous lesions, including erythema, petechiae, and ecchymoses, are sometimes noted, resembling those seen in erysipelas and other Gram-negative septicemias [10].
The respiratory form is characterized by cough, dyspnea, tachypnea, and nasal discharge. Affected pigs may adopt a dog-sitting posture to facilitate breathing. Morbidity can reach 20-30% in affected groups, with mortality varying from 2% to 50% in untreated septicemic outbreaks.
Pathogenetically, A. suis invades the tonsillar crypts and multiplies in the upper respiratory tract. Under immunosuppressive conditions or in the presence of the porcine respiratory disease complex (PRDC), the organism breaches mucosal barriers and enters the bloodstream, causing septicemia with disseminated intravascular coagulation and multi-organ failure. The RTX toxins (ApxI, ApxII) induce pore formation in host cell membranes, leading to necrosis of alveolar macrophages and pulmonary endothelium, resulting in pulmonary edema, fibrinous pleuritis, and hemorrhagic necrosis [2]. Lipid profiling of A. suis during co-infection with PRRSV shows altered host lipid metabolism, potentially enhancing bacterial survival [11].
Pathology
Gross lesions in septicemic cases include generalized vascular congestion, petechial hemorrhages on serosal surfaces, epicardium, and kidneys, as well as fibrinonecrotic pneumonia. Serosanguinous pleural effusion and fibrinous polyserositis (pleuritis, pericarditis, peritonitis) are common. The lungs often exhibit cranioventral consolidation with multifocal necrotic foci. In the respiratory form, lesions are predominantly in the cranial and middle lung lobes, with purulent bronchopneumonia and interlobular edema. Histologically, there is necrotizing bronchopneumonia with intense infiltration of neutrophils, fibrin exudation, and thrombosis of pulmonary vessels. These lesions are difficult to differentiate from those caused by A. pleuropneumoniae and other Pasteurellaceae without ancillary diagnostic testing [12, 13].
Diagnosis
Accurate diagnosis of A. suis infection requires a combination of clinical, pathological, and laboratory methods.
Sample Collection
Suitable specimens include: tonsil swabs, nasal swabs, bronchoalveolar lavage fluid, lung tissue (affected and unaffected), pleural fluid, and blood from acutely febrile pigs. Swabs should be placed in transport medium (e.g., Amies with charcoal) and shipped refrigerated. Postmortem samples should include fresh lung and tonsil for culture and molecular testing.
Bacteriological Culture
A. suis grows on chocolate agar or blood agar supplemented with NAD (V factor) when incubated at 37°C in 5% CO2. Satellite growth around a Staphylococcus aureus nurse streak (which provides NAD) is a classical isolation method. Colonies appear after 24-48 hours as small, grayish, translucent, and non-hemolytic (or weakly hemolytic on bovine blood agar). Biochemical identification reveals urease-positive, oxidase-positive, indole-negative, and fermentation of glucose, sucrose, and mannitol. Matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS) provides rapid and accurate identification [1].
Molecular Diagnostics
Polymerase chain reaction (PCR) assays targeting the apxIV gene (conserved among Actinobacillus species) or the 16S rRNA gene are used for species-specific detection. Multiplex real-time PCR panels allow simultaneous detection of A. suis alongside other PRDC pathogens such as Streptococcus suis, Glaesserella parasuis, and A. pleuropneumoniae [14, 15, 16, 17]. A triplex real-time PCR for S. suis, G. parasuis, and A. pleuropneumoniae has been validated [14]; incorporating A. suis in similar panels is recommended for diagnostic efficiency.
Targeted next-generation sequencing (tNGS) panels offer a comprehensive approach, enabling simultaneous detection and differentiation of multiple bacterial and viral respiratory pathogens, including A. suis, with high sensitivity and specificity [18]. This technology can be particularly useful for polymicrobial PRDC cases.
Serological assays, such as ELISA, are available for population-level screening to detect exposure, but are not generally used for individual diagnosis.
Mermaid Diagnostic Workflow Diagram
flowchart TD
A[Acute respiratory disease / sudden death in pigs 3-12 wks], > B{Postmortem examination}
B, > C[Gross lesions: fibrinous pneumonia, serosanguinous pleural effusion, petechiae]
C, > D[Select fresh lung, tonsil, pleural fluid]
D, > E{Primary laboratory methods}
E, > F[Gram stain: Gram-negative coccobacilli]
E, > G[Culture on NAD-supplemented agar (chocolate / satellite growth)]
G, > H[Biochemical / MALDI-TOF identification]
E, > I[Molecular detection: real-time multiplex PCR (apxIV / 16S rRNA) or tNGS panel]
H, > J[Positive: Actinobacillus suis]
I, > J
J, > K{Antimicrobial susceptibility testing}
K, > L[MIC determination by broth microdilution or disk diffusion]
L, > M[Select appropriate antimicrobial based on susceptibility profile]
Treatment
Antimicrobial therapy should be initiated immediately based on clinical suspicion, ideally guided by antimicrobial susceptibility testing (AST) of the isolated strain. A. suis is generally susceptible to beta-lactams, cephalosporins, fluoroquinolones, trimethoprim-sulfonamides, and certain macrolides, but resistance patterns vary geographically.
Commonly Used Antimicrobials
Beta-lactams: Ceftiofur (third-generation cephalosporin) is frequently used for parenteral treatment of septicemic and respiratory infections. Pharmacokinetic/pharmacodynamic (PK/PD) modeling supports optimized dosing regimens for ceftiofur to achieve target attainment against porcine respiratory pathogens [19]. Amoxicillin administered in drinking water has been evaluated for respiratory pathogens, but time-restricted dosing may be necessary to improve exposure and efficacy; predicted PK/PD inadequacy has been noted for amoxicillin in water against A. suis [20, 21].
Fluoroquinolones: Pradofloxacin and marbofloxacin exhibit strong bactericidal activity against A. suis and other swine respiratory pathogens [22, 23]. Resistance is still low in European isolates (less than 5%) [24].
Macrolides: Gamithromycin administered intramuscularly has shown efficacy against bacterial swine respiratory disease, though data specific to A. suis are limited [25].
Other classes: Trimethoprim-sulfonamides, tetracyclines, and florfenicol are also options, but susceptibility must be confirmed. European surveillance of respiratory pathogens from pigs (2009-2012; 2018-2021) showed that A. suis maintains high susceptibility to ceftiofur, tiamulin, and florfenicol, while resistance to tetracyclines and sulfonamides is more common [26, 27, 28, 24].
The emergence of multidrug resistance (MDR) is a concern. Applying standardized definitions for MDR, extensive drug resistance (XDR), and pandrug resistance (PDR) to veterinary pathogens [29] is important for monitoring. In vitro mixed biofilm formation between A. suis and other porcine respiratory bacteria (e.g., S. suis) can reduce antimicrobial susceptibility [30], complicating therapy in polymicrobial infections.
Control and Prevention
Control of A. suis infections relies on a combination of management practices, vaccination, and prudent antimicrobial use.
Biosecurity and Management
All-in/all-out production and strict hygiene reduce pathogen load and transmission. Segregated early weaning can reduce exposure to older carrier dams [31]. Minimizing stressors such as overcrowding, poor ventilation, and temperature fluctuations is critical. Good ventilation systems help reduce ammonia levels and aerosolized bacteria.
Biosecurity measures include quarantine of incoming stock, footbaths, and dedicated farm clothing. Feral swine should be excluded from domestic pig premises, as they may carry A. suis and other bacterial pathogens [3, 4].
Vaccination
Commercial vaccines for A. suis are not widely available in all regions. Autogenous (bacterin) vaccines are sometimes used in endemically infected herds. The efficacy of bacterial vaccines for swine respiratory disease has been systematically reviewed, but evidence specific to A. suis is limited [32]. Vaccination combined with other control measures may reduce the severity of outbreaks.
Antimicrobial Stewardship
Prophylactic antimicrobial use should be minimized and replaced with metaphylaxis targeting affected groups. Controlled time-restricted dosing of amoxicillin in water has been proposed to improve exposure while reducing overall antimicrobial consumption [20]. Regular surveillance of antimicrobial resistance patterns (e.g., through national programmes) is essential to guide empirical therapy [26, 28].
Role of Coinfections
Given the importance of viral triggers (especially PRRSV), control of primary viral infections through vaccination and biosecurity indirectly reduces the incidence of secondary A. suis disease [8, 9]. The impact of feed additives on clinical symptoms and the microbiome has been studied in PRRSV/S. suis co-infections [33]; similar approaches may benefit A. suis control, although data are scarce.
In summary, A. suis is an important but sometimes overlooked component of the porcine respiratory disease complex and a cause of acute septicemia in young pigs. Accurate diagnosis using culture and molecular methods, combined with AST-guided therapy and robust preventive management, is essential to reduce losses. Continued surveillance and research into vaccine development and antimicrobial alternatives are warranted.
References
[1] Silva APSP, Almeida M, Michael A, et al. Detection and disease diagnosis trends (2017-2022) for Streptococcus suis, Glaesserella parasuis, Mycoplasma hyorhinis, Actinobacillus suis and Mycoplasma hyosynoviae at Iowa State University Veterinary Diagnostic Laboratory. BMC Vet Res. 2023. https://pubmed.ncbi.nlm.nih.gov/38087358/
[2] Kulathunga DGRS, Fakher AA, Costa MO. Actinobacillus suis isolated from diseased pigs are phylogenetically related but harbour different number of toxin gene copies in their genomes. Vet Rec Open. 2022. https://pubmed.ncbi.nlm.nih.gov/36213600/
[3] Cleveland CA, DeNicola A, Dubey JP, et al. Survey for selected pathogens in wild pigs (Sus scrofa) from Guam, Marianna Islands, USA. Vet Microbiol. 2017. https://pubmed.ncbi.nlm.nih.gov/28622856/
[4] Baroch JA, Gagnon CA, Lacouture S, et al. Exposure of feral swine (Sus scrofa) in the United States to selected pathogens. Can J Vet Res. 2015. https://pubmed.ncbi.nlm.nih.gov/25673913/
[5] Bujold AR, MacInnes JI. Attachment of Actinobacillus suis H91-0380 and Its Isogenic Adhesin Mutants to Extracellular Matrix Components of the Tonsils of the Soft Palate of Swine. Infect Immun. 2016. https://pubmed.ncbi.nlm.nih.gov/27481253/
[6] Guo Y, Li Y, Wen Z, et al. Epidemiology of Major Bacterial Pathogens Associated with Porcine Respiratory Disease Complex: A Cross-Sectional Study from Intensive Swine Farms in Xinjiang, China (2024-2025). Vet Sci. 2026. https://pubmed.ncbi.nlm.nih.gov/42076738/
[7] Rao J, Wei X, Li H, et al. Novel Multiplex PCR Assay and Its Application in Detecting Prevalence and Antibiotic Susceptibility of Porcine Respiratory Bacterial Pathogens in Guangxi, China. Microbiol Spectr. 2023. https://pubmed.ncbi.nlm.nih.gov/36916923/
[8] Museau L, Hervet C, Saade G, et al. Prospecting potential links between PRRSV infection susceptibility of alveolar macrophages and other respiratory infectious agents present in conventionally reared pigs. Vet Immunol Immunopathol. 2020. https://pubmed.ncbi.nlm.nih.gov/32905850/
[9] Brockmeier SL, Loving CL, Palmer MV, et al. Comparison of Asian porcine high fever disease isolates of porcine reproductive and respiratory syndrome virus to United States isolates for their ability to cause disease and secondary bacterial infection in swine. Vet Microbiol. 2017. https://pubmed.ncbi.nlm.nih.gov/28619168/
[10] Papatsiros VG, Athanasiou LV, Psalla D, et al. Erythema Multiforme Associated with Respiratory Disease in a Commercial Breeding Pig Herd. Viral Immunol. 2015. https://pubmed.ncbi.nlm.nih.gov/26266696/
[11] Zhao Z, Chu Y, Gong J, et al. Lipid profiling of the secondary infecting bacteria with porcine reproductive and respiratory syndrome virus, including Streptococcus suis serotype 2, and their hosts. Microb Pathog. 2026. https://pubmed.ncbi.nlm.nih.gov/42114787/
[12] Piñeyro PE, Burrough ER, Almeida M, et al. Overview of porcine interstitial and bronchointerstitial pneumonia: infectious and non-infectious causes. J Vet Diagn Invest. 2026. https://pubmed.ncbi.nlm.nih.gov/42116569/
[13] Petri FAM, Ferreira GC, Arruda LP, et al. Associations between Pleurisy and the Main Bacterial Pathogens of the Porcine Respiratory Diseases Complex (PRDC). Animals (Basel). 2023. https://pubmed.ncbi.nlm.nih.gov/37174529/
[14] Zhuang H, Kang S, Zheng C, et al. A Triplex Real-Time PCR Assay for Simultaneous Detection of Streptococcus suis, Glaesserella parasuis, and Actinobacillus pleuropneumoniae. Transbound Emerg Dis. 2026. https://pubmed.ncbi.nlm.nih.gov/42181751/
[15] Goto Y, Fukunari K, Tada S, et al. A multiplex real-time RT-PCR system to simultaneously diagnose 16 pathogens associated with swine respiratory disease. J Appl Microbiol. 2023. https://pubmed.ncbi.nlm.nih.gov/37951290/
[16] Yi L, Jin M, Gao M, et al. Specific quantitative detection of Streptococcus suis and Actinobacillus pleuropneumoniae in co-infection and mixed biofilms. Front Cell Infect Microbiol. 2022. https://pubmed.ncbi.nlm.nih.gov/35992166/
[17] Sunaga F, Tsuchiaka S, Kishimoto M, et al. Development of a one-run real-time PCR detection system for pathogens associated with porcine respiratory diseases. J Vet Med Sci. 2020. https://pubmed.ncbi.nlm.nih.gov/31866601/
[18] Elshafie NO, Wilkes RP. Analytic and Diagnostic Validation of a Targeted Next-Generation Sequencing Panel for Common and Emerging Swine Respiratory Pathogens. Microorganisms. 2026. https://pubmed.ncbi.nlm.nih.gov/42197544/
[19] Zhang C, Liu H, Kou Z, et al. Optimizing the Dosage of Ceftiofur to Treat Porcine Respiratory Infection Based on a Population Pharmacokinetic Model and Monte Carlo Simulation. J Vet Pharmacol Ther. 2026. https://pubmed.ncbi.nlm.nih.gov/41235752/
[20] Lacampagne M, Hervé G, Claustre L, et al. Predicted PK/PD inadequacy of amoxicillin delivered in drinking water against respiratory pathogens in post-weaning piglets: can time-restricted dosing improve animals' exposure and potential treatment efficacy? Vet Anim Sci. 2026. https://pubmed.ncbi.nlm.nih.gov/42221906/
[21] Burch DGS, Sperling D. Amoxicillin-current use in swine medicine. J Vet Pharmacol Ther. 2018. https://pubmed.ncbi.nlm.nih.gov/29352469/
[22] Blondeau JM, Fitch SD. Bactericidal Activity of Pradofloxacin and Other Antimicrobials Against Swine Respiratory Bacterial Pathogens. Pathogens. 2025. https://pubmed.ncbi.nlm.nih.gov/41305407/
[23] El Garch F, Kroemer S, Galland D, et al. Survey of susceptibility to marbofloxacin in bacteria isolated from diseased pigs in Europe. Vet Rec. 2017. https://pubmed.ncbi.nlm.nih.gov/28348142/
[24] El Garch F, de Jong A, Simjee S, et al. Monitoring of antimicrobial susceptibility of respiratory tract pathogens isolated from diseased cattle and pigs across Europe, 2009-2012: VetPath results. Vet Microbiol. 2016. https://pubmed.ncbi.nlm.nih.gov/27102206/
[25] Xiao T, Yang Y, Zhang Y, et al. Efficacy of gamithromycin injection administered intramuscularly against bacterial swine respiratory disease. Res Vet Sci. 2020. https://pubmed.ncbi.nlm.nih.gov/31778852/
[26] de Jong A, Temmerman R, Rose M, et al. Pan-European analysis shows stable, low antimicrobial resistance in most bovine and porcine respiratory tract pathogens. Front Microbiol. 2026. https://pubmed.ncbi.nlm.nih.gov/41809594/
[27] Somogyi Z, Mag P, Simon R, et al. Susceptibility of Actinobacillus pleuropneumoniae, Pasteurella multocida and Streptococcus suis Isolated from Pigs in Hungary between 2018 and 2021. Antibiotics (Basel). 2023. https://pubmed.ncbi.nlm.nih.gov/37627719/
[28] de Jong A, Morrissey I, Rose M, et al. Antimicrobial susceptibility among respiratory tract pathogens isolated from diseased cattle and pigs from different parts of Europe. J Appl Microbiol. 2023. https://pubmed.ncbi.nlm.nih.gov/37391360/
[29] Sweeney MT, Lubbers BV, Schwarz S, et al. Applying definitions for multidrug resistance, extensive drug resistance and pandrug resistance to clinically significant livestock and companion animal bacterial pathogens. J Antimicrob Chemother. 2018. https://pubmed.ncbi.nlm.nih.gov/29481657/
[30] Wang Y, Gong S, Dong X, et al. In vitro Mixed Biofilm of Streptococcus suis and Actinobacillus pleuropneumoniae Impacts Antibiotic Susceptibility and Modulates Virulence Factor Gene Expression. Front Microbiol. 2020. https://pubmed.ncbi.nlm.nih.gov/32373078/
[31] Assavacheep P, Thanawongnuwech R. Porcine respiratory disease complex: Dynamics of polymicrobial infections and management strategies after the introduction of the African swine fever. Front Vet Sci. 2022. https://pubmed.ncbi.nlm.nih.gov/36504860/
[32] Sargeant JM, Deb B, Bergevin MD, et al. Efficacy of bacterial vaccines to prevent respiratory disease in swine: a systematic review and network meta-analysis. Anim Health Res Rev. 2019. https://pubmed.ncbi.nlm.nih.gov/32081115/
[33] Tran HT, Mercado AJ, Lahoti MM, et al. Effects of a novel feed additive on clinical symptoms and the nasal and cecal microbiome in nursery pigs challenged with PRRSV and Streptococcus suis. Transl Anim Sci. 2026. https://pubmed.ncbi.nlm.nih.gov/42211862/
[34] de Oliveira IMF, Fredriksen S, Gutiérrez MF, et al. Culturomics of the pig tonsil microbiome identifies new species and an untapped source of novel antimicrobials. Microbiome. 2025. https://pubmed.ncbi.nlm.nih.gov/40158143/
[35] Miguélez-Pérez R, Mencía-Ares O, Gutiérrez-Martín CB, et al. Biofilm formation in Streptococcus suis: in vitro impact of serovars and assessment of coinfections with other porcine respiratory disease complex bacterial pathogens. Vet Res. 2024. https://pubmed.ncbi.nlm.nih.gov/39617946/
[36] Meléndez A, Tejedor MT, Mitjana O, et al. Perception about the Major Health Challenges in Different Swine Production Stages in Spain. Vet Sci. 2024. https://pubmed.ncbi.nlm.nih.gov/38393102/
[37] Zhang K, Sun Z, Shi K, et al. RPA-CRISPR/Cas12a-Based Detection of Haemophilus parasuis. Animals (Basel). 2023. https://pubmed.ncbi.nlm.nih.gov/37958075/
[38] Siteavu MI, Drugea RI, Pitoiu E, et al. Antimicrobial Resistance of Actinobacillus pleuropneumoniae, Streptococcus suis, and Pasteurella multocida Isolated from Romanian Swine Farms. Microorganisms. 2023. https://pubmed.ncbi.nlm.nih.gov/37894069/
[39] Renzhammer R, Auer A, Loncaric I, et al. Retrospective Analysis of the Detection of Pathogens Associated with the Porcine Respiratory Disease Complex in Routine Diagnostic Samples from Austrian Swine Stocks. Vet Sci. 2023. https://pubmed.ncbi.nlm.nih.gov/37888553/
[40] Hennig-Pauka I, Hartmann M, Merkel J, et al. Coinfections and Phenotypic Antimicrobial Resistance in Actinobacillus pleuropneumoniae Strains Isolated From Diseased Swine in North Western Germany-Temporal Patterns in Samples From Routine Laboratory Practice From 2006 to 2020. Front Vet Sci. 2021. https://pubmed.ncbi.nlm.nih.gov/35155648/
[41] Zhu H, Chang X, Zhou J, et al. Co-infection analysis of bacterial and viral respiratory pathogens from clinically healthy swine in Eastern China. Vet Med Sci. 2021. https://pubmed.ncbi.nlm.nih.gov/34032016/
[42] Hayer SS, Rovira A, Olsen K, et al. Prevalence and time trend analysis of antimicrobial resistance in respiratory bacterial pathogens collected from diseased pigs in USA between 2006-2016. Res Vet Sci. 2020. https://pubmed.ncbi.nlm.nih.gov/31785428/
[43] LeBel G, Vaillancourt K, Bercier P, et al. Antibacterial activity against porcine respiratory bacterial pathogens and in vitro biocompatibility of essential oils. Arch Microbiol. 2019. https://pubmed.ncbi.nlm.nih.gov/30955056/
[44] Dione M, Masembe C, Akol J, et al. The importance of on-farm biosecurity: Sero-prevalence and risk factors of bacterial and viral pathogens in smallholder pig systems in Uganda. Acta Trop. 2018. https://pubmed.ncbi.nlm.nih.gov/29949731/
[45] Wallgren P, Nörregård E, Molander B, et al. Serological patterns of Actinobacillus pleuropneumoniae, Mycoplasma hyopneumoniae, Pasteurella multocida and Streptococcus suis in pig herds affected by pleuritis. Acta Vet Scand. 2016. https://pubmed.ncbi.nlm.nih.gov/27716292/
[46] Cheong Y, Oh C, Lee K, et al. Survey of porcine respiratory disease complex-associated pathogens among commercial pig farms in Korea via oral fluid method. J Vet Sci. 2017. https://pubmed.ncbi.nlm.nih.gov/27586468/
[47] Loera-Muro A, Jacques M, Avelar-González FJ, et al. Auxotrophic Actinobacillus pleurpneumoniae grows in multispecies biofilms without the need for nicotinamide-adenine dinucleotide (NAD) supplementation. BMC Microbiol. 2016. https://pubmed.ncbi.nlm.nih.gov/27349384/
[48] Curran DM, Adamiak PJ, Fegan JE, et al. Sequence and structural diversity of transferrin receptors in Gram-negative porcine pathogens. Vaccine. 2015. https://pubmed.ncbi.nlm.nih.gov/26263196/