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Topic:Macrophages

Macrophages are a type of white blood cell found in horses that play a significant role in the immune system. They are involved in the detection, phagocytosis, and destruction of bacteria and other harmful organisms. Additionally, macrophages contribute to tissue homeostasis and repair by removing dead cells and stimulating tissue regeneration. In horses, macrophages are distributed throughout various tissues, including the liver, lungs, and lymph nodes, where they help orchestrate the body's response to infection and injury. This page compiles peer-reviewed research studies and scholarly articles that explore the function, regulation, and clinical implications of macrophages in equine health.
Immunophenotypic characterization and depletion of pulmonary intravascular macrophages of horses.
Veterinary research    April 22, 2004   Volume 35, Issue 1 39-51 doi: 10.1051/vetres:2003041
Parbhakar OP, Duke T, Townsend HG, Singh B.Pulmonary intravascular macrophages (PIMs) are present in horses and are believed to increase their sensitivity to endotoxin-induced cardio-pulmonary shock. However, owing to a lack of a marker for PIMs and the inability to isolate them, their precise contributions in the horse remain unknown. We designed this study to identify an immuno-phenotypic marker for PIMs and to develop a protocol for their transient depletion with gadolinium chloride (GC). GC is a lanthanide that has been used to deplete liver and lung macrophages. The horses (N = 15) were divided into control (n = 5) and GC-treated ...
PU.1 binding to ets motifs within the equine infectious anemia virus long terminal repeat (LTR) enhancer: regulation of LTR activity and virus replication in macrophages.
Journal of virology    March 16, 2004   Volume 78, Issue 7 3407-3418 doi: 10.1128/jvi.78.7.3407-3418.2004
Hines R, Sorensen BR, Shea MA, Maury W.Binding of the transcription factor PU.1 to its DNA binding motif regulates the expression of a number of B-cell- and myeloid-specific genes. The long terminal repeat (LTR) of macrophage-tropic strains of equine infectious anemia virus (EIAV) contains three PU.1 binding sites, namely an invariant promoter-proximal site as well as two upstream sites. We have previously shown that these sites are important for EIAV LTR activity in primary macrophages (W. Maury, J. Virol. 68:6270-6279, 1994). Since the sequences present in these three binding motifs are not identical, we sought to determine the r...
Moxifloxacin pharmacokinetics in horses and disposition into phagocytes after oral dosing.
Journal of veterinary pharmacology and therapeutics    March 5, 2004   Volume 27, Issue 1 57-60 doi: 10.1046/j.0140-7783.2003.00529.x
Gardner SY, Davis JL, Jones SL, LaFevers DH, Hoskins MS, McArver EM, Papich MG.No abstract available
Influence of long terminal repeat and env on the virulence phenotype of equine infectious anemia virus.
Journal of virology    February 14, 2004   Volume 78, Issue 5 2478-2485 doi: 10.1128/jvi.78.5.2478-2485.2004
Payne SL, Pei XF, Jia B, Fagerness A, Fuller FJ.The molecular clones pSPeiav19 and p19/wenv17 of equine infectious anemia virus (EIAV) differ in env and long terminal repeats (LTRs) and produce viruses (EIAV(19) and EIAV(17), respectively) of dramatically different virulence phenotypes. These constructs were used to generate a series of chimeric clones to test the individual contributions of LTR, surface (SU), and transmembrane (TM)/Rev regions to the disease potential of the highly virulent EIAV(17). The LTRs of EIAV(19) and EIAV(17) differ by 16 nucleotides in the transcriptional enhancer region. The two viruses differ by 30 amino acids i...
Generation and characterisation of an equine macrophage cell line (e-CAS cells) derived from equine bone marrow cells.
Veterinary immunology and immunopathology    January 1, 2004   Volume 97, Issue 1-2 65-76 doi: 10.1016/j.vetimm.2003.08.012
Werners AH, Bull S, Fink-Gremmels J, Bryant CE.Macrophages play a pivotal role in the pathophysiology of many diseases by mediating the host immune response to infections and intoxications. The species-specific activation of macrophages and the differential response in cytokine production impedes the extrapolation of results between species. Therefore, the aim of this study was to isolate and immortalise macrophages from equine bone marrow (BM) cells in order to study equine-specific signalling pathways. The isolated BM-derived macrophages (referred to as e-CAS cells) showed proliferation kinetics similar to that of standardised cell lines...
Virulent and avirulent strains of equine arteritis virus induce different quantities of TNF-alpha and other proinflammatory cytokines in alveolar and blood-derived equine macrophages.
Virology    October 14, 2003   Volume 314, Issue 2 662-670 doi: 10.1016/s0042-6822(03)00506-3
Moore BD, Balasuriya UB, Watson JL, Bosio CM, MacKay RJ, MacLachlan NJ.Equine arteritis virus (EAV) infects endothelial cells (ECs) and macrophages in horses, and many of the clinical manifestations of equine viral arteritis (EVA) reflect vascular injury. To further evaluate the potential role of EAV-induced, macrophage-derived cytokines in the pathogenesis of EVA, we infected cultured equine alveolar macrophages (AMphi), blood monocyte-derived macrophages (BMphi), and pulmonary artery ECs with either a virulent (KY84) or an avirulent (CA95) strain of EAV. EAV infection of equine AMphi, BMphi, and ECs resulted in their activation with increased transcription of g...
Differences in polymorphonucleocyte function and local inflammatory response between horses and ponies.
Equine veterinary journal    October 1, 2003   Volume 35, Issue 6 561-569 doi: 10.2746/042516403775467234
Wilmink JM, Veenman JN, van den Boom R, Rutten VP, Niewold TA, Broekhuisen-Davies JM, Lees R, Armstrong S, van Weeren PR, Barneveld A.Wound healing proceeds faster in ponies than in horses and complications during healing, such as wound infection, occur less frequently in ponies. Earlier studies suggested that this difference might be related to differences in the initial post traumatic inflammatory response. Objective: That polymorphonuclear leucocyte (PMN) function and profiles of humoral factors in local inflammatory processes are different in horses and ponies. Methods: PMNs were isolated from venous blood of horses and ponies. Chemotaxis and reactive oxygen species (ROS) production was determined. Tissue cages were impl...
Effects of midazolam on equine innate immune response: a flow cytometric study.
Veterinary immunology and immunopathology    September 13, 2003   Volume 95, Issue 1-2 11-19 doi: 10.1016/s0165-2427(03)00097-7
Massoco C, Palermo-Neto J.Benzodiazepines (BDZ) are among the most frequently used class of psychotropic drugs employed in veterinary medicine in Brazil and worldwide due to their anxiolytic, muscle relaxant and anticonvulsant effects [J. Clin. Pharmacol. 33 (1993) 124]. Peripheral benzodiazepine receptor (PBR) sites were described in peripheral organs, endocrine steroidogenic tissues and immune organs and cells. Midazolam is a mixed-type agonist of PBRs. The present study is focused on the effects of midazolam on equine peripheral blood neutrophils, peritoneal macrophages and cortisol levels in plasma. Adult horses we...
An improved Pythium insidiosum-vaccine formulation with enhanced immunotherapeutic properties in horses and dogs with pythiosis.
Vaccine    June 12, 2003   Volume 21, Issue 21-22 2797-2804 doi: 10.1016/s0264-410x(03)00225-1
Mendoza L, Mandy W, Glass R.The immunotherapeutic properties of a new Pythium insidiosum-vaccine formulation (PIV), was evaluated in 18 horses and 6 dogs with proven pythiosis from different enzootic areas in the United States. All injected horses but one responded with a weak (=29 mm, n=3), a mild (30-90 mm, n=7) or a strong (=100 mm, n=7) inflammatory reactions at the site of injection. Three equines with weak or negative reactions at the injection site were not cured. Seven equines with strong reactions at their injection sites, however, were cured. Six of the eight horses with mild reactions were also cured. The rema...
Alveolar fibrosis and changes in equine lung morphometry in response to intrapulmonary blood.
Equine veterinary journal. Supplement    October 31, 2002   Issue 34 451-458 doi: 10.1111/j.2042-3306.2002.tb05465.x
McKane SA, Slocombe RF.Necropsy studies of horses suffering exercise-induced pulmonary haemorrhage (EIPH) have identified mild inflammatory lesions with evidence of alveolar fibrosis and bronchiolitis. These lesions were thought to be the result of viral infections that predisposed the affected regions of the lung to EIPH. We have shown previously that during erythrophagocytosis in the alveolar space, there is a prolonged period of macrophage influx and activation. This present study used morphometric analysis to quantify the effects of macrophage activity during erythrophagocytosis, on the alveolar cell population ...
Cytological analysis of equine bronchoalveolar lavage fluid. Part 2: Comparison of smear and cytocentrifuged preparations.
Equine veterinary journal    July 11, 2002   Volume 34, Issue 3 292-296 doi: 10.2746/042516402776186155
Pickles K, Pirie RS, Rhind S, Dixon PM, McGorum BC.The aim of this study was to develop a diagnostically useful smear method for preparation of equine bronchoalveolar lavage fluid (BALF) for use by practitioners. A smear method for equine BALF preparation which included the addition of serum was developed, and cell morphology, differential cell counts (DCC) and repeatability of counting DCC compared with those of cytocentrifuged BALF preparations. BALF samples (n = 21) were collected from 5 control horses and 5 heaves-susceptible horses. Smear preparations of BALF produced smaller, darker, staining cells, making cytological identification more...
Cytological analysis of equine bronchoalveolar lavage fluid. Part 1: Comparison of sequential and pooled aliquots.
Equine veterinary journal    July 11, 2002   Volume 34, Issue 3 288-291 doi: 10.2746/042516402776186137
Pickles K, Pirie RS, Rhind S, Dixon PM, McGorum BC.The aim of this study was to investigate whether initial equine bronchoalveolar lavage fluid (BALF) aliquots were more representative of bronchial cytology that bronchiolar and alveolar cytology. Cell viability and total nucleated (TCC), differential (DCC) and absolute cell counts of cytocentrifuged preparations of 3 sequentially collected BALF aliquots (Aliquots 1-3) were compared with those of pooled BALF (Aliquot 4) to assess whether all aliquots were representative of the lavaged lung segment. BALF samples (n = 21) were collected from control horses (n = 5) or heaves-affected horses (n = 5...
A novel lipoarabinomannan from the equine pathogen Rhodococcus equi. Structure and effect on macrophage cytokine production.
The Journal of biological chemistry    June 18, 2002   Volume 277, Issue 35 31722-31733 doi: 10.1074/jbc.M203008200
Garton NJ, Gilleron M, Brando T, Dan HH, Giguère S, Puzo G, Prescott JF, Sutcliffe IC.Rhodococcus equi is a major cause of foal morbidity and mortality. We have investigated the presence of lipoglycan in this organism as closely related bacteria, notably Mycobacterium tuberculosis, produce lipoarabinomannans (LAM) that may play multiple roles as virulence determinants. The lipoglycan was structurally characterized by gas chromatography-mass spectrometry following permethylation, capillary electrophoresis after chemical degradation, and (1)H and (31)P and two-dimensional heteronuclear nuclear magnetic resonance studies. Key structural features of the lipoglycan are a linear alph...
Alveolar macrophage graded hemosiderin score from bronchoalveolar lavage in horses with exercise-induced pulmonary hemorrhage and controls.
Journal of veterinary internal medicine    June 4, 2002   Volume 16, Issue 3 281-286 doi: 10.1892/0891-6640(2002)016<0281:amghsf>2.3.co;2
Doucet MY, Viel L.The objective of this study was to determine if a quantitative scoring system for evaluation of hemosiderin content of alveolar macrophages obtained by bronchoalevolar lavage provides a more sensitive test for the detection of exercise-induced pulmonary hemorrhage (EIPH) in horses than does endoscopy of the lower airways. A sample population composed of 74 Standardbred racehorses aged 2-5 years was used. Horses were grouped as either control (EIPH-negative) or EIPH-positive based on history and repeated postexertional endoscopic evaluation of the bronchial airways. Bronchoalveolar lavage was p...
Temporal localization of immunoreactive transforming growth factor beta1 in normal equine skin and in full-thickness dermal wounds.
Veterinary surgery : VS    May 8, 2002   Volume 31, Issue 3 274-280 doi: 10.1053/jvet.2002.32397
Theoret CL, Barber SM, Gordon JR.To describe the localization of immunoreactive transforming growth factor (TGF)-beta1 in both normal skin and full-thickness dermal wounds of the limb and the thorax of the horse. Methods: Six full-thickness excisional wounds were created on the lateral aspect of one metacarpal region and on the midthoracic area of each horse. Sequentially collected tissue specimens from wound margins were assessed for TGF-beta1 expression by immunohistochemistry. Methods: Four horses (2 to 4 years of age). Methods: A neutralizing monoclonal anti-human TGF-beta1 antibody was used to detect the spatial expressi...
Histoplasmosis in the lung of a race horse with yersiniosis.
The Journal of veterinary medical science    January 5, 2002   Volume 63, Issue 11 1229-1231 doi: 10.1292/jvms.63.1229
Katayama Y, Kuwano A, Yoshihara T.A 4-year-old female thoroughbred race horse died of acute peritonitis caused by necrotizing granulomatous duodenitis. Yersinia enterocolitica was immunohistochemically demonstrated in macrophages in granulomas developed in the duodenum, lung, liver and abdominal lymph nodes. The yeast-like fungi were found in the cytoplasmic vacuoles of macrophages in the lung that infiltrated into the granulomas and surrounding alveoli with congestive edema. The yeast-like fungi were positively stained by Gomori-Grocott chromic acid methenamine silver stain and immuno-histochemically stained with anti-histopl...
Idiopathic systemic granulomatous disease and macrophage expression of PTHrP in a miniature pony.
Journal of comparative pathology    October 2, 2001   Volume 125, Issue 2-3 214-218 doi: 10.1053/jcpa.2001.0484
Sellers RS, Toribio RE, Blomme EA.Idiopathic systemic granulomatous disease, which has been reported in horses, cattle and human beings, is characterized by perivascular granulomatous and lymphoplasmacytic inflammation in many organ systems. Diagnosis is based on the exclusion of possible viral, fungal or bacterial causes. The disease was identified in a miniature pony with widespread lymphoplasmacytic and granulomatous inflammation, special staining techniques having revealed no evidence of any aetiological agent. Skin lesions, which were severe, consisted of hyperkeratosis and serocellular crust formation, with inflammatory ...
Evaluation of collagenase activity, matrix metalloproteinase-8, and matrix metalloproteinase-13 in horses with chronic obstructive pulmonary disease.
American journal of veterinary research    July 17, 2001   Volume 62, Issue 7 1142-1148 doi: 10.2460/ajvr.2001.62.1142
Raulo SM, Sorsa TA, Kiili MT, Maisi PS.To determine collagenase activity and evaluate matrix metalloproteinase (MMP)-8 and MMP-13 in horses with chronic obstructive pulmonary disease (COPD). Methods: 12 horses with COPD and 12 healthy control horses. Methods: Collagenase activity was determined by use of an assay for degradation of type-I collagen. Western immunoblot analysis was used to identify interstitial collagenases MMP-8 and MMP-13 in tracheal epithelial lining fluid (TELF). Immunocytochemistry and in situ hybridization were used to determine cellular expression of these 2 collagenases in cells in bronchoalveolar lavage flui...
DH82 cells: a macrophage cell line for the replication and study of equine infectious anemia virus.
Journal of virological methods    May 30, 2001   Volume 95, Issue 1-2 47-56 doi: 10.1016/s0166-0934(01)00288-9
Hines R, Maury W.In vivo, tissue macrophages have been implicated as an important cell for the replication of equine infectious anemia virus (EIAV). Laboratory investigations of EIAV/macrophage interactions, however, have been hampered by the laborious blood monocyte isolation procedures. In addition, adherent equine macrophage cultures generally have poor long-term viability and are resistant to transfection. This report describes an adherent canine macrophage-like cell line, DH82, that supports the replication of EIAV. This cell line was easily transfectable and supported EIAV Tat transactivation of the LTR....
Effects of training on resting peripheral blood and BAL-derived leucocyte function in horses.
Equine veterinary journal    May 16, 2001   Volume 33, Issue 3 238-243 doi: 10.2746/042516401776249651
Raidal SL, Rose RJ, Love DN.In this study, the effects of prolonged, high intensity training on aspects of peripheral blood and bronchoalveolar lavage (BAL)-derived leucocyte function were evaluated in 8 horses. All horses undertook a 7 week endurance training programme, followed by 5 weeks of high intensity training (HIT). Thereafter, horses were divided into control (C) and overtraining (OT) groups. The frequency and intensity of training were increased more substantially for horses in the OT group. Training was terminated in week 32 when horses in the OT group demonstrated a significant performance reduction. Peripher...
Fatal nonneurological EHV-1 infection in a yearling filly.
Veterinary pathology    December 6, 2000   Volume 37, Issue 6 672-676 doi: 10.1354/vp.37-6-672
Del Piero F, Wilkins PA, Timoney PJ, Kadushin J, Vogelbacker H, Lee JW, Berkowitz SJ, La Perle KM.A case of fatal nonneurological equine herpesvirus 1 (EHV-1) infection in a yearling filly is described. Gross lesions included extensive pulmonary edema, prominent laryngeal lymphoid follicles, and congestion and edema of the dorsal third ventricle choroid plexus. Histologically, there was vasculitis, hemorrhage, and edema in the lungs and dorsal third ventricle choroid plexus as well as mild intestinal crypt necrosis with occasional intranuclear inclusion bodies. The perivascular and vascular inflammatory infiltrates were comprised mainly of T lymphocytes and macrophages. EHV-1 antigen was i...
Immunohistochemical detection of virulence-associated Rhodococcus equi antigens in pulmonary and intestinal lesions in horses.
Journal of comparative pathology    October 18, 2000   Volume 123, Issue 2-3 186-189 doi: 10.1053/jcpa.2000.0392
Mariotti F, Cuteri V, Takai S, Renzoni G, Pascucci L, Vitellozzi G.Two horses with Rhodococcus equi infection were examined post mortem by an immunohistochemical method (peroxidase-antiperoxidase; PAP) with a monoclonal antibody (Mab 10G5) to the 15-17 kDa antigen of R. equi. One of the horses was also examined bacteriologically, R. equi being isolated in culture. Immunolabelling with this Mab was marked and widespread. On the other hand, the immunohistochemical reactivity of infected macrophages with a polyclonal antibody specific for lysozyme was slight. Thus, Mab 10G5 would appear to be a useful diagnostic reagent in R. equi infection, with or without cult...
Mutations occurring during serial passage of Japanese equine infectious anemia virus in primary horse macrophages.
Virus research    August 10, 2000   Volume 68, Issue 1 93-98 doi: 10.1016/s0168-1702(00)00147-7
Zheng YH, Sentsui H, Kono Y, Ikuta K.An attenuated equine infectious anemia virus (EIAV), named V26, was previously obtained after 50 passages of the Japanese virulent strain V70 in primary macrophage culture. To clarify the differences between both viruses, their full-length sequences were determined. There were higher mutations in S2 (6.15% amino acid difference) and LTR (10.7% nucleotide difference). The presumed initiation codon of the S2 gene was absent from the sequence of V26. There was a large insertion within the long-terminal repeat (LTR) U3 hypervariable region of V26. In addition, there were minor mutations in gag (1....
Equid herpesvirus 1: platelets and alveolar macrophages are potential sources of activated TGF-B1 in the horse.
Veterinary immunology and immunopathology    July 13, 2000   Volume 75, Issue 1-2 71-79 doi: 10.1016/s0165-2427(00)00187-2
Chesters PM, Hughes A, Edington N.Cell mediated responses to Equid herpesvirus 1 (EHV-1) are of short duration in vivo and require considerable expansion to be detected in vitro. Raised serum levels of active transforming growth factor B (TGF-B1) have been shown to depress proliferative T cell responses in experimental infections with EHV-1 in ponies. The present work indicates that latent transforming growth factor B (TGF-B1) is present in circulating platelets, lymph node, bronchial epithelium and alveolar macrophages. Activation of platelets in vitro by thrombin resulted in the release of latent TGF-B1 from platelets, with ...
The effect of high intensity exercise on the functional capacity of equine pulmonary alveolar macrophages and BAL-derived lymphocytes.
Research in veterinary science    July 6, 2000   Volume 68, Issue 3 249-253 doi: 10.1053/rvsc.1999.0373
Raidal SL, Love DN, Bailey GD, Rose RJ.The effect of strenuous exercise on the functional capacity of pulmonary alveolar macrophages (PAM) and bronchoalveolar lavage-derived lymphocytes was determined in eight horses prior to and after 7 weeks of training. Strenuous exercise had no effect on the total cell count or the percentage of live cells in bronchoalveolar lavage (BAL) samples prior to or following training. However, training was associated with a significant increase in the total cell count of pre-exercise BAL samples and a significant reduction in the percentage of live cells in post-exercise samples. Strenuous exercise was...
Immunomodulation in horses.
The Veterinary clinics of North America. Equine practice    April 7, 2000   Volume 16, Issue 1 183-viii doi: 10.1016/s0749-0739(17)30126-8
Rush BR, Flaminio MJ.Nonspecific immunomodulators are substances that induce non-antigen-specific enhancement of the body's native or acquired defense mechanisms. Immunomodulant preparations are most often used for treatment of chronic, viral, or bacterial infection with evidence of secondary immunosuppression. The proposed mechanism of action of these products is macrophage activation and subsequent release of cytokines that enhance humoral and cell-mediated immunity. In equine medicine, nonspecific immunostimulant products are used for prevention and treatment of infectious respiratory disease and treatment of s...
Characterization of peripheral blood and pulmonary leukocyte function in healthy foals.
Veterinary immunology and immunopathology    March 14, 2000   Volume 73, Issue 3-4 267-285 doi: 10.1016/s0165-2427(00)00149-5
Flaminio MJ, Rush BR, Davis EG, Hennessy K, Shuman W, Wilkerson MJ.Studies in infants and foals indicate an age-dependent maturation of peripheral lymphocyte subsets. The age-dependent relationship for maturation of cellular immune responses, such as phagocytosis and lymphocyte responses of the peripheral and pulmonary-derived leukocytes, has not been characterized in foals. Lymphocyte subpopulations, mitogen stimulation response of lymphocytes, lymphokine-activated killing cell activity, phagocytosis and oxidative burst activity, and serum immunoglobulin (Ig) classes G and M concentrations were determined in developing foals. This study illustrates age-depen...
Tissue sites of persistent infection and active replication of equine infectious anemia virus during acute disease and asymptomatic infection in experimentally infected equids.
Journal of virology    March 9, 2000   Volume 74, Issue 7 3112-3121 doi: 10.1128/jvi.74.7.3112-3121.2000
Harrold SM, Cook SJ, Cook RF, Rushlow KE, Issel CJ, Montelaro RC.Equine infectious anemia virus (EIAV) infection of horses is characterized by recurring cycles of disease and viremia that typically progress to an inapparent infection in which clinical symptoms are absent as host immune responses maintain control of virus replication indefinitely. The dynamics of EIAV viremia and its association with disease cycles have been well characterized, but there has been to date no comprehensive quantitative analyses of the specific tissue sites of EIAV infection and replication in experimentally infected equids during acute disease episodes and during asymptomatic ...
Immune cell populations in the equine corpus luteum throughout the oestrous cycle and early pregnancy: an immunohistochemical and flow cytometric study.
Journal of reproduction and fertility    February 26, 2000   Volume 117, Issue 2 281-290 doi: 10.1530/jrf.0.1170281
Lawler DF, Hopkins J, Watson ED.Recent evidence indicates that the cells of the immune system and their large network of secretory products, or cytokines, play an active role in the ovary throughout the oestrous cycle. In the present study, immune cell populations (T and B lymphocytes, macrophages, granulocytes and eosinophils) and expression of major histocompatibility complex (MHC) class II were investigated in corpora lutea from mares in early (days 2-4), mid- (days 7-10) and late (days 12-14) dioestrus, the post-luteolytic phase (days 16-17) and early pregnancy. The number of T lymphocytes within the corpus luteum increa...
Equine macrophage identification with an antibody (Ki-M6) to human CD68 and a new monoclonal antibody (JB10).
Journal of comparative pathology    February 24, 2000   Volume 122, Issue 2-3 145-154 doi: 10.1053/jcpa.1999.0351
Siedek EM, Honnah-Symns N, Fincham SC, Mayall S, Hamblin AS.Monoclonal antibodies (mAbs) recognizing equine macrophages are scarce. The present study compared the immunocytochemical staining of various equine tissues (lymphoid tissue, lung, liver, small intestine, skin and blood leucocytes) by an antibody, Ki-M6, which detects CD68 in human macrophages and dendritic cells, and by a new anti-equine mAb, JB10, with staining produced by two previously described anti-equine macrophage mAbs, CZ2.2 and CZ3.3. Ki-M6 was shown to identify equine macrophages, which had a distribution different from those identified by CZ2.2 and CZ3.3. JB10 identified equine mac...
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