Monocyte

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Monocyte
Blausen 0649 Monocyte (crop).png
3D rendering of a monocyte
Monocytes, a type of white blood cell (Giemsa stained).jpg
Monocytes under a light microscope from a peripheral blood smear surrounded by red blood cells
Details
System Immune system
Identifiers
MeSH D009000
TH H2.00.04.1.02010
FMA 62864
Anatomical terms of microanatomy

Monocytes are a type of leukocyte or white blood cell. They are the largest type of leukocyte in blood and can differentiate into macrophages and monocyte-derived dendritic cells. As a part of the vertebrate innate immune system monocytes also influence adaptive immune responses and exert tissue repair functions. There are at least three subclasses of monocytes in human blood based on their phenotypic receptors.

Contents

Structure

Monocytes are amoeboid in appearance, and have nongranulated cytoplasm. [1] Thus they are classified as agranulocytes, although they might occasionally display some azurophil granules and/or vacuoles. With a diameter of 15–22 μm, monocytes are the largest cell type in peripheral blood. [2] [3] Monocytes are mononuclear cells and the ellipsoidal nucleus is often lobulated/indented, causing a bean-shaped or kidney-shaped appearance. [4] Monocytes compose 2% to 10% of all leukocytes in the human body.

Development

Comparison of monoblast, promonocyte and monocyte. Monoblast, promonocyte and monocyte.jpg
Comparison of monoblast, promonocyte and monocyte.

Monocytes are produced by the bone marrow from precursors called monoblasts, bipotent cells that differentiated from hematopoietic stem cells. [5] Monocytes circulate in the bloodstream for about one to three days and then typically migrate into tissues throughout the body where they differentiate into macrophages and dendritic cells.

Subpopulations

In humans

The first clear description of monocyte subsets by flow cytometry dates back to the late 1980s, when a population of CD16-positive monocytes was described. [6] [7] Today, three types of monocytes are recognized in human blood: [8]

  1. The classical monocyte is characterized by high level expression of the CD14 cell surface receptor (CD14++ CD16− monocyte)
  2. The non-classical monocyte shows low level expression of CD14 and additional co-expression of the CD16 receptor (CD14+CD16++ monocyte). [9]
  3. The intermediate monocyte expresses high levels of CD14 and low levels of CD16 (CD14++CD16+ monocytes).

While in humans the level of CD14 expression can be used to differentiate non-classical and intermediate monocytes, the slan (6-Sulfo LacNAc) cell surface marker was shown to give an unequivocal separation of the two cell types. [10] [11]

Ghattas et al. state that the "intermediate" monocyte population is likely to be a unique subpopulation of monocytes, as opposed to a developmental step, due to their comparatively high expression of surface receptors involved in reparative processes (including vascular endothelial growth factor receptors type 1 and 2, CXCR4, and Tie-2) as well as evidence that the "intermediate" subset is specifically enriched in the bone marrow. [12]

In mice

In mice, monocytes can be divided in two subpopulations. Inflammatory monocytes (CX3CR1 low, CCR2 pos, Ly6C high, PD-L1 neg), which are equivalent to human classical CD14++ CD16− monocytes and resident monocytes (CX3CR1 high, CCR2 neg, Ly6C low, PD-L1 pos), which are equivalent to human non-classical CD14+ CD16+ monocytes. Resident monocytes have the ability to patrol along the endothelium wall in the steady state and under inflammatory conditions. [13] [14] [15] [16]

Function

Monocytes are mechanically active cells [17] and migrate from blood to an inflammatory site to perform their functions. As explained before, they can differentiate into macrophages and dendritic cells, but the different monocyte subpopulations can also exert specific functions on their own. In general, monocytes and their macrophage and dendritic cell progeny serve three main functions in the immune system. These are phagocytosis, antigen presentation, and cytokine production. Phagocytosis is the process of uptake of microbes and particles followed by digestion and destruction of this material. Monocytes can perform phagocytosis using intermediary (opsonising) proteins such as antibodies or complement that coat the pathogen, as well as by binding to the microbe directly via pattern recognition receptors that recognize pathogens. Monocytes are also capable of killing infected host cells via antibody-dependent cell-mediated cytotoxicity. Vacuolization may be present in a cell that has recently phagocytized foreign matter.

Differentiation into other effector cells

Monocytes can migrate into tissues and replenish resident macrophage populations. Macrophages have a high antimicrobial and phagocytic activity and thereby protect tissues from foreign substances. They are cells that possess a large smooth nucleus, a large area of cytoplasm, and many internal vesicles for processing foreign material. Although they can be derived from monocytes, a large proportion is already formed prenatally in the yolk sac and foetal liver. [18]

In vitro, monocytes can differentiate into dendritic cells by adding the cytokines granulocyte macrophage colony-stimulating factor (GM-CSF) and interleukin 4. [19] Such monocyte-derived cells do, however, retain the signature of monocytes in their transcriptome and they cluster with monocytes and not with bona fide dendritic cells. [20]

Specific functions of monocyte subpopulations

Aside from their differentiation capacity, monocytes can also directly regulate immune responses. As explained before, they are able to perform phagocytosis. Cells of the classical subpopulation are the most efficient phagocytes and can additionally secrete inflammation-stimulating factors. The intermediate subpopulation is important for antigen presentation and T lymphocyte stimulation. [21] Briefly, antigen presentation describes a process during which microbial fragments that are present in the monocytes after phagocytosis are incorporated into MHC molecules. They are then trafficked to the cell surface of the monocytes (or macrophages or dendritic cells) and presented as antigens to activate T lymphocytes, which then mount a specific immune response against the antigen. Non-classical monocytes produce high amounts of pro-inflammatory cytokines like tumor necrosis factor and interleukin-12 after stimulation with microbial products. Furthermore, a monocyte patrolling behavior has been demonstrated in humans both for the classical and the non-classical monocytes, meaning that they slowly move along the endothelium to examine it for pathogens. [22] Said et al. showed that activated monocytes express high levels of PD-1 which might explain the higher expression of PD-1 in CD14+CD16++ monocytes as compared to CD14++CD16− monocytes. Triggering monocytes-expressed PD-1 by its ligand PD-L1 induces IL-10 production, which activates CD4 Th2 cells and inhibits CD4 Th1 cell function. [23] Many factors produced by other cells can regulate the chemotaxis and other functions of monocytes. These factors include most particularly chemokines such as monocyte chemotactic protein-1 (CCL2) and monocyte chemotactic protein-3 (CCL7); certain arachidonic acid metabolites such as leukotriene B4 and members of the 5-hydroxyicosatetraenoic acid and 5-oxo-eicosatetraenoic acid family of OXE1 receptor agonists (e.g., 5-HETE and 5-oxo-ETE); and N-Formylmethionine leucyl-phenylalanine and other N-formylated oligopeptides which are made by bacteria and activate the formyl peptide receptor 1. [24] Other microbial products can directly activate monocytes and this leads to production of pro-inflammatory and, with some delay, of anti-inflammatory cytokines. Typical cytokines produced by monocytes are TNF, IL-1, and IL-12.

Clinical significance

A scanning electron microscope (SEM) image of normal circulating human blood. One can see red blood cells, several knobby white blood cells including lymphocytes, a monocyte, a neutrophil, and many small disc-shaped platelets. SEM blood cells.jpg
A scanning electron microscope (SEM) image of normal circulating human blood. One can see red blood cells, several knobby white blood cells including lymphocytes, a monocyte, a neutrophil, and many small disc-shaped platelets.

A monocyte count is part of a complete blood count and is expressed either as a percentage of monocytes among all white blood cells or as absolute numbers. Both may be useful, but these cells became valid diagnostic tools only when monocyte subsets are determined. Monocytic cells may contribute to the severity and disease progression in COVID-19 patients. [25]

Monocytosis

Monocytosis is the state of excess monocytes in the peripheral blood. It may be indicative of various disease states. Examples of processes that can increase a monocyte count include:

A high count of CD14+CD16++ monocytes is found in severe infection (sepsis). [30]

In the field of atherosclerosis, high numbers of the CD14++CD16+ intermediate monocytes were shown to be predictive of cardiovascular events in populations at risk. [31] [32]

CMML is characterized by a persistent monocyte count of > 1000/microL of blood. Analysis of monocyte subsets has demonstrated predominance of classical monocytes and absence of CD14lowCD16+ monocytes. [33] [34] The absence of non-classical monocytes can assist in diagnosis of the disease and the use of slan as a marker can improve specificity. [35]

Monocytopenia

Monocytopenia is a form of leukopenia associated with a deficiency of monocytes. A very low count of these cells is found after therapy with immuno-suppressive glucocorticoids. [36]

Also, non-classical slan+ monocytes are strongly reduced in patients with hereditary diffuse leukoencephalopathy with spheroids, a neurologic disease associated with mutations in the macrophage colony-stimulating factor receptor gene. [10]

Blood content

Reference ranges for blood tests of white blood cells, comparing monocyte amount (shown in green) with other cells. Reference ranges for blood tests - white blood cells.png
Reference ranges for blood tests of white blood cells, comparing monocyte amount (shown in green) with other cells.

See also

Further reading

Related Research Articles

<span class="mw-page-title-main">Dendritic cell</span> Accessory cell of the mammalian immune system

A dendritic cell (DC) is an antigen-presenting cell of the mammalian immune system. A DC's main function is to process antigen material and present it on the cell surface to the T cells of the immune system. They act as messengers between the innate and adaptive immune systems.

<span class="mw-page-title-main">Macrophage</span> Type of white blood cell

Macrophages are a type of white blood cell of the innate immune system that engulf and digest pathogens, such as cancer cells, microbes, cellular debris, and foreign substances, which do not have proteins that are specific to healthy body cells on their surface. This process is called phagocytosis, which acts to defend the host against infection and injury.

<span class="mw-page-title-main">CD32</span> Surface receptor glycoprotein

CD32, also known as FcγRII or FCGR2, is a surface receptor glycoprotein belonging to the Ig gene superfamily. CD32 can be found on the surface of a variety of immune cells. CD32 has a low-affinity for the Fc region of IgG antibodies in monomeric form, but high affinity for IgG immune complexes. CD32 has two major functions: cellular response regulation, and the uptake of immune complexes. Cellular responses regulated by CD32 include phagocytosis, cytokine stimulation, and endocytic transport. Dysregulated CD32 is associated with different forms of autoimmunity, including systemic lupus erythematosus. In humans, there are three major CD32 subtypes: CD32A, CD32B, and CD32C. While CD32A and CD32C are involved in activating cellular responses, CD32B is inhibitory.

<span class="mw-page-title-main">Phagocyte</span> Cells that ingest harmful matter within the body

Phagocytes are cells that protect the body by ingesting harmful foreign particles, bacteria, and dead or dying cells. Their name comes from the Greek phagein, "to eat" or "devour", and "-cyte", the suffix in biology denoting "cell", from the Greek kutos, "hollow vessel". They are essential for fighting infections and for subsequent immunity. Phagocytes are important throughout the animal kingdom and are highly developed within vertebrates. One litre of human blood contains about six billion phagocytes. They were discovered in 1882 by Ilya Ilyich Mechnikov while he was studying starfish larvae. Mechnikov was awarded the 1908 Nobel Prize in Physiology or Medicine for his discovery. Phagocytes occur in many species; some amoebae behave like macrophage phagocytes, which suggests that phagocytes appeared early in the evolution of life.

<span class="mw-page-title-main">Kupffer cell</span> Macrophages located in the liver

Kupffer cells, also known as stellate macrophages and Kupffer–Browicz cells, are specialized cells localized in the liver within the lumen of the liver sinusoids and are adhesive to their endothelial cells which make up the blood vessel walls. Kupffer cells comprise the largest population of tissue-resident macrophages in the body. Gut bacteria, bacterial endotoxins, and microbial debris transported to the liver from the gastrointestinal tract via the portal vein will first come in contact with Kupffer cells, the first immune cells in the liver. It is because of this that any change to Kupffer cell functions can be connected to various liver diseases such as alcoholic liver disease, viral hepatitis, intrahepatic cholestasis, steatohepatitis, activation or rejection of the liver during liver transplantation and liver fibrosis. They form part of the mononuclear phagocyte system.

A histiocyte is a vertebrate cell that is part of the mononuclear phagocyte system. The mononuclear phagocytic system is part of the organism's immune system. The histiocyte is a tissue macrophage or a dendritic cell. Part of their job is to clear out neutrophils once they've reached the end of their lifespan.

<span class="mw-page-title-main">Fc receptor</span> Surface protein important to the immune system

In immunology, an Fc receptor is a protein found on the surface of certain cells – including, among others, B lymphocytes, follicular dendritic cells, natural killer cells, macrophages, neutrophils, eosinophils, basophils, human platelets, and mast cells – that contribute to the protective functions of the immune system. Its name is derived from its binding specificity for a part of an antibody known as the Fc region. Fc receptors bind to antibodies that are attached to infected cells or invading pathogens. Their activity stimulates phagocytic or cytotoxic cells to destroy microbes, or infected cells by antibody-mediated phagocytosis or antibody-dependent cell-mediated cytotoxicity. Some viruses such as flaviviruses use Fc receptors to help them infect cells, by a mechanism known as antibody-dependent enhancement of infection.

<span class="mw-page-title-main">CD14</span> Mammalian protein found in Homo sapiens

CD14 is a human protein made mostly by macrophages as part of the innate immune system. It helps to detect bacteria in the body by binding lipopolysaccharide (LPS), a pathogen-associated molecular pattern (PAMP).

<span class="mw-page-title-main">Integrin alpha M</span> Mammalian protein found in Homo sapiens

Integrin alpha M (ITGAM) is one protein subunit that forms heterodimeric integrin alpha-M beta-2 (αMβ2) molecule, also known as macrophage-1 antigen (Mac-1) or complement receptor 3 (CR3). ITGAM is also known as CR3A, and cluster of differentiation molecule 11B (CD11B). The second chain of αMβ2 is the common integrin β2 subunit known as CD18, and integrin αMβ2 thus belongs to the β2 subfamily integrins.

<span class="mw-page-title-main">Monoblast</span>

Monoblasts are the committed progenitor cells that differentiated from a committed macrophage or dendritic cell precursor (MDP) in the process of hematopoiesis. They are the first developmental stage in the monocyte series leading to a macrophage. Their myeloid cell fate is induced by the concentration of cytokines they are surrounded by during development. These cytokines induce the activation of transcription factors which push completion of the monoblast's myeloid cell fate. Monoblasts are normally found in bone marrow and do not appear in the normal peripheral blood. They mature into monocytes which, in turn, develop into macrophages. They then are seen as macrophages in the normal peripheral blood and many different tissues of the body. Macrophages can produce a variety of effector molecules that initiate local, systemic inflammatory responses. These monoblast differentiated cells are equipped to fight off foreign invaders using pattern recognition receptors to detect antigen as part of the innate immune response.

<span class="mw-page-title-main">Alveolar macrophage</span>

An alveolar macrophage, pulmonary macrophage, is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls.

<span class="mw-page-title-main">CCL18</span> Mammalian protein found in Homo sapiens

Chemokine ligand 18 (CCL18) is a small cytokine belonging to the CC chemokine family. The functions of CCL18 have been well studied in laboratory settings, however the physiological effects of the molecule in living organisms have been difficult to characterize because there is no similar protein in rodents that can be studied. The receptor for CCL18 has been identified in humans only recently, which will help scientists understand the molecule's role in the body.

<span class="mw-page-title-main">CD68</span> Mammalian protein found in Homo sapiens

CD68 is a protein highly expressed by cells in the monocyte lineage, by circulating macrophages, and by tissue macrophages.

CD16, also known as FcγRIII, is a cluster of differentiation molecule found on the surface of natural killer cells, neutrophils, monocytes, macrophages, and certain T cells. CD16 has been identified as Fc receptors FcγRIIIa (CD16a) and FcγRIIIb (CD16b), which participate in signal transduction. The most well-researched membrane receptor implicated in triggering lysis by NK cells, CD16 is a molecule of the immunoglobulin superfamily (IgSF) involved in antibody-dependent cellular cytotoxicity (ADCC). It can be used to isolate populations of specific immune cells through fluorescent-activated cell sorting (FACS) or magnetic-activated cell sorting, using antibodies directed towards CD16.

Macrophage-1 antigen is a complement receptor ("CR3") consisting of CD11b and CD18.

Gamma delta T cells are T cells that have a γδ T-cell receptor (TCR) on their surface. Most T cells are αβ T cells with TCR composed of two glycoprotein chains called α (alpha) and β (beta) TCR chains. In contrast, γδ T cells have a TCR that is made up of one γ (gamma) chain and one δ (delta) chain. This group of T cells is usually less common than αβ T cells. Their highest abundance is in the gut mucosa, within a population of lymphocytes known as intraepithelial lymphocytes (IELs).

<span class="mw-page-title-main">MARCO</span> Protein-coding gene in the species Homo sapiens

Macrophage receptor with collagenous structure (MARCO) is a protein that in humans is encoded by the MARCO gene. MARCO is a class A scavenger receptor that is found on particular subsets of macrophages. Scavenger receptors are pattern recognition receptors (PRRs) found most commonly on immune cells. Their defining feature is that they bind to polyanions and modified forms of a type of cholesterol called low-density lipoprotein (LDL). MARCO is able to bind and phagocytose these ligands and pathogen-associated molecular patterns (PAMPs), leading to the clearance of pathogens and cell signaling events that lead to inflammation. As part of the innate immune system, MARCO clears, or scavenges, pathogens, which leads to inflammatory responses. The scavenger receptor cysteine-rich (SRCR) domain at the end of the extracellular side of MARCO binds ligands to activate the subsequent immune responses. MARCO expression on macrophages has been associated with tumor development and also with Alzheimer's disease, via decreased responses of cells when ligands bind to MARCO.

B10 cells are a sub-class of regulatory B-cells that are involved in inhibiting immune responses in both humans and mice. B10 cells are named for their ability to produce inhibitory interleukin: Interleukin-10 (IL-10). One of their unique abilities is that they suppress the innate and adaptive immune signals, making them important for regulating the inflammatory response. Like the B cell, the B10 cell requires antigen specific binding to the surface of CD5 receptor to illicit a response from the T-cell. Once an antigen binds to the CD19 receptor, immediate downregulation in B-cell receptor (BCR) signal expression occurs and mediates the release of IL-10 cytokines. In mice and humans, B10 cells are distinguishable in their expression of measurable IL-10 due to the lack of unique cell surface markers expressed by regulatory B cells. However, IL-10 competence is not limited to any one subset of B cells. B10 cells do not possess unique phenotypic markers or transcription factors for further identification. B10 cells predominantly localize in the spleen, though they are also found in the blood, lymph nodes, Peyer's patches, intestinal tissues, central nervous system, and peritoneal cavity. B10 cells proliferate during inflammatory and disease responses.

<span class="mw-page-title-main">Gwendalyn J. Randolph</span> American immunologist

Gwendalyn J. Randolph is an American immunologist, the Emil R. Unanue Distinguished Professor in the Department of Immunology and Pathology at Washington University School of Medicine where she is currently co-director of the Immunology Graduate Program. During her postdoctoral work, Randolph characterized monocyte differentiation to dendritic cells and macrophages and made advances in our understanding of dendritic cell trafficking and the fate of monocytes recruited to sites of inflammation. Her lab has contributed to the Immunological Genome Project by characterizing macrophage gene expression. Her work now focuses on the immunological mechanisms driving atherosclerosis and inflammatory bowel disease (IBD) by exploring lymphatic function and lipoprotein trafficking.

<span class="mw-page-title-main">Dermal macrophage</span> Skin macrophages used for wound repair and hair growth

Dermal macrophages are macrophages in the skin that facilitate skin homeostasis by mediating wound repair, hair growth, and salt balance. Their functional role in these processes is the mediator of inflammation. They can acquire an M1 or M2 phenotype to promote or suppress an inflammatory response, thereby influencing other cells' activity via the production of pro-inflammatory or anti-inflammatory cytokines. Dermal macrophages' ability to acquire pro-inflammatory properties also potentiates them in cancer defence. M1 macrophages can suppress tumour growth in the skin by their pro-inflammatory properties. However, M2 macrophages support tumour growth and invasion by the production of Th2 cytokines such as TGFβ and IL-10. Thus, the exact contribution of each phenotype to cancer defence and the skin's homeostasis is still unclear.

References

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  20. ↑ Robbins, Scott H; Walzer, Thierry; Dembélé, Doulaye; Thibault, Christelle; Defays, Axel; Bessou, Gilles; Xu, Huichun; Vivier, Eric; Sellars, MacLean; Pierre, Philippe; Sharp, Franck R; Chan, Susan; Kastner, Philippe; Dalod, Marc (2008). "Novel insights into the relationships between dendritic cell subsets in human and mouse revealed by genome-wide expression profiling". Genome Biology . 9 (1): R17. doi: 10.1186/gb-2008-9-1-r17 . PMC   2395256 . PMID   18218067.
  21. ↑ Wong, K. L. et al. Gene expression profiling reveals the defining features of the classical, intermediate, and nonclassical human monocyte subsets. Blood 118, e16-31, doi:10.1182/blood-2010-12-326355 (2011)
  22. ↑ Collison, Joanna L.; Carlin, Leo M.; Eichmann, Martin; Geissmann, Frederic; Peakman, Mark (1 August 2015). "Heterogeneity in the Locomotory Behavior of Human Monocyte Subsets over Human Vascular Endothelium In Vitro". The Journal of Immunology . 195 (3): 1162–1170. doi: 10.4049/jimmunol.1401806 . PMID   26085686.
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  26. ↑ Hoyer, FF; Zhang, X; Coppin, E; Vasamsetti, SB; Modugu, G; Schloss, MJ; Rohde, D; McAlpine, CS; Iwamoto, Y; Libby, P; Naxerova, K; Swirski, F; Dutta, P; Nahrendorf, P (April 2020). "Bone Marrow Endothelial Cells Regulate Myelopoiesis in Diabetes". Circulation. 142 (3): 244–258. doi:10.1161/CIRCULATIONAHA.120.046038. PMC   7375017 . PMID   32316750.
  27. ↑ Heidt, Timo; Sager, Hendrik B; Courties, Gabriel; Dutta, Partha; Iwamoto, Yoshiko; Zaltsman, Alex; von zur Muhlen, Constantin; Bode, Christoph; Fricchione, Gregory L; Denninger, John; Lin, Charles P; Vinegoni, Claudio; Libby, Peter; Swirski, Filip K; Weissleder, Ralph; Nahrendorf, Matthias (July 2014). "Chronic variable stress activates hematopoietic stem cells". Nature Medicine. 20 (7): 754–758. doi:10.1038/nm.3589. PMC   4087061 . PMID   24952646.
  28. ↑ Swirski, Filip K.; Libby, Peter; Aikawa, Elena; Alcaide, Pilar; Luscinskas, F. William; Weissleder, Ralph; Pittet, Mikael J. (2 January 2007). "Ly-6Chi monocytes dominate hypercholesterolemia-associated monocytosis and give rise to macrophages in atheromata". Journal of Clinical Investigation . 117 (1): 195–205. doi:10.1172/JCI29950. PMC   1716211 . PMID   17200719.
  29. ↑ "Old and "hangry" monocytes turn from friend to foe under assault". doi:10.1016/j.immuni.2023.03.013.{{cite journal}}: Cite journal requires |journal= (help)
  30. ↑ Fingerle, G; Pforte, A; Passlick, B; Blumenstein, M; Strobel, M; Ziegler- Heitbrock, Hw (15 November 1993). "The novel subset of CD14+/CD16+ blood monocytes is expanded in sepsis patients". Blood. 82 (10): 3170–3176. doi: 10.1182/blood.v82.10.3170.3170 . PMID   7693040.
  31. ↑ Heine, G.H.; Ulrich, C.; Seibert, E.; Seiler, S.; Marell, J.; Reichart, B.; Krause, M.; Schlitt, A.; Köhler, H.; Girndt, M. (March 2008). "CD14++CD16+ monocytes but not total monocyte numbers predict cardiovascular events in dialysis patients". Kidney International . 73 (5): 622–629. doi: 10.1038/sj.ki.5002744 . PMID   18160960.
  32. ↑ Rogacev, Kyrill S.; Cremers, Bodo; Zawada, Adam M.; Seiler, Sarah; Binder, Nadine; Ege, Philipp; Große-Dunker, Gunnar; Heisel, Isabel; Hornof, Florian; Jeken, Jana; Rebling, Niko M.; Ulrich, Christof; Scheller, Bruno; Böhm, Michael; Fliser, Danilo; Heine, Gunnar H. (October 2012). "CD14++CD16+ Monocytes Independently Predict Cardiovascular Events". Journal of the American College of Cardiology . 60 (16): 1512–1520. doi: 10.1016/j.jacc.2012.07.019 . PMID   22999728.
  33. ↑ Vuckovic, S.; Fearnley, D. B.; Gunningham, S.; Spearing, R. L.; Patton, W. N.; Hart, D. N. J. (June 1999). "Dendritic cells in chronic myelomonocytic leukaemia: Dendritic Cells in CMML". British Journal of Haematology . 105 (4): 974–985. doi: 10.1046/j.1365-2141.1999.01431.x . PMID   10554809. S2CID   22571555.
  34. ↑ Selimoglu-Buet, Dorothée; Wagner-Ballon, Orianne; Saada, Véronique; Bardet, Valérie; Itzykson, Raphaël; Bencheikh, Laura; Morabito, Margot; Met, Elisabeth; Debord, Camille; Benayoun, Emmanuel; Nloga, Anne-Marie; Fenaux, Pierre; Braun, Thorsten; Willekens, Christophe; Quesnel, Bruno; Adès, Lionel; Fontenay, Michaela; Rameau, Philippe; Droin, Nathalie; Koscielny, Serge; Solary, Eric (4 June 2015). "Characteristic repartition of monocyte subsets as a diagnostic signature of chronic myelomonocytic leukemia". Blood. 125 (23): 3618–3626. doi:10.1182/blood-2015-01-620781. PMC   4497970 . PMID   25852055.
  35. ↑ Tarfi, Sihem; Badaoui, Bouchra; Freynet, Nicolas; Morabito, Margot; Lafosse, Jeffie; Toma, Andréa; Etienne, Gabriel; Micol, Jean-Baptiste; Sloma, Ivan; Fenaux, Pierre; Solary, Eric; Selimoglu-Buet, Dorothée; Wagner-Ballon, Orianne (April 2020). "Disappearance of slan-positive non-classical monocytes for diagnosis of chronic myelomonocytic leukemia with an associated inflammatory state". Haematologica . 105 (4): e147–e152. doi: 10.3324/haematol.2019.219782 . PMID   31413091. S2CID   199663779.
  36. ↑ Fingerle-Rowson, G; Angstwurm, M; Andreesen, R; Ziegler-Heitbrock, HW (June 1998). "Selective depletion of CD14 + CD16 + monocytes by glucocorticoid therapy: Depletion of CD14+ CD16+ monocytes by glucocorticoids". Clinical & Experimental Immunology . 112 (3): 501–506. doi:10.1046/j.1365-2249.1998.00617.x. PMC   1904988 . PMID   9649222.
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