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Neutrophil granulocyte
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"Neutrophil" redirects here. For organisms that grow in neutral pH environments, see

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neutrophile.

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Neutrophil granulocytes (also known as

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neutrophils) are the most abundant (40% to 75%)

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type of white blood cells in mammals and form an

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essential part of the innate immune system. They are

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formed from stem cells in the bone marrow. They are

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short-lived and highly motile. Neutrophils may be

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subdivided into segmented neutrophils (or segs)

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and banded neutrophils (or bands). They form

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part of the polymorphonuclear cell family (PMNs)

together with basophils and eosinophils.[1][2][3]

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The name neutrophil derives from staining

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characteristics on hematoxylin and eosin (H&E)

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histological or cytological preparations. Whereas

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basophilic white blood cells stain dark blue and

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eosinophilic white blood cells stain bright red,

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Neutrophil granulocyte

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Neutrophils with a segmented nuclei

surrounded by erythrocytes, the intra-cellular
granules are visible in the cytoplasm (Giemsa
stained)
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TH H2.00.04.1.02012

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eosinophilic white blood cells stain bright red,

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neutrophils stain a neutral pink. Normally, neutrophils

Neutrophils are a type of phagocyte and are normally

found in the bloodstream. During the beginning (acute)

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phase of inflammation, particularly as a result of bacterial

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infection, environmental exposure,[4] and some

cancers,[5][6] neutrophils are one of the first-responders of

inflammatory cells to migrate towards the site of

inflammation. They migrate through the blood vessels,

Bosanski

then through interstitial tissue, following chemical signals

Catal
etina
Deutsch

such as Interleukin-8 (IL-8), C5a, fMLP and Leukotriene B4

in a process called chemotaxis. They are the predominant
cells in pus, accounting for its whitish/yellowish

Eesti

Espaol
Euskara

Neutrophils are recruited to the site of injury within minutes following trauma, and are the hallmark
of acute inflammation.[7]
Contents [hide]

Franais
Galego

1 Characteristics

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2 Life span

3 Chemotaxis

4 Anti-microbial function

Bahasa Indonesia

4.1 Phagocytosis

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4.2 Degranulation

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3D Rendering of a Neutrophil

appearance.

Anatomical terminology

contain a nucleus divided into 25 lobes.

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TH H2.00.04.1.02012

4.3 Neutrophil extracellular traps (NETs)

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Basa Jawa

5 Role in disease

6 Neutrophil antigens

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7 Video

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8 Additional images

9 References

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Polski

Characteristics

[edit]

Neutrophil granulocytes have an average diameter of 12-

Portugus
Romn

15 micrometers (m) in peripheral blood smears. When

analyzing a pure neutrophil suspension on an automated

Shqip

cell counter, neutrophils have an average diameter of 8

Simple English

9 m.

Slovenina
/ srpski

With the eosinophil and the basophil, they form the class of

Srpskohrvatski /

polymorphonuclear cells, named for the nucleus'

Svenska

multilobulated shape (as compared to lymphocytes and

monocytes, the other types of white cells). The nucleus

Trke

has a characteristic lobed appearance, the separate lobes

connected by chromatin. The nucleolus disappears as the

Ting Vit

neutrophil matures, which is something that happens in

only a few other types of nucleated cells.[8]:168 In the

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cytoplasm, the Golgi apparatus is small, mitochondria and

ribosomes are sparse, and the rough endoplasmic
reticulum is absent.[8]:170 The cytoplasm also contains
about 200 granules, of which a third are azurophilic.[8]:170

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Neutrophil granulocyte migrates

from the blood vessel to the matrix,
secreting proteolytic enzymes, in order
to dissolve intercellular connections (to
the improvement of its mobility) and
envelop bacteria through phagocytosis.

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A minor difference is found between the neutrophils from a

male subject and a female subject. The cell nucleus of a
neutrophil from a female subject shows a small additional X
chromosome structure, known as a "neutrophil
drumstick".[8]:174
Neutrophils will show hypersegmentation (many segments
of nucleus) as they mature.
Neutrophils are the most abundant white blood cells in
humans (approximately 1011 are produced daily); they
account for approximately 50-70% of all white blood cells
(leukocytes). The stated normal range for human blood

Hypersegmented neutrophil.

counts varies between laboratories, but a neutrophil count

of 2.57.5 x 109/L is a
standard normal range.
People of African and Middle
Eastern descent may have
lower counts, which are still
Reference ranges for blood tests of white blood cells, comparing
neutrophil amount (shown in pink) with that of other cells.

normal. A report may divide

neutrophils into segmented
neutrophils and bands.

When circulating in the bloodstream and unactivated, neutrophils are spherical. Once activated,
they change shape and become more amorphous or amoeba-like and can extend pseudopods as
they hunt for antigens.[9]
Neutrophils have a preference to engulf refined carbohydrate[10][11][12] (glucose, fructose, sucrose,
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honey and orange juice[10]) over bacteria.[10] In 1973 Sanchez et al. found that the neutrophil
phagocytic capacity to engulf bacteria is affected when simple sugars are digested,[10] and that
fasting strengthens the neutrophils' phagocytic capacity to engulf bacteria.[10] However, the
digestion of normal starches has no effect. It was concluded that the function, and not the number,
of phagocytes in engulfing bacteria was altered by the ingestion of sugars.[10] In 2007 researchers
at the Whitehead Institute of Biomedical Research found that even amongst just a selection of
sugars, neutrophils can again be picky and engulf preferred sugars.[11][12]

Life span

[edit]

The average lifespan of (nonactivated human) neutrophils

in the circulation is about 5.4
days.[13] Upon activation, they
marginate (position
themselves adjacent to the
blood vessel endothelium),
and undergo selectindependent capture followed
by integrin-dependent
adhesion in most cases, after
which they migrate into
tissues, where they survive for
12 days.[14]
Neutrophils are much more
numerous than the longeropen in browser PRO version

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HSC=Hematopoietic stem cell, Progenitor=Progenitor cell, Lblast=lymphoblast, Lymphocyte, Mo-blast=Monoblast, Monocyte,

Myeloblast, Pro-M=Promyelocyte, Myelocyte, Meta-M=Metamyelocyte,
Neutrophil, Eosinophil, Basophil, Pro-E=Proerythroblast, Baso-

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numerous than the longerlived monocyte/macrophage

phagocytes. A pathogen

E=Basophilic erythroblast, poly-e=Polychromatic erythroblast, OrthoE=orthochromatic erythroblast, Erythrocyte, Promegakaryocyte,

megakaryocyte, Platelet

(disease-causing
microorganism or virus) is likely to first encounter a neutrophil. Some experts hypothesize that the
short lifetime of neutrophils is an evolutionary adaptation. The short lifetime of neutrophils
minimizes propagation of those pathogens that parasitize phagocytes because the more time such
parasites spend outside a host cell, the more likely they will be destroyed by some component of
the body's defenses. Also, because neutrophil antimicrobial products can also damage host
tissues, their short life limits damage to the host during inflammation.[14]
Neutrophils will often be phagocytosed by macrophages after digestion of pathogens. PECAM-1
and phosphatidylserine on the cell surface are involved in this process.

Chemotaxis

[edit]

Neutrophils undergo a process called chemotaxis, which allows them to migrate toward sites of
infection or inflammation. Cell surface receptors allow neutrophils to detect chemical gradients of
molecules such as interleukin-8 (IL-8), interferon gamma (IFN-gamma), C3a, C5a, and Leukotriene
B4, which these cells use to direct the path of their migration.
Neutrophils have a variety of specific receptors, including complement receptors, cytokine
receptors for interleukins and interferon gamma (IFN-gamma), receptors for chemokines,
receptors to detect and adhere to endothelium, receptors for lectins and proteins, and Fc
receptors for opsonin.[15]

Anti-microbial function
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Being highly motile, neutrophils quickly congregate at a focus of infection, attracted by cytokines
expressed by activated endothelium, mast cells, and macrophages. Neutrophils express[16] and
release cytokines, which in turn amplify inflammatory reactions by several other cell types.
In addition to recruiting and activating other cells of the immune system, neutrophils play a key role
in the front-line defence against invading pathogens. Neutrophils have three methods for directly
attacking micro-organisms: phagocytosis (ingestion), release of soluble anti-microbials (including
granule proteins), and generation of neutrophil extracellular traps (NETs).[17]

Phagocytosis [edit]
Neutrophils are phagocytes, capable of ingesting microorganisms or particles. For targets to be
recognised, they must be coated in opsoninsa process known as antibody opsonization.[9] They
can internalize and kill many microbes, each phagocytic event resulting in the formation of a
phagosome into which reactive oxygen species and hydrolytic enzymes are secreted. The
consumption of oxygen during the generation of reactive oxygen species has been termed the
"respiratory burst", although unrelated to respiration or energy production.
The respiratory burst involves the activation of the enzyme NADPH oxidase, which produces large
quantities of superoxide, a reactive oxygen species. Superoxide decays spontaneously or is
broken down via enzymes known as superoxide dismutases (Cu/ZnSOD and MnSOD), to hydrogen
peroxide, which is then converted to hypochlorous acid HClO, by the green heme enzyme
myeloperoxidase. It is thought that the bactericidal properties of HClO are enough to kill bacteria
phagocytosed by the neutrophil, but this may instead be a step necessary for the activation of
proteases.[18]

Degranulation [edit]
Neutrophils also release an assortment of proteins in three types of granules by a process called
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degranulation. The contents of these granules have antimicrobial properties, and help combat
infection.
Granule type
azurophilic granules
(or "primary
granules")
specific granules (or

Protein
myeloperoxidase, bactericidal/permeability-increasing protein (BPI),
defensins, and the serine proteases neutrophil elastase and cathepsin G
alkaline phosphatase, lysozyme, NADPH oxidase, collagenase, lactoferrin

"secondary granules") and cathelicidin

tertiary granules

cathepsin and gelatinase

Neutrophil extracellular traps (NETs) [edit]

In 2004, Brinkmann and colleagues described a striking observation that activation of neutrophils
causes the release of web-like structures of DNA; this represents a third mechanism for killing
bacteria.[19] These neutrophil extracellular traps (NETs) comprise a web of fibers composed of
chromatin and serine proteases that trap and kill microbes extracellularly. It is suggested that NETs
provide a high local concentration of antimicrobial components and bind, disarm, and kill microbes
independent of phagocytic uptake. In addition to their possible antimicrobial properties, NETs may
serve as a physical barrier that prevents further spread of pathogens. Trapping of bacteria may be
a particularly important role for NETs in sepsis, where NETs are formed within blood vessels.[20]
Recently, NETs have been shown to play a role in inflammatory diseases, as NETs could be
detected in preeclampsia, a pregnancy-related inflammatory disorder in which neutrophils are
known to be activated.[21] In addition, NETs are known to exhibit pro-thrombotic effects both in
vitro[22] and in vivo.[23][24]
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Role in disease

[edit]

Low neutrophil counts are termed neutropenia. This can be congenital (genetic disorder) or it can
develop later, as in the case of aplastic anemia or some kinds of leukemia. It can also be a sideeffect of medication, most prominently chemotherapy. Neutropenia makes an individual highly
susceptible to infections. Neutropenia can be the result of colonization by intracellular neutrophilic
parasites.
In alpha 1-antitrypsin deficiency, the important neutrophil enzyme elastase is not adequately
inhibited by alpha 1-antitrypsin, leading to excessive tissue damage in the presence of
inflammation the most prominent one being pulmonary emphysema.
In Familial Mediterranean fever (FMF), a mutation in the pyrin (or marenostrin) gene, which is
expressed mainly in neutrophil granulocytes, leads to a constitutively active acute-phase response
and causes attacks of fever, arthralgia, peritonitis, and eventually amyloidosis.[25]

Neutrophil antigens

[edit]

There are five (HNA 1-5) sets of neutrophil antigen recognised.[26] The three HNA-1 antigens (a-c)
are located on the low affinity Fc- receptor IIIb (FCGR3B :CD16b) The single known HNA-2a
antigen is located on CD177. The HNA-3 antigen system has two antigens (3a and 3b) which are
located on the seventh exon of the CLT2 gene (SLC44A2). The HNA-4 and HNA-5 antigen
systems each have two known antigens (a and b) and are located in the 2 integrin. HNA-4 is
located on the M chain (CD11b) and HNA-5 is located on the L integrin unit (CD11a).

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Sorry, your browser either has

JavaScript disabled or does not have
any supported player.
You can download the clip or download
a player to play the clip in your browser.

A rapidly moving neutrophil can be seen taking up

several conidia over an imaging time of 2 hours with one
frame every 30 seconds.

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A neutrophil can be seen here selectively taking up

several Candida yeasts (fluorescently labeled in green)
despite several contacts with Aspergillus fumigatus
conidia (unlabeled, white/clear) in a 3-D collagen matrix.
Imaging time was 2 hours with one frame every 30
seconds.

[1]

Neutrophils display highly directional amoeboid motility in infected footpad and phalanges.

Intravital imaging was performed in the footpad path of LysM-eGFP mice 20 minutes after infection
with Listeria monocytogenes.[27]
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Additional images

A scanning electron

[edit]

Blood cell lineage

microscope image of a

More complete lineages

(very large)

single neutrophil (yellow),

engulfing anthrax
bacteria (orange)

References

[edit]

1. ^ Witko-Sarsat, V; Rieu P, Descamps-Latscha B, Lesavre P, Halbwachs-Mecarelli L (2000).

"Neutrophils: molecules, functions and pathophysiological aspects"

. Lab Invest 80 (5): 61753.

doi:10.1038/labinvest.3780067 . PMID 10830774 .

2. ^ Klebanoff, SJ; Clark, RA (1978). "The Neutrophil: Function and Clinical Disorders". Elsevier/NorthHolland Amsterdam. ISBN 0-444-80020-4.
3. ^ Nathan, C (Mar 2006). "Neutrophils and immunity: challenges and opportunities"

. Nature

Reviews Immunology 6 (March): 17382. doi:10.1038/nri1785 . ISSN 1474-1733 .

PMID 16498448 .

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4. ^ Jacobs, L; Nawrot, Tim S; De Geus, Bas; Meeusen, Romain; Degraeuwe, Bart; Bernard, Alfred;
Sughis, Muhammad; Nemery, Benoit; Panis, Luc (Oct 2010). "Subclinical responses in healthy
cyclists briefly exposed to traffic-related air pollution"

. Environmental Health 9 (64): 64.

doi:10.1186/1476-069X-9-64 . PMC 2984475 . PMID 20973949 .

5. ^ Waugh, DJ; Wilson, C. (Nov 2008). "The interleukin-8 pathway in cancer"

. Clinical Cancer

Research 14 (21): 673541. doi:10.1158/1078-0432.CCR-07-4843 . ISSN 1078-0432 .

PMID 18980965 .
6. ^ De Larco, JE; Wuertz, BR; Furcht, LT (Aug 2004). "The Potential Role of Neutrophils in Promoting
the Metastatic Phenotype of Tumors Releasing Interleukin-8"

. Clinical Cancer Research 10 (15):

4895900. doi:10.1158/1078-0432.CCR-03-0760 . ISSN 1078-0432 . PMID 15297389 .

7. ^ Cohen, Stephen; Burns, Richard C. (2002). Pathways of the Pulp (8th ed.). St. Louis: Mosby.
p. 465.
8. ^ a

bc d

Zucker-Franklin, Dorothy; Greaves, M. F.; Grossi, C. E.; Marmont, A. M. (1988).

"Neutrophils". Atlas of Blood Cells: Function and Pathology 1 (2nd ed.). Philadelphia: Lea &
Ferbiger. ISBN 0-8121-1094-3.
9. ^ a

Edwards, Steven W. (1994). Biochemistry and physiology of the neutrophil. Cambridge

University Press. p. 6. ISBN 0-521-41698-1.

10. ^ a

bc de f

Albert Sanchez, J. L. Reeser, H. S. Lau, P. Y. Yahiku, R. E. Willard, P. J. McMillan, S.

Y. Cho, A. R. Magie, and U. D. Register (1973). "Role of sugars in human neutrophilic

phagocytosis"

. The American Society for Clinical Nutrition. Retrieved 2013-09-08. "These data

suggest that the function and not the number of phagocytes was altered by ingestion of sugars. This
implicates glucose and other simple carbohydrates in the control of phagocytosis and shows that
the effects last for at least 5 hr. On the other hand, a fast of 36 or 60 hr significantly increased (P <
0.001) the phagocytic index."
11. ^ a

Rubin-Bejerano, I.; Abeijon, C.; Magnelli, P.; Grisafi, P.; Fink, G. R. (July 2007). "Phagocytosis

by human neutrophils is stimulated by a unique fungal cell wall component"

. Cell Host Microbe 2

(1): 5567. doi:10.1016/j.chom.2007.06.002 . PMC 2083279 . PMID 18005717 .

^a
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12. ^ a

Kneller, Alyssa (2007). "White blood cells are picky about sugar"

. Whitehead Institute.

Retrieved 2013-08-09.
13. ^ Pillay J, den Braber I, Vrisekoop N, Kwast LM, de Boer RJ, Borghans JA, Tesselaar K,
Koenderman L. (29 Jul 2010). "In vivo labeling with 2 H2 O reveals a human neutrophil lifespan of 5.4
days"
14. ^ a

text

. Blood 116 (4): 625627. PMID 20410504 .

Wheater, Paul R.; Stevens, Alan (2002). Wheater's basic histopathology: a colour atlas and
(PDF). Edinburgh: Churchill Livingstone. ISBN 0-443-07001-6.

15. ^ Serhan, Charles N.; Ward, Peter A.; Gilroy, Derek W. (2010). Fundamentals of Inflammation .
Cambridge University Press. pp. 5354. ISBN 0-521-88729-1.
16. ^ Ear, T; McDonald, PP (2008). "Cytokine generation, promoter activation, and oxidant-independent
NF-kappaB activation in a transfectable human neutrophilic cellular model"

. BMC Immunol. 9: 14.

doi:10.1186/1471-2172-9-14 . PMC 2322942 . PMID 18405381 .

17. ^ Hickey, MJ; Kubes, P (2009). "Intravascular immunity: the hostpathogen encounter in blood
vessels". Nature Reviews Immunology (Nature Publishing Group) 9 (5): 36475.
doi:10.1038/nri2532 . PMID 19390567 .
18. ^ Segal, AW (2005). "How neutrophils kill microbes"

. Annu Rev Immunol 9 (5): 197223.

doi:10.1146/annurev.immunol.23.021704.115653 . PMC 2092448 . PMID 15771570 .

19. ^ Brinkmann, Volker; Ulrike Reichard, Christian Goosmann, Beatrix Fauler, Yvonne Uhlemann, David
S. Weiss, Yvette Weinrauch, Arturo Zychlinsky (5 March 2004). "Neutrophil Extracellular Traps Kill
Bacteria"

. Science (AAAS) 303 (5663): 15321535. doi:10.1126/science.1092385 . ISSN 0036-

8075 . PMID 15001782 . Retrieved 2007-04-09.

20. ^ Clark SR, Ma AC, Tavener AS, McDonald B, Goodarzi Z, Kelly MM, Patel KD, Chakrabarti S,
McAvoy E, Sinclair GD, Keys EM, Allen-Vercoe E, DeVinney R, Doig CJ, Green FHY and Kubes P
(Apr 2007). "Platelet Toll-Like Receptor-4 Activates Neutrophil Extracellular Traps to Ensnare
Bacteria in Endotoxemic and Septic Blood"

. Nature Medicine (Nature Publishing Group) 13 (4):

4639. doi:10.1038/nm1565 . ISSN 1078-8956 . PMID 17384648 .

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21. ^ Gupta, AK; Hasler, P; Holzgreve, W; Hahn, S (Jun 2007). "Neutrophil NETs: a novel contributor to
preeclampsia-associated placental hypoxia?". Semin Immunopathol 29 (2): 1637.
doi:10.1007/s00281-007-0073-4 . ISSN 1863-2297 . PMID 17621701 .
22. ^ Fuchs, TA; Brill, A, Duerschmied, D, Schatzberg, D, Monestier, M, Myers DD, Jr, Wrobleski, SK,
Wakefield, TW, Hartwig, JH, Wagner, DD (Sep 7, 2010). "Extracellular DNA traps promote
thrombosis"

. Proceedings of the National Academy of Sciences of the United States of America

107 (36): 158805. doi:10.1073/pnas.1005743107 . PMC 2936604 . PMID 20798043 .

23. ^ Brill, A; Fuchs, TA, Savchenko, A, Thomas, GM, Martinod, K, De Meyer, SF, Bhandari, AA,
Wagner, DD (Nov 1, 2011). "Neutrophil Extracellular Traps Promote Deep Vein Thrombosis in
Mice"

. Journal of thrombosis and haemostasis : JTH 10 (1): 13644. doi:10.1111/j.1538-

7836.2011.04544.x

. PMC 3319651 . PMID 22044575 .

24. ^ Borissoff, JI; ten Cate, H (September 2011). "From neutrophil extracellular traps release to
thrombosis: an overshooting host-defense mechanism?". Journal of thrombosis and haemostasis :
JTH 9 (9): 17914. doi:10.1111/j.1538-7836.2011.04425.x

. PMID 21718435 .

25. ^ Ozen, S (Jul 2004). "Familial mediterranean fever: revisiting an ancient disease"
Journal of Pediatrics 162 (78): 44954. doi:10.1007/s00431-003-1223-x

. European

. ISSN 0340-6199 .

PMID 12751000 .
26. ^ Chu HT, Lin H, Tsao TT, Chang CF, Hsiao WW, Yeh TJ, Chang CM, Liu YW, Wang TY, Yang KC,
Chen TJ, Chen JC, Chen KC, Kao CY (2013). "Genotyping of human neutrophil antigens (HNA) from
whole genome sequencing data"

. BMC Med Genomics 6 (1): 31. PMID 24028078 .

27. ^ Graham D.B., Zinselmeyer B.H., Mascarenhas F., Delgado R., Miller M.J., Swat W.; Zinselmeyer;
Mascarenhas; Delgado; Miller; Swat (2009). Unutmaz, Derya, ed. "ITAM signaling by Vav family Rho
guanine nucleotide exchange factors regulates interstitial transit rates of neutrophils in vivo"

. PLoS

ONE 4 (2): e4652. doi:10.1371/journal.pone.0004652 . PMC 2645696 . PMID 19247495 .

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V T E

Myeloid lineage - Blood (WBC and RBC)

[show]

V T E

Immunology: Lymphocytic adaptive immune system and

complement

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Categories: Cell biology

Granulocytes

Phagocytes

Human cells

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