neutrophil granulocyte

8/9/2019 Neutrophil Granulocyte

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Neutrophil granulocyte migratesfrom the blood vess el to the matrix,secreting proteolytic enzymes , in orderto dissolve intercellular connections (tothe improvement of its m obility) andenvelop bacteria through phagocytosis .

5 Role in disease6 Neutrophil antigens7 Video8 Additional images9 References

Characteristics [edit]Neutrophil granulocytes have an average diameter of 12-15 micrometers (m) inperipheral blood smears . Whenanalyzing a pure neutrophil suspension on an automatedcell counter, neutrophils have an average diameter of 89 m.

With the eosinophil and the basophil, they form the class of polymorphonuclear cells , named for the nucleus 'multilobulated shape (as compared to lymphocytes andmonocytes , the other types of white cells). The nucleushas a characteristic lobed appearance, the separate lobesconnected by chromatin. The nucleolus disappears as the

neutrophil matures, which is something that happens inonly a few other types of nucleated cells. [8]:168 In thecytoplasm, the Golgi apparatus is small, mitochondria andribosomes are sparse, and the rough endoplasmicreticulum is absent. [8]:170 The cytoplasm also containsabout 200 granules, of which a third are azurophilic.[8]:170

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Hypersegmented neutrophil.

Reference ranges for blood tests of white blood cells, comparingneutrophil amount (shown in pink) with that of other cells.

A minor difference is found between the neutrophils from amale subject and a female subject. The cell nucleus of aneutrophil from a female subject shows a small additional Xchromosome structure, known as a "neutrophildrumstick".[8]:174

Neutrophils will show hypersegmentation (many segmentsof nucleus) as they mature.

Neutrophils are the most abundant white blood cells inhumans (approximately 10 11 are produced daily); theyaccount for approximately 50-70% of all white blood cells(leukocytes). The stated normal range for human bloodcounts varies between laboratories, but a neutrophil count

of 2.57.5 x 109/L is astandard normal range.People of African and MiddleEastern descent may havelower counts, which are stillnormal. A report may divideneutrophils into segmentedneutrophils 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 asthey hunt for antigens. [9]

Neutrophils have a preference to engulf refined carbohydrate [10][11][12] (glucose, fructose, sucrose,
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HSC=Hematopoietic stem cell , Progenitor=Progenitor cell, L-blast=lymphoblast, Lymphocyte, Mo-blast=Monoblast, Monocyte,Myeloblast, Pro-M=Promyelocyte, Myelocyte, Meta-M=Metamyelocyte,Neutrophil, Eosinophil, Basophil, Pro-E=Proerythroblast, Baso-

honey and orange juice [10]) over bacteria. [10] In 1973 Sanchez et al. found that the neutrophilphagocytic capacity to engulf bacteria is affected when simple sugars are digested, [10] and thatfasting strengthens the neutrophils' phagocytic capacity to engulf bacteria. [10] However, thedigestion 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 researchersat the Whitehead Institute of Biomedical Research found that even amongst just a selection ofsugars, neutrophils can again be picky and engulf preferred sugars. [11][12]

Life span [edit]The average lifespan of (non-activated human) neutrophilsin the circulation is about 5.4

days. [13] Upon activation, theymarginate (positionthemselves adjacent to theblood vessel endothelium),and undergo selectin-dependent capture followedby integrin-dependentadhesion in most cases, after

which they migrate intotissues, where they survive for12 days. [14]

Neutrophils are much more -
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E=Basophilic erythroblast, poly-e=Polychromatic erythroblast, Ortho-E=orthochromatic erythroblas t, Erythrocyte, Promegakaryocyte ,megakaryocyte , Platelet

lived monocyte/macrophagephagocytes. A pathogen(disease-causingmicroorganism or virus) is likely to first encounter a neutrophil. Some experts hypothesize that theshort lifetime of neutrophils is an evolutionary adaptation. The short lifetime of neutrophils

minimizes propagation of those pathogens that parasitize phagocytes because the more time suchparasites spend outside a host cell, the more likely they will be destroyed by some component ofthe body's defenses. Also, because neutrophil antimicrobial products can also damage hosttissues , their short life limits damage to the host during inflammation.[14]

Neutrophils will often bephagocytosed by macrophages after digestion of pathogens. PECAM-1and 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 ofinfection or inflammation. Cell surface receptors allow neutrophils to detect chemical gradients ofmolecules such as interleukin-8 (IL-8),interferon gamma (IFN-gamma), C3a, C5a, and LeukotrieneB4, which these cells use to direct the path of their migration.

Neutrophils have a variety of specific receptors, including complement receptors, cytokinereceptors for interleukins and interferon gamma (IFN-gamma), receptors for chemokines,receptors to detect and adhere to endothelium, receptors for lectins and proteins, and Fcreceptors for opsonin.[15]

Anti-microbial function [edit]
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Being highlymotile, neutrophils quickly congregate at a focus of infection, attracted by cytokinesexpressed by activated endothelium, mast cells , and macrophages . Neutrophils express [16] andrelease 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 rolein the front-line defence against invading pathogens. Neutrophils have three methods for directlyattacking micro-organisms: phagocytosis (ingestion), release of soluble anti-microbials (includinggranule proteins), and generation of neutrophil extracellular traps (NETs).[17]

Phagocytosis [edit]

Neutrophils are phagocytes, capable of ingesting microorganisms or particles. For targets to berecognised, they must be coated in opsonins a process known as antibody opsonization.[9] Theycan 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. Theconsumption 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 largequantities of superoxide , a reactive oxygen species. Superoxide decays spontaneously or isbroken down via enzymes known as superoxide dismutases (Cu/ZnSOD and MnSOD), to hydrogenperoxide, which is then converted to hypochlorous acid HClO, by the green heme enzymemyeloperoxidase. It is thought that the bactericidal properties of HClO are enough to kill bacteriaphagocytosed by the neutrophil, but this may instead be a step necessary for the activation ofproteases. [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 combatinfection.

Granule type Protein

azurophilic granules(or "primarygranules")

myeloperoxidase, bactericidal/permeability-increasing protein (BPI),

defensins , and the serine proteases neutrophil elastase and cathepsin G

specific granules (or"secondary granules")

alkaline phosphatase , lysozyme, NADPH oxidase, collagenase , lactoferrinand cathelicidin

tertiary granules cathepsin and gelatinase

Neutrophil extracellular traps (NETs) [edit]

In 2004, Brinkmann and colleagues described a striking observation that activation of neutrophilscauses the release of web-like structures of DNA; this represents a third mechanism for killingbacteria.[19] These neutrophil extracellular traps (NETs) comprise a web of fibers composed ofchromatin and serine proteases that trap and kill microbes extracellularly. It is suggested that NETsprovide a high local concentration of antimicrobial components and bind, disarm, and kill microbesindependent 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 bea particularly important role for NETs insepsis , where NETs are formed within blood vessels. [20]

Recently, NETs have been shown to play a role in inflammatory diseases, as NETs could bedetected in preeclampsia , a pregnancy-related inflammatory disorder in which neutrophils areknown to be activated. [21] In addition, NETs are known to exhibit pro-thrombotic effects both invitro [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 candevelop later, as in the case of aplastic anemia or some kinds of leukemia. It can also be a side-effect of medication, most prominently chemotherapy . Neutropenia makes an individual highlysusceptible 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 adequatelyinhibited byalpha 1-antitrypsin, leading to excessive tissue damage in the presence ofinflammation the most prominent one being pulmonary emphysema .

In Familial Mediterranean fever (FMF), a mutation in the pyrin (or marenostrin ) gene, which isexpressed 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-2aantigen 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 antigensystems each have two known antigens (a and b) and are located in the 2 integrin. HNA-4 islocated on the M chain ( CD11b) and HNA-5 is located on the L integrin unit (CD11a).

Video [edit]
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A rapidly moving neutrophil can be seen taking upseveral conidia over an imaging time of 2 hours with oneframe every 30 seconds.

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A neutrophil can be seen here selectively taking upseveral Candida yeasts (fluorescently labeled in green)despite several contacts with Aspergillus fumigatusconidia (unlabeled, white/clear) in a 3-D collagen matrix.Imaging time was 2 hours with one frame every 30seconds.

[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 [edit]

A scanning electronmicroscope image of asingle neutrophil (yellow),engulfing anthraxbacteria (orange)

Blood cell lineage

More complete lineages(very large)

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/North-Holland Amsterdam. ISBN 0-444-80020-4.

3. ^ Nathan, C (Mar 2006). "Neutrophils and immunity: challenges and opportunities" . NatureReviews 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 healthycyclists 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 Promotingthe 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 b c 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 b Edwards, Steven W. (1994). Biochemistry and physiology of the neutrophil . CambridgeUniversity Press. p. 6. ISBN 0-521-41698-1.

10. ^ a b c d e 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 neutrophilicphagocytosis" . The American Society for Clinical Nutrition. Retrieved 2013-09-08. "These datasuggest 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 thatthe effects last for at least 5 hr. On the other hand, a fast of 36 or 60 hr significantly increased (P

neutrophil granulocyte

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