Logo of biochemjBJ Latest papers and much more!
Biochem J. Mar 1, 1991; 274(Pt 2): 381–386.
PMCID: PMC1150148

Ectocytosis caused by sublytic autologous complement attack on human neutrophils. The sorting of endogenous plasma-membrane proteins and lipids into shed vesicles.


During sublytic complement attack on human neutrophils, plasma-membrane vesicles are shed from the cell surface as a cell-protection mechanism. By using surface-iodinated neutrophils it was found that less than 2% of surface label was recovered in shed vesicles under conditions where 40% of complement component C9 was shed. SDS/PAGE of 125I-labelled shed vesicles and plasma membranes showed differences in iodination pattern, demonstrating the sorting of membrane proteins into the shed vesicles. Analysis of 32P-labelled phospholipids after labeling of neutrophils with [32P]Pi before sublytic complement attack showed the presence of phosphatidic acid, phosphatidylcholine, phosphatidyl-ethanolamine, phosphatidylinositol and polyphosphoinositides in shed vesicles. Quantitative analysis using [3H]acetic anhydride-labelling method showed that the molar proportions of phosphatidylethanolamine, phosphatidylinositol, phosphatidylserine and sphingomyelin were the same in shed vesicles as in plasma membranes. In contrast, the molar proportions of cholesterol and diacylglycerol relative to sphingomyelin were almost twice those found in plasma membranes. The data demonstrate the existence of protein and lipid sorting mechanisms during the formation of shed vesicles when neutrophils are subject to sublytic complement attack. The term 'ectocytosis' is proposed to describe triggered shedding of right-side-out membrane vesicles from the surface of eukaryotic cells.

Full text

Full text is available as a scanned copy of the original print version. Get a printable copy (PDF file) of the complete article (1.8M), or click on a page image below to browse page by page. Links to PubMed are also available for Selected References.

Images in this article

Click on the image to see a larger version.

Selected References

These references are in PubMed. This may not be the complete list of references from this article.
  • Allan D, Cockcroft S. The fatty acid composition of 1,2-diacylglycerol and polyphosphoinositides from human erythrocyte membranes. Biochem J. 1983 Aug 1;213(2):555–557. [PMC free article] [PubMed]
  • Allan D, Michell RH. Accumulation of 1,2-diacylglycerol in the plasma membrane may lead to echinocyte transformation of erythrocytes. Nature. 1975 Nov 27;258(5533):348–349. [PubMed]
  • Allan D, Michell RH. The possible role of lipids in control of membrane fusion during secretion. Symp Soc Exp Biol. 1979;33:323–336. [PubMed]
  • Baron MD, Pope B, Luzio JP. The membrane topography of ecto-5'-nucleotidase in rat hepatocytes. Biochem J. 1986 Jun 1;236(2):495–502. [PMC free article] [PubMed]
  • Bennett JP, Cockcroft S, Caswell AH, Gomperts BD. Plasma-membrane location of phosphatidylinositol hydrolysis in rabbit neutrophils stimulated with formylmethionyl-leucylphenylalanine. Biochem J. 1982 Dec 15;208(3):801–808. [PMC free article] [PubMed]
  • Beverley PC, Linch D, Delia D. Isolation of human haematopoietic progenitor cells using monoclonal antibodies. Nature. 1980 Sep 25;287(5780):332–333. [PubMed]
  • Black PH. Shedding from the cell surface of normal and cancer cells. Adv Cancer Res. 1980;32:75–199. [PubMed]
  • BLIGH EG, DYER WJ. A rapid method of total lipid extraction and purification. Can J Biochem Physiol. 1959 Aug;37(8):911–917. [PubMed]
  • Boyum A. Separation of blood leucocytes, granulocytes and lymphocytes. Tissue Antigens. 1974;4(4):269–274. [PubMed]
  • Branch WJ, Mullock BM, Luzio JP. Rapid subcellular fractionation of the rat liver endocytic compartments involved in transcytosis of polymeric immunoglobulin A and endocytosis of asialofetuin. Biochem J. 1987 Jun 1;244(2):311–315. [PMC free article] [PubMed]
  • Campbell AK, Luzio JP. Intracellular free calcium as a pathogen in cell damage initiated by the immune system. Experientia. 1981 Oct 15;37(10):1110–1112. [PubMed]
  • Campbell AK, Morgan BP. Monoclonal antibodies demonstrate protection of polymorphonuclear leukocytes against complement attack. Nature. 1985 Sep 12;317(6033):164–166. [PubMed]
  • Carney DF, Hammer CH, Shin ML. Elimination of terminal complement complexes in the plasma membrane of nucleated cells: influence of extracellular Ca2+ and association with cellular Ca2+. J Immunol. 1986 Jul 1;137(1):263–270. [PubMed]
  • Carney DF, Lang TJ, Shin ML. Multiple signal messengers generated by terminal complement complexes and their role in terminal complement complex elimination. J Immunol. 1990 Jul 15;145(2):623–629. [PubMed]
  • FOLCH J, LEES M, SLOANE STANLEY GH. A simple method for the isolation and purification of total lipides from animal tissues. J Biol Chem. 1957 May;226(1):497–509. [PubMed]
  • Goldstein JL, Anderson RG, Brown MS. Coated pits, coated vesicles, and receptor-mediated endocytosis. Nature. 1979 Jun 21;279(5715):679–685. [PubMed]
  • Hauser G, Eichberg J, Gonzalez-Sastre F. Regional distribution of polyphosphoinositides in rat brain. Biochim Biophys Acta. 1971 Oct 5;248(1):87–95. [PubMed]
  • Hoerl BJ, Scott RE. Plasma membrane vesiculation: a cellular response to injury. Virchows Arch B Cell Pathol. 1978 Jun 19;27(4):335–345. [PubMed]
  • Holdsworth G, Coleman R. Plasma-membrane components can be removed from isolated lymphocytes by the bile salts glycocholate and taurocholate without cell lysis. Biochem J. 1976 Aug 15;158(2):493–495. [PMC free article] [PubMed]
  • Laemmli UK. Cleavage of structural proteins during the assembly of the head of bacteriophage T4. Nature. 1970 Aug 15;227(5259):680–685. [PubMed]
  • Luzio JP, Stanley KK. The isolation of endosome-derived vesicles from rat hepatocytes. Biochem J. 1983 Oct 15;216(1):27–36. [PMC free article] [PubMed]
  • Maguire GA, Docherty K, Hales CN. Sugar transport in rat liver lysosomes. Direct demonstration by using labelled sugars. Biochem J. 1983 Apr 15;212(1):211–218. [PMC free article] [PubMed]
  • Morgan BP. Non-lethal complement-membrane attack on human neutrophils: transient cell swelling and metabolic depletion. Immunology. 1988 Jan;63(1):71–77. [PMC free article] [PubMed]
  • Morgan BP. Complement membrane attack on nucleated cells: resistance, recovery and non-lethal effects. Biochem J. 1989 Nov 15;264(1):1–14. [PMC free article] [PubMed]
  • Morgan BP, Campbell AK. The recovery of human polymorphonuclear leucocytes from sublytic complement attack is mediated by changes in intracellular free calcium. Biochem J. 1985 Oct 1;231(1):205–208. [PMC free article] [PubMed]
  • Morgan BP, Daw RA, Siddle K, Luzio JP, Campbell AK. Immunoaffinity purification of human complement component C9 using monoclonal antibodies. J Immunol Methods. 1983 Nov 25;64(3):269–281. [PubMed]
  • Morgan BP, Luzio JP, Campbell AK. Intracellular Ca2+ and cell injury: a paradoxical role of Ca2+ in complement membrane attack. Cell Calcium. 1986 Dec;7(5-6):399–411. [PubMed]
  • Morgan BP, Dankert JR, Esser AF. Recovery of human neutrophils from complement attack: removal of the membrane attack complex by endocytosis and exocytosis. J Immunol. 1987 Jan 1;138(1):246–253. [PubMed]
  • Müller-Eberhard HJ. Molecular organization and function of the complement system. Annu Rev Biochem. 1988;57:321–347. [PubMed]
  • Newby AC. Role of adenosine deaminase, ecto-(5'-nucleotidase) and ecto-(non-specific phosphatase) in cyanide-induced adenosine monophosphate catabolism in rat polymorphonuclear leucocytes. Biochem J. 1980 Mar 15;186(3):907–918. [PMC free article] [PubMed]
  • Richardson PJ, Luzio JP. Complement-mediated production of plasma-membrane vesicles from rat fat-cells. Biochem J. 1980 Mar 15;186(3):897–906. [PMC free article] [PubMed]
  • Salomon Y, Londos C, Rodbell M. A highly sensitive adenylate cyclase assay. Anal Biochem. 1974 Apr;58(2):541–548. [PubMed]
  • Scott RE, Maercklein PB. Plasma membrane vesiculation in 3T3 and SV3T3 cells. II. Factors affecting the process of vesiculation. J Cell Sci. 1979 Feb;35:245–252. [PubMed]
  • Scott RE, Perkins RG, Zschunke MA, Hoerl BJ, Maercklein PB. Plasma membrane vesiculation in 3T3 and SV3T3 cells. I. Morphological and biochemical characterization. J Cell Sci. 1979 Feb;35:229–243. [PubMed]
  • Shirley PS, Wang P, DeChatelet LR, Waite M. Absence of the membrane marker enzyme 5'-nucleotidase in human polymorphonuclear leukocytes. Biochem Med. 1976 Jun;15(3):289–295. [PubMed]
  • Shukla SD, Berriman J, Coleman R, Finean JB, Michell RH. Membrane protein segregation during release of microvesicles from human erythrocytes. FEBS Lett. 1978 Jun 15;90(2):289–292. [PubMed]
  • Sims PJ, Faioni EM, Wiedmer T, Shattil SJ. Complement proteins C5b-9 cause release of membrane vesicles from the platelet surface that are enriched in the membrane receptor for coagulation factor Va and express prothrombinase activity. J Biol Chem. 1988 Dec 5;263(34):18205–18212. [PubMed]
  • Stein JM, Smith GA, Luzio JP. An acetylation method for the quantification of membrane lipids, including phospholipids, polyphosphoinositides and cholesterol. Biochem J. 1991 Mar 1;274(Pt 2):375–379. [PMC free article] [PubMed]
  • van Meer G. Lipid traffic in animal cells. Annu Rev Cell Biol. 1989;5:247–275. [PubMed]
  • Wiedmer T, Shattil SJ, Cunningham M, Sims PJ. Role of calcium and calpain in complement-induced vesiculation of the platelet plasma membrane and in the exposure of the platelet factor Va receptor. Biochemistry. 1990 Jan 23;29(3):623–632. [PubMed]

Articles from Biochemical Journal are provided here courtesy of The Biochemical Society


Related citations in PubMed

See reviews...See all...

Cited by other articles in PMC

See all...


  • Compound
    PubChem Compound links
  • MedGen
    Related information in MedGen
  • PubMed
    PubMed citations for these articles
  • Substance
    PubChem Substance links

Recent Activity

Your browsing activity is empty.

Activity recording is turned off.

Turn recording back on

See more...