913:
its role in the pathogenesis of higher organisms, more recent studies suggested a broader physiological role in defense against simple eukaryotic predators and its role in inter-bacteria interactions. The Type VI secretion system gene clusters contain from 15 to more than 20 genes, two of which, Hcp and VgrG, have been shown to be nearly universally secreted substrates of the system. Structural analysis of these and other proteins in this system bear a striking resemblance to the tail spike of the T4 phage, and the activity of the system is thought to functionally resemble phage infection.
17:
106:
855:
528:
468:
428:
398:
933:. Vesicles from a number of bacterial species have been found to contain virulence factors, some have immunomodulatory effects, and some can directly adhere to and intoxicate host cells. release of vesicles has been demonstrated as a general response to stress conditions, the process of loading cargo proteins seems to be selective.
912:
led to decreased virulence and pathogenicity. Since then, Type VI secretion systems have been found in a quarter of all proteobacterial genomes, including animal, plant, human pathogens, as well as soil, environmental or marine bacteria. While most of the early studies of Type VI secretion focused on
404:
Type I secretion is a chaperone dependent secretion system employing the Hly and Tol gene clusters. The process begins as a leader sequence on the protein to be secreted is recognized by HlyA and binds HlyB on the membrane. This signal sequence is extremely specific for the ABC transporter. The HlyAB
869:
with their C-terminus which inserts into the outer membrane, allowing the rest of the peptide (the passenger domain) to reach the outside of the cell. Often, autotransporters are cleaved, leaving the beta-barrel domain in the outer membrane and freeing the passenger domain. Some researchers believe
449:. Once there, they pass through the outer membrane via a multimeric (12–14 subunits) complex of pore forming secretin proteins. In addition to the secretin protein, 10–15 other inner and outer membrane proteins compose the full secretion apparatus, many with as yet unknown function. Gram-negative
405:
complex stimulates HlyD which begins to uncoil and reaches the outer membrane where TolC recognizes a terminal molecule or signal on HlyD. HlyD recruits TolC to the inner membrane and HlyA is excreted outside of the outer membrane via a long-tunnel protein channel.
498:) can inject proteins into eukaryotic cells. The low Ca concentration in the cytosol opens the gate that regulates T3SS. One such mechanism to detect low calcium concentration has been illustrated by the lcrV (Low Calcium Response) antigen utilized by
504:, which is used to detect low calcium concentrations and elicits T3SS attachment. The Hrp system in plant pathogens inject harpins and pathogen effector proteins through similar mechanisms into plants. This secretion system was first discovered in
812:. T4SS are cell envelope-spanning complexes, or, in other words, 11–13 core proteins that form a channel through which DNA and proteins can travel from the cytoplasm of the donor cell to the cytoplasm of the recipient cell. T4SS also secrete
1694:
Thanassi DG, Stathopoulos C, Karkal A, Li H (2005). "Protein secretion in the absence of ATP: the autotransporter, two-partner secretion and chaperone/usher pathways of gram-negative bacteria (review)".
453:
use a modified version of the type II system for their biogenesis, and in some cases certain proteins are shared between a pilus complex and type II system within a single bacterial species.
239:
gradient: the pH of the cytosol is 7.4, the ER's pH is 7.0, and the cis-golgi has a pH of 6.5. Secretory vesicles have pHs ranging between 5.0 and 6.0; some secretory vesicles evolve into
97:) of a bacterial cell to its exterior. Secretion is a very important mechanism in bacterial functioning and operation in their natural surrounding environment for adaptation and survival.
925:. Portions of the outer membrane pinch off, forming nano-scale spherical structures made of a lipopolysaccharide-rich lipid bilayer enclosing periplasmic materials, and are deployed for
381:, one of two translocation systems, which requires the presence of an N-terminal signal peptide on the secreted protein. Others are translocated across the cytoplasmic membrane by the
804:
The Type IV secretion system (T4SS) is the general mechanism by which bacterial cells secrete or take up macromolecules. Their precise mechanism remains unknown. T4SS is encoded on
510:
and showed that toxins could be injected directly from the bacterial cytoplasm into the cytoplasm of its host's cells rather than simply be secreted into the extracellular medium.
713:, which uses this system to introduce the T-DNA portion of the Ti plasmid into the plant host, which in turn causes the affected area to develop into a crown gall (tumor).
740:
408:
Type I secretion system transports various molecules, from ions, drugs, to proteins of various sizes (20 – 900 kDa). The molecules secreted vary in size from the small
445:
Proteins secreted through the type II system, or main terminal branch of the general secretory pathway, depend on the Sec or Tat system for initial transport into the
663:
1261:"Structural features of the Pseudomonas fluorescens biofilm adhesin LapA required for LapG-dependent cleavage, biofilm formation, and cell surface localization"
2218:
An optimal set of features for predicting type IV secretion system effector proteins for a subset of species based on a multi-level feature selection approach
921:
In addition to the use of the multiprotein complexes listed above, Gram-negative bacteria possess another method for release of material: the formation of
171:
1840:
Mougous JD, Cuff ME, Raunser S, Shen A, Zhou M, Gifford CA, Goodman AL, Joachimiak G, Ordoñez CL, Lory S, Walz T, Joachimiak A, Mekalanos JJ (June 2006).
263:(IL1) etc. which do not have a signal sequence. They do not use the classical ER-Golgi pathway. These are secreted through various nonclassical pathways.
45:
is the removal of certain substances or waste products from a cell or organism. The classical mechanism of cell secretion is via secretory portals at the
389:
have two membranes, thus making secretion topologically more complex. There are at least six specialized secretion systems in Gram-negative bacteria.
849:
374:(ABC) type transporters are common to the three domains of life. Some secreted proteins are translocated across the cytoplasmic membrane by the
57:
structures embedded in the cell membrane, where secretory vesicles transiently dock and fuse to release intra-vesicular contents from the cell.
619:
607:
2340:
1539:"Structural and functional characterization of the VirB5 protein from the type IV secretion system encoded by the conjugative plasmid pKM101"
474:
It is homologous to the basal body in bacterial flagella. It is like a molecular syringe through which a bacterium (e.g. certain types of
424:
and the lipases. Type I secretion is also involved in export of non-proteinaceous substrates like cyclic β-glucans and polysaccharides.
2806:
840:, which stands for ankyrin repeat domain-containing protein A. Additionally, T4SS coupling proteins are VirD4, which bind to VirE2.
875:
1738:
Gerlach RG, Hensel M (October 2007). "Protein secretion systems and adhesins: the molecular armory of Gram-negative pathogens".
1775:"Identification of a conserved bacterial protein secretion system in Vibrio cholerae using the Dictyostelium host model system"
2801:
2272:
2249:
1243:
575:
683:
2605:
288:
In addition, proteins can be released from cells by mechanical or physiological wounding and through non-lethal, transient
874:
which form similar beta-barrel structures. A common example of an autotransporter that uses this secretion system is the
382:
1113:
Nickel W, Seedorf M (2008). "Unconventional mechanisms of protein transport to the cell surface of eukaryotic cells".
2333:
1316:
969:
922:
2217:
828:
As shown in the above figure, TraC, in particular consists of a three helix bundle and a loose globular appendage.
836:
T4SS has two effector proteins: firstly, ATS-1, which stands for
Anaplasma translocated substrate 1, and secondly
1773:
Pukatzki S, Ma AT, Sturtevant D, Krastins B, Sarracino D, Nelson WC, Heidelberg JF, Mekalanos JJ (January 2006).
194:
2313:
1898:
671:
974:
2326:
979:
926:
266:
At least four nonclassical (unconventional) protein secretion pathways have been described. They include:
930:
955:
species, the accessory secretory system handles the export of highly repetitive adhesion glycoproteins.
964:
942:
789:
768:
667:
365:
271:
60:
865:
system for crossing the inner membrane. Proteins which use this pathway have the capability to form a
817:
773:
588:
462:
2306:
887:
696:
440:
2170:"Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response"
1185:"Protein release through nonlethal oncotic pores as an alternative nonclassical secretory pathway"
816:
factor proteins directly into host cells as well as taking up DNA from the medium during natural
416:
2349:
2415:
1148:
McNeil PL, Steinhardt RA (2003). "Plasma membrane disruption: repair, prevention, adaptation".
904:
805:
735:
386:
317:
2235:
2837:
2832:
2500:
2475:
2455:
1161:
1126:
793:
702:
305:
147:
780:
Protein members of this family are components of the type IV secretion system. They mediate
370:
Secretion is not unique to eukaryotes – it is also present in bacteria and archaea as well.
205:
to the edge of the cell. More modification can occur in the secretory vesicles (for example
2720:
2530:
2480:
1853:
1786:
1550:
1453:
725:
650:
371:
771:
into its eukaryotic host. The prototypic Type IVA secretion system is the VirB complex of
8:
2842:
2635:
2505:
715:
276:
159:
80:
2095:
1857:
1790:
1554:
1356:
1329:
2827:
2796:
2750:
2730:
2715:
2540:
2535:
2194:
2169:
2104:
2079:
2014:
1989:
1965:
1940:
1874:
1841:
1817:
1804:
1774:
1720:
1671:
1647:"Type IV secretion in the obligatory intracellular bacterium Anaplasma phagocytophilum"
1646:
1568:
1511:
1486:
1462:
1437:
1413:
1388:
1369:
1285:
1260:
1259:
Boyd CD, Smith TJ, El-Kirat-Chatel S, Newell PD, DufrĂŞne YF, O'Toole GA (August 2014).
1211:
1184:
1090:
1081:
1065:
1041:
1016:
612:
167:
30:
2302:
1622:
1605:
1581:
1538:
2735:
2615:
2470:
2268:
2261:
2245:
2199:
2185:
2150:
2109:
2060:
2019:
1970:
1921:
1879:
1822:
1755:
1712:
1676:
1662:
1627:
1586:
1516:
1467:
1418:
1361:
1346:
1312:
1290:
1239:
1216:
1165:
1130:
1095:
1046:
989:
658:
580:
183:
139:
135:
556:
2460:
2189:
2181:
2140:
2099:
2091:
2050:
2009:
2001:
1960:
1952:
1913:
1869:
1861:
1842:"A virulence locus of Pseudomonas aeruginosa encodes a protein secretion apparatus"
1812:
1794:
1747:
1724:
1704:
1666:
1658:
1617:
1576:
1558:
1506:
1498:
1457:
1449:
1408:
1400:
1373:
1351:
1341:
1280:
1272:
1206:
1196:
1157:
1122:
1085:
1077:
1036:
1028:
706:
646:
410:
321:
313:
301:
289:
624:
292:
in the plasma membrane induced by washing cells with serum-free media or buffers.
193:
In the Golgi apparatus, the glycosylation of the proteins is modified and further
2745:
2710:
2055:
2039:"The Type VI secretion system - a widespread and versatile cell targeting system"
2038:
1502:
994:
898:
871:
730:
600:
568:
506:
500:
345:
337:
309:
187:
163:
151:
85:
2220:”, PLOS ONE Journal, 2018, 13, e0197041. (doi.org/10.1371/journal.pone.0197041.)
1032:
723:
into gastric epithelial cells, which is associated with gastric carcinogenesis.
2600:
2410:
2239:
1779:
Proceedings of the
National Academy of Sciences of the United States of America
1751:
1543:
Proceedings of the
National Academy of Sciences of the United States of America
1438:"Biogenesis, architecture, and function of bacterial type IV secretion systems"
893:
837:
217:
123:
34:
2005:
1917:
1708:
1487:"Type IV secretion: the Agrobacterium VirB/D4 and related conjugation systems"
2821:
2680:
2575:
2465:
2440:
2435:
892:
Type VI secretion systems were originally identified in 2006 by the group of
861:
Also called the autotransporter system, type V secretion involves use of the
785:
781:
260:
221:
155:
76:
46:
1865:
1799:
1563:
1436:
Christie PJ, Atmakuri K, Krishnamoorthy V, Jakubowski S, Cascales E (2005).
1201:
709:. It is capable of transporting both DNA and proteins. It was discovered in
2725:
2705:
2670:
2665:
2655:
2650:
2630:
2610:
2520:
2490:
2450:
2430:
2420:
2400:
2395:
2390:
2385:
2203:
2154:
2113:
2064:
2023:
1974:
1956:
1925:
1883:
1826:
1759:
1716:
1680:
1631:
1590:
1520:
1471:
1422:
1365:
1294:
1220:
1169:
1134:
1099:
1050:
325:
202:
1236:
Bacterial
Secreted Proteins: Secretory Mechanisms and Role in Pathogenesis
584:
450:
29:
is the movement of material from one point to another, such as a secreted
2775:
2770:
2740:
2695:
2640:
2580:
2565:
2560:
2525:
2485:
2375:
2365:
2360:
2234:
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P, eds. (2002).
1066:"Discovery of the 'porosome'; the universal secretory machinery in cells"
984:
866:
232:
127:
63:
means the transport or translocation of effector molecules. For example:
54:
2145:
2128:
1808:
1435:
1276:
896:
at the
Harvard Medical School (Boston, USA) in two bacterial pathogens,
739:, the causing agent of legionellosis (Legionnaires' disease) utilizes a
2760:
2755:
2700:
2685:
2625:
2590:
2585:
2555:
2515:
2495:
2425:
2405:
1572:
1404:
1330:"Helicobacter pylori CagA: a new paradigm for bacterial carcinogenesis"
476:
378:
333:
270:
direct protein translocation across the plasma membrane likely through
225:
210:
179:
2318:
850:
Trimeric autotransporter adhesin § Type V secretion system (T5SS)
420:
cell adhesion protein LapA of 520 kDa. The best characterized are the
150:(ER). As they are synthesized, these proteins translocate into the ER
105:
16:
2675:
2660:
2645:
2595:
2570:
2129:"Bacterial outer membrane vesicles and the host–pathogen interaction"
813:
446:
421:
131:
120:
90:
50:
42:
908:. These were identified when mutations in the Hcp and VrgG genes in
2765:
2690:
2620:
2550:
2545:
2370:
809:
563:
488:
482:
304:
have the ability to be secretory cells. They have a well-developed
240:
198:
143:
64:
2780:
2510:
2380:
1017:"Neuronal porosome proteome: Molecular dynamics and architecture"
206:
175:
94:
68:
1015:
Lee JS, Jeremic A, Shin L, Cho WJ, Chen X, Jena BP (July 2012).
2445:
678:
494:
375:
353:
349:
2077:
1258:
201:
are then moved into secretory vesicles which travel along the
1537:
Yeo HJ, Yuan Q, Beck MR, Baron C, Waksman G (December 2003).
1491:
Biochimica et
Biophysica Acta (BBA) - Molecular Cell Research
341:
72:
38:
2216:
Z. Esna Ashari, N. Dasgupta, K. Brayton & S. Broschat, “
1772:
1693:
720:
640:
595:
551:
456:
329:
256:
252:
197:, including cleavage and functionalization, may occur. The
2080:"Structure and regulation of the type VI secretion system"
2030:
513:
2283:
1606:"F factor conjugation is a true type IV secretion system"
392:
2078:
Silverman JM, Brunet YR, Cascales E, Mougous JD (2012).
1604:
Lawley TD, Klimke WA, Gubbins MJ, Frost LS (July 2003).
1603:
2233:
854:
527:
467:
427:
397:
284:
release via exosomes derived from multivesicular bodies
236:
235:
control is maintained over this sequence by usage of a
186:
containing the properly folded proteins then enter the
2071:
1839:
729:, the causative agent of whooping cough, secretes the
228:, dumping its contents out of the cell's environment.
916:
170:
are usually identified here and retrotranslocated by
1990:"What is type VI secretion doing in all those bugs?"
23:
Controlled release of substances by cells or tissues
1389:"The versatile bacterial type IV secretion systems"
2260:
1987:
1896:
881:
870:remnants of the autotransporters gave rise to the
767:rafficking genes) system, to translocate numerous
434:
1897:Bingle LE, Bailey CM, Pallen MJ (February 2008).
1327:
1014:
843:
2819:
1988:Schwarz S, Hood RD, Mougous JD (December 2010).
1536:
1386:
1147:
1532:
1530:
1150:Annual Review of Cell and Developmental Biology
1115:Annual Review of Cell and Developmental Biology
2263:The Physiology and Biochemistry of Prokaryotes
1644:
2334:
1740:International Journal of Medical Microbiology
1737:
1112:
2167:
1932:
1890:
1833:
1766:
1731:
1687:
1638:
1597:
1527:
1478:
1429:
1380:
1309:Bacterial Pathogenesis: A Molecular Approach
1008:
936:
359:
2244:(4th ed.). New York: Garland Science.
1645:Rikihisa Y, Lin M, Niu H (September 2010).
1321:
719:uses a type IV secretion system to deliver
2341:
2327:
2126:
2036:
1307:Salyers, A. A. & Whitt, D. D. (2002).
1233:
1227:
1070:Journal of Cellular and Molecular Medicine
526:
2807:Timeline of biology and organic chemistry
2305:at the U.S. National Library of Medicine
2267:(2nd ed.). Oxford University Press.
2193:
2144:
2103:
2054:
2013:
1964:
1873:
1816:
1798:
1670:
1621:
1580:
1562:
1510:
1461:
1412:
1387:Cascales E, Christie PJ (November 2003).
1355:
1345:
1328:Hatakeyama M, Higashi H (December 2005).
1284:
1210:
1200:
1089:
1040:
246:
1938:
1484:
1311:, 2nd ed., Washington, D.C.: ASM Press.
1162:10.1146/annurev.cellbio.19.111301.140101
1127:10.1146/annurev.cellbio.24.110707.175320
1063:
457:Type III secretion system (T3SS or TTSS)
104:
15:
2348:
2312:T5SS / Autotransporter illustration at
1899:"Type VI secretion: a beginner's guide"
1182:
1057:
929:to manipulate environment or invade at
514:Type IV secretion system (T4SS or TFSS)
2820:
1454:10.1146/annurev.micro.58.030603.123630
393:Type I secretion system (T1SS or TOSS)
100:
2802:Nobel Prize in Physiology or Medicine
2322:
2258:
2168:McBroom AJ, Kuehn MJ (January 2007).
53:. Porosomes are permanent cup-shaped
2127:Kuehn MJ, Kesty NC (November 2005).
316:that produce secretions include the
2096:10.1146/annurev-micro-121809-151619
733:partly through the type IV system.
414:peptide colicin V, (10 kDa) to the
383:twin-arginine translocation pathway
295:
13:
2226:
1082:10.1111/j.1582-4934.2006.tb00294.x
917:Release of outer membrane vesicles
853:
466:
426:
396:
356:to lubricate and protect the eye.
224:at porosomes, by a process called
14:
2854:
2296:
970:Bacterial outer membrane vesicles
923:bacterial outer membrane vesicles
876:Trimeric Autotransporter Adhesins
2281:
2186:10.1111/j.1365-2958.2006.05522.x
1663:10.1111/j.1462-5822.2010.01500.x
1347:10.1111/j.1349-7006.2005.00130.x
344:to lubricate the skin and hair.
195:post-translational modifications
2210:
2161:
2120:
1981:
1941:"The type VI secretion toolkit"
1906:Current Opinion in Microbiology
882:Type VI secretion system (T6SS)
831:
792:ancestrally related to that of
435:Type II secretion system (T2SS)
178:, where they are degraded by a
134:process of secretion. Proteins
1301:
1252:
1176:
1141:
1106:
844:Type V secretion system (T5SS)
61:Secretion in bacterial species
1:
2241:Molecular Biology of the Cell
2084:Annual Review of Microbiology
1623:10.1016/S0378-1097(03)00430-0
1485:Christie PJ (November 2004).
1442:Annual Review of Microbiology
1000:
635:Available protein structures:
251:There are many proteins like
2056:10.1016/j.resmic.2013.03.017
1503:10.1016/j.bbamcr.2004.02.013
1393:Nature Reviews. Microbiology
980:Membrane vesicle trafficking
927:membrane vesicle trafficking
823:
213:in the secretory vesicles).
115:
89:) from across the interior (
7:
1234:Wooldridge, K, ed. (2009).
1183:Chirico WJ (October 2011).
1033:10.1016/j.jprot.2012.05.017
958:
799:
705:machinery of bacteria, the
272:membrane transport proteins
10:
2859:
1939:Cascales E (August 2008).
1752:10.1016/j.ijmm.2007.03.017
1697:Molecular Membrane Biology
1238:. Caister Academic Press.
965:Bacterial effector protein
943:Bacterial secretion system
940:
885:
847:
694:
460:
438:
366:Bacterial secretion system
363:
312:to fulfill this function.
243:, which have a pH of 4.8.
2789:
2356:
2006:10.1016/j.tim.2010.09.001
1918:10.1016/j.mib.2008.01.006
1709:10.1080/09687860500063290
1610:FEMS Microbiology Letters
975:Host–pathogen interaction
937:In gram-positive bacteria
774:Agrobacterium tumefaciens
741:type IVB secretion system
711:Agrobacterium tumefaciens
677:
657:
639:
634:
630:
618:
606:
594:
574:
562:
550:
542:
537:
525:
520:
463:Type III secretion system
360:In gram-negative bacteria
172:ER-associated degradation
2307:Medical Subject Headings
2043:Research in Microbiology
888:Type VI secretion system
808:conjugative elements in
743:, known as the icm/dot (
697:Type IV secretion system
532:Type IV secretion system
441:Type II secretion system
2133:Genes & Development
1866:10.1126/science.1128393
1800:10.1073/pnas.0510322103
1564:10.1073/pnas.2535211100
1265:Journal of Bacteriology
1202:10.1186/1471-2121-12-46
931:host–pathogen interface
417:Pseudomonas fluorescens
2416:Biological engineering
2174:Molecular Microbiology
2037:Coulthurst SJ (2013).
1994:Trends in Microbiology
1957:10.1038/embor.2008.131
905:Pseudomonas aeruginosa
858:
736:Legionella pneumophila
471:
431:
401:
387:Gram-negative bacteria
318:gastrointestinal tract
247:Nonclassical secretion
112:
20:
2501:Developmental biology
2476:Computational biology
2456:Cellular microbiology
1651:Cellular Microbiology
1021:Journal of Proteomics
857:
794:bacterial conjugation
470:
430:
400:
306:endoplasmic reticulum
216:Eventually, there is
148:endoplasmic reticulum
108:
19:
2721:Reproductive biology
2606:Mathematical biology
2531:Evolutionary biology
2481:Conservation biology
1064:Anderson LL (2006).
726:Bordetella pertussis
701:It is homologous to
372:ATP binding cassette
158:and where molecular
146:docked to the rough
138:for the outside are
2506:Ecological genetics
2350:Branches of biology
2236:"Search: Secretion"
2146:10.1101/gad.1299905
1858:2006Sci...312.1526M
1791:2006PNAS..103.1528P
1555:2003PNAS..10015947Y
1277:10.1128/JB.01629-14
716:Helicobacter pylori
281:lysosomal secretion
101:In eukaryotic cells
81:pathogenic bacteria
2797:History of biology
2731:Structural biology
2716:Relational biology
2541:Generative biology
2536:Freshwater biology
1405:10.1038/nrmicro753
990:Secretory proteins
859:
472:
432:
402:
168:Misfolded proteins
113:
31:chemical substance
21:
2815:
2814:
2736:Synthetic biology
2616:Molecular biology
2471:Cognitive biology
2274:978-0-19-512579-5
2251:978-0-8153-3218-3
1852:(5779): 1526–30.
1245:978-1-904455-42-4
769:effector proteins
693:
692:
689:
688:
684:structure summary
322:digestive enzymes
154:, where they are
2850:
2461:Chemical biology
2343:
2336:
2329:
2320:
2319:
2291:
2278:
2266:
2259:White D (2000).
2255:
2221:
2214:
2208:
2207:
2197:
2165:
2159:
2158:
2148:
2124:
2118:
2117:
2107:
2075:
2069:
2068:
2058:
2034:
2028:
2027:
2017:
1985:
1979:
1978:
1968:
1936:
1930:
1929:
1903:
1894:
1888:
1887:
1877:
1837:
1831:
1830:
1820:
1802:
1770:
1764:
1763:
1735:
1729:
1728:
1691:
1685:
1684:
1674:
1642:
1636:
1635:
1625:
1601:
1595:
1594:
1584:
1566:
1549:(26): 15947–52.
1534:
1525:
1524:
1514:
1482:
1476:
1475:
1465:
1433:
1427:
1426:
1416:
1384:
1378:
1377:
1359:
1349:
1325:
1319:
1305:
1299:
1298:
1288:
1256:
1250:
1249:
1231:
1225:
1224:
1214:
1204:
1189:BMC Cell Biology
1180:
1174:
1173:
1145:
1139:
1138:
1110:
1104:
1103:
1093:
1061:
1055:
1054:
1044:
1012:
755:ultiplication /
707:conjugative pili
632:
631:
530:
518:
517:
411:Escherichia coli
346:Meibomian glands
338:sebaceous glands
302:human cell types
296:In human tissues
209:is cleaved from
130:, have a highly
2858:
2857:
2853:
2852:
2851:
2849:
2848:
2847:
2818:
2817:
2816:
2811:
2785:
2746:Systems biology
2711:Quantum biology
2352:
2347:
2299:
2294:
2275:
2252:
2229:
2227:Further reading
2224:
2215:
2211:
2166:
2162:
2139:(22): 2645–55.
2125:
2121:
2076:
2072:
2035:
2031:
1986:
1982:
1937:
1933:
1901:
1895:
1891:
1838:
1834:
1771:
1767:
1736:
1732:
1692:
1688:
1643:
1639:
1602:
1598:
1535:
1528:
1497:(1–3): 219–34.
1483:
1479:
1434:
1430:
1385:
1381:
1326:
1322:
1306:
1302:
1271:(15): 2775–88.
1257:
1253:
1246:
1232:
1228:
1181:
1177:
1146:
1142:
1111:
1107:
1062:
1058:
1027:(13): 3952–62.
1013:
1009:
1003:
995:Secretor status
961:
945:
939:
919:
910:Vibrio cholerae
899:Vibrio cholerae
890:
884:
852:
846:
834:
826:
802:
731:pertussis toxin
699:
608:OPM superfamily
533:
516:
507:Yersinia pestis
501:Yersinia pestis
465:
459:
443:
437:
395:
368:
362:
320:which secretes
310:Golgi apparatus
298:
249:
188:Golgi apparatus
164:protein folding
118:
103:
86:Vibrio cholerae
47:plasma membrane
41:. In contrast,
24:
12:
11:
5:
2856:
2846:
2845:
2840:
2835:
2830:
2813:
2812:
2810:
2809:
2804:
2799:
2793:
2791:
2787:
2786:
2784:
2783:
2778:
2773:
2768:
2763:
2758:
2753:
2748:
2743:
2738:
2733:
2728:
2723:
2718:
2713:
2708:
2703:
2698:
2693:
2688:
2683:
2678:
2673:
2668:
2663:
2658:
2653:
2648:
2643:
2638:
2633:
2628:
2623:
2618:
2613:
2608:
2603:
2601:Marine biology
2598:
2593:
2588:
2583:
2578:
2573:
2568:
2563:
2558:
2553:
2548:
2543:
2538:
2533:
2528:
2523:
2518:
2513:
2508:
2503:
2498:
2493:
2488:
2483:
2478:
2473:
2468:
2463:
2458:
2453:
2448:
2443:
2438:
2433:
2428:
2423:
2418:
2413:
2411:Bioinformatics
2408:
2403:
2398:
2393:
2388:
2383:
2378:
2373:
2368:
2363:
2357:
2354:
2353:
2346:
2345:
2338:
2331:
2323:
2317:
2316:
2310:
2298:
2297:External links
2295:
2293:
2292:
2279:
2273:
2256:
2250:
2230:
2228:
2225:
2223:
2222:
2209:
2160:
2119:
2070:
2029:
1980:
1931:
1889:
1832:
1785:(5): 1528–33.
1765:
1730:
1703:(1–2): 63–72.
1686:
1657:(9): 1213–21.
1637:
1596:
1526:
1477:
1428:
1379:
1340:(12): 835–43.
1334:Cancer Science
1320:
1300:
1251:
1244:
1226:
1175:
1140:
1105:
1056:
1006:
1002:
999:
998:
997:
992:
987:
982:
977:
972:
967:
960:
957:
949:Staphylococcus
941:Main article:
938:
935:
918:
915:
894:John Mekalanos
886:Main article:
883:
880:
845:
842:
833:
830:
825:
822:
818:transformation
801:
798:
786:macromolecules
695:Main article:
691:
690:
687:
686:
681:
675:
674:
661:
655:
654:
644:
637:
636:
628:
627:
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598:
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566:
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559:
554:
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547:
544:
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539:
535:
534:
531:
523:
522:
515:
512:
461:Main article:
458:
455:
439:Main article:
436:
433:
394:
391:
364:Main article:
361:
358:
340:which secrete
332:which secrete
297:
294:
286:
285:
282:
279:
274:
248:
245:
218:vesicle fusion
117:
114:
102:
99:
22:
9:
6:
4:
3:
2:
2855:
2844:
2841:
2839:
2836:
2834:
2831:
2829:
2826:
2825:
2823:
2808:
2805:
2803:
2800:
2798:
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2792:
2788:
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2779:
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2729:
2727:
2724:
2722:
2719:
2717:
2714:
2712:
2709:
2707:
2704:
2702:
2699:
2697:
2694:
2692:
2689:
2687:
2684:
2682:
2681:Phylogenetics
2679:
2677:
2674:
2672:
2669:
2667:
2664:
2662:
2659:
2657:
2654:
2652:
2649:
2647:
2644:
2642:
2639:
2637:
2634:
2632:
2629:
2627:
2624:
2622:
2619:
2617:
2614:
2612:
2609:
2607:
2604:
2602:
2599:
2597:
2594:
2592:
2589:
2587:
2584:
2582:
2579:
2577:
2576:Human biology
2574:
2572:
2569:
2567:
2564:
2562:
2559:
2557:
2554:
2552:
2549:
2547:
2544:
2542:
2539:
2537:
2534:
2532:
2529:
2527:
2524:
2522:
2519:
2517:
2514:
2512:
2509:
2507:
2504:
2502:
2499:
2497:
2494:
2492:
2489:
2487:
2484:
2482:
2479:
2477:
2474:
2472:
2469:
2467:
2466:Chronobiology
2464:
2462:
2459:
2457:
2454:
2452:
2449:
2447:
2444:
2442:
2441:Biotechnology
2439:
2437:
2436:Biostatistics
2434:
2432:
2429:
2427:
2424:
2422:
2419:
2417:
2414:
2412:
2409:
2407:
2404:
2402:
2399:
2397:
2394:
2392:
2389:
2387:
2384:
2382:
2379:
2377:
2374:
2372:
2369:
2367:
2364:
2362:
2359:
2358:
2355:
2351:
2344:
2339:
2337:
2332:
2330:
2325:
2324:
2321:
2315:
2311:
2308:
2304:
2301:
2300:
2289:
2285:
2280:
2276:
2270:
2265:
2264:
2257:
2253:
2247:
2243:
2242:
2237:
2232:
2231:
2219:
2213:
2205:
2201:
2196:
2191:
2187:
2183:
2180:(2): 545–58.
2179:
2175:
2171:
2164:
2156:
2152:
2147:
2142:
2138:
2134:
2130:
2123:
2115:
2111:
2106:
2101:
2097:
2093:
2089:
2085:
2081:
2074:
2066:
2062:
2057:
2052:
2049:(6): 640–54.
2048:
2044:
2040:
2033:
2025:
2021:
2016:
2011:
2007:
2003:
2000:(12): 531–7.
1999:
1995:
1991:
1984:
1976:
1972:
1967:
1962:
1958:
1954:
1951:(8): 735–41.
1950:
1946:
1942:
1935:
1927:
1923:
1919:
1915:
1911:
1907:
1900:
1893:
1885:
1881:
1876:
1871:
1867:
1863:
1859:
1855:
1851:
1847:
1843:
1836:
1828:
1824:
1819:
1814:
1810:
1806:
1801:
1796:
1792:
1788:
1784:
1780:
1776:
1769:
1761:
1757:
1753:
1749:
1746:(6): 401–15.
1745:
1741:
1734:
1726:
1722:
1718:
1714:
1710:
1706:
1702:
1698:
1690:
1682:
1678:
1673:
1668:
1664:
1660:
1656:
1652:
1648:
1641:
1633:
1629:
1624:
1619:
1615:
1611:
1607:
1600:
1592:
1588:
1583:
1578:
1574:
1570:
1565:
1560:
1556:
1552:
1548:
1544:
1540:
1533:
1531:
1522:
1518:
1513:
1508:
1504:
1500:
1496:
1492:
1488:
1481:
1473:
1469:
1464:
1459:
1455:
1451:
1447:
1443:
1439:
1432:
1424:
1420:
1415:
1410:
1406:
1402:
1399:(2): 137–49.
1398:
1394:
1390:
1383:
1375:
1371:
1367:
1363:
1358:
1353:
1348:
1343:
1339:
1335:
1331:
1324:
1318:
1317:1-55581-171-X
1314:
1310:
1304:
1296:
1292:
1287:
1282:
1278:
1274:
1270:
1266:
1262:
1255:
1247:
1241:
1237:
1230:
1222:
1218:
1213:
1208:
1203:
1198:
1194:
1190:
1186:
1179:
1171:
1167:
1163:
1159:
1155:
1151:
1144:
1136:
1132:
1128:
1124:
1120:
1116:
1109:
1101:
1097:
1092:
1087:
1083:
1079:
1076:(1): 126–31.
1075:
1071:
1067:
1060:
1052:
1048:
1043:
1038:
1034:
1030:
1026:
1022:
1018:
1011:
1007:
1005:
996:
993:
991:
988:
986:
983:
981:
978:
976:
973:
971:
968:
966:
963:
962:
956:
954:
953:Streptococcus
950:
944:
934:
932:
928:
924:
914:
911:
907:
906:
901:
900:
895:
889:
879:
877:
873:
868:
864:
856:
851:
841:
839:
829:
821:
819:
815:
811:
807:
806:Gram-negative
797:
796:machineries.
795:
791:
787:
783:
782:intracellular
778:
776:
775:
770:
766:
762:
758:
754:
750:
746:
742:
738:
737:
732:
728:
727:
722:
718:
717:
712:
708:
704:
698:
685:
682:
680:
676:
673:
669:
665:
662:
660:
656:
652:
648:
645:
642:
638:
633:
629:
626:
623:
621:
617:
614:
611:
609:
605:
602:
599:
597:
593:
590:
586:
582:
579:
577:
573:
570:
567:
565:
561:
558:
555:
553:
549:
545:
541:
536:
529:
524:
519:
511:
509:
508:
503:
502:
497:
496:
491:
490:
485:
484:
479:
478:
469:
464:
454:
452:
448:
442:
429:
425:
423:
419:
418:
413:
412:
406:
399:
390:
388:
384:
380:
377:
373:
367:
357:
355:
351:
347:
343:
339:
335:
331:
327:
323:
319:
315:
311:
307:
303:
293:
291:
290:oncotic pores
283:
280:
278:
275:
273:
269:
268:
267:
264:
262:
261:interleukin-1
258:
254:
244:
242:
238:
234:
229:
227:
223:
222:cell membrane
219:
214:
212:
208:
204:
200:
196:
191:
189:
185:
181:
177:
173:
169:
165:
161:
157:
153:
149:
145:
141:
137:
133:
129:
125:
122:
111:
107:
98:
96:
92:
88:
87:
82:
78:
77:cholera toxin
74:
70:
66:
62:
58:
56:
52:
48:
44:
40:
36:
32:
28:
18:
2838:Cell biology
2833:Biochemistry
2726:Sociobiology
2706:Protistology
2671:Photobiology
2666:Pharmacology
2656:Parasitology
2651:Paleontology
2631:Neuroscience
2611:Microbiology
2521:Epidemiology
2491:Cytogenetics
2451:Cell biology
2431:Biosemiotics
2421:Biomechanics
2401:Biogeography
2396:Biochemistry
2391:Bacteriology
2386:Astrobiology
2288:Cells alive!
2287:
2262:
2240:
2212:
2177:
2173:
2163:
2136:
2132:
2122:
2087:
2083:
2073:
2046:
2042:
2032:
1997:
1993:
1983:
1948:
1945:EMBO Reports
1944:
1934:
1909:
1905:
1892:
1849:
1845:
1835:
1782:
1778:
1768:
1743:
1739:
1733:
1700:
1696:
1689:
1654:
1650:
1640:
1613:
1609:
1599:
1546:
1542:
1494:
1490:
1480:
1445:
1441:
1431:
1396:
1392:
1382:
1337:
1333:
1323:
1308:
1303:
1268:
1264:
1254:
1235:
1229:
1192:
1188:
1178:
1153:
1149:
1143:
1118:
1114:
1108:
1073:
1069:
1059:
1024:
1020:
1010:
1004:
952:
948:
946:
920:
909:
903:
897:
891:
862:
860:
835:
832:Interactions
827:
803:
784:transfer of
779:
772:
764:
760:
756:
752:
748:
744:
734:
724:
714:
710:
700:
505:
499:
493:
487:
481:
475:
473:
451:type IV pili
444:
415:
409:
407:
403:
369:
326:gastric acid
299:
287:
265:
250:
230:
215:
203:cytoskeleton
192:
156:glycosylated
126:, including
119:
109:
84:
59:
26:
25:
2776:Xenobiology
2771:Virophysics
2741:Systematics
2696:Primatology
2641:Ornithology
2581:Ichthyology
2566:Herpetology
2561:Gerontology
2526:Epigenetics
2486:Cryobiology
2376:Agrostology
2366:Aerobiology
2361:Abiogenesis
2314:Uni MĂĽnster
2284:"Home page"
1616:(1): 1–15.
1156:: 697–731.
1121:: 287–308.
985:Secretomics
867:beta-barrel
703:conjugation
620:OPM protein
538:Identifiers
334:surfactants
233:biochemical
140:synthesized
128:human cells
55:lipoprotein
2843:Physiology
2822:Categories
2761:Toxicology
2756:Teratology
2701:Proteomics
2686:Physiology
2626:Neontology
2591:Lipidology
2586:Immunology
2556:Geobiology
2516:Embryology
2496:Dendrology
2426:Biophysics
2406:Biogeology
2303:Secretions
2090:: 453–72.
1912:(1): 3–8.
1448:: 451–85.
1001:References
848:See also:
647:structures
477:Salmonella
422:RTX toxins
379:translocon
226:exocytosis
211:proinsulin
180:proteasome
160:chaperones
121:Eukaryotic
2828:Secretion
2676:Phycology
2661:Pathology
2646:Osteology
2636:Nutrition
2596:Mammalogy
2571:Histology
824:Structure
814:virulence
790:mechanism
763:rganelle
759:efect in
569:IPR012991
447:periplasm
241:lysosomes
220:with the
144:ribosomes
116:Mechanism
91:cytoplasm
75:(such as
51:porosomes
43:excretion
27:Secretion
2790:See also
2766:Virology
2751:Taxonomy
2691:Pomology
2621:Mycology
2551:Genomics
2546:Genetics
2371:Agronomy
2282:Avon D.
2204:17163978
2155:16291643
2114:22746332
2065:23542428
2024:20961764
1975:18617888
1926:18289922
1884:16763151
1827:16432199
1809:30048406
1760:17482513
1717:16092525
1681:20670295
1632:12855161
1591:14673074
1521:15546668
1472:16153176
1423:15035043
1366:16367902
1357:11159386
1295:24837291
1221:22008609
1170:14570587
1135:18590485
1100:16563225
1051:22659300
959:See also
947:In some
810:bacteria
800:Function
751:ellular
664:RCSB PDB
564:InterPro
489:Yersinia
483:Shigella
385:(Tat).
352:secrete
277:blebbing
259:(bFGF),
255:(aFGF),
199:proteins
184:vesicles
136:targeted
110:Porosome
65:proteins
2781:Zoology
2511:Ecology
2381:Anatomy
2195:1868505
2105:3595004
2015:2991376
1966:2515208
1875:2800167
1854:Bibcode
1846:Science
1818:1345711
1787:Bibcode
1725:2708575
1672:3598623
1573:3149111
1551:Bibcode
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