424:
the proteins and DNA in the endospore. Next the peptidoglycan cortex forms between the two layers and the bacterium adds a spore coat to the outside of the forespore. In the final stages of endospore formation the newly forming endospore is dehydrated and allowed to mature before being released from the mother cell. The cortex is what makes the endospore so resistant to temperature. The cortex contains an inner membrane known as the core. The inner membrane that surrounds this core leads to the endospore's resistance against UV light and harsh chemicals that would normally destroy microbes. Sporulation is now complete, and the mature endospore will be released when the surrounding vegetative cell is degraded.
45:
27:
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202:
543:. Even if an endospore is located in plentiful nutrients, it may fail to germinate unless activation has taken place. This may be triggered by heating the endospore. Germination involves the dormant endospore starting metabolic activity and thus breaking hibernation. It is commonly characterised by rupture or absorption of the spore coat, swelling of the endospore, an increase in metabolic activity, and loss of resistance to environmental stress.
618:. The powder found in contaminated postal letters consisted of anthrax endospores. This intentional distribution led to 22 known cases of anthrax (11 inhalation and 11 cutaneous). The case fatality rate among those patients with inhalation anthrax was 45% (5/11). The six other individuals with inhalation anthrax and all the individuals with cutaneous anthrax recovered. Had it not been for antibiotic therapy, many more might have been stricken.
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554:. It is believed that the macromolecule accountable for maintaining the dormant state has a protein coat rich in cystine, stabilized by S-S linkages. A reduction in these linkages has the potential to change the tertiary structure, causing the protein to unfold. This conformational change in the protein is thought to be responsible for exposing active enzymatic sites necessary for endospore germination.
395:
233:
496:, are used as probes to verify that an autoclaved item has been rendered truly sterile: a small capsule containing the spores is put into the autoclave with the items; after the cycle the content of the capsule is cultured to check if anything will grow from it. If nothing will grow, then the spores were destroyed and the sterilization was successful.
562:. The bee fossilized in amber was dated to being about 25 million years old. The spores germinated when the amber was cracked open and the material from the gut of the bee was extracted and placed in nutrient medium. After the spores were analyzed by microscopy, it was determined that the cells were very similar to
557:
Endospores can stay dormant for a very long time. For instance, endospores were found in the tombs of the
Egyptian pharaohs. When placed in appropriate medium, under appropriate conditions, they were able to be reactivated. In 1995, Raul Cano of California Polytechnic State University found bacterial
507:
Bacterial endospores are resistant to antibiotics, most disinfectants, and physical agents such as radiation, boiling, and drying. The impermeability of the spore coat is thought to be responsible for the endospore's resistance to chemicals. The heat resistance of endospores is due to a variety of
423:
of the cell surrounds this wall and pinches off to leave a double membrane around the DNA, and the developing structure is now known as a forespore. Calcium dipicolinate, the calcium salt of dipicolinic acid, is incorporated into the forespore during this time. The dipicolinic acid helps stabilize
186:
said "There are viable bacterial spores that have been found that are 40 million years old on Earth—and we know they're very hardened to radiation." Common antibacterial agents that work by destroying vegetative cell walls do not affect endospores. Endospores are commonly found in soil and water,
503:
are killed by low-temperature, and non-corrosive, ethylene oxide sterilizers. Ethylene oxide is the only low-temperature sterilant to stop outbreaks on these instruments. In contrast, "high level disinfection" does not kill endospores but is used for instruments such as a colonoscope that do not
276:
endospores indicates the presence of a component with a regular periodic structure, which Kadota and Iijima speculated might be formed from a keratin-like protein. However, after further studies this group concluded that the structure of the spore coat protein was different from keratin. When the
362:
The position of the endospore differs among bacterial species and is useful in identification. The main types within the cell are terminal, subterminal, and centrally placed endospores. Terminal endospores are seen at the poles of cells, whereas central endospores are more or less in the middle.
327:
within the core, which is thought to stabilize the DNA. Dipicolinic acid could be responsible for the heat resistance of the spore, and calcium may aid in resistance to heat and oxidizing agents. However, mutants resistant to heat but lacking dipicolinic acid have been isolated, suggesting other
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Sporulation requires the presence of free oxygen. In the natural situation, this means the vegetative cycles occur within the low oxygen environment of the infected host and, within the host, the organism is exclusively in the vegetative form. Once outside the host, sporulation commences upon
470:
at a temperature exceeding the boiling point of water, 100 °C. Endospores are able to survive at 100 °C for hours, although the larger the number of hours the fewer that will survive. An indirect way to destroy them is to place them in an environment that reactivates them to their
1757:
463:) must be in contact with the spores for at least several minutes; a very small proportion of spores can survive longer than 10 minutes in such a solution. Higher concentrations of bleach are not more effective, and can cause some types of bacteria to aggregate and thus survive.
331:
Visualising endospores under light microscopy can be difficult due to the impermeability of the endospore wall to dyes and stains. While the rest of a bacterial cell may stain, the endospore is left colourless. To combat this, a special stain technique called a
328:
mechanisms contributing to heat resistance are also at work. Small acid-soluble proteins (SASPs) are found in endospores. These proteins tightly bind and condense the DNA, and are in part responsible for resistance to UV light and DNA-damaging chemicals.
214:
The bacterial life cycle does not necessarily include sporulation. Sporulation is usually triggered by adverse environmental conditions, so as to help the survival of the bacterium. Endospores exhibit no signs of life and can thus be described as
546:
Outgrowth follows germination and involves the core of the endospore manufacturing new chemical components and exiting the old spore coat to develop into a fully functional vegetative bacterial cell, which can divide to produce more cells.
641:
spores are useful for the expression of recombinant proteins and in particular for the surface display of peptides and proteins as a tool for fundamental and applied research in the fields of microbiology, biotechnology and vaccination.
363:
Subterminal endospores are those between these two extremes, usually seen far enough towards the poles but close enough to the center so as not to be considered either terminal or central. Lateral endospores are seen occasionally.
515:
Small acid-soluble proteins (SASPs) saturate the endospore's DNA and protect it from heat, drying, chemicals, and radiation. They also function as a carbon and energy source for the development of a vegetative bacterium during
471:
vegetative state. They will germinate within a day or two with the right environmental conditions, and then the vegetative cells, not as hardy as endospores, can be straightforwardly destroyed. This indirect method is called
90:
to lie dormant for extended periods, even centuries. There are many reports of spores remaining viable over 10,000 years, and revival of spores millions of years old has been claimed. There is one report of viable spores of
625:
sporulates when it sees oxygen instead of the carbon dioxide present in mammal blood; this signals to the bacteria that it has reached the end of the animal, and an inactive dispersable morphology is useful.
411:
When a bacterium detects environmental conditions are becoming unfavourable it may start the process of endosporulation, which takes about eight hours. The DNA is replicated and a membrane wall known as a
223:
under the appropriate conditions. Endospores have survived thousands of years until environmental stimuli trigger germination. They have been characterized as the most durable cells produced in nature.
1448:
519:
The cortex may osmotically remove water from the interior of the endospore and the dehydration that results is thought to be very important in the endospore's resistance to heat and radiation.
82:(i.e., not an offspring). It is a stripped-down, dormant form to which the bacterium can reduce itself. Endospore formation is usually triggered by a lack of nutrients, and usually occurs in
156:. Dipicolinic acid is a spore-specific chemical that appears to help in the ability for endospores to maintain dormancy. This chemical accounts for up to 10% of the spore's dry weight.
179:
growth on cheese after boiling the cheese. His notion of spores being the reproductive mechanism for the growth was a large blow to the previous suggestions of spontaneous generation.
272:
endospores, the spore coat is estimated to contain more than 70 coat proteins, which are organized into an inner and an outer coat layer. The X-ray diffraction pattern of purified
1863:
408:
Under conditions of starvation, especially the lack of carbon and nitrogen sources, a single endospore forms within some of the bacteria through a process called sporulation.
1897:
2091:
431:, endospores are the result of a morphological differentiation process triggered by nutrient limitation (starvation) in the environment; endosporulation is initiated by
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becomes more favorable, the endospore can reactivate itself into a vegetative state. Most types of bacteria cannot change to the endospore form. Examples of bacterial
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2011:
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Madigan, Michael T.; Bender, Kelly S.; Buckley, Daniel H.; Sattley, W. Matthew; Stahl, David A. (2018). "Microbial Cell
Structure and Function".
894:
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is used. That allows the endospore to show up as red, while the rest of the cell stains blue. Another staining technique for endospores is the
236:
Variations in endospore morphology: (1, 4) central endospore; (2, 3, 5) terminal endospore; (6) lateral endospore
1097:
957:
908:
1987:
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198:, instead of endospores. Exospores and endospores are two kinds of "hibernating" or dormant stages seen in some classes of microorganisms.
1867:
475:. It was the usual method for a while in the late 19th century before the introduction of inexpensive autoclaves. Prolonged exposure to
187:
where they may survive for long periods of time. A variety of different microorganisms form "spores" or "cysts", but the endospores of
550:
Endospores possess five times more sulfur than vegetative cells. This excess sulfur is concentrated in spore coats as an amino acid,
1500:
1889:
1314:
1373:
Cano, RJ; Borucki, MK (1995). "Revival and identification of bacterial spores in 25- to 40-million-year-old
Dominican amber".
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1981:
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86:. In endospore formation, the bacterium divides within its cell wall, and one side then engulfs the other. Endospores enable
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1453:
1890:"Ethylene Oxide Sterilization | Disinfection & Sterilization Guidelines | Guidelines Library | Infection Control |CDC"
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Bacteria produce a single endospore internally. The spore is sometimes surrounded by a thin covering known as the
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Endospores are resistant to most agents that would normally kill the vegetative cells they formed from. Unlike
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Finally, DNA repair enzymes contained within the endospore are able to repair damaged DNA during germination.
440:
382:. Sometimes the endospore can be so large the cell can be distended around the endospore. This is typical of
340:, which stains endospores green and bacterial bodies red. The arrangement of spore layers is as follows:
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564:
93:
2501:
2424:
2387:
2183:
1562:
Henriques AO, Moran CP Jr (2007). "Structure, assembly, and function of the spore surface layers".
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Calcium dipicolinate, abundant within the endospore, may stabilize and protect the endospore's DNA.
305:-like proteins known as SASPs (small acid-soluble spore proteins), that protect the spore DNA from
1968:
1628:
Hiragi Y, Iijima K, and Kadota H (1967). "Hexagonal single crystal pattern from the spore coat of
580:, the molecular details of endospore formation have been extensively studied, specifically in the
466:
While significantly resistant to heat and radiation, endospores can be destroyed by burning or by
504:
enter sterile bodily cavities. This latter method uses only warm water, enzymes, and detergents.
44:
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exposure to the air and the spore forms are essentially the exclusive phase in the environment.
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125:
98:
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Heninger, Sara; Christine A. Anderson; Gerald Beltz; Andrew B. Onderdonk (January 1, 2009).
1813:
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2332:
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1350:. Cornell University College of Agriculture and Life Sciences, Department of Microbiology.
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8:
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within the "starving" population.Most disinfectants such as household cleaning products,
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Reactivation of the endospore occurs when conditions are more favourable and involves
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36:
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1816:"Decontamination of Bacillus anthracis Spores: Evaluation of Various Disinfectants"
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of the endospore. The core contains the spore chromosomal DNA which is encased in
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1936:
1425:
1321:. Washington DC: National Council for Science and the Environment. Archived from
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1083:
1006:
880:
590:
589:. These studies have contributed much to our understanding of the regulation of
420:
378:
107:
490:
The endospores of certain types of (typically non-pathogenic) bacteria, such as
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radiation and heat. The core also contains normal cell structures, such as
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spores in the gut of a fossilized bee trapped in amber from a tree in the
26:
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Kadota H, Iijima K (1965). "The X-ray diffraction pattern of spores of
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1034:
845:
448:
444:
294:
241:
219:. Endospores retain viability indefinitely and they can germinate into
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1653:
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455:(ETO), and 10% bleach are effective against endospores. To kill most
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281:
genome was sequenced, no ortholog of human keratin was detected. The
257:
188:
149:
71:
1919:
Keynan, A.; Evenchik, Z.; Halvorson, H. O.; Hastings, J. W. (1964).
2168:
2145:
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endospores to extreme terrestrial and extraterrestrial environments
775:
719:
551:
436:
310:
253:
232:
87:
74:. The name "endospore" is suggestive of a spore or seed-like form (
67:
63:
1713:
1684:
499:
In hospitals, endospores on delicate invasive instruments such as
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936:
456:
373:
321:
314:
261:
137:
102:
1793:(10th ed.). Burlington, MA: Jones & Bartlett Learning.
1297:
194:
Some classes of bacteria can turn into exospores, also known as
159:
Endospores can survive without nutrients. They are resistant to
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2399:
1918:
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394:
97:
in salt crystals approximately 25 million years old. When the
2572:
1685:"The complete genome sequence of the gram-positive bacterium
480:
249:
205:
Formation of an endospore through the process of sporulation.
79:
1465:
650:
Examples of endospore-forming bacteria include the genera:
40:
showing endospores as green and the vegetative cell as red
1496:
320:
Up to 20% of the dry weight of the endospore consists of
171:. Thermo-resistant endospores were first hypothesized by
145:
66:, tough, and non-reproductive structure produced by some
1555:
568:
which is found in bees in the
Dominican Republic today.
419:
begins to form between it and the rest of the cell. The
366:
Examples of bacteria having terminal endospores include
376:. Bacteria having a centrally placed endospore include
306:
2097:
Malachite green – Endospore staining technique (video)
2012:"OIE Listed Diseases and Other Diseases of Importance"
1627:
1493:"Impacts 'more likely' to have spread life from Earth"
447:
have little effect on endospores. However, sterilant
2049:
Bacterial Spores: Current
Research and Applications
191:are by far the most resistant to harsh conditions.
1590:
167:, high temperature, extreme freezing and chemical
2712:
1621:
1416:Ringo, John (2004). "Reproduction of Bacteria".
1312:
2046:
1784:
1782:
1780:
1778:
1561:
2126:
1523:Introduction to Bacteria and Their Ecobiology
645:
2040:
1596:
1317:. In Sidney Draggan; C.J. Cleveland (eds.).
459:spores, standard household bleach (with 10%
389:
285:lies beneath the spore coat and consists of
1788:
1775:
1520:
1372:
1298:Murray, Patrick R.; Ellen Jo Baron (2003).
2140:
2133:
2119:
293:lies beneath the cortex and surrounds the
144:The endospore consists of the bacterium's
2092:Endospores – Brief microbiology text page
1944:
1839:
1712:
1490:
1342:
1340:
1338:
1336:
1334:
1332:
252:that excludes large toxic molecules like
209:
135:. Endospore formation is not found among
1514:
1409:
393:
231:
200:
43:
25:
1484:
621:According to WHO veterinary documents,
372:, the pathogen that causes the disease
16:Protective structure formed by bacteria
2713:
1576:10.1146/annurev.micro.61.080706.093224
1329:
78:means 'within'), but it is not a true
2114:
1900:from the original on 17 November 2019
1682:
1415:
1302:. Vol. 1. Washington, D.C.: ASM.
54:imaged with phase-contrast microscopy
2695:
2079:Marise A. Hussey and Anne Zayaitz –
1921:"Activation of bacterial endospores"
1521:Doetsch, R. N.; Cook, T. M. (1973).
1459:
248:. The spore coat, which acts like a
1882:
1503:from the original on 24 August 2011
317:, but is not metabolically active.
13:
1354:from the original on June 15, 2018
487:, will also kill most endospores.
404:Bacterial morphological plasticity
14:
2732:
2072:
105:that can form endospores include
2694:
2685:
2684:
2085:American Society of Microbiology
1789:Pommerville, Jeffrey C. (2014).
633:
2383:Bacterial cellular morphologies
2004:
1993:from the original on 2012-10-23
1961:
1912:
1856:
1807:
1750:
1729:
1676:
1468:Brock Biology of Microorganisms
1300:Manual of Clinical Microbiology
526:
1615:10.1080/00021369.1965.10858352
1442:
1366:
1306:
1291:
493:Geobacillus stearothermophilus
189:low G+C gram-positive bacteria
1:
1970:Anthrax in humans and animals
1683:Kunst F, et al. (1997).
1285:
571:
441:quaternary ammonium compounds
398:Endospore formation and cycle
256:, is resistant to many toxic
2047:Abel-Santos, E, ed. (2012).
1937:10.1128/JB.88.2.313-318.1964
1791:Fundamentals of microbiology
1491:BBC Staff (23 August 2011).
1454:Dorland's Medical Dictionary
1426:10.1017/CBO9780511807022.018
608:Endospores of the bacterium
227:
7:
1976:(4th ed.). OIE. 2008.
1739:, Wm. C. Brown Publishers,
1273:
357:
48:Phase-bright endospores of
10:
2737:
2626:Bacteria (classifications)
2348:Primary nutritional groups
1832:10.1177/153567600901400103
646:Endospore-forming bacteria
576:As a simplified model for
401:
18:
2680:
2611:
2591:
2543:
2432:
2423:
2375:
2268:
2206:
2192:
2152:
1531:10.1007/978-94-015-1135-3
1313:C. Michael Hogan (2010).
565:Lysinibacillus sphaericus
390:Formation and destruction
2502:Bacterial outer membrane
2081:Endospore stain protocol
578:cellular differentiation
19:Not to be confused with
1925:Journal of Bacteriology
1395:10.1126/science.7538699
2497:Gram-negative bacteria
2476:Gram-positive bacteria
2053:Caister Academic Press
2021:. 2012. Archived from
1348:"Bacterial Endospores"
399:
338:Schaeffer-Fulton stain
237:
210:Life cycle of bacteria
206:
120:Bacillus thuringiensis
84:gram-positive bacteria
55:
41:
2352:Substrate preference
1735:Prescott, L. (1993).
1319:Encyclopedia of Earth
595:transcription factors
402:Further information:
397:
264:that are involved in
260:and may also contain
244:, which overlies the
235:
204:
161:ultraviolet radiation
152:and large amounts of
126:Clostridium botulinum
47:
29:
2333:Microbial metabolism
2083:– Microbe Library (
1420:. pp. 153–160.
1418:Fundamental Genetics
1238:Thermoflavimicrobium
1210:Thermoalkalibacillus
769:Clostridiisalibacter
616:2001 anthrax attacks
94:Bacillus marismortui
2583:Non-motile bacteria
2179:Pathogenic bacteria
1705:1997Natur.390..249K
1646:1967Natur.215..154H
1387:1995Sci...268.1060C
1381:(5213): 1060–1064.
461:sodium hypochlorite
51:Paenibacillus alvei
21:Endospory in plants
2512:Lipopolysaccharide
2028:on August 12, 2016
2019:Terrestrial Manual
1762:Biology LibreTexts
1758:"2.4E: Endospores"
1564:Annu Rev Microbiol
1280:Endospore staining
1217:Thermoanaerobacter
1119:Sporolactobacillus
611:Bacillus anthracis
560:Dominican Republic
477:ionising radiation
400:
384:Clostridium tetani
369:Clostridium tetani
238:
207:
132:Clostridium tetani
114:Bacillus anthracis
56:
42:
2708:
2707:
2607:
2606:
2553:Bacterial capsule
2519:Periplasmic space
2486:Lipoteichoic acid
2371:
2370:
2343:Microbial ecology
2338:Nitrogen fixation
2062:978-1-908230-00-3
1983:978-92-4-154753-6
1820:Applied Biosafety
1687:Bacillus subtilis
1630:Bacillus subtilis
1599:Bacillus subtilis
1540:978-94-015-1137-7
1224:Thermoanaeromonas
1203:Thermoactinomyces
1196:Thermoacetogenium
1091:Sporoanaerobacter
1063:Salsuginibacillus
979:Ornithinibacillus
818:Desulfosporosinus
797:Dendrosporobacter
789:Coxiella burnetii
639:Bacillus subtilis
614:were used in the
586:Bacillus subtilis
270:Bacillus subtilus
184:Steinn Sigurdsson
177:Bacillus subtilis
37:Bacillus subtilis
2728:
2698:
2697:
2688:
2687:
2636:Former groupings
2430:
2429:
2281:Human microbiome
2204:
2203:
2135:
2128:
2121:
2112:
2111:
2067:
2066:
2044:
2038:
2037:
2035:
2033:
2027:
2016:
2008:
2002:
2001:
1999:
1998:
1992:
1975:
1965:
1959:
1958:
1948:
1916:
1910:
1909:
1907:
1905:
1896:. 4 April 2019.
1886:
1880:
1879:
1877:
1875:
1870:on March 3, 2016
1866:. Archived from
1860:
1854:
1853:
1843:
1811:
1805:
1804:
1786:
1773:
1772:
1770:
1769:
1754:
1748:
1733:
1727:
1726:
1716:
1699:(6657): 249–56.
1680:
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1673:
1654:10.1038/215154a0
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1406:
1370:
1364:
1363:
1361:
1359:
1344:
1327:
1326:
1310:
1304:
1303:
1295:
1189:Thalassobacillus
811:Desulfosporomusa
804:Desulfotomaculum
734:Caldanaerobacter
706:Aneurinibacillus
221:vegetative cells
154:dipicolinic acid
2736:
2735:
2731:
2730:
2729:
2727:
2726:
2725:
2711:
2710:
2709:
2704:
2676:
2631:Bacterial phyla
2615:
2603:
2587:
2545:
2539:
2530:Arabinogalactan
2435:
2419:
2367:
2264:
2208:
2196:
2188:
2174:Lysogenic cycle
2155:
2148:
2139:
2075:
2070:
2063:
2045:
2041:
2031:
2029:
2025:
2014:
2010:
2009:
2005:
1996:
1994:
1990:
1984:
1973:
1967:
1966:
1962:
1917:
1913:
1903:
1901:
1888:
1887:
1883:
1873:
1871:
1864:"The Autoclave"
1862:
1861:
1857:
1812:
1808:
1801:
1787:
1776:
1767:
1765:
1756:
1755:
1751:
1734:
1730:
1681:
1677:
1640:(5097): 154–5.
1626:
1622:
1603:Agric Biol Chem
1595:
1591:
1560:
1556:
1541:
1519:
1515:
1506:
1504:
1489:
1485:
1478:
1464:
1460:
1447:
1443:
1436:
1414:
1410:
1371:
1367:
1357:
1355:
1346:
1345:
1330:
1311:
1307:
1296:
1292:
1288:
1276:
1271:
1266:Vulcanobacillus
1245:Thermovenabulum
1112:Sporohalobacter
1084:Sporacetigenium
1007:Paraliobacillus
881:Gracilibacillus
648:
636:
591:gene expression
574:
529:
451:agents such as
429:persister cells
421:plasma membrane
406:
392:
379:Bacillus cereus
360:
230:
212:
196:microbial cysts
175:after studying
108:Bacillus cereus
32:endospore stain
24:
17:
12:
11:
5:
2734:
2724:
2723:
2706:
2705:
2703:
2702:
2692:
2681:
2678:
2677:
2675:
2674:
2673:
2672:
2667:
2662:
2657:
2647:
2642:
2633:
2628:
2622:
2620:
2609:
2608:
2605:
2604:
2602:
2601:
2595:
2593:
2589:
2588:
2586:
2585:
2580:
2575:
2570:
2565:
2560:
2555:
2549:
2547:
2541:
2540:
2538:
2537:
2532:
2521:
2516:
2515:
2514:
2509:
2493:
2488:
2483:
2472:
2471:
2470:
2465:
2460:
2446:
2440:
2438:
2427:
2421:
2420:
2418:
2417:
2412:
2407:
2402:
2397:
2396:
2395:
2390:
2388:cell structure
2379:
2377:
2373:
2372:
2369:
2368:
2366:
2365:
2364:
2363:
2361:Saccharophilic
2358:
2350:
2345:
2340:
2335:
2330:
2329:
2328:
2323:
2318:
2313:
2303:
2298:
2293:
2288:
2278:
2272:
2270:
2266:
2265:
2263:
2262:
2257:
2252:
2250:Microaerophile
2247:
2246:
2245:
2240:
2230:
2229:
2228:
2223:
2212:
2210:
2201:
2190:
2189:
2187:
2186:
2181:
2176:
2171:
2166:
2160:
2158:
2150:
2149:
2138:
2137:
2130:
2123:
2115:
2109:
2108:
2102:Resistance of
2099:
2094:
2089:
2074:
2073:External links
2071:
2069:
2068:
2061:
2039:
2003:
1982:
1960:
1931:(2): 313–318.
1911:
1881:
1855:
1806:
1800:978-1449688615
1799:
1774:
1749:
1728:
1675:
1620:
1589:
1554:
1539:
1513:
1483:
1476:
1470:. p. 92.
1458:
1441:
1434:
1408:
1365:
1328:
1325:on 2011-05-11.
1305:
1289:
1287:
1284:
1283:
1282:
1275:
1272:
1270:
1269:
1262:
1255:
1252:Tuberibacillus
1248:
1241:
1234:
1231:Thermobacillus
1227:
1220:
1213:
1206:
1199:
1192:
1185:
1178:
1171:
1164:
1161:Syntrophospora
1157:
1154:Syntrophomonas
1150:
1147:Sporotomaculum
1143:
1136:
1129:
1122:
1115:
1108:
1105:Sporobacterium
1101:
1094:
1087:
1080:
1073:
1066:
1059:
1056:Salinibacillus
1052:
1045:
1038:
1031:
1024:
1017:
1010:
1003:
996:
989:
982:
975:
968:
965:Oceanobacillus
961:
954:
947:
940:
933:
930:Lysinibacillus
926:
919:
912:
905:
902:Heliobacterium
898:
891:
884:
877:
874:Geosporobacter
870:
863:
856:
849:
842:
839:Desulfurispora
835:
832:Desulfunispora
828:
825:Desulfovirgula
821:
814:
807:
800:
793:
779:
772:
765:
758:
755:Cerasibacillus
751:
744:
737:
730:
723:
716:
709:
702:
695:
688:
681:
674:
671:Alkalibacillus
667:
660:
652:
647:
644:
635:
632:
603:RNA polymerase
582:model organism
573:
570:
528:
525:
524:
523:
520:
517:
513:
473:Tyndallization
453:ethylene oxide
433:quorum sensing
391:
388:
359:
356:
355:
354:
351:
348:
345:
229:
226:
211:
208:
181:Astrophysicist
173:Ferdinand Cohn
70:in the phylum
15:
9:
6:
4:
3:
2:
2733:
2722:
2719:
2718:
2716:
2701:
2693:
2691:
2683:
2682:
2679:
2671:
2668:
2666:
2663:
2661:
2658:
2656:
2653:
2652:
2651:
2648:
2646:
2643:
2641:
2640:Schizomycetes
2637:
2634:
2632:
2629:
2627:
2624:
2623:
2621:
2619:
2614:
2610:
2600:
2597:
2596:
2594:
2590:
2584:
2581:
2579:
2576:
2574:
2571:
2569:
2566:
2564:
2561:
2559:
2556:
2554:
2551:
2550:
2548:
2542:
2536:
2533:
2531:
2528:
2526:
2522:
2520:
2517:
2513:
2510:
2508:
2505:
2504:
2503:
2500:
2498:
2494:
2492:
2489:
2487:
2484:
2482:
2481:Teichoic acid
2479:
2477:
2473:
2469:
2466:
2464:
2461:
2459:
2456:
2455:
2454:
2453:Peptidoglycan
2450:
2447:
2445:
2444:Cell membrane
2442:
2441:
2439:
2437:
2431:
2428:
2426:
2422:
2416:
2413:
2411:
2408:
2406:
2403:
2401:
2398:
2394:
2391:
2389:
2386:
2385:
2384:
2381:
2380:
2378:
2374:
2362:
2359:
2357:
2354:
2353:
2351:
2349:
2346:
2344:
2341:
2339:
2336:
2334:
2331:
2327:
2324:
2322:
2319:
2317:
2314:
2311:
2307:
2304:
2302:
2299:
2297:
2294:
2292:
2289:
2287:
2284:
2283:
2282:
2279:
2277:
2274:
2273:
2271:
2267:
2261:
2258:
2256:
2253:
2251:
2248:
2244:
2241:
2239:
2236:
2235:
2234:
2231:
2227:
2224:
2222:
2219:
2218:
2217:
2214:
2213:
2211:
2205:
2202:
2200:
2195:
2191:
2185:
2182:
2180:
2177:
2175:
2172:
2170:
2167:
2165:
2162:
2161:
2159:
2157:
2151:
2147:
2143:
2136:
2131:
2129:
2124:
2122:
2117:
2116:
2113:
2107:
2105:
2100:
2098:
2095:
2093:
2090:
2088:
2086:
2082:
2077:
2076:
2064:
2058:
2054:
2050:
2043:
2024:
2020:
2013:
2007:
1989:
1985:
1979:
1972:
1971:
1964:
1956:
1952:
1947:
1942:
1938:
1934:
1930:
1926:
1922:
1915:
1899:
1895:
1891:
1885:
1869:
1865:
1859:
1851:
1847:
1842:
1837:
1833:
1829:
1825:
1821:
1817:
1810:
1802:
1796:
1792:
1785:
1783:
1781:
1779:
1763:
1759:
1753:
1746:
1745:0-697-01372-3
1742:
1738:
1732:
1724:
1720:
1715:
1714:10.1038/36786
1710:
1706:
1702:
1698:
1694:
1690:
1688:
1679:
1671:
1667:
1663:
1659:
1655:
1651:
1647:
1643:
1639:
1635:
1631:
1624:
1616:
1612:
1608:
1604:
1600:
1593:
1585:
1581:
1577:
1573:
1569:
1565:
1558:
1550:
1546:
1542:
1536:
1532:
1528:
1524:
1517:
1502:
1498:
1494:
1487:
1479:
1477:9781292235103
1473:
1469:
1462:
1456:
1455:
1450:
1445:
1437:
1435:9780511807022
1431:
1427:
1423:
1419:
1412:
1404:
1400:
1396:
1392:
1388:
1384:
1380:
1376:
1369:
1353:
1349:
1343:
1341:
1339:
1337:
1335:
1333:
1324:
1320:
1316:
1309:
1301:
1294:
1290:
1281:
1278:
1277:
1268:
1267:
1263:
1261:
1260:
1259:Virgibacillus
1256:
1254:
1253:
1249:
1247:
1246:
1242:
1240:
1239:
1235:
1233:
1232:
1228:
1226:
1225:
1221:
1219:
1218:
1214:
1212:
1211:
1207:
1205:
1204:
1200:
1198:
1197:
1193:
1191:
1190:
1186:
1184:
1183:
1182:Terribacillus
1179:
1177:
1176:
1172:
1170:
1169:
1168:Tenuibacillus
1165:
1163:
1162:
1158:
1156:
1155:
1151:
1149:
1148:
1144:
1142:
1141:
1137:
1135:
1134:
1130:
1128:
1127:
1123:
1121:
1120:
1116:
1114:
1113:
1109:
1107:
1106:
1102:
1100:
1099:
1095:
1093:
1092:
1088:
1086:
1085:
1081:
1079:
1078:
1074:
1072:
1071:
1067:
1065:
1064:
1060:
1058:
1057:
1053:
1051:
1050:
1049:Propionispora
1046:
1044:
1043:
1042:Pontibacillus
1039:
1037:
1036:
1032:
1030:
1029:
1028:Piscibacillus
1025:
1023:
1022:
1021:Pelotomaculum
1018:
1016:
1015:
1011:
1009:
1008:
1004:
1002:
1001:
1000:Paenibacillus
997:
995:
994:
990:
988:
987:
983:
981:
980:
976:
974:
973:
969:
967:
966:
962:
960:
959:
955:
953:
952:
948:
946:
945:
944:Metabacterium
941:
939:
938:
934:
932:
931:
927:
925:
924:
923:Lentibacillus
920:
918:
917:
913:
911:
910:
906:
904:
903:
899:
897:
896:
892:
890:
889:
885:
883:
882:
878:
876:
875:
871:
869:
868:
864:
862:
861:
857:
855:
854:
850:
848:
847:
843:
841:
840:
836:
834:
833:
829:
827:
826:
822:
820:
819:
815:
813:
812:
808:
806:
805:
801:
799:
798:
794:
791:
790:
785:
784:
780:
778:
777:
773:
771:
770:
766:
764:
763:
759:
757:
756:
752:
750:
749:
745:
743:
742:
738:
736:
735:
731:
729:
728:
727:Brevibacillus
724:
722:
721:
717:
715:
714:
713:Anoxybacillus
710:
708:
707:
703:
701:
700:
696:
694:
693:
689:
687:
686:
685:Amphibacillus
682:
680:
679:
675:
673:
672:
668:
666:
665:
661:
659:
658:
654:
653:
651:
643:
640:
634:Biotechnology
631:
627:
624:
619:
617:
613:
612:
606:
604:
600:
596:
592:
588:
587:
583:
579:
569:
567:
566:
561:
555:
553:
548:
544:
542:
538:
534:
521:
518:
514:
511:
510:
509:
505:
502:
497:
495:
494:
488:
486:
482:
478:
474:
469:
464:
462:
458:
454:
450:
446:
442:
438:
434:
430:
425:
422:
418:
417:
409:
405:
396:
387:
385:
381:
380:
375:
371:
370:
364:
352:
349:
346:
343:
342:
341:
339:
335:
334:Moeller stain
329:
326:
323:
318:
316:
312:
308:
304:
300:
296:
292:
288:
287:peptidoglycan
284:
280:
275:
271:
267:
263:
259:
255:
251:
247:
243:
234:
225:
222:
218:
203:
199:
197:
192:
190:
185:
182:
178:
174:
170:
169:disinfectants
166:
162:
157:
155:
151:
147:
142:
140:
139:
134:
133:
128:
127:
122:
121:
116:
115:
110:
109:
104:
100:
96:
95:
89:
85:
81:
77:
73:
69:
65:
61:
53:
52:
46:
39:
38:
33:
28:
22:
2721:Bacteriology
2670:Mendosicutes
2655:Gracilicutes
2635:
2535:Mycolic acid
2525:Mycobacteria
2523:
2495:
2490:
2474:
2410:Coccobacilli
2310:in pregnancy
2276:Extremophile
2260:Aerotolerant
2194:Biochemistry
2156:microbiology
2142:Microbiology
2103:
2084:
2080:
2048:
2042:
2030:. Retrieved
2023:the original
2018:
2006:
1995:. Retrieved
1969:
1963:
1928:
1924:
1914:
1902:. Retrieved
1893:
1884:
1872:. Retrieved
1868:the original
1858:
1823:
1819:
1809:
1790:
1766:. Retrieved
1764:. 2016-03-02
1761:
1752:
1737:Microbiology
1736:
1731:
1696:
1692:
1686:
1678:
1637:
1633:
1629:
1623:
1609:(1): 80–81.
1606:
1602:
1598:
1592:
1567:
1563:
1557:
1522:
1516:
1505:. Retrieved
1486:
1467:
1461:
1452:
1444:
1417:
1411:
1378:
1374:
1368:
1356:. Retrieved
1323:the original
1318:
1308:
1299:
1293:
1264:
1257:
1250:
1243:
1236:
1229:
1222:
1215:
1208:
1201:
1194:
1187:
1180:
1175:Tepidibacter
1173:
1166:
1159:
1152:
1145:
1138:
1133:Sporosarcina
1131:
1124:
1117:
1110:
1103:
1096:
1089:
1082:
1075:
1068:
1061:
1054:
1047:
1040:
1033:
1026:
1019:
1012:
1005:
998:
991:
984:
977:
970:
963:
956:
949:
942:
935:
928:
921:
914:
907:
900:
895:Halonatronum
893:
888:Halobacillus
886:
879:
872:
865:
858:
853:Filobacillus
851:
844:
837:
830:
823:
816:
809:
802:
795:
787:
781:
774:
767:
760:
753:
746:
739:
732:
725:
718:
711:
704:
697:
692:Anaerobacter
690:
683:
678:Ammoniphilus
676:
669:
662:
655:
649:
638:
637:
628:
623:B. anthracis
622:
620:
609:
607:
601:subunits of
599:sigma factor
584:
575:
563:
556:
549:
545:
540:
536:
532:
530:
527:Reactivation
516:germination.
506:
498:
491:
489:
465:
426:
413:
410:
407:
383:
377:
367:
365:
361:
350:Spore cortex
330:
325:dipicolinate
319:
298:
290:
282:
278:
273:
269:
245:
239:
217:cryptobiotic
213:
193:
176:
158:
143:
136:
130:
124:
118:
112:
106:
92:
75:
59:
57:
49:
35:
34:of the cell
2558:Slime layer
2238:Facultative
2226:Facultative
1894:www.cdc.gov
1826:(1): 7–10.
1570:: 555–588.
1449:"endospore"
1358:October 21,
1098:Sporobacter
1077:Shimazuella
986:Oxalophagus
958:Natroniella
909:Heliophilum
867:Geobacillus
762:Clostridium
748:Caminicella
741:Caloramator
699:Anaerospora
664:Actinomyces
537:germination
468:autoclaving
279:B. subtilis
274:B. subtilis
266:germination
165:desiccation
99:environment
2665:Mollicutes
2660:Firmicutes
2650:Prokaryota
2568:Glycocalyx
2393:plasticity
2356:Lipophilic
2209:preference
2184:Resistance
1997:2013-08-22
1904:11 October
1768:2019-12-30
1507:2011-08-24
1315:"Bacteria"
1286:References
1140:Sporotalea
1070:Seinonella
1035:Planifilum
846:Filifactor
597:, and the
572:Importance
533:activation
501:endoscopes
485:gamma rays
479:, such as
449:alkylating
445:detergents
347:Spore coat
344:Exosporium
313:and other
295:protoplast
246:spore coat
242:exosporium
2618:evolution
2592:Composite
2491:Endospore
2449:Cell wall
2425:Structure
2316:Placental
2255:Nanaerobe
2233:Anaerobic
2164:Infection
1126:Sporomusa
1014:Pelospora
993:Oxobacter
916:Laceyella
657:Acetonema
541:outgrowth
508:factors:
353:Core wall
311:ribosomes
303:chromatin
291:core wall
258:molecules
228:Structure
150:ribosomes
72:Bacillota
60:endospore
2715:Category
2690:Category
2613:Taxonomy
2546:envelope
2436:envelope
2326:Salivary
2243:Obligate
2221:Obligate
2169:Exotoxin
2146:Bacteria
2104:bacillus
2032:June 18,
1988:Archived
1955:14203345
1898:Archived
1874:June 18,
1850:20967138
1584:18035610
1549:46703605
1501:Archived
1352:Archived
1274:See also
951:Moorella
783:Coxiella
776:Cohnella
720:Bacillus
552:cysteine
437:alcohols
358:Location
254:lysozyme
88:bacteria
68:bacteria
2700:Commons
2599:Biofilm
2578:Fimbria
2563:S-layer
2544:Outside
2405:Bacilli
2321:Uterine
2306:Vaginal
2216:Aerobic
2199:ecology
2154:Medical
1841:2957119
1723:9384377
1701:Bibcode
1670:4160084
1662:4963432
1642:Bibcode
1403:7538699
1383:Bibcode
1375:Science
937:Mahella
457:anthrax
374:tetanus
322:calcium
315:enzymes
262:enzymes
138:Archaea
103:species
64:dormant
2645:Monera
2415:Spiral
2207:Oxygen
2059:
1980:
1953:
1946:277301
1943:
1848:
1838:
1797:
1743:
1721:
1693:Nature
1668:
1660:
1634:Nature
1582:
1547:
1537:
1474:
1432:
1401:
972:Orenia
860:Gelria
786:(i.e.
539:, and
481:x-rays
416:septum
414:spore
289:. The
283:cortex
129:, and
2573:Pilus
2527:only:
2507:Porin
2499:only:
2478:only:
2400:Cocci
2376:Shape
2296:Mouth
2269:Other
2026:(PDF)
2015:(PDF)
1991:(PDF)
1974:(PDF)
1666:S2CID
1545:S2CID
268:. In
250:sieve
80:spore
62:is a
2616:and
2434:Cell
2301:Skin
2291:Lung
2197:and
2057:ISBN
2034:2016
1978:ISBN
1951:PMID
1906:2019
1876:2016
1846:PMID
1795:ISBN
1741:ISBN
1719:PMID
1658:PMID
1580:PMID
1535:ISBN
1472:ISBN
1430:ISBN
1399:PMID
1360:2018
483:and
443:and
299:core
76:endo
2468:DAP
2463:NAG
2458:NAM
2286:Gut
1941:PMC
1933:doi
1836:PMC
1828:doi
1709:doi
1697:390
1650:doi
1638:215
1632:".
1611:doi
1601:".
1572:doi
1527:doi
1497:BBC
1451:at
1422:doi
1391:doi
1379:268
297:or
146:DNA
58:An
30:An
2717::
2638::
2451::
2144::
2055:.
2051:.
2017:.
1986:.
1949:.
1939:.
1929:88
1927:.
1923:.
1892:.
1844:.
1834:.
1824:14
1822:.
1818:.
1777:^
1760:.
1717:.
1707:.
1695:.
1691:.
1664:.
1656:.
1648:.
1636:.
1607:29
1605:.
1578:.
1568:61
1566:.
1543:.
1533:.
1525:.
1499:.
1495:.
1428:.
1397:.
1389:.
1377:.
1331:^
605:.
593:,
535:,
439:,
386:.
307:UV
163:,
148:,
141:.
123:,
117:,
111:,
2312:)
2308:(
2134:e
2127:t
2120:v
2087:)
2065:.
2036:.
2000:.
1957:.
1935::
1908:.
1878:.
1852:.
1830::
1803:.
1771:.
1747:.
1725:.
1711::
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1672:.
1652::
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1617:.
1613::
1586:.
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1551:.
1529::
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1438:.
1424::
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1393::
1385::
1362:.
792:)
23:.
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