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Chemostat

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concentration of the limiting nutrient. In turn, this will reduce the specific growth rate of the cells, which will lead to a decline in the number of cells as they keep being removed from the system with the outflow. This results in a steady state. Due to self-regulation, the steady state is stable. This enables the experimenter to control the specific growth rate of the microorganisms by changing the speed of the pump feeding fresh medium into the vessel.
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Chemostats in research are used for investigations in cell biology, as a source for large volumes of uniform cells or protein. The chemostat is often used to gather steady state data about an organism in order to generate a mathematical model relating to its metabolic processes. Chemostats are also
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Even though the limiting substrate concentration in the chemostat is usually very low, and is maintained by discrete highly concentrated influx pulses, in practice the temporal variation in the concentration within the chemostat is small (a few percent or less) and can thus be viewed as quasi-steady
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Enclosed chemostat vessel with a continuous and adjustable inflow of medium and outflow of effluent, used for controlled growth of microorganisms. The system maintains a constant volume and level of aeration. The growth rate of the microorganism is controlled by manipulation of the inflow of fresh
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However, since the concentration of the limiting nutrient in the chemostat cannot exceed the concentration in the feed, the specific growth rate that the cells can reach in the chemostat is usually slightly lower than the maximal specific growth rate because specific growth rate usually increases
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and all culture parameters remain constant (culture volume, dissolved oxygen concentration, nutrient and product concentrations, pH, cell density, etc.). In addition, environmental conditions can be controlled by the experimenter. Microorganisms growing in chemostats usually reach a steady state
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These other mutations are expected only through successive sweeps of mutants with a fitness advantage. One can only expect multiple mutants to arise if each mutation is independently beneficial, and not in cases where the mutations are individually neutral but together advantageous. Successive
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Another important feature of chemostats and other continuous culture systems is that they are well-mixed so that environmental conditions are homogenous or uniform and microorganisms are randomly dispersed and encounter each other randomly. Therefore, competition and other interactions in the
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with the inflowing fresh medium. The limiting nutrient is a nutrient essential for growth, present in the medium at a limiting concentration (all other nutrients are usually supplied in surplus). However, the higher the number of cells becomes, the more nutrient is consumed, lowering the
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to which fresh medium is continuously added, while culture liquid containing left over nutrients, metabolic end products and microorganisms is continuously removed at the same rate to keep the culture volume constant. By changing the rate with which medium is added to the bioreactor the
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When the conditions are right (a large enough population, and multiple targets in the genome for simple advantageous mutations) multiple strains are expected to successively takeover the population, and to do so in a relatively timed and paced manner. The timing depends on the type of
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A strain that requires two specific SNPs where only their combination gives a fitness advantage (whereas each one separately is neutral), is likely to appear only if the target size (the number of different SNP locations that give rise to an advantageous mutation) for each SNP is very
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There are cases where successive takeovers occur so rapidly that it is very difficult to differentiate between strains, even when examining allele frequency. Thus, a lineage of multiple takeovers of consecutive strains might appear as the takeover of a single strain with a cohort of
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The limiting substrate is usually at such low concentrations that it is undetectable. As a result, the concentration of the limiting substrate can vary greatly over time (percentage-wise) as different strains takeover the population, even if resulting changes in OD are too small to
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because of a negative feedback between growth rate and nutrient consumption: if a low number of cells are present in the bioreactor, the cells can grow at growth rates higher than the dilution rate as they consume little nutrient so growth is less limited by the addition of
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In a takeover succession, even if the selective improvement of each of the strains stays constant (e.g. each new strain is better than the previous strain by a constant factor) – the takeover rate does not stay constant, but rather diminishes from strain to
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The time it takes for the cell density (OD) to converge to a steady-state value (overshoot/undershoot) will often be long (multiple chemostat turnovers), especially when the initial inoculum is large. But, the time can be minimized with proper parameter
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For characteristically large chemostats, a hyper-mutating strain does not give enough of an advantage to warrant use. Also, it does not have enough of a selective advantage to be expected to always appear through random mutation and take over the
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By abruptly lowering the influx limiting substrate concentration it is possible to temporarily subject the cells to relatively harsher conditions, until the chemostat stabilizes back to the steady state (on the time order of the dilution rate
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medium, while the population density is regulated through changing the concentration of the limiting nutrient. This open system allows researchers to maintain the exponential growth phase of cells for use in physiological experiments.
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occurs to be on the order of the chemostat turnover time. Note, this is usually substantially shorter than the time for an advantageous strain to take over the chemostat population. This is not necessarily so in a small
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Competition for single and multiple resources, the evolution of resource acquisition and utilization pathways, cross-feeding/symbiosis, antagonism, predation, and competition among predators have all been studied in
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Van Hulle SW, Van Den Broeck S, Maertens J, Villez K, Schelstraete G, Volcke EI, Vanrolleghem PA (2003). "Practical experiences with start-up and operation of a continuously aerated lab-scale SHARON reactor".
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A “pulsed” chemostat (with very large influx pulses) has a substantially lower selective capacity than a standard quasi-continuous chemostat, for a mutant strain with increased fitness in limiting conditions.
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A chemostat might appear to be in steady state, but mutant strain takeovers can occur continuously, even though they are not detectable by monitoring macro scale parameters like OD or product concentrations.
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The steady state concentration of the limiting substrate in the chemostat is independent of the influx concentration. The influx concentration will affect the cell concentration and thus the steady state
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to render them hydrophobic. However, cells will be selected for attachment to the walls since those that do will not be removed from the system. Those bacteria that stick firmly to the walls forming a
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The seemingly extreme scenario where we require every possible single SNP to co-exist at least once in the chemostat is actually quite likely. A large chemostat is very likely to reach this state.
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Bacteria travel upstream quite easily. They will reach the reservoir of sterile medium quickly unless the liquid path is interrupted by an air break in which the medium falls in drops through air.
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de Crécy E, Metzgar D, Allen C, Pénicaud M, Lyons B, Hansen CJ, de Crécy-Lagard V (2007). "Development of a novel continuous culture device for experimental evolution of bacterial populations".
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The time for takeover of a superior mutant will be quite constant across a range of operation parameters. For characteristic operation values the take over time is on the order of days to weeks.
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A slow dilution rate creates a selection pressure for a mutant strain which can grow in media with no limiting substrate (presumably by consuming a different substrate present in the media);
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Dripping the media into the chamber actually results in small pulses of nutrients and thus oscillations in concentrations, again upsetting the "static" property of the chemostat.
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in ecology and evolutionary biology. In the one case, mutation/selection is a nuisance, in the other case, it is the desired process under study. Chemostats can also be used to
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Zhang Z, Boccazzi P, Choi HG, Perozziello G, Sinskey AJ, Jensen KF (2006). "Microchemostat-microbial continuous culture in a polymer-based, instrumented microbioreactor".
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for further scientific study. Variations in the dilution rate permit the study of the metabolic strategies pursued by the organisms at different growth rates.
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that can increase fitness it should appear in the population after only few chemostat doublings, for characteristically large chemostats (e.g. 10^11
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Different dilution rates selectively favor different mutant strains to take over the chemostat population, if such a strain exists. For example:
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Continuous efforts to remedy each defect lead to variations on the basic chemostat quite regularly. Examples in the literature are numerous.
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Other types of mutant strains (e.g. two SNPs with a small target size, more SNPs or in smaller chemostats) are highly unlikely to appear.
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Cells may grow on the walls or adhere to other surfaces, which may be overcome by treating the glass walls of the vessel with a
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A mid-range dilution rate creates a selection pressure for a mutant strain with a higher affinity to the limiting substrate;
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Becks L, Hilker FM, Malchow H, JĂĽrgens K, Arndt H (2005). "Experimental demonstration of chaos in a microbial food web".
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Wides A, Milo R (2018). "Understanding the Dynamics and Optimizing the Performance of Chemostat Selection Experiments".
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Furthermore, the time until mutation appearance is independent of genome size, but dependent on per-BP mutation rate.
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https://web.archive.org/web/20060504172359/http://www.rpi.edu/dept/chem-eng/Biotech-Environ/Contin/chemosta.htm
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A fast dilution rate creates a selection pressure for a mutant strain with a raised maximal growth rate;
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One of the most important features of chemostats is that microorganisms can be grown in a physiological
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The above points are expected to be the same across different asexually reproductive species (
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Dykhuizen DE, Hartl DL (May 1981). "Evolution of Competitive Ability in Escherichia coli".
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is inversely related to the time it takes the biomass to double, called doubling time 
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under constant environmental conditions. In this steady state, growth occurs at a constant
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Pfeiffer T, Bonhoeffer S (2004). "Evolution of cross-feeding in microbial populations".
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Each microorganism growing on a particular substrate has a maximal specific growth rate
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Jones LE, Ellner SP (2007). "Effects of rapid prey evolution on predator-prey cycles".
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Comprehensive chemostat manual (Dunham lab). Procedures and principles are general.
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A final thesis including mathematical models of the chemostat and other bioreactors
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Mixing may not truly be uniform, upsetting the "static" property of the chemostat.
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Dripping can be made less drastic with smaller droplets and larger vessel volumes
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Bonomi A, Fredrickson AG (1976). "Protozoan feeding and bacterial wall growth".
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takeovers are the only reliable way for evolution to proceed in a chemostat.
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Fernandez-de-Cossio-Diaz, Jorge; Leon, Kalet; Mulet, Roberto (2017-11-13).
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Foaming results in overflow with the volume of liquid not exactly constant.
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Various applications use paddles, bubbling, or other mechanisms for mixing
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is the half-saturation constant (this equation assumes Monod kinetics).
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The takeover time is predictable given the relevant strain parameters.
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The rate of nutrient exchange is expressed as the dilution rate 
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A chemostat diagram featuring inflow (feed) and outflow (effluent).
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Chemostats are frequently used in the industrial manufacturing of
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is the substrate or nutrient concentration in the chemostat and
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with nutrient concentration as described by the kinetics of the
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Fermentation setups closely related to the chemostats are the
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For a large chemostat the expected time until an advantageous
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Novick A, Szilard L (1950). "Description of the Chemostat".
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http://www.pererikstrandberg.se/examensarbete/chemostat.pdf
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for specific types of bacterial mutants in culture such as
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James TW (1961). "Continuous Culture of Microorganisms".
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of the micro-organism is equal to the dilution rate 
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Schlegel HG, Jannasch HW (1967). "Enrichment cultures".
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Many approaches have been taken to reduce wall growth
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Many improvements target the threat of contamination
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The highest specific growth' rates ( 301:becomes a function of dilution rate  3312:Instruments used in medical laboratories 2793: 2001: 1580: 1385:Varma, A.; Palsson, B. O. (1994-10-01). 999: 627:Relevant discussion may be found on the 349:{\displaystyle t_{d}={\frac {\ln 2}{D}}} 98:can be easily controlled within limits. 63: 2828:Inductively coupled plasma (ICP) device 1917: 1915: 1913: 1911: 1909: 1842: 1798: 1754: 1151: 931: 840: 3323: 2876:Transmission electron microscope (TEM) 1391:Applied and Environmental Microbiology 1290: 1256:Wick LM, Weilenmann H, Egli T (2002). 1112: 908:E. coli long-term evolution experiment 903:Continuous stirred-tank reactor (CSTR) 721: 3153: 2972: 2792: 2452: 2013: 1975: 1448: 1446: 1061: 536: 134:chemostat are global, in contrast to 1906: 1669:Dykhuizen DE, Hartl DL (June 1983). 1249: 604: 1364:10.1146/annurev.mi.21.100167.000405 1014:10.1146/annurev.mi.15.100161.000331 13: 2871:Scanning electron microscope (SEM) 1443: 811: 601:Experimental design considerations 14: 3342: 1940: 2916:Thermogravimetric analyzer (TGA) 2726:Nuclear magnetic resonance (NMR) 1411:10.1128/aem.60.10.3724-3731.1994 620:relies largely or entirely on a 609: 141: 27: 2973: 1888:Commun. Agric. Appl. Biol. Sci. 1711: 1662: 1623: 1378: 1343: 934:Brock Biology of Microorganisms 501:or those that are resistant to 478: 106: 1530:Daughton CG, Hsieh DP (1977). 1207:Dykhuizen DE, Dean AM (2004). 925: 1: 2453: 1556:10.1128/aem.34.2.175-184.1977 1002:Annual Review of Microbiology 936:. Pearson. pp. 152–153. 918: 861: 292:Therefore, the doubling time 170:of culture in the bioreactor 128: 1808:Appl. Microbiol. Biotechnol. 1687:10.1128/mr.47.2.150-168.1983 1490:10.1371/journal.pcbi.1005835 1137:10.1016/0025-5564(92)90002-E 979:10.1126/science.112.2920.715 101: 7: 2014: 1275:10.1099/00221287-148-9-2889 1225:10.1534/genetics.103.025205 1176:10.1534/genetics.104.034488 881: 744: 524: 483: 10: 3347: 3154: 1459:PLOS Computational Biology 701:Parameter choice and setup 227:Specific growth rate  3309: 3260: 3210: 3177: 3164: 3160: 3149: 3120: 3089: 3071:Time-domain reflectometer 3033: 2985: 2981: 2968: 2924: 2884: 2861: 2838:Liquid chromatograph (LC) 2805: 2801: 2788: 2749: 2706: 2655: 2624: 2561: 2530: 2507: 2491: 2465: 2461: 2448: 2331: 2265: 2177: 2102: 2024: 2020: 2009: 1820:10.1007/s00253-007-1168-5 1671:"Selection in chemostats" 1321:10.1007/s00285-007-0094-6 1046:10.1099/00221287-14-3-601 932:Madigan, Michael (2015). 48: 38: 26: 21: 1536:Appl. Environ. Microbiol 2906:Melting-point apparatus 2287:Cryogenic storage dewar 893:Biochemical engineering 166:, over the volume  150:. At steady state, the 2843:Mass spectrometer (MS) 2833:Gas chromatograph (GC) 453: 350: 283: 213: 70: 43:Biological engineering 3218:Acid-resistant gloves 2899:differential scanning 1776:10.1002/bit.260180209 454: 351: 284: 214: 67: 2795:Analytical chemistry 2297:Laminar flow cabinet 2003:Laboratory equipment 1352:Annu. Rev. Microbiol 841:Successive takeovers 633:improve this article 519:evolutionary biology 402: 312: 245: 177: 152:specific growth rate 117:specific growth rate 92:specific growth rate 81:ical environment is 16:A type of bioreactor 3167:Personal protective 2076:Meker–Fisher burner 1764:Biotechnol. Bioeng. 1548:1977ApEnM..34..175D 1481:2017PLSCB..13E5835F 1403:1994ApEnM..60.3724V 1090:10.1038/nature03627 1082:2005Natur.435.1226B 971:1950Sci...112..715N 722:Steady state growth 3010:Function generator 2993:Bench power supply 2932:Analytical balance 2693:Ostwald viscometer 2688:Graduated cylinder 2427:Inoculation needle 1648:10.1007/BF00275997 1632:G. S. K. Wolkowicz 1268:(Pt 9): 2889–902. 537:Technical concerns 521:using chemostats. 449: 346: 279: 209: 71: 3318: 3317: 3305: 3304: 3301: 3300: 3278:Fire extinguisher 3268:Biosafety cabinet 3256: 3255: 3145: 3144: 3141: 3140: 3076:Transistor tester 3066:Spectrum analyzer 2964: 2963: 2960: 2959: 2784: 2783: 2780: 2779: 2656:Measuring devices 2478:Soxhlet extractor 2444: 2443: 2440: 2439: 2392:Spectrophotometer 2387:Pipeclay triangle 2139:Mortar and pestle 1894:(2 Pt A): 77–84. 1397:(10): 3724–3731. 1033:J. Gen. Microbiol 943:978-0-321-89739-8 698: 697: 683: 444: 344: 305:in steady state: 277: 207: 194: 193: 190: 122:limiting nutrient 62: 61: 3338: 3175: 3174: 3162: 3161: 3151: 3150: 3056:Network analyzer 2983: 2982: 2970: 2969: 2803: 2802: 2790: 2789: 2463: 2462: 2450: 2449: 2432:Inoculation loop 2302:Microtiter plate 2242:Test tube holder 2134:Magnetic stirrer 2022: 2021: 2011: 2010: 1996: 1989: 1982: 1973: 1972: 1934: 1933: 1931: 1919: 1904: 1903: 1881: 1875: 1874: 1863:10.1039/b518396k 1846: 1840: 1839: 1802: 1796: 1795: 1758: 1752: 1751: 1715: 1709: 1708: 1698: 1666: 1660: 1659: 1627: 1621: 1620: 1584: 1578: 1577: 1567: 1527: 1521: 1520: 1510: 1492: 1474: 1465:(11): e1005835. 1450: 1441: 1440: 1430: 1382: 1376: 1375: 1347: 1341: 1340: 1314: 1294: 1288: 1287: 1277: 1253: 1247: 1246: 1236: 1204: 1198: 1197: 1187: 1155: 1149: 1148: 1116: 1110: 1109: 1076:(7046): 1226–9. 1065: 1059: 1058: 1048: 1024: 1018: 1017: 997: 991: 990: 954: 948: 947: 929: 888:Bacterial growth 693: 690: 684: 682: 641: 613: 605: 458: 456: 455: 450: 445: 443: 436: 435: 422: 420: 419: 355: 353: 352: 347: 345: 340: 329: 324: 323: 288: 286: 285: 280: 278: 276: 275: 266: 255: 218: 216: 215: 210: 208: 200: 195: 191: 189:medium flow rate 188: 187: 31: 19: 18: 3346: 3345: 3341: 3340: 3339: 3337: 3336: 3335: 3321: 3320: 3319: 3314: 3297: 3293:Solvent cabinet 3252: 3223:Eyewash station 3206: 3171: 3169:equipment (PPE) 3168: 3156: 3137: 3116: 3085: 3029: 3020:Pulse generator 2986:Control devices 2977: 2956: 2920: 2886:Thermochemistry 2880: 2857: 2797: 2776: 2745: 2702: 2668:Conical measure 2651: 2620: 2557: 2526: 2503: 2487: 2457: 2436: 2417:Test tube brush 2327: 2312:Picotiter plate 2275: 2261: 2257:Lab drying rack 2216:Extension clamp 2192: 2173: 2112: 2098: 2034: 2016: 2005: 2000: 1943: 1938: 1937: 1920: 1907: 1882: 1878: 1847: 1843: 1803: 1799: 1759: 1755: 1732:10.2307/2408204 1716: 1712: 1667: 1663: 1628: 1624: 1585: 1581: 1528: 1524: 1451: 1444: 1383: 1379: 1348: 1344: 1295: 1291: 1254: 1250: 1205: 1201: 1156: 1152: 1117: 1113: 1066: 1062: 1025: 1021: 998: 994: 965:(2920): 715–6. 955: 951: 944: 930: 926: 921: 884: 864: 843: 814: 812:Single takeover 747: 724: 703: 694: 688: 685: 642: 640: 626: 614: 603: 539: 527: 486: 481: 474: 431: 427: 426: 421: 415: 411: 403: 400: 399: 393: 387: 372: 365: 330: 328: 319: 315: 313: 310: 309: 300: 271: 267: 256: 254: 246: 243: 242: 237: 225: 199: 186: 178: 175: 174: 144: 131: 109: 104: 34: 17: 12: 11: 5: 3344: 3334: 3333: 3316: 3315: 3310: 3307: 3306: 3303: 3302: 3299: 3298: 3296: 3295: 3290: 3288:Safety cabinet 3285: 3280: 3275: 3270: 3264: 3262: 3258: 3257: 3254: 3253: 3251: 3250: 3248:Safety goggles 3245: 3243:Safety glasses 3240: 3238:Nitrile gloves 3235: 3233:Medical gloves 3230: 3225: 3220: 3214: 3212: 3208: 3207: 3205: 3204: 3199: 3194: 3189: 3184: 3178: 3172: 3165: 3158: 3157: 3147: 3146: 3143: 3142: 3139: 3138: 3136: 3135: 3130: 3128:Alligator clip 3124: 3122: 3118: 3117: 3115: 3114: 3109: 3104: 3102:Soldering iron 3099: 3093: 3091: 3087: 3086: 3084: 3083: 3078: 3073: 3068: 3063: 3058: 3053: 3048: 3046:Logic analyzer 3043: 3037: 3035: 3031: 3030: 3028: 3027: 3022: 3017: 3012: 3007: 3006: 3005: 3003:Voltage source 3000: 2998:Current source 2989: 2987: 2979: 2978: 2966: 2965: 2962: 2961: 2958: 2957: 2955: 2954: 2949: 2944: 2939: 2937:Colony counter 2934: 2928: 2926: 2922: 2921: 2919: 2918: 2913: 2908: 2903: 2902: 2901: 2890: 2888: 2882: 2881: 2879: 2878: 2873: 2867: 2865: 2859: 2858: 2856: 2855: 2850: 2845: 2840: 2835: 2830: 2825: 2820: 2815: 2809: 2807: 2799: 2798: 2786: 2785: 2782: 2781: 2778: 2777: 2775: 2774: 2769: 2764: 2759: 2753: 2751: 2747: 2746: 2744: 2743: 2738: 2733: 2728: 2723: 2718: 2712: 2710: 2704: 2703: 2701: 2700: 2695: 2690: 2685: 2680: 2675: 2670: 2665: 2659: 2657: 2653: 2652: 2650: 2649: 2644: 2639: 2634: 2628: 2626: 2622: 2621: 2619: 2618: 2613: 2608: 2603: 2598: 2593: 2588: 2583: 2578: 2576:Vacuum (Dewar) 2573: 2567: 2565: 2559: 2558: 2556: 2555: 2550: 2545: 2540: 2534: 2532: 2528: 2527: 2525: 2524: 2519: 2513: 2511: 2505: 2504: 2502: 2501: 2495: 2493: 2489: 2488: 2486: 2485: 2480: 2475: 2469: 2467: 2459: 2458: 2446: 2445: 2442: 2441: 2438: 2437: 2435: 2434: 2429: 2424: 2419: 2414: 2409: 2404: 2399: 2394: 2389: 2384: 2379: 2374: 2369: 2364: 2359: 2354: 2349: 2346: 2341: 2335: 2333: 2329: 2328: 2326: 2325: 2322: 2319: 2314: 2309: 2304: 2299: 2294: 2289: 2284: 2278: 2276: 2274: 2273: 2270: 2266: 2263: 2262: 2260: 2259: 2254: 2249: 2247:Test tube rack 2244: 2239: 2236: 2231: 2228: 2223: 2222:Funnel support 2220: 2217: 2214: 2209: 2204: 2199: 2195: 2193: 2191: 2190: 2187: 2182: 2178: 2175: 2174: 2172: 2171: 2166: 2161: 2156: 2151: 2146: 2141: 2136: 2131: 2129:Liquid whistle 2126: 2121: 2115: 2113: 2111: 2110: 2107: 2103: 2100: 2099: 2097: 2096: 2094:Vacuum dry box 2091: 2086: 2081: 2078: 2073: 2068: 2063: 2058: 2056:Heating mantle 2053: 2048: 2043: 2041:Alcohol burner 2037: 2035: 2033: 2032: 2029: 2025: 2018: 2017: 2007: 2006: 1999: 1998: 1991: 1984: 1976: 1970: 1969: 1964: 1959: 1954: 1949: 1942: 1941:External links 1939: 1936: 1935: 1905: 1876: 1841: 1797: 1753: 1710: 1675:Microbiol. Rev 1661: 1630:G. J. Butler; 1622: 1601:10.1086/383593 1595:(6): E126–35. 1579: 1522: 1442: 1377: 1342: 1289: 1248: 1219:(4): 2015–26. 1199: 1150: 1111: 1060: 1039:(3): 601–622. 1019: 992: 949: 942: 923: 922: 920: 917: 916: 915: 910: 905: 900: 895: 890: 883: 880: 863: 860: 859: 858: 854: 850: 842: 839: 838: 837: 834: 833: 832: 829: 826: 820: 813: 810: 809: 808: 804: 801: 790: 782: 779: 778: 777: 770: 769: 768: 764: 755:If there is a 746: 743: 742: 741: 737: 734: 730: 723: 720: 719: 718: 714: 710: 702: 699: 696: 695: 631:. Please help 617: 615: 608: 602: 599: 598: 597: 594: 591: 588: 585: 582: 575: 574: 571: 568: 565: 554: 543: 538: 535: 526: 523: 507:bacteriophages 485: 482: 480: 477: 470: 460: 459: 448: 442: 439: 434: 430: 425: 418: 414: 410: 407: 391: 385: 379:Monod equation 370: 363: 357: 356: 343: 339: 336: 333: 327: 322: 318: 296: 290: 289: 274: 270: 265: 262: 259: 253: 250: 235: 224: 221: 220: 219: 206: 203: 198: 192:culture volume 185: 182: 143: 140: 130: 127: 108: 105: 103: 100: 60: 59: 50: 46: 45: 40: 36: 35: 32: 24: 23: 15: 9: 6: 4: 3: 2: 3343: 3332: 3329: 3328: 3326: 3313: 3308: 3294: 3291: 3289: 3286: 3284: 3281: 3279: 3276: 3274: 3271: 3269: 3266: 3265: 3263: 3259: 3249: 3246: 3244: 3241: 3239: 3236: 3234: 3231: 3229: 3226: 3224: 3221: 3219: 3216: 3215: 3213: 3209: 3203: 3202:Safety shower 3200: 3198: 3195: 3193: 3190: 3188: 3185: 3183: 3180: 3179: 3176: 3173: 3170: 3163: 3159: 3152: 3148: 3134: 3131: 3129: 3126: 3125: 3123: 3119: 3113: 3112:Wire stripper 3110: 3108: 3105: 3103: 3100: 3098: 3095: 3094: 3092: 3088: 3082: 3079: 3077: 3074: 3072: 3069: 3067: 3064: 3062: 3059: 3057: 3054: 3052: 3049: 3047: 3044: 3042: 3039: 3038: 3036: 3032: 3026: 3023: 3021: 3018: 3016: 3013: 3011: 3008: 3004: 3001: 2999: 2996: 2995: 2994: 2991: 2990: 2988: 2984: 2980: 2976: 2971: 2967: 2953: 2950: 2948: 2945: 2943: 2942:Spiral plater 2940: 2938: 2935: 2933: 2930: 2929: 2927: 2923: 2917: 2914: 2912: 2909: 2907: 2904: 2900: 2897: 2896: 2895: 2892: 2891: 2889: 2887: 2883: 2877: 2874: 2872: 2869: 2868: 2866: 2864: 2860: 2854: 2851: 2849: 2846: 2844: 2841: 2839: 2836: 2834: 2831: 2829: 2826: 2824: 2821: 2819: 2816: 2814: 2811: 2810: 2808: 2806:Compositional 2804: 2800: 2796: 2791: 2787: 2773: 2770: 2768: 2765: 2763: 2760: 2758: 2755: 2754: 2752: 2748: 2742: 2739: 2737: 2734: 2732: 2729: 2727: 2724: 2722: 2719: 2717: 2714: 2713: 2711: 2709: 2705: 2699: 2696: 2694: 2691: 2689: 2686: 2684: 2681: 2679: 2676: 2674: 2671: 2669: 2666: 2664: 2661: 2660: 2658: 2654: 2648: 2645: 2643: 2640: 2638: 2635: 2633: 2630: 2629: 2627: 2623: 2617: 2614: 2612: 2609: 2607: 2604: 2602: 2599: 2597: 2594: 2592: 2589: 2587: 2584: 2582: 2579: 2577: 2574: 2572: 2569: 2568: 2566: 2564: 2560: 2554: 2551: 2549: 2546: 2544: 2541: 2539: 2536: 2535: 2533: 2529: 2523: 2520: 2518: 2515: 2514: 2512: 2510: 2506: 2500: 2497: 2496: 2494: 2490: 2484: 2481: 2479: 2476: 2474: 2471: 2470: 2468: 2464: 2460: 2456: 2451: 2447: 2433: 2430: 2428: 2425: 2423: 2420: 2418: 2415: 2413: 2410: 2408: 2405: 2403: 2400: 2398: 2395: 2393: 2390: 2388: 2385: 2383: 2380: 2378: 2375: 2373: 2370: 2368: 2365: 2363: 2360: 2358: 2355: 2353: 2350: 2348:Balance brush 2347: 2345: 2342: 2340: 2337: 2336: 2334: 2330: 2324:Weighing dish 2323: 2321:Weighing boat 2320: 2318: 2315: 2313: 2310: 2308: 2305: 2303: 2300: 2298: 2295: 2293: 2290: 2288: 2285: 2283: 2280: 2279: 2277: 2271: 2268: 2267: 2264: 2258: 2255: 2253: 2250: 2248: 2245: 2243: 2240: 2237: 2235: 2232: 2229: 2227: 2224: 2221: 2218: 2215: 2213: 2212:Burette clamp 2210: 2208: 2205: 2203: 2200: 2197: 2196: 2194: 2188: 2186: 2183: 2180: 2179: 2176: 2170: 2167: 2165: 2162: 2160: 2157: 2155: 2152: 2150: 2147: 2145: 2142: 2140: 2137: 2135: 2132: 2130: 2127: 2125: 2122: 2120: 2117: 2116: 2114: 2108: 2105: 2104: 2101: 2095: 2092: 2090: 2087: 2085: 2082: 2079: 2077: 2074: 2072: 2069: 2067: 2064: 2062: 2059: 2057: 2054: 2052: 2049: 2047: 2046:Bunsen burner 2044: 2042: 2039: 2038: 2036: 2030: 2027: 2026: 2023: 2019: 2012: 2008: 2004: 1997: 1992: 1990: 1985: 1983: 1978: 1977: 1974: 1968: 1965: 1963: 1960: 1958: 1955: 1953: 1950: 1948: 1945: 1944: 1930: 1925: 1918: 1916: 1914: 1912: 1910: 1901: 1897: 1893: 1890: 1889: 1880: 1872: 1868: 1864: 1860: 1857:(7): 906–13. 1856: 1852: 1845: 1837: 1833: 1829: 1825: 1821: 1817: 1814:(2): 489–96. 1813: 1810: 1809: 1801: 1793: 1789: 1785: 1781: 1777: 1773: 1770:(2): 239–52. 1769: 1766: 1765: 1757: 1749: 1745: 1741: 1737: 1733: 1729: 1726:(3): 581–94. 1725: 1721: 1714: 1706: 1702: 1697: 1692: 1688: 1684: 1681:(2): 150–68. 1680: 1676: 1672: 1665: 1657: 1653: 1649: 1645: 1642:(2): 67–191. 1641: 1637: 1633: 1626: 1618: 1614: 1610: 1606: 1602: 1598: 1594: 1590: 1583: 1575: 1571: 1566: 1561: 1557: 1553: 1549: 1545: 1542:(2): 175–84. 1541: 1537: 1533: 1526: 1518: 1514: 1509: 1504: 1500: 1496: 1491: 1486: 1482: 1478: 1473: 1468: 1464: 1460: 1456: 1449: 1447: 1438: 1434: 1429: 1424: 1420: 1416: 1412: 1408: 1404: 1400: 1396: 1392: 1388: 1381: 1373: 1369: 1365: 1361: 1357: 1353: 1346: 1338: 1334: 1330: 1326: 1322: 1318: 1313: 1312:q-bio/0609032 1308: 1305:(4): 541–73. 1304: 1300: 1293: 1285: 1281: 1276: 1271: 1267: 1263: 1259: 1252: 1244: 1240: 1235: 1230: 1226: 1222: 1218: 1214: 1210: 1203: 1195: 1191: 1186: 1181: 1177: 1173: 1169: 1165: 1161: 1154: 1146: 1142: 1138: 1134: 1130: 1126: 1122: 1121:Kevrekidis IG 1115: 1107: 1103: 1099: 1095: 1091: 1087: 1083: 1079: 1075: 1071: 1064: 1056: 1052: 1047: 1042: 1038: 1034: 1030: 1023: 1015: 1011: 1007: 1003: 996: 988: 984: 980: 976: 972: 968: 964: 960: 953: 945: 939: 935: 928: 924: 914: 911: 909: 906: 904: 901: 899: 896: 894: 891: 889: 886: 885: 879: 877: 873: 869: 855: 851: 847: 846: 845: 835: 830: 827: 824: 823: 821: 818: 817: 816: 805: 802: 799: 798:S. cerevisiae 795: 791: 787: 783: 780: 774: 773: 771: 765: 762: 758: 754: 753: 751: 750: 749: 738: 735: 731: 728: 727: 726: 715: 711: 707: 706: 705: 692: 689:December 2019 681: 678: 674: 671: 667: 664: 660: 657: 653: 650: â€“  649: 645: 644:Find sources: 638: 634: 630: 624: 623: 622:single source 618:This section 616: 612: 607: 606: 595: 592: 589: 586: 583: 580: 579: 578: 572: 569: 566: 563: 559: 555: 552: 548: 544: 541: 540: 534: 532: 522: 520: 516: 510: 508: 504: 500: 496: 492: 476: 473: 469: 465: 446: 440: 437: 432: 428: 423: 412: 408: 405: 398: 397: 396: 395: 384: 380: 374: 369: 362: 341: 337: 334: 331: 325: 320: 316: 308: 307: 306: 304: 299: 295: 272: 268: 263: 260: 257: 251: 248: 241: 240: 239: 234: 230: 204: 201: 196: 183: 180: 173: 172: 171: 169: 165: 161: 157: 153: 149: 142:Dilution rate 139: 137: 126: 123: 118: 114: 99: 97: 96:microorganism 93: 88: 84: 80: 76: 66: 58: 54: 51: 47: 44: 41: 37: 30: 25: 20: 3273:Fire blanket 3211:Eye and hand 3197:Rubber apron 3061:Oscilloscope 3025:Potentiostat 2952:Plate reader 2848:pH indicator 2818:CHN analyzer 2813:AutoAnalyzer 2606:Round-bottom 2499:Boston round 2362:Filter paper 2317:Refrigerator 2234:Retort stand 2202:Clamp holder 2198:Beaker clamp 2164:Vortex mixer 2159:Stirring rod 2154:Static mixer 2118: 2084:Teclu burner 1891: 1886: 1879: 1854: 1850: 1844: 1811: 1806: 1800: 1767: 1762: 1756: 1723: 1719: 1713: 1678: 1674: 1664: 1639: 1635: 1625: 1592: 1588: 1582: 1539: 1535: 1525: 1462: 1458: 1394: 1390: 1380: 1355: 1351: 1345: 1302: 1298: 1292: 1265: 1262:Microbiology 1261: 1251: 1216: 1212: 1202: 1170:(1): 19–31. 1167: 1163: 1153: 1128: 1124: 1114: 1073: 1069: 1063: 1036: 1032: 1022: 1005: 1001: 995: 962: 958: 952: 933: 927: 865: 844: 815: 748: 725: 704: 686: 676: 669: 662: 655: 643: 619: 576: 528: 511: 487: 479:Applications 471: 467: 463: 461: 389: 382: 375: 367: 360: 358: 302: 297: 293: 291: 232: 228: 226: 167: 163: 159: 155: 147: 145: 132: 113:steady state 110: 107:Steady state 82: 78: 74: 72: 3331:Bioreactors 3261:Other items 3187:Face shield 3034:Measurement 3015:Galvanostat 2975:Electronics 2925:Other items 2911:Thermometer 2894:Calorimeter 2823:Colorimeter 2767:Gas syringe 2750:Other items 2678:Eye dropper 2553:Watch glass 2538:Evaporating 2517:Cold finger 2332:Other items 2238:Screw clamp 2230:Pinch clamp 2219:Flask clamp 2169:Wash bottle 2124:Homogenizer 1636:J Math Biol 1299:J Math Biol 1131:(1): 1–55. 1125:Math Biosci 876:retentostat 868:turbidostat 648:"Chemostat" 503:antibiotics 49:Application 3192:Respirator 3133:Test probe 3051:Multimeter 2863:Microscopy 2683:Eudiometer 2647:Separatory 2616:Volumetric 2581:Erlenmeyer 2509:Condensers 2473:Dean–Stark 2422:Wire brush 2382:Microscope 2377:Centrifuge 2352:Cork borer 2307:Petri dish 2282:Agar plate 2269:Containers 2252:Wire gauze 2089:Water bath 2051:Desiccator 1929:1806.00272 1472:1705.09708 1119:Pavlou S, 919:References 898:Changestat 862:Variations 857:mutations. 849:mutations. 807:chemostat. 789:chemostat. 659:newspapers 499:auxotrophs 491:microcosms 129:Well-mixed 87:bioreactor 3283:Fume hood 3228:Glove box 3081:Voltmeter 2466:Apparatus 2455:Glassware 2344:Autoclave 2339:Aspirator 2292:Incubator 2226:Iron ring 2149:Sonicator 2119:Chemostat 2061:Hot plate 1720:Evolution 1656:120858390 1499:1553-7358 1419:0099-2240 1358:: 49–70. 1008:: 27–46. 913:Fed-batch 629:talk page 547:agitation 413:μ 335:⁡ 261:⁡ 249:μ 102:Operation 85:ic) is a 75:chemostat 22:Chemostat 3325:Category 3182:Lab coat 3107:Tweezers 3097:Heat gun 2853:pH meter 2762:Bell jar 2642:Dropping 2596:Florence 2586:Fernbach 2548:Syracuse 2407:Scoopula 2357:Crucible 2066:Lab oven 1900:15296140 1871:16804595 1851:Lab Chip 1836:25787277 1828:17896105 1792:41343643 1748:28563589 1617:31110741 1609:15266392 1517:29131817 1337:16927689 1329:17483952 1284:12213934 1243:15342537 1213:Genetics 1194:15687276 1164:Genetics 1098:15988524 1055:13346021 987:14787503 882:See also 874:and the 872:auxostat 800:, etc.). 786:mutation 745:Mutation 551:aeration 525:Industry 489:used as 484:Research 136:biofilms 57:Industry 53:Research 39:Industry 3121:General 3041:Ammeter 2741:Thistle 2698:Pipette 2673:Cuvette 2663:Burette 2632:BĂĽchner 2625:Funnels 2611:Schlenk 2591:Fleaker 2571:BĂĽchner 2492:Bottles 2412:Spatula 2402:Stopper 2372:Forceps 2272:Storage 2189:Holders 2109:Shakers 2080:Striker 2028:Heaters 2015:General 1784:1267931 1740:2408204 1705:6308409 1589:Am. Nat 1544:Bibcode 1508:5703580 1477:Bibcode 1437:7986045 1399:Bibcode 1372:4860267 1234:1470984 1185:1449705 1145:1550993 1106:4380653 1078:Bibcode 967:Bibcode 959:Science 853:strain. 794:E. coli 763:cells). 761:E. coli 733:detect. 717:choice. 673:scholar 562:biofilm 531:ethanol 515:ecology 94:of the 3155:Safety 2757:Beaker 2736:Thiele 2721:Cragie 2716:Drying 2637:Hirsch 2601:Retort 2563:Flasks 2531:Dishes 2522:Liebig 2483:Kipp's 2397:Splint 2207:Tripod 2185:Clamps 2181:Stands 2144:Shaker 2106:Mixers 2031:Dryers 1898:  1869:  1834:  1826:  1790:  1782:  1746:  1738:  1703:  1696:281569 1693:  1654:  1615:  1607:  1574:410368 1572:  1565:242618 1562:  1515:  1505:  1497:  1435:  1428:201879 1425:  1417:  1370:  1335:  1327:  1282:  1241:  1231:  1192:  1182:  1143:  1104:  1096:  1070:Nature 1053:  985:  940:  870:, the 767:large. 713:state. 675:  668:  661:  654:  646:  558:silane 495:enrich 462:where 238:, by: 154:  77:(from 3090:Tools 2708:Tubes 2543:Petri 1924:arXiv 1832:S2CID 1788:S2CID 1736:JSTOR 1652:S2CID 1613:S2CID 1467:arXiv 1333:S2CID 1307:arXiv 1102:S2CID 680:JSTOR 666:books 2772:Vial 2731:Test 2367:File 2071:Kiln 1896:PMID 1867:PMID 1824:PMID 1780:PMID 1744:PMID 1701:PMID 1605:PMID 1570:PMID 1513:PMID 1495:ISSN 1433:PMID 1415:ISSN 1368:PMID 1325:PMID 1280:PMID 1239:PMID 1190:PMID 1141:PMID 1094:PMID 1051:PMID 983:PMID 938:ISBN 652:news 549:and 517:and 83:stat 79:chem 55:and 1859:doi 1816:doi 1772:doi 1728:doi 1691:PMC 1683:doi 1644:doi 1597:doi 1593:163 1560:PMC 1552:doi 1503:PMC 1485:doi 1423:PMC 1407:doi 1360:doi 1317:doi 1270:doi 1266:148 1229:PMC 1221:doi 1217:167 1180:PMC 1172:doi 1168:170 1133:doi 1129:108 1086:doi 1074:435 1041:doi 1010:doi 975:doi 963:112 757:SNP 740:D). 709:OD. 635:by 505:or 417:max 386:max 371:max 364:max 3327:: 1908:^ 1892:68 1865:. 1853:. 1830:. 1822:. 1812:77 1786:. 1778:. 1768:18 1742:. 1734:. 1724:35 1722:. 1699:. 1689:. 1679:47 1677:. 1673:. 1650:. 1640:24 1638:. 1611:. 1603:. 1591:. 1568:. 1558:. 1550:. 1540:34 1538:. 1534:. 1511:. 1501:. 1493:. 1483:. 1475:. 1463:13 1461:. 1457:. 1445:^ 1431:. 1421:. 1413:. 1405:. 1395:60 1393:. 1389:. 1366:. 1356:21 1354:. 1331:. 1323:. 1315:. 1303:55 1301:. 1278:. 1264:. 1260:. 1237:. 1227:. 1215:. 1211:. 1188:. 1178:. 1166:. 1162:. 1139:. 1127:. 1100:. 1092:. 1084:. 1072:. 1049:. 1037:14 1035:. 1031:. 1006:15 1004:. 981:. 973:. 961:. 796:, 394:): 390:D' 332:ln 258:ln 138:. 73:A 1995:e 1988:t 1981:v 1932:. 1926:: 1902:. 1873:. 1861:: 1855:6 1838:. 1818:: 1794:. 1774:: 1750:. 1730:: 1707:. 1685:: 1658:. 1646:: 1619:. 1599:: 1576:. 1554:: 1546:: 1519:. 1487:: 1479:: 1469:: 1439:. 1409:: 1401:: 1374:. 1362:: 1339:. 1319:: 1309:: 1286:. 1272:: 1245:. 1223:: 1196:. 1174:: 1147:. 1135:: 1108:. 1088:: 1080:: 1057:. 1043:: 1016:. 1012:: 989:. 977:: 969:: 946:. 691:) 687:( 677:· 670:· 663:· 656:· 639:. 625:. 553:. 472:S 468:K 464:S 447:, 441:S 438:+ 433:S 429:K 424:S 409:= 406:D 392:c 383:ÎĽ 368:ÎĽ 361:ÎĽ 342:D 338:2 326:= 321:d 317:t 303:D 298:d 294:t 273:d 269:t 264:2 252:= 236:d 233:t 229:ÎĽ 205:V 202:F 197:= 184:= 181:D 168:V 164:F 160:D 156:ÎĽ 148:D

Index

Chemostat schematic
Biological engineering
Research
Industry

bioreactor
specific growth rate
microorganism
steady state
specific growth rate
limiting nutrient
biofilms
specific growth rate
Monod equation
microcosms
enrich
auxotrophs
antibiotics
bacteriophages
ecology
evolutionary biology
ethanol
agitation
aeration
silane
biofilm

single source
talk page
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