470:
secondary male mates are chosen through female choice. Scientists believe multiple paternity has evolved in response to virgin insemination by low quality secondary male mates who have not undergone selection through intrasexual fighting. Females have developed a mechanism for sperm precedence to retain control over offspring paternity and increase offspring fitness. Further examination of female genitalia has supported this hypothesis. The sierra dome spider exhibits this behavior as a form of genetic bet hedging, reducing the risk of producing low quality offspring and contracting venereal disease. This form of bet hedging is notably different than most other forms of bet hedging, as it has not arisen in response to environmental conditions, but rather it has arisen as a result of the species mating system.
412:, where it is thought that individuals are more likely to mate with individuals whose MHC is less similar to their own in order to produce variable offspring. In accordance with the bet hedging model, it has been found that the reproductive success of mating pairs of Atlantic salmon is environmentally dependent, where certain MHC constructs are only advantageous under specific environmental circumstances. Thus, this supports the evidence that MHC diversity is crucial for the long-term reproductive success of the parents, as the tradeoff for an initial decrease in short-term reproductive fitness is mediated by the survival of a few of their offspring in a variable environment.
343:. The daughter cells display either low PHB levels or high PHB levels, which are better suited to short and long-term starvation, respectively. It has been reported that the low-PHB must compete effectively for resources in order to survive, whereas the high-PHB cells can survive for over a year without food. In this example, the PHB phenotype is being ‘bet-hedged’, as the survivability of the offspring largely depends on their environment, where only one phenotype is likely to survive under specific conditions.
512:, variation exists in the distribution of growth rates among yeast micro-colonies and that slow growth is a predictor of resistance to heat. Tsl1 is one gene that was determined as a factor in this resistance. The abundance of this gene was shown to correlate with heat and stress resistance, and thus survival of the yeast micro-colonies under harsh conditions by using bet hedging. This illustrates that by using bet hedging, pathogenic strains of this yeast that are harmful to humans are more difficult to treat.
168:
28:
113:
this strategy actually invest in several different strategies at once, resulting in low variation in long-term success. This could be demonstrated by a clutch of eggs of different sizes, each optimal for one potential environment of the offspring. While this means that offspring specialized for another environment are less likely to survive to adulthood, it also protects against the possibility of no offspring surviving to the next year.
73:
1999:
321:
are able to display this phenomenon quite nicely due to their ability to reproduce quickly enough to track evolution in a single population over a short period of time. This rapid rate of reproduction has allowed for the study of bet hedging in labs through experimental evolution models. These models
291:
The good year specialist has the highest fitness during a good year but does very poorly during a bad year, while the reverse is true for a bad year specialist. The conservative bet hedger does equally well in all years and the diversified bet hedger in this example uses the two specialist strategies
99:
In conservative bet hedging, individuals lower their expected fitness in exchange for a lower variance in fitness. The idea of this strategy is for an organism to "always play it safe" by using the same successful low-risk strategy regardless of environmental conditions. An example of this would be
31:
Variance in egg size is an example of bet-hedging. Fitness may be maximized by producing many, small eggs and thus many offspring. However, larger eggs may help offspring survive stressful conditions. Producing a range of egg sizes can both ensure that some offspring survive stressful conditions, and
538:
The classic example of bet hedging, delayed seed germination, has been extensively studied in desert annuals. One four-year field study found that populations in historically worse (drier) environments had lower germination rates. They also found a large range of germination dates and flexibility in
478:
Bet hedging is employed in fungi similarly to bacteria, but in fungi, it is more complex. This phenomenon is beneficial to fungi, but in some cases, it has harmful effects on humans, illustrating that bet hedging has clinical importance. One study suggests that bet hedging may even contribute to the
429:
cycles have been shown to mediate the timing of torpor and reproduction, and in mice have been shown to mediate this process entirely, heedless to the environment. In the marsupial species, however, an adaptive coin flipping mechanism is employed where neither torpor nor reproduction are affected by
146:
Bet hedging is understood to be a mode of response to environmental change. Adaptations that allow organisms to survive in fluctuating environmental conditions provide an evolutionary advantage. While a bet hedging trait may not be optimal for any one environment, this is outweighed by the benefits
112:
In contrast to conservative bet hedging, diversified bet hedging occurs when individuals lower their expected fitness in a given year while also increasing the variance of survival between offspring. This strategy uses the idea of not "putting all of your eggs in a basket." Individuals implementing
52:
is maximized for that year if all of its seeds germinate. However, if a drought occurs that kills germinated plants, but not ungerminated seeds, plants with seeds remaining in the seed bank will have a fitness advantage. Therefore, it can be advantageous for plants to "hedge their bets" in case of a
449:
an aquatic crustacean species found in many ponds of the
Northeast United States, is one of the most well-studied examples of bet hedging. This species uses a form of diversified bet hedging called germ banking, in which emergence timing among offspring from a single clutch is highly variable. This
163:
A common example used when describing bet hedging is comparing the arithmetic and geometric fitness between specialist and bet hedging genotypes. The table below shows the relative fitness of four phenotypes in 'good' and 'bad' years and their respective means if 'good' years occur 75% of the time
121:
An individual using this type of bet hedging chooses what strategy to use based on a prediction of what the environment will be like. Organisms using this form of bet hedging make these predictions and select strategies annually. For example, an organism may produce clutches of different egg sizes
458:
production of dormant eggs peaks just before the annual dry season in June when ponds levels decrease. In permanent ponds, dormant egg production increases in March, just before an annual increase in feeding activity of sunfish. This example demonstrates that germ banking may take different forms
299:
It is also important to realize that the fitness of any strategy is dependent on a large number of factors, such as the ratio of good to bad years and its relative fitness between good and bad years. Small changes in the strategies or environment having a large impact on which is optimal. In the
295:
In this example, fitness is approximately equal within the specialist and bet hedger strategies, with the bet hedgers having a significantly higher fitness than the specialists. While the good year specialist' has the highest arithmetic mean, the bet hedging strategies are still preferred due to
482:
One way fungi use bet hedging is by displaying different colony morphologies when grown on agar plates. This variation allows for colonies with different morphologies, including resistances that allow them to survive, to thrive and reproduce in different conditions or environments. As a result,
469:
are polyandrous, mating with secondary males in order to compensate for uncertainty regarding the quality of the primary mate. Primary male mates are considered to be of higher fitness than secondary males, as primary mates must overcome intrasexual fighting prior to mating with a female, while
364:
Prokaryotic persistence as a method of bet hedging is thus of importance to the field of medicine due to bacterial persistence. Because bet hedging produces phenotypically diverse offspring randomly in order to survive catastrophic conditions, it is difficult to develop treatments for bacterial
76:
The evolution of an allele that is deleterious in a normal environment (white) but advantageous in an alternate environment (grey). The bet-hedging allele arises twice due to mutation. The first occurrence is lost before the environment changes, but the second mutant reaches fixation due to the
373:
Eukaryotic bet hedging models, unlike prokaryotic models, tend to be used to study more complex evolutionary processes. In the context of eukaryotes, bet hedging is best used as a way to analyze complex environmental influences affecting the selective pressures underlying the principle of bet
546:
Bet hedging through a seed bank has also been implicated in the persistence of weeds. One study of twenty weed species showed that the percentage of viable seeds after 5 years increased with soil depth, and germination rates decreased with soil depth (although specific numbers varied between
534:
Plants provide simple examples for studying bet hedging in wildlife, allowing for field studies but without as many confounding factors as animals. Studying closely related plant species can help us understand more about the circumstances under which bet hedging evolves.
300:
above example, the diversified bet hedger outweighs the conservative bet hedger if it uses the good year specialist strategy more often. In contrast, if the relative fitness of the good year specialist was 0.35 in a bad year, it becomes the optimal strategy.
122:
from year to year, increasing variation in offspring success between clutches. Unlike conservative and diversified bet hedging strategies, adaptive coin flipping is not concerned with minimizing the variation in fitness between years.
130:
To determine if a bet hedging allele is favored, the long-term fitness of each allele must be compared. Particularly in highly variable environments where bet hedging is likely to evolve, long-term fitness is best measured using the
408:(MHC)-dependent mating systems, which have been shown in other species to be important for determining disease resistance among offspring. Namely, there is evidence that selection for increased MHC diversity is a strong influence on
361:, they will produce both cells with normal cell growth and another population of persisters to continue this cycle as the case may be. The ability to switch between the persister and normal phenotype is a form of bet-hedging.
543:. Other studies of desert annuals have also found a relationship between temporal variation and lower germination rates. One of these studies also found the density of seeds in the seed bank to affect germination rates.
356:
exist with the ability to arrest their growth, which leaves them unaffected by dramatic changes to the environment. Once the persister cells grow to form another population of its species, which may or may not be
39:
occurs when organisms suffer decreased fitness in their typical conditions in exchange for increased fitness in stressful conditions. Biological bet hedging was originally proposed to explain the observation of a
525:
with variation in their dormancy because non-dormant ascospores can be killed by heat, but dormant ascospores will survive. The only con is that it will take longer for the dormant ascospores to be germinated.
1317:
McAllan, B. M.; Feay, N.; Bradley, A. J.; Geiser, F. (2012). "The influence of reproductive hormones on the torpor patterns of the marsupial sminthopsis macroura: Bet-hedging in an unpredictable environment".
155:
long enough for alternative environments, in which the bet hedger has an advantage over genotypes adapted to the previous environment, to occur. Over many subsequent environmental alternations, selection may
143:
result in it having an expected geometric mean of zero. This makes it appropriate for circumstances where a single genotype may have variable fitness depending on environmental circumstances.
559:
Collectively, these findings do provide evidence for bet hedging in plants, but also show the importance of competition and phenotypic plasticity that simple bet hedging models often ignore.
313:
Experiments in bet hedging using prokaryotic model organisms provide some of the most simplified views of the evolution of bet hedging. As bet hedging involves a stochastic switching between
550:
In bet-hedging species, seed dormancy appears correlated "to a higher polyphenol (flavonoid) content in seed coats, resulting in darker morphs (Gianella et al., 2021)." In barrel medick (
556:) four flavonoid controlling genes, in addition to peroxidases and thio/peroxiredoxins, "have been associated with differential dormancy along an aridity gradient (Renzi et al., 2020)."
171:
Graph depicting the relative fitness of the strategies shown in the table based on the probability of a bad year. Note that any strategy may be the most fit depending on this value.
454:, germ banking occurs when parents produce dormant eggs prior to annual environmental shifts that yield increased risk for developing offspring. For example, in temporary ponds,
502:
While bet hedging in fungi is important, not much is known about the mechanisms for the different strategies employed by different species. Researchers have studied
1798:
Gremer, Jennifer R.; Venable, D. Lawrence (2014-03-01). "Bet hedging in desert winter annual plants: optimal germination strategies in a variable environment".
459:
within a species depending on the environmental risk presented. Bet hedging through variable egg hatching patterns are seen in other crustaceans as well.
1742:
Philippi, Thomas (1993-09-01). "Bet-Hedging
Germination of Desert Annuals: Variation Among Populations and Maternal Effects in Lepidium lasiocarpum".
151:
tend to be favored in more variable environments. In order for a bet hedging allele to spread, it must persist in the typical environment through
331:
stores carbon and energy in a compound known as poly-3-hydroxybutyrate (PHB) in order to withstand carbon-deficient environments. When starved,
2024:
1841:
Roberts, H. A.; Feast, Patricia M. (1972-12-01). "Fate of Seeds of Some Annual Weeds in
Different Depths of Cultivated and Undisturbed Soil".
53:
drought by producing some seeds that germinate immediately and other seeds that lie dormant. Other examples of biological bet hedging include
91:
There are three categories (strategies) of bet-hedging: "conservative" bet-hedging, "diversified" bet-hedging, and "adaptive coin flipping."
1496:
Watson, Paul J. (1991-02-01). "Multiple paternity as genetic bet-hedging in female sierra dome spiders, Linyphia litigiosa (Linyphiidae)".
547:
species). This indicates that weeds will engage in bet hedging at higher rates in circumstances where the costs of bet hedging are lower.
1270:"Mate choice for major histocompatibility complex genetic divergence as a bet-hedging strategy in the atlantic salmon (salmo salar)"
434:
species makes a more active decision about when to use torpor that is better-suited to the uncertain environment in which it lives.
405:
1353:
Evans, Margaret E K; Dennehy, John J (2005-12-01). "Germ
Banking: Bet‐Hedging and Variable Release from Egg and Seed Dormancy".
2003:
1688:
Clauss, M. J.; Venable, D. L. (2000-02-01). "Seed
Germination in Desert Annuals: An Empirical Test of Adaptive Bet Hedging".
462:
Invertebrate bet-hedging has also been observed in the mating systems of some species of spider. Female sierra dome spiders (
104:
with a constant egg size that may not be optimal for any environmental condition, but result in the lowest overall variance.
595:
many years after the original infection. The delay with which shingles emerges has been explained as a form of bet hedging.
483:
fungal infections may be more difficult to treat if bet hedging is involved. For example, pathogenic strains of yeast like
1108:
Beaumont, H. J. E.; Kost, C.; Rainey, P. B.; Gallie, J.; Ferguson, G. C. (2009). "Experimental evolution of bet hedging".
2019:
450:
reduces the potential costs of a catastrophic event during a particularly vulnerable time in offspring development. In
292:
each 50% of the time; they perform better than the conservative bet hedger in good years, but worse during a bad year.
365:
infections, as bet hedging may ensure the survival of its species within its host, heedless to the antibiotic used.
348:
358:
1457:"Ectoparasitic Argulus coregoni (Crustacea: Branchiura) Hedge Their Bets: Studies on Egg Hatching Dynamics"
1362:
584:
327:
1638:"Experimental evolution of bet hedging under manipulated environmental uncertainty in Neurospora crassa"
540:
139:. The geometric mean is highly sensitive to small values. Even rare occurrences of zero fitness for a
1219:
1093:
Seger, Jon; Brockmann, H. Jane (1987). "What is bet-hedging?". In Harvey, P. H.; Partridge, L. (eds.).
374:
hedging. However, because
Eukarya is a broad category, this section has been subdivided into kingdoms
1941:
1117:
623:
416:
1367:
659:"Female multiple mating as a genetic bet-hedging strategy when mate choice criteria are unreliable"
552:
515:
A group of researchers studied another way bet hedging is used by looking at the ascomycete fungus
66:
54:
479:
failure of chemotherapy in cancer due to mechanisms similar to that of bet hedging used in fungi.
1775:
1721:
1513:
1478:
1396:
1250:
1200:
1151:
784:
735:
686:
495:
using this strategy will resist treatments. These fungi are known to cause an infection known as
425:
strategy in order to reduce their metabolic rate to survive environmental changes. Reproductive
27:
1977:
1959:
1910:
1858:
1823:
1815:
1767:
1759:
1713:
1705:
1667:
1618:
1564:
1437:
1416:"Resistance of dormant stages of planktonic invertebrates to adverse environmental conditions"
1388:
1380:
1335:
1299:
1242:
1192:
1143:
1075:
1040:
991:
973:
954:"Modes of response to environmental change and the elusive empirical evidence for bet hedging"
934:
926:
891:
833:
776:
727:
678:
639:
517:
340:
325:
Within prokarya, there are a multitude of bet hedging examples. In one example, the bacterium
49:
1967:
1949:
1900:
1892:
1850:
1807:
1751:
1697:
1657:
1649:
1608:
1598:
1554:
1544:
1505:
1468:
1427:
1372:
1327:
1289:
1281:
1234:
1182:
1133:
1125:
1067:
1030:
1022:
981:
965:
918:
881:
873:
823:
815:
766:
717:
670:
631:
491:
485:
1238:
1138:
399:
395:
353:
157:
136:
72:
1587:"Bet Hedging in Yeast by Heterogeneous, Age-Correlated Expression of a Stress Protectant"
1945:
1121:
627:
1905:
1880:
1854:
1662:
1637:
1613:
1586:
1559:
1532:
1294:
1269:
1035:
1010:
986:
953:
886:
861:
828:
803:
132:
41:
1972:
1929:
1509:
614:
Cohen, Dan (1966-09-01). "Optimizing reproduction in a randomly varying environment".
2013:
1473:
1456:
1071:
674:
635:
508:
to determine the mechanism of bet hedging in this species. It was determined that in
504:
336:
152:
101:
17:
1517:
1204:
788:
739:
690:
1725:
1400:
1254:
1155:
45:
1779:
167:
1603:
1549:
1331:
1058:
Philippi, Tom; Seger, Jon (1989). "Hedging one's evolutionary bets, revisited".
922:
862:"Bet-hedging as an evolutionary game: the trade-off between egg size and number"
819:
496:
409:
1187:
1170:
1026:
771:
754:
722:
705:
588:
539:
germination for drier populations when exposed to rain, a phenomenon known as
522:
318:
1963:
1862:
1819:
1763:
1709:
1441:
1384:
977:
930:
682:
442:
Many invertebrate species are known to exhibit various forms of bet hedging.
1432:
1415:
444:
431:
314:
147:
of higher fitness across a variety of environments. Therefore, bet hedging
58:
1981:
1954:
1914:
1827:
1771:
1717:
1671:
1653:
1622:
1568:
1392:
1339:
1303:
1285:
1246:
1196:
1147:
1079:
1044:
995:
969:
938:
909:
Dempster, Everett R. (1955-01-01). "Maintenance of
Genetic Heterogeneity".
895:
877:
837:
780:
731:
1998:
643:
592:
464:
140:
62:
1129:
1636:
Graham, Jeffrey K.; Smith, Myron L.; Simons, Andrew M. (22 July 2014).
1482:
568:
426:
1896:
1811:
1455:
Hakalahti, Teija; Häkkinen, Heli; Valtonen, E. Tellervo (2004-01-01).
1171:"Individual-level bet hedging in the bacterium sinorhizobium meliloti"
755:"Individual-Level Bet Hedging in the Bacterium Sinorhizobium meliloti"
335:
populations begin to display bet hedging by forming two non-identical
1097:. Vol. 4. Oxford, UK: Oxford University Press. pp. 182–211.
804:"Bacterial Persistence: A Model of Survival in Changing Environments"
580:
422:
383:
375:
148:
44:, or a reservoir of ungerminated seeds in the soil. For example, an
1928:
Stumpf, Michael P. H.; Laidlaw, Zoe; Jansen, Vincent A. A. (2002).
1755:
1701:
1376:
658:
567:
Thus far, research on bet hedging involving species in the domain
379:
322:
have been used to deduce the evolutionary origins of bet hedging.
71:
26:
1268:
Evans, M. L.; Dionne, M.; Miller, K. M.; Bernatchez, L. (2012).
706:"Diversity of speed-accuracy strategies benefits social insects"
1011:"The evolution of bet-hedging adaptations to rare scenarios"
415:
A second example among vertebrates is the marsupial species
1585:
Levy, Sasha F.; Ziv, Naomi; Siegal, Mark L. (8 May 2012).
1885:
New
Phytologist: International Journal of Plant Science
1642:
Proceedings of the Royal
Society B: Biological Sciences
1220:"Bistability, epigenetics, and bet-hedging in bacteria"
866:
Proceedings of the Royal
Society B: Biological Sciences
135:, which is multiplicative instead of additive like the
753:
Ratcliff, William C.; Denison, R. Ford (2010-10-12).
1879:
Huss, Jessica C.; Gierlinger, Notburga (June 2021).
579:
Bet hedging has been used to explain the latency of
32:
that many offspring are produced in good conditions.
1934:
Proceedings of the National Academy of Sciences USA
911:
Cold Spring Harbor Symposia on Quantitative Biology
1218:Veening, J.; Smits, W. K.; Kuipers, O. P. (2008).
860:Olofsson, H.; Ripa, J.; Jonzen, N. (27 May 2009).
430:manipulation of hormones, suggesting that this
1009:King, Oliver D.; Masel, Joanna (2007-12-01).
521:. It was observed that this species produces
8:
855:
853:
851:
849:
847:
352:. In a given population of this bacteria,
1971:
1953:
1904:
1661:
1612:
1602:
1558:
1548:
1472:
1431:
1366:
1293:
1186:
1137:
1034:
985:
885:
827:
770:
721:
346:Another example of bet hedging arises in
1169:Ratcliff, W. C.; Denison, R. F. (2010).
174:
166:
1874:
1872:
603:
1239:10.1146/annurev.micro.62.081307.163002
1095:Oxford surveys in evolutionary biology
77:presence of the alternate environment.
1793:
1791:
1789:
1737:
1735:
1683:
1681:
1580:
1578:
1533:"Yeast Survive by Hedging Their Bets"
1320:General and Comparative Endocrinology
7:
609:
607:
1855:10.1111/j.1365-3180.1972.tb01226.x
25:
1930:"Herpes Viruses Hedge Their Bets"
1060:Trends in Ecology & Evolution
591:at first infection and can cause
404:) have been hypothesized to have
164:and 'bad' years 25% of the time.
1997:
1474:10.1111/j.0030-1299.2004.13213.x
1274:Proceedings: Biological Sciences
958:Proceedings: Biological Sciences
952:Simons, Andrew M. (2011-06-07).
675:10.1046/j.1440-1703.2001.00423.x
571:has not been easily accessible.
406:major histocompatibility complex
1881:"Functional packaging of seeds"
1355:The Quarterly Review of Biology
802:Kussell, E. (31 January 2005).
69:in the presence of antibiotics.
1414:Radzikowski, J. (2013-07-01).
1139:11858/00-001M-0000-002A-07D3-B
1015:Theoretical Population Biology
616:Journal of Theoretical Biology
1:
2025:Evolutionary biology concepts
1531:Meadows, Robin (8 May 2012).
1510:10.1016/S0003-3472(05)80486-5
1227:Annual Review of Microbiology
296:their higher geometric mean.
1604:10.1371/journal.pbio.1001325
1550:10.1371/journal.pbio.1001327
1420:Journal of Plankton Research
1072:10.1016/0169-5347(89)90138-9
636:10.1016/0022-5193(66)90188-3
1332:10.1016/j.ygcen.2012.08.024
923:10.1101/SQB.1955.020.01.005
820:10.1534/genetics.104.035352
657:Yasui, Yukio (2001-12-01).
2041:
349:Mycobacterium tuberculosis
1188:10.1016/j.cub.2010.08.036
1027:10.1016/j.tpb.2007.08.006
772:10.1016/j.cub.2010.08.036
723:10.1016/j.cub.2008.08.028
179:
704:Burns, James G. (2008).
160:the allele to fixation.
95:Conservative bet hedging
1744:The American Naturalist
1690:The American Naturalist
587:, for instance, causes
108:Diversified bet hedging
57:, foraging behavior in
1955:10.1073/pnas.232546899
1654:10.1098/rspb.2014.0706
1286:10.1098/rspb.2011.0909
970:10.1098/rspb.2011.0176
878:10.1098/rspb.2009.0500
585:Varicella Zoster Virus
328:Sinorhizobium meliloti
172:
117:Adaptive coin flipping
100:an organism producing
78:
61:, nutrient storage in
55:female multiple mating
37:Biological bet hedging
33:
2004:Bet hedging (biology)
1433:10.1093/plankt/fbt032
541:phenotypic plasticity
170:
75:
67:bacterial persistence
30:
18:Bet-hedging (biology)
2006:at Wikimedia Commons
456:Diaptomus sanguineus
452:Diaptomus sanguineus
445:Diaptomus sanguineus
359:antibiotic resistant
317:across generations,
2020:Biological theorems
1946:2002PNAS...9915234S
1940:(23): 15234–15237.
1130:10.1038/nature08504
1122:2009Natur.462...90B
964:(1712): 1601–1609.
872:(1669): 2963–2969.
663:Ecological Research
628:1966JThBi..12..119C
553:Medicago truncatula
418:Sminthopsis macrour
1648:(1787): 20140706.
465:Linyphia litigiosa
173:
79:
34:
2002:Media related to
1897:10.1111/nph.17299
1812:10.1111/ele.12241
1280:(1727): 379–386.
1181:(19): 1740–1744.
765:(19): 1740–1744.
716:(20): R953–R954.
518:Neurospora crassa
394:In example, West
289:
288:
16:(Redirected from
2032:
2001:
1986:
1985:
1975:
1957:
1925:
1919:
1918:
1908:
1891:(6): 2154–2163.
1876:
1867:
1866:
1838:
1832:
1831:
1795:
1784:
1783:
1739:
1730:
1729:
1685:
1676:
1675:
1665:
1633:
1627:
1626:
1616:
1606:
1582:
1573:
1572:
1562:
1552:
1528:
1522:
1521:
1498:Animal Behaviour
1493:
1487:
1486:
1476:
1452:
1446:
1445:
1435:
1411:
1405:
1404:
1370:
1350:
1344:
1343:
1314:
1308:
1307:
1297:
1265:
1259:
1258:
1224:
1215:
1209:
1208:
1190:
1166:
1160:
1159:
1141:
1105:
1099:
1098:
1090:
1084:
1083:
1055:
1049:
1048:
1038:
1006:
1000:
999:
989:
949:
943:
942:
906:
900:
899:
889:
857:
842:
841:
831:
814:(4): 1807–1814.
799:
793:
792:
774:
750:
744:
743:
725:
701:
695:
694:
654:
648:
647:
611:
492:Candida glabrata
486:Candida albicans
175:
21:
2040:
2039:
2035:
2034:
2033:
2031:
2030:
2029:
2010:
2009:
1994:
1989:
1927:
1926:
1922:
1878:
1877:
1870:
1840:
1839:
1835:
1800:Ecology Letters
1797:
1796:
1787:
1741:
1740:
1733:
1687:
1686:
1679:
1635:
1634:
1630:
1597:(5): e1001325.
1584:
1583:
1576:
1543:(5): e1001327.
1530:
1529:
1525:
1495:
1494:
1490:
1454:
1453:
1449:
1413:
1412:
1408:
1368:10.1.1.587.7117
1352:
1351:
1347:
1316:
1315:
1311:
1267:
1266:
1262:
1222:
1217:
1216:
1212:
1175:Current Biology
1168:
1167:
1163:
1116:(7269): 90–93.
1107:
1106:
1102:
1092:
1091:
1087:
1057:
1056:
1052:
1008:
1007:
1003:
951:
950:
946:
908:
907:
903:
859:
858:
845:
801:
800:
796:
759:Current Biology
752:
751:
747:
710:Current Biology
703:
702:
698:
656:
655:
651:
613:
612:
605:
601:
577:
565:
532:
476:
440:
396:Atlantic salmon
392:
371:
354:persister cells
311:
306:
253:Arithmetic mean
137:arithmetic mean
128:
119:
110:
97:
89:
84:
23:
22:
15:
12:
11:
5:
2038:
2036:
2028:
2027:
2022:
2012:
2011:
2008:
2007:
1993:
1992:External links
1990:
1988:
1987:
1920:
1868:
1849:(4): 316–324.
1833:
1806:(3): 380–387.
1785:
1756:10.1086/285551
1750:(3): 488–507.
1731:
1702:10.1086/303314
1696:(2): 168–186.
1677:
1628:
1574:
1523:
1504:(2): 343–360.
1488:
1467:(2): 295–302.
1447:
1426:(4): 707–723.
1406:
1377:10.1086/498282
1361:(4): 431–451.
1345:
1326:(2): 265–276.
1309:
1260:
1233:(1): 193–210.
1210:
1161:
1100:
1085:
1050:
1021:(4): 560–575.
1001:
944:
901:
843:
794:
745:
696:
669:(4): 605–616.
649:
622:(1): 119–129.
602:
600:
597:
576:
573:
564:
561:
531:
528:
475:
472:
439:
436:
421:, which use a
391:
388:
370:
367:
341:binary fission
337:daughter cells
310:
307:
305:
302:
287:
286:
283:
280:
277:
274:
272:Geometric mean
268:
267:
264:
261:
258:
255:
249:
248:
245:
242:
239:
236:
230:
229:
226:
223:
220:
217:
211:
210:
204:
198:
192:
186:
182:
181:
178:
133:geometric mean
127:
124:
118:
115:
109:
106:
96:
93:
88:
85:
83:
80:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2037:
2026:
2023:
2021:
2018:
2017:
2015:
2005:
2000:
1996:
1995:
1991:
1983:
1979:
1974:
1969:
1965:
1961:
1956:
1951:
1947:
1943:
1939:
1935:
1931:
1924:
1921:
1916:
1912:
1907:
1902:
1898:
1894:
1890:
1886:
1882:
1875:
1873:
1869:
1864:
1860:
1856:
1852:
1848:
1844:
1843:Weed Research
1837:
1834:
1829:
1825:
1821:
1817:
1813:
1809:
1805:
1801:
1794:
1792:
1790:
1786:
1781:
1777:
1773:
1769:
1765:
1761:
1757:
1753:
1749:
1745:
1738:
1736:
1732:
1727:
1723:
1719:
1715:
1711:
1707:
1703:
1699:
1695:
1691:
1684:
1682:
1678:
1673:
1669:
1664:
1659:
1655:
1651:
1647:
1643:
1639:
1632:
1629:
1624:
1620:
1615:
1610:
1605:
1600:
1596:
1592:
1588:
1581:
1579:
1575:
1570:
1566:
1561:
1556:
1551:
1546:
1542:
1538:
1534:
1527:
1524:
1519:
1515:
1511:
1507:
1503:
1499:
1492:
1489:
1484:
1480:
1475:
1470:
1466:
1462:
1458:
1451:
1448:
1443:
1439:
1434:
1429:
1425:
1421:
1417:
1410:
1407:
1402:
1398:
1394:
1390:
1386:
1382:
1378:
1374:
1369:
1364:
1360:
1356:
1349:
1346:
1341:
1337:
1333:
1329:
1325:
1321:
1313:
1310:
1305:
1301:
1296:
1291:
1287:
1283:
1279:
1275:
1271:
1264:
1261:
1256:
1252:
1248:
1244:
1240:
1236:
1232:
1228:
1221:
1214:
1211:
1206:
1202:
1198:
1194:
1189:
1184:
1180:
1176:
1172:
1165:
1162:
1157:
1153:
1149:
1145:
1140:
1135:
1131:
1127:
1123:
1119:
1115:
1111:
1104:
1101:
1096:
1089:
1086:
1081:
1077:
1073:
1069:
1065:
1061:
1054:
1051:
1046:
1042:
1037:
1032:
1028:
1024:
1020:
1016:
1012:
1005:
1002:
997:
993:
988:
983:
979:
975:
971:
967:
963:
959:
955:
948:
945:
940:
936:
932:
928:
924:
920:
916:
912:
905:
902:
897:
893:
888:
883:
879:
875:
871:
867:
863:
856:
854:
852:
850:
848:
844:
839:
835:
830:
825:
821:
817:
813:
809:
805:
798:
795:
790:
786:
782:
778:
773:
768:
764:
760:
756:
749:
746:
741:
737:
733:
729:
724:
719:
715:
711:
707:
700:
697:
692:
688:
684:
680:
676:
672:
668:
664:
660:
653:
650:
645:
641:
637:
633:
629:
625:
621:
617:
610:
608:
604:
598:
596:
594:
590:
586:
583:viruses. The
582:
574:
572:
570:
562:
560:
557:
555:
554:
548:
544:
542:
536:
529:
527:
524:
520:
519:
513:
511:
510:S. cerevisiae
507:
506:
505:S. cerevisiae
500:
498:
494:
493:
488:
487:
480:
473:
471:
468:
466:
460:
457:
453:
448:
446:
438:Invertebrates
437:
435:
433:
428:
424:
420:
419:
413:
411:
407:
403:
402:
397:
389:
387:
385:
381:
377:
368:
366:
362:
360:
355:
351:
350:
344:
342:
338:
334:
330:
329:
323:
320:
316:
308:
303:
301:
297:
293:
284:
281:
278:
275:
273:
270:
269:
265:
262:
259:
256:
254:
251:
250:
246:
243:
240:
237:
235:
232:
231:
227:
224:
221:
218:
216:
213:
212:
209:
205:
203:
200:Conservative
199:
197:
193:
191:
187:
184:
183:
177:
176:
169:
165:
161:
159:
154:
153:genetic drift
150:
144:
142:
138:
134:
125:
123:
116:
114:
107:
105:
103:
94:
92:
86:
81:
74:
70:
68:
64:
60:
56:
51:
47:
43:
38:
29:
19:
1937:
1933:
1923:
1888:
1884:
1846:
1842:
1836:
1803:
1799:
1747:
1743:
1693:
1689:
1645:
1641:
1631:
1594:
1591:PLOS Biology
1590:
1540:
1537:PLOS Biology
1536:
1526:
1501:
1497:
1491:
1464:
1460:
1450:
1423:
1419:
1409:
1358:
1354:
1348:
1323:
1319:
1312:
1277:
1273:
1263:
1230:
1226:
1213:
1178:
1174:
1164:
1113:
1109:
1103:
1094:
1088:
1066:(2): 41–44.
1063:
1059:
1053:
1018:
1014:
1004:
961:
957:
947:
914:
910:
904:
869:
865:
811:
807:
797:
762:
758:
748:
713:
709:
699:
666:
662:
652:
619:
615:
578:
566:
558:
551:
549:
545:
537:
533:
516:
514:
509:
503:
501:
490:
484:
481:
477:
463:
461:
455:
451:
443:
441:
417:
414:
400:
393:
372:
363:
347:
345:
332:
326:
324:
312:
304:In organisms
298:
294:
290:
271:
252:
233:
214:
207:
206:Diversified
201:
195:
189:
162:
145:
129:
120:
111:
98:
90:
46:annual plant
36:
35:
497:candidiasis
410:mate choice
401:Salmo salar
390:Vertebrates
333:S. meliloti
319:prokaryotes
208:Bet hedger
202:Bet hedger
196:Specialist
190:Specialist
59:bumble bees
2014:Categories
599:References
589:chickenpox
523:ascospores
315:phenotypes
188:Good Year
185:Year Type
180:Phenotype
87:Categories
1964:0027-8424
1863:1365-3180
1820:1461-0248
1764:0003-0147
1710:0003-0147
1442:0142-7873
1385:0033-5770
1363:CiteSeerX
978:1471-2954
931:0091-7451
917:: 25–32.
683:1440-1703
432:marsupial
194:Bad Year
126:Evolution
42:seed bank
1982:12409612
1915:33629369
1828:24393387
1772:19425988
1718:10686159
1672:24870047
1623:22589700
1569:22589702
1518:53152243
1393:16519139
1340:22974513
1304:21697172
1247:18537474
1205:16856229
1197:20869244
1148:19890329
1080:21227310
1045:17915273
996:21411456
939:13433552
896:19474039
838:15687275
808:Genetics
789:16856229
781:20869244
740:16696224
732:18957249
691:34683958
593:shingles
376:Animalia
309:Prokarya
141:genotype
102:clutches
82:Overview
63:rhizobia
1942:Bibcode
1906:8252473
1726:4439415
1663:4071552
1614:3348152
1560:3348148
1483:3548212
1401:2343748
1295:3223684
1255:3747871
1156:4369450
1118:Bibcode
1036:2118055
987:3081777
887:2817213
829:1449587
644:6015423
624:Bibcode
575:Viruses
569:Archaea
563:Archaea
530:Plantae
427:hormone
380:Plantae
369:Eukarya
339:during
149:alleles
50:fitness
1980:
1973:137573
1970:
1962:
1913:
1903:
1861:
1826:
1818:
1780:498615
1778:
1770:
1762:
1724:
1716:
1708:
1670:
1660:
1621:
1611:
1567:
1557:
1516:
1481:
1440:
1399:
1391:
1383:
1365:
1338:
1302:
1292:
1253:
1245:
1203:
1195:
1154:
1146:
1110:Nature
1078:
1043:
1033:
994:
984:
976:
937:
929:
894:
884:
836:
826:
787:
779:
738:
730:
689:
681:
642:
581:Herpes
423:torpor
382:, and
285:0.763
282:0.763
279:0.707
276:0.707
266:0.768
263:0.763
260:0.723
257:0.813
247:0.625
244:0.763
228:0.815
225:0.763
65:, and
1776:S2CID
1722:S2CID
1514:S2CID
1479:JSTOR
1461:Oikos
1397:S2CID
1251:S2CID
1223:(PDF)
1201:S2CID
1152:S2CID
785:S2CID
736:S2CID
687:S2CID
474:Fungi
384:Fungi
238:0.25
222:0.63
158:sweep
1978:PMID
1960:ISSN
1911:PMID
1859:ISSN
1824:PMID
1816:ISSN
1768:PMID
1760:ISSN
1714:PMID
1706:ISSN
1668:PMID
1619:PMID
1565:PMID
1438:ISSN
1389:PMID
1381:ISSN
1336:PMID
1300:PMID
1243:PMID
1193:PMID
1144:PMID
1076:PMID
1041:PMID
992:PMID
974:ISSN
935:PMID
927:ISSN
892:PMID
834:PMID
777:PMID
728:PMID
679:ISSN
640:PMID
241:1.0
219:1.0
215:Good
1968:PMC
1950:doi
1901:PMC
1893:doi
1889:230
1851:doi
1808:doi
1752:doi
1748:142
1698:doi
1694:155
1658:PMC
1650:doi
1646:281
1609:PMC
1599:doi
1555:PMC
1545:doi
1506:doi
1469:doi
1465:107
1428:doi
1373:doi
1328:doi
1324:179
1290:PMC
1282:doi
1278:279
1235:doi
1183:doi
1134:hdl
1126:doi
1114:462
1068:doi
1031:PMC
1023:doi
982:PMC
966:doi
962:278
919:doi
882:PMC
874:doi
870:276
824:PMC
816:doi
812:169
767:doi
718:doi
671:doi
632:doi
489:or
234:Bad
48:'s
2016::
1976:.
1966:.
1958:.
1948:.
1938:99
1936:.
1932:.
1909:.
1899:.
1887:.
1883:.
1871:^
1857:.
1847:12
1845:.
1822:.
1814:.
1804:17
1802:.
1788:^
1774:.
1766:.
1758:.
1746:.
1734:^
1720:.
1712:.
1704:.
1692:.
1680:^
1666:.
1656:.
1644:.
1640:.
1617:.
1607:.
1595:10
1593:.
1589:.
1577:^
1563:.
1553:.
1541:10
1539:.
1535:.
1512:.
1502:41
1500:.
1477:.
1463:.
1459:.
1436:.
1424:35
1422:.
1418:.
1395:.
1387:.
1379:.
1371:.
1359:80
1357:.
1334:.
1322:.
1298:.
1288:.
1276:.
1272:.
1249:.
1241:.
1231:62
1229:.
1225:.
1199:.
1191:.
1179:20
1177:.
1173:.
1150:.
1142:.
1132:.
1124:.
1112:.
1074:.
1062:.
1039:.
1029:.
1019:72
1017:.
1013:.
990:.
980:.
972:.
960:.
956:.
933:.
925:.
915:20
913:.
890:.
880:.
868:.
864:.
846:^
832:.
822:.
810:.
806:.
783:.
775:.
763:20
761:.
757:.
734:.
726:.
714:18
712:.
708:.
685:.
677:.
667:16
665:.
661:.
638:.
630:.
620:12
618:.
606:^
499:.
386:.
378:,
1984:.
1952::
1944::
1917:.
1895::
1865:.
1853::
1830:.
1810::
1782:.
1754::
1728:.
1700::
1674:.
1652::
1625:.
1601::
1571:.
1547::
1520:.
1508::
1485:.
1471::
1444:.
1430::
1403:.
1375::
1342:.
1330::
1306:.
1284::
1257:.
1237::
1207:.
1185::
1158:.
1136::
1128::
1120::
1082:.
1070::
1064:4
1047:.
1025::
998:.
968::
941:.
921::
898:.
876::
840:.
818::
791:.
769::
742:.
720::
693:.
673::
646:.
634::
626::
467:)
447:,
398:(
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
