3507:
3061:
3502:{\displaystyle {\begin{aligned}{\frac {1}{N_{e}^{(F)}}}={\frac {1}{4T}}\left\{{\frac {1}{N_{0}^{f}}}+{\frac {1}{N_{0}^{m}}}+\sum _{i}\left(\ell _{i+1}^{f}\right)^{2}\left(v_{i+1}^{f}\right)^{2}\left({\frac {1}{\ell _{i+1}^{f}}}-{\frac {1}{\ell _{i}^{f}}}\right)\right.\,\,\,\,\,\,\,\,&\\\left.{}+\sum _{i}\left(\ell _{i+1}^{m}\right)^{2}\left(v_{i+1}^{m}\right)^{2}\left({\frac {1}{\ell _{i+1}^{m}}}-{\frac {1}{\ell _{i}^{m}}}\right)\right\}.&\end{aligned}}}
1095:
948:
36:
324:) determined the effective-to-census population size ratio for haploid (mitochondrial DNA, Y chromosomal DNA), and diploid (autosomal DNA) loci separately: the ratio of the effective to the census population size was estimated as 0.6–0.7 for autosomal and X-chromosomal DNA, 0.7–0.9 for mitochondrial DNA and 0.5 for Y-chromosomal DNA.
231:
as the population under consideration". More generally, an effective population size may be defined as the number of individuals in an idealised population that has a value of any given population genetic quantity that is equal to the value of that quantity in the population of interest. The two
3585:
s ≫ 1/N, and largely determined by neutral genetic drift if s ≪ 1/N. In real populations, the cutoff value of s may depend instead on local recombination rates. This limit to selection in a real population may be captured in a toy Wright-Fisher simulation through the appropriate choice of Ne.
3568:
The coalescent effective size may have little relationship to the number of individuals physically present in a population. Measured coalescent effective population sizes vary between genes in the same population, being low in genome areas of low recombination and high in genome areas of high
316:
ratios. Seven different estimation methods were used in the surveyed studies. Accordingly, the ratios ranged widely from 10 for
Pacific oysters to 0.994 for humans, with an average of 0.34 across the examined species. Based on these data they subsequently estimated more comprehensive ratios,
1090:{\displaystyle ={{\begin{matrix}{\frac {1}{10}}\end{matrix}}+{\begin{matrix}{\frac {1}{100}}\end{matrix}}+{\begin{matrix}{\frac {1}{50}}\end{matrix}}+{\begin{matrix}{\frac {1}{80}}\end{matrix}}+{\begin{matrix}{\frac {1}{20}}\end{matrix}}+{\begin{matrix}{\frac {1}{500}}\end{matrix}} \over 6}}
3016:
779:
In the following examples, one or more of the assumptions of a strictly idealised population are relaxed, while other assumptions are retained. The variance effective population size of the more relaxed population model is then calculated with respect to the strict model.
299:
populations of census size 16, the variance effective population size has been measured as equal to 11.5. This measurement was achieved through studying changes in the frequency of a neutral allele from one generation to another in over 100 replicate populations.
2195:
121:. In an idealised diploid population with no selection at any locus, the expectation of the coalescence time in generations is equal to twice the census population size. The effective population size is measured from genetic data as within-species
112:
The same real population could have a different effective population size for different properties of interest, such as genetic drift over one generation vs. over many generations, and a different effective population size at different
2588:
769:
1450:
886:
1850:
2305:
470:
2483:
1670:
1360:
1300:
3026:
Similarly, the inbreeding effective number can be calculated for a diploid population with discrete age structure. This was first given by
Johnson, but the notation more closely resembles Emigh and Pollak.
609:
522:
2616:
Assume a haploid population with discrete age structure. An example might be an organism that can survive several discrete breeding seasons. Further, define the following age structure characteristics:
2849:
2385:
1539:
However, in natural populations the variance is often larger than this. The vast majority of individuals may have no offspring, and the next generation stems only from a small number of individuals, so
2837:
1534:
3066:
652:
1958:
1579:
3573:
can cause neutral mutations to have sojourn times proportional to log(N): this may explain the relationship between measured effective population size and the local recombination rate.
2064:
as the size of the idealized population that has the same change in average inbreeding coefficient as the population under consideration. The presentation follows
Kempthorne (1957).
102:
that would experience the same rate of genetic drift as the real population, whereby the latter, due partly to the limited proportion of breeding individuals, has a normally larger
2073:
4707:
1993:
1130:
109:. Idealised populations are based on unrealistic but convenient assumptions including random mating, simultaneous birth of each new generation, and constant population size.
942:
1158:
560:
4503:
2705:
1692:. In the extreme case of a population experiencing no variation in family size, in a laboratory population in which the number of offspring is artificially controlled,
54:
363:
2759:
2648:
176:
4553:
1905:
1212:
1188:
3563:
3543:
146:
5737:
4990:
2021:
4664:
2725:
2671:
672:
386:
3521:, a neutral allele remains in a population for Ne generations, where Ne is the effective population size. An idealised diploid population will have a pairwise
3565:
is the mutation rate. The sojourn effective population size can therefore be estimated empirically by dividing the nucleotide diversity by the mutation rate.
5881:
5951:
2494:
680:
5488:
3569:
recombination. Sojourn times are proportional to N in neutral theory, but for alleles under selection, sojourn times are proportional to log(N).
1395:
5961:
5689:
4957:
4815:
4490:
806:
3581:
In an idealised Wright-Fisher model, the fate of an allele, beginning at an intermediate frequency, is largely determined by selection if the
4712:
4525:
4407:
3518:
1736:
5966:
2210:
5523:
2396:
4184:
Emigh TH, Pollak E (1979). "Fixation probabilities and effective population numbers in diploid populations with overlapping generations".
1590:
6154:
4546:
1311:
1251:
5571:
3030:
Assume the same basic parameters for the life table as given for the haploid case, but distinguishing between male and female, such as
394:
317:
accounting for fluctuations in population size, variance in family size and unequal sex-ratio. These ratios average to only 0.10-0.11.
5907:
5730:
4983:
3011:{\displaystyle N_{e}^{(F)}={\frac {N_{0}T}{1+\sum _{i}\ell _{i+1}^{2}v_{i+1}^{2}({\frac {1}{\ell _{i+1}}}-{\frac {1}{\ell _{i}}})}}.}
5243:
4657:
72:
3586:
Populations with different selection effective population sizes are predicted to evolve profoundly different genome architectures.
524:
denote the same, typically larger, variance in the actual population under consideration. The variance effective population size
2316:
565:
478:
303:
For coalescent effective population sizes, a survey of publications on 102 mostly wildlife animal and plant species yielded 192
5986:
5699:
5566:
5278:
4752:
4511:
6557:
6244:
4757:
4539:
4024:
Karlin, Samuel (1968-09-01). "Rates of
Approach to Homozygosity for Finite Stochastic Models with Variable Population Size".
232:
population genetic quantities identified by Wright were the one-generation increase in variance across replicate populations
6371:
6016:
2775:
2650:
182:, the coalescent effective population size may not reflect the census population size at all, or may reflect its logarithm.
1486:
6417:
5971:
5723:
4976:
5849:
4687:
6542:
4650:
1911:
6552:
6547:
6206:
5097:
4732:
614:
6271:
5991:
5195:
5092:
1546:
4853:
6452:
6064:
5956:
5814:
5799:
5185:
4727:
3595:
6211:
1225:
of the population size, which in this example is 126.7. The harmonic mean tends to be dominated by the smallest
6442:
6437:
6407:
5674:
5556:
4905:
4590:
3619:
2604:
When organisms live longer than one breeding season, effective population sizes have to take into account the
2190:{\displaystyle F_{t}={\frac {1}{N}}\left({\frac {1+F_{t-2}}{2}}\right)+\left(1-{\frac {1}{N}}\right)F_{t-1}.}
6286:
6149:
6059:
5927:
5809:
5779:
5636:
5601:
5321:
5288:
5263:
4717:
6432:
6376:
6311:
6174:
6109:
6044:
5704:
5606:
5394:
5102:
5082:
4702:
4526:
https://web.archive.org/web/20050524144622/http://www.kursus.kvl.dk/shares/vetgen/_Popgen/genetics/3/6.htm
3801:
2052:
179:
3647:
562:
is defined as the size of an idealized population with the same variance. This is found by substituting
6336:
6281:
6144:
6129:
5912:
5869:
5859:
5854:
5611:
5591:
5447:
5437:
5379:
5374:
5210:
5062:
4944:
4838:
4833:
4737:
4566:
3600:
3582:
1873:
the number of females. For example, with 80 males and 20 females (an absolute population size of 100):
1226:
280:
1968:
1105:
6462:
6427:
6422:
6346:
6341:
6296:
6194:
6164:
6159:
6011:
5874:
5864:
5409:
5248:
5037:
4790:
4626:
4585:
4440:
4193:
3793:
3624:
3522:
1465:
913:
337:
333:
256:
216:
122:
99:
4968:
4494:
6512:
6487:
6351:
6321:
6266:
6179:
6069:
6054:
6001:
5834:
5769:
5651:
5581:
5112:
4934:
4810:
4697:
4673:
4616:
3806:
3570:
186:
5715:
1140:
6523:
6472:
6467:
6276:
6239:
5937:
5902:
5759:
5684:
5586:
5518:
5508:
5442:
5389:
5200:
5145:
5107:
5032:
4924:
4920:
4780:
4329:
4139:
4088:
4049:
3924:
3861:
3827:
3655:
1242:
527:
207:
Depending on the quantity of interest, effective population size can be defined in several ways.
5981:
2680:
6412:
6381:
6169:
5996:
5804:
5669:
5646:
5503:
5384:
5160:
5072:
5057:
5042:
5022:
4915:
4910:
4848:
4775:
4767:
4742:
4468:
4403:
4380:
4321:
4283:
4242:
4166:
4115:
4041:
4006:
3889:
3819:
3763:
3702:
3682:
252:
248:
149:
118:
6366:
6229:
6221:
6139:
6021:
6006:
5942:
5922:
5839:
5829:
5824:
5789:
5621:
5561:
5432:
5233:
5175:
5087:
5047:
4874:
4785:
4458:
4448:
4370:
4360:
4313:
4273:
4232:
4201:
4156:
4148:
4105:
4097:
4033:
3996:
3988:
3955:
3916:
3879:
3871:
3811:
3753:
3745:
3711:
3672:
3664:
2734:
2623:
366:
220:
155:
114:
1883:
1194:
1166:
117:
in the genome. The effective population size is most commonly measured with respect to the
6502:
6361:
6331:
6326:
6316:
6249:
6234:
6114:
6094:
5976:
5844:
5750:
5641:
5551:
5493:
5478:
5077:
5003:
4692:
3548:
3528:
2766:
1369:
represents dioeciousness and may take the value 0 (for not dioecious) or 1 for dioecious.
1222:
247:
Today, the effective population size is usually estimated empirically with respect to the
131:
103:
2067:
For the idealized population, the inbreeding coefficients follow the recurrence equation
2003:
1460:
If population size is to remain constant, each individual must contribute on average two
4444:
4197:
3797:
343:
6482:
6306:
6259:
6189:
6184:
6079:
5946:
5819:
5626:
5616:
5596:
5473:
5399:
5364:
5303:
5180:
5135:
5027:
4939:
4929:
4843:
4722:
4595:
4463:
4428:
4375:
4348:
4237:
4220:
4161:
4134:
4110:
4084:"Inbreeding and variance effective numbers in populations with overlapping generations"
4083:
4001:
3976:
3884:
3849:
3758:
3733:
3700:
Wright S (1938). "Size of population and breeding structure in relation to evolution".
3677:
2710:
2656:
657:
371:
2051:
Alternatively, the effective population size may be defined by noting how the average
199:. Some versions of the effective population size are used in wildlife conservation.
6536:
6507:
5483:
5457:
5414:
5404:
5359:
5346:
5326:
5218:
5052:
5007:
4879:
4869:
4825:
4278:
4261:
4205:
3729:
1727:
900:
797:
284:
260:
224:
212:
208:
196:
190:
126:
4333:
4053:
3831:
17:
6492:
6477:
6134:
6104:
6049:
5932:
5897:
5774:
5273:
4895:
4747:
4580:
4424:
4301:
241:
4531:
3715:
4365:
2583:{\displaystyle N_{e}^{(F)}={\frac {1}{2\left(1-{\frac {P_{t+1}}{P_{t}}}\right)}}}
764:{\displaystyle N_{e}^{(v)}={p(1-p) \over 2{\widehat {\operatorname {var} }}(p)}.}
5784:
5513:
5331:
5293:
5268:
5258:
5223:
5170:
5150:
4304:(2012). "Rethinking Hardy–Weinberg and genetic drift in undergraduate biology".
4152:
4101:
3875:
3749:
2597:
2204:) instead of inbreeding coefficient, we get the approximate recurrence equation
3668:
6497:
6074:
6039:
5679:
5631:
5576:
5546:
5452:
5369:
5313:
5190:
5140:
3960:
3944:"Effective population size/adult population size ratios in wildlife: a review"
3943:
3850:"Genetic Draft and Quasi-Neutrality in Large Facultatively Sexual Populations"
2605:
793:
228:
193:
4045:
3044:
for the number of newborn females and males, respectively (notice lower case
1464:
to the next generation. An idealized population assumes that this follows a
6402:
6356:
6084:
5528:
5498:
5298:
5253:
5228:
5165:
5155:
5130:
5122:
5067:
4900:
4453:
3907:
Buri, P (1956). "Gene frequency in small populations of mutant
Drosophila".
3815:
1723:
1238:
4472:
4384:
4325:
4317:
4287:
4246:
4170:
4010:
3992:
3893:
3823:
3767:
3686:
1445:{\displaystyle N_{e}=N+{\begin{matrix}{\frac {1}{2}}\approx N\end{matrix}}}
4429:"Evolution of molecular error rates and the consequences for evolvability"
4119:
6457:
6386:
5917:
5424:
5336:
5283:
5238:
4631:
4621:
4562:
1469:
881:{\displaystyle {1 \over N_{e}}={1 \over t}\sum _{i=1}^{t}{1 \over N_{i}}}
185:
The concept of effective population size was introduced in the field of
6447:
6254:
6124:
6119:
5746:
5694:
5354:
4999:
3928:
1845:{\displaystyle N_{e}^{(v)}=N_{e}^{(F)}={4N_{m}N_{f} \over N_{m}+N_{f}}}
4642:
2300:{\displaystyle 1-F_{t}=P_{t}=P_{0}\left(1-{\frac {1}{2N}}\right)^{t}.}
2478:{\displaystyle {\frac {P_{t+1}}{P_{t}}}=1-{\frac {1}{2N_{e}^{(F)}}}.}
1461:
321:
3920:
3784:
Lynch, M.; Conery, J.S. (2003). "The origins of genome complexity".
4349:"Limits to the Rate of Adaptive Substitution in Sexual Populations"
4037:
1665:{\displaystyle N_{e}^{(v)}={4N-2D \over 2+\operatorname {var} (k)}}
215:
originally defined it as "the number of breeding individuals in an
3866:
1355:{\displaystyle N_{e}=N+{\begin{matrix}{\frac {D}{2}}\end{matrix}}}
1295:{\displaystyle N_{e}=N+{\begin{matrix}{\frac {1}{2}}\end{matrix}}}
465:{\displaystyle \operatorname {var} (p'\mid p)={p(1-p) \over 2N}.}
4221:"Is the population size of a species relevant to its evolution?"
3977:"Generation time and effective population size in Polar Eskimos"
1477:
5719:
4972:
4646:
4535:
2600:
example: overlapping generations and age-structured populations
29:
1584:
The effective population size is then smaller, and given by:
236:
and the one-generation change in the inbreeding coefficient
3333:
3316:
2380:{\displaystyle {\frac {P_{t+1}}{P_{t}}}=1-{\frac {1}{2N}}.}
2055:
changes from one generation to the next, and then defining
604:{\displaystyle {\widehat {\operatorname {var} }}(p'\mid p)}
517:{\displaystyle {\widehat {\operatorname {var} }}(p'\mid p)}
178:. In a population with selection at many loci and abundant
4135:"Inbreeding in populations with overlapping generations"
2593:
although researchers rarely use this equation directly.
50:
2832:{\displaystyle T=\sum _{i=0}^{\infty }\ell _{i}v_{i}=}
2761:
The number of newborn individuals per breeding season.
2390:
The inbreeding effective size can be found by solving
1419:
1335:
1275:
1064:
1043:
1022:
1001:
980:
959:
3551:
3531:
3064:
2852:
2778:
2737:
2713:
2683:
2659:
2626:
2497:
2399:
2319:
2213:
2076:
2006:
1971:
1914:
1886:
1739:
1593:
1549:
1529:{\displaystyle \operatorname {var} (k)={\bar {k}}=2.}
1489:
1398:
1314:
1254:
1197:
1169:
1143:
1108:
951:
916:
809:
788:
Population size varies over time. Suppose there are
683:
660:
617:
568:
530:
481:
397:
374:
346:
267:. Another important effective population size is the
158:
134:
4510:. The University of British Columbia. Archived from
6395:
6295:
6220:
6093:
6030:
5890:
5758:
5660:
5539:
5466:
5423:
5345:
5312:
5209:
5121:
5015:
4888:
4862:
4824:
4799:
4766:
4680:
4609:
4573:
1730:1:1 ratio, effective population size is given by:
320:
A genealogical analysis of human hunter-gatherers (
45:
may be too technical for most readers to understand
4262:"Toward a selection theory of molecular evolution"
3734:"Wright and Fisher on Inbreeding and Random Drift"
3557:
3537:
3501:
3010:
2843:Then, the inbreeding effective population size is
2831:
2753:
2719:
2699:
2665:
2642:
2582:
2477:
2379:
2299:
2189:
2015:
1987:
1953:{\displaystyle ={4\times 80\times 20 \over 80+20}}
1952:
1899:
1844:
1664:
1573:
1528:
1444:
1354:
1294:
1206:
1182:
1152:
1124:
1089:
936:
880:
763:
666:
646:
603:
554:
516:
464:
380:
357:
170:
140:
899:= 6). Then the effective population size is the
796:, then effective population size is given by the
240:. These two are closely linked, and derived from
219:that would show the same amount of dispersion of
1389:, so this is usually trivial and often ignored:
895:= 10, 100, 50, 80, 20, 500 for six generations (
4347:Daniel B. Weissman; Nicholas H. Barton (2012).
283:at which selection becomes more important than
148:, because in such an idealised population, the
647:{\displaystyle \operatorname {var} (p'\mid p)}
5731:
4984:
4658:
4547:
2707:The chance an individual will survive to age
1574:{\displaystyle \operatorname {var} (k)>2.}
8:
3843:
3841:
3779:
3777:
203:Overview: Types of effective population size
4493:. University of Connecticut. Archived from
2200:Using Panmictic Index (1 −
5952:Latitudinal gradients in species diversity
5738:
5724:
5716:
4991:
4977:
4969:
4665:
4651:
4643:
4554:
4540:
4532:
4462:
4452:
4374:
4364:
4277:
4236:
4160:
4109:
4000:
3959:
3883:
3865:
3805:
3757:
3676:
3550:
3530:
3473:
3468:
3459:
3448:
3437:
3428:
3417:
3407:
3396:
3381:
3371:
3360:
3345:
3336:
3326:
3325:
3324:
3323:
3322:
3321:
3320:
3319:
3302:
3297:
3288:
3277:
3266:
3257:
3246:
3236:
3225:
3210:
3200:
3189:
3174:
3159:
3154:
3145:
3134:
3129:
3120:
3100:
3083:
3078:
3069:
3065:
3063:
2991:
2982:
2965:
2956:
2947:
2936:
2926:
2915:
2905:
2884:
2877:
2862:
2857:
2851:
2820:
2810:
2800:
2789:
2777:
2742:
2736:
2712:
2688:
2682:
2658:
2631:
2625:
2564:
2548:
2542:
2522:
2507:
2502:
2496:
2457:
2452:
2439:
2422:
2406:
2400:
2398:
2359:
2342:
2326:
2320:
2318:
2288:
2268:
2250:
2237:
2224:
2212:
2172:
2153:
2117:
2104:
2090:
2081:
2075:
2005:
1975:
1970:
1918:
1913:
1891:
1885:
1833:
1820:
1808:
1798:
1788:
1773:
1768:
1749:
1744:
1738:
1618:
1603:
1598:
1592:
1548:
1509:
1508:
1488:
1422:
1418:
1403:
1397:
1338:
1334:
1319:
1313:
1278:
1274:
1259:
1253:
1196:
1174:
1168:
1142:
1112:
1107:
1067:
1063:
1046:
1042:
1025:
1021:
1004:
1000:
983:
979:
962:
958:
955:
950:
926:
917:
915:
891:For example, say the population size was
870:
861:
855:
844:
830:
819:
810:
808:
735:
734:
708:
693:
688:
682:
659:
616:
570:
569:
567:
540:
535:
529:
483:
482:
480:
427:
396:
373:
345:
157:
133:
73:Learn how and when to remove this message
57:, without removing the technical details.
5850:Predator–prey (Lotka–Volterra) equations
5489:Tritrophic interactions in plant defense
334:Wright-Fisher idealized population model
5882:Random generalized Lotka–Volterra model
3611:
2839:average age of a reproducing individual
5690:Herbivore adaptations to plant defense
4958:Index of evolutionary biology articles
1472:of the number of gametes contributed,
238:(inbreeding effective population size)
4069:An Introduction to Genetic Statistics
3519:neutral theory of molecular evolution
55:make it understandable to non-experts
7:
5705:Predator avoidance in schooling fish
3648:"Evolution in Mendelian populations"
3048:for females, compared to upper case
265:coalescent effective population size
234:(variance effective population size)
6155:Intermediate disturbance hypothesis
3055:The inbreeding effective number is
269:selection effective population size
5908:Ecological effects of biodiversity
4528:— on Københavns Universitet.
4400:The Origins of Genome Architecture
4238:10.1111/j.0014-3820.2001.tb00732.x
3848:R.A. Neher; B.I. Shraiman (2011).
2801:
1229:that the population goes through.
255:, estimated as the within-species
25:
5244:Generalist and specialist species
4574:Concepts in Quantitative Genetics
3975:S. Matsumura; P. Forster (2008).
2310:The difference per generation is
1988:{\displaystyle ={6400 \over 100}}
1125:{\displaystyle ={0.1945 \over 6}}
5967:Occupancy–abundance relationship
4508:Biology 434: Population Genetics
4279:10.1111/j.1558-5646.2007.00308.x
1726:of a population varies from the
1456:Variance in reproductive success
34:
5987:Relative abundance distribution
5700:Plant defense against herbivory
5567:Competitive exclusion principle
5279:Mesopredator release hypothesis
4504:"The Effective Population Size"
937:{\displaystyle {1 \over N_{e}}}
388:in the previous generation, is
5572:Consumer–resource interactions
4758:Constructive neutral evolution
4489:Holsinger, Kent (2008-08-26).
4186:Theoretical Population Biology
4071:. Iowa State University Press.
3090:
3084:
2999:
2953:
2869:
2863:
2514:
2508:
2464:
2458:
1780:
1774:
1756:
1750:
1656:
1650:
1610:
1604:
1562:
1556:
1514:
1502:
1496:
752:
746:
726:
714:
700:
694:
641:
624:
598:
581:
547:
541:
511:
494:
445:
433:
421:
404:
244:, but they are not identical.
1:
6418:Biological data visualization
6245:Environmental niche modelling
5972:Population viability analysis
3716:10.1126/science.87.2263.425-a
1675:Note that if the variance of
784:Variations in population size
279:is the critical value of the
5903:Density-dependent inhibition
4708:Fisher's fundamental theorem
4366:10.1371/journal.pgen.1002740
4206:10.1016/0040-5809(79)90028-5
1480:number contributed, i.e. 2:
1153:{\displaystyle =0.032416667}
6372:Liebig's law of the minimum
6207:Resource selection function
5098:Metabolic theory of ecology
4733:Coefficient of relationship
4491:"Effective Population Size"
3876:10.1534/genetics.111.128876
3750:10.1534/genetics.109.110023
3620:"Effective population size"
2651:Fisher's reproductive value
1864:is the number of males and
1221:Note this is less than the
555:{\displaystyle N_{e}^{(v)}}
6574:
6272:Niche apportionment models
5992:Relative species abundance
5196:Primary nutritional groups
5093:List of feeding behaviours
4502:Whitlock, Michael (2008).
2700:{\displaystyle \ell _{i}=}
125:divided by four times the
6521:
6453:Ecosystem based fisheries
6065:Interspecific competition
5957:Minimum viable population
5815:Maximum sustainable yield
5800:Intraspecific competition
5795:Effective population size
5675:Anti-predator adaptations
5186:Photosynthetic efficiency
4953:
4728:Coefficient of inbreeding
4601:Effective population size
4260:Hahn, Matthew W. (2008).
4153:10.1093/genetics/87.3.581
4102:10.1093/genetics/68.4.581
3961:10.1017/S0016672300034455
3596:Minimum viable population
3513:Coalescent effective size
2047:Inbreeding effective size
800:of the population sizes:
87:effective population size
6443:Ecological stoichiometry
6408:Alternative stable state
4906:Evolutionary game theory
4688:Hardy–Weinberg principle
4591:Quantitative trait locus
3669:10.1093/genetics/16.2.97
3577:Selection effective size
1718:Non-Fisherian sex-ratios
340:of the allele frequency
6287:Ontogenetic niche shift
6150:Ideal free distribution
6060:Ecological facilitation
5810:Malthusian growth model
5780:Consumer-resource model
5637:Paradox of the plankton
5602:Energy systems language
5322:Chemoorganoheterotrophy
5289:Optimal foraging theory
5264:Heterotrophic nutrition
4718:Shifting balance theory
4454:10.1073/pnas.1012918108
4398:Lynch, Michael (2007).
4026:The American Naturalist
3816:10.1126/science.1089370
2030:Again, this results in
328:Variance effective size
6433:Ecological forecasting
6377:Marginal value theorem
6175:Landscape epidemiology
6110:Cross-boundary subsidy
6045:Biological interaction
5395:Microbial intelligence
5083:Green world hypothesis
4703:Linkage disequilibrium
4402:. Sinauer Associates.
4318:10.1002/bies.201100178
4219:Gillespie, JH (2001).
4082:Felsenstein J (1971).
3993:10.1098/rspb.2007.1724
3559:
3539:
3503:
3012:
2833:
2805:
2755:
2754:{\displaystyle N_{0}=}
2721:
2701:
2667:
2644:
2643:{\displaystyle v_{i}=}
2584:
2479:
2381:
2301:
2191:
2053:inbreeding coefficient
2017:
1989:
1954:
1901:
1846:
1666:
1575:
1530:
1446:
1356:
1296:
1208:
1184:
1154:
1126:
1091:
938:
882:
860:
765:
668:
648:
605:
556:
518:
466:
382:
359:
291:Empirical measurements
227:or the same amount of
180:linkage disequilibrium
172:
171:{\displaystyle 4N\mu }
142:
104:census population size
6558:Quantitative genetics
6438:Ecological humanities
6337:Ecological energetics
6282:Niche differentiation
6145:Habitat fragmentation
5913:Ecological extinction
5860:Small population size
5612:Feed conversion ratio
5592:Ecological succession
5524:San Francisco Estuary
5438:Ecological efficiency
5380:Microbial cooperation
4945:Quantitative genetics
4854:Balding–Nichols model
4839:Population bottleneck
4834:Small population size
4738:Selection coefficient
4567:Quantitative genetics
4067:Kempthorne O (1957).
3601:Small population size
3583:selection coefficient
3560:
3540:
3504:
3013:
2834:
2785:
2756:
2722:
2702:
2668:
2645:
2585:
2480:
2382:
2302:
2192:
2018:
1990:
1955:
1902:
1900:{\displaystyle N_{e}}
1847:
1667:
1576:
1531:
1447:
1385:approximately equals
1357:
1297:
1209:
1207:{\displaystyle =30.8}
1185:
1183:{\displaystyle N_{e}}
1155:
1127:
1092:
939:
883:
840:
766:
669:
649:
606:
557:
519:
467:
383:
360:
281:selection coefficient
173:
143:
6463:Evolutionary ecology
6428:Ecological footprint
6423:Ecological economics
6347:Ecological threshold
6342:Ecological indicator
6212:Source–sink dynamics
6165:Land change modeling
6160:Insular biogeography
6012:Species distribution
5751:Modelling ecosystems
5410:Microbial metabolism
5249:Intraguild predation
5038:Biogeochemical cycle
5004:Modelling ecosystems
4816:Background selection
4803:on genomic variation
4801:Effects of selection
4753:Population structure
4627:Evolutionary biology
3942:R. Frankham (1995).
3625:Blackwell Publishing
3558:{\displaystyle \mu }
3549:
3538:{\displaystyle \mu }
3529:
3523:nucleotide diversity
3062:
2850:
2776:
2735:
2711:
2681:
2657:
2624:
2495:
2397:
2317:
2211:
2074:
2004:
1969:
1912:
1884:
1737:
1591:
1547:
1487:
1466:Poisson distribution
1396:
1312:
1252:
1195:
1167:
1141:
1106:
949:
914:
807:
775:Theoretical examples
681:
658:
615:
566:
528:
479:
395:
372:
344:
338:conditional variance
217:idealised population
156:
141:{\displaystyle \mu }
132:
100:idealised population
98:) is the size of an
18:Effective population
6543:Population genetics
6513:Theoretical ecology
6488:Natural environment
6352:Ecosystem diversity
6322:Ecological collapse
6312:Bateman's principle
6267:Limiting similarity
6180:Landscape limnology
6002:Species homogeneity
5840:Population modeling
5835:Population dynamics
5652:Trophic state index
4935:Population genomics
4811:Genetic hitchhiking
4698:Identity by descent
4674:Population genetics
4617:Population genetics
4445:2011PNAS..108.1082R
4198:1979TPBio..15...86E
4133:Johnson DL (1977).
3987:(1642): 1501–1508.
3798:2003Sci...302.1401L
3792:(5649): 1401–1404.
3571:Genetic hitchhiking
3478:
3453:
3412:
3376:
3307:
3282:
3241:
3205:
3164:
3139:
3094:
2952:
2931:
2873:
2518:
2468:
2016:{\displaystyle =64}
1784:
1760:
1614:
1305:or more generally,
1241:, i.e. there is no
1237:If a population is
704:
551:
187:population genetics
6553:Ecological metrics
6548:Population ecology
6524:Outline of ecology
6473:Industrial ecology
6468:Functional ecology
6332:Ecological deficit
6277:Niche construction
6240:Ecosystem engineer
6017:Species–area curve
5938:Introduced species
5753:: Other components
5685:Deimatic behaviour
5587:Ecological network
5519:North Pacific Gyre
5504:hydrothermal vents
5443:Ecological pyramid
5390:Microbial food web
5201:Primary production
5146:Foundation species
4921:Landscape genetics
3555:
3535:
3499:
3497:
3464:
3433:
3392:
3356:
3350:
3293:
3262:
3221:
3185:
3179:
3150:
3125:
3074:
3008:
2932:
2911:
2910:
2853:
2829:
2751:
2717:
2697:
2663:
2640:
2580:
2498:
2475:
2448:
2377:
2297:
2187:
2013:
1985:
1950:
1897:
1842:
1764:
1740:
1662:
1594:
1571:
1526:
1442:
1440:
1352:
1350:
1292:
1290:
1243:self-fertilisation
1204:
1180:
1150:
1122:
1087:
1079:
1058:
1037:
1016:
995:
974:
934:
903:of these, giving:
878:
761:
684:
664:
644:
601:
552:
531:
514:
462:
378:
358:{\displaystyle p'}
355:
221:allele frequencies
168:
138:
27:Ecological concept
6530:
6529:
6413:Balance of nature
6170:Landscape ecology
6055:Community ecology
5997:Species diversity
5933:Indicator species
5928:Gradient analysis
5805:Logistic function
5713:
5712:
5670:Animal coloration
5647:Trophic mutualism
5385:Microbial ecology
5176:Photoheterotrophs
5161:Myco-heterotrophy
5073:Ecosystem ecology
5058:Carrying capacity
5023:Abiotic component
4966:
4965:
4916:Genetic genealogy
4911:Fitness landscape
4640:
4639:
4409:978-0-87893-484-3
4231:(11): 2161–2169.
3948:Genetics Research
3710:(2263): 430–431.
3646:Wright S (1931).
3517:According to the
3479:
3454:
3341:
3308:
3283:
3170:
3165:
3140:
3113:
3095:
3052:for inbreeding).
3003:
2997:
2977:
2901:
2769:is calculated as
2720:{\displaystyle i}
2666:{\displaystyle i}
2608:for the species.
2578:
2570:
2470:
2428:
2372:
2348:
2281:
2161:
2133:
2098:
2026:
2025:
1983:
1948:
1840:
1679:is less than 2,
1660:
1517:
1430:
1346:
1286:
1217:
1216:
1120:
1085:
1075:
1054:
1033:
1012:
991:
970:
932:
876:
838:
825:
756:
743:
667:{\displaystyle N}
578:
491:
457:
381:{\displaystyle p}
257:genetic diversity
123:genetic diversity
83:
82:
75:
16:(Redirected from
6565:
6230:Ecological niche
6202:selection theory
6022:Umbrella species
6007:Species richness
5943:Invasive species
5923:Flagship species
5830:Population cycle
5825:Overexploitation
5790:Ecological yield
5740:
5733:
5726:
5717:
5622:Mesotrophic soil
5562:Climax community
5494:Marine food webs
5433:Biomagnification
5234:Chemoorganotroph
5088:Keystone species
5048:Biotic component
4993:
4986:
4979:
4970:
4875:J. B. S. Haldane
4667:
4660:
4653:
4644:
4556:
4549:
4542:
4533:
4522:
4520:
4519:
4498:
4477:
4476:
4466:
4456:
4439:(3): 1082–1087.
4420:
4414:
4413:
4395:
4389:
4388:
4378:
4368:
4344:
4338:
4337:
4298:
4292:
4291:
4281:
4257:
4251:
4250:
4240:
4216:
4210:
4209:
4181:
4175:
4174:
4164:
4130:
4124:
4123:
4113:
4079:
4073:
4072:
4064:
4058:
4057:
4032:(927): 443–455.
4021:
4015:
4014:
4004:
3972:
3966:
3965:
3963:
3939:
3933:
3932:
3904:
3898:
3897:
3887:
3869:
3845:
3836:
3835:
3809:
3781:
3772:
3771:
3761:
3726:
3720:
3719:
3697:
3691:
3690:
3680:
3652:
3643:
3637:
3636:
3634:
3632:
3616:
3564:
3562:
3561:
3556:
3544:
3542:
3541:
3536:
3508:
3506:
3505:
3500:
3498:
3495:
3490:
3486:
3485:
3481:
3480:
3477:
3472:
3460:
3455:
3452:
3447:
3429:
3422:
3421:
3416:
3411:
3406:
3386:
3385:
3380:
3375:
3370:
3349:
3337:
3328:
3318:
3315:
3314:
3310:
3309:
3306:
3301:
3289:
3284:
3281:
3276:
3258:
3251:
3250:
3245:
3240:
3235:
3215:
3214:
3209:
3204:
3199:
3178:
3166:
3163:
3158:
3146:
3141:
3138:
3133:
3121:
3114:
3112:
3101:
3096:
3093:
3082:
3070:
3017:
3015:
3014:
3009:
3004:
3002:
2998:
2996:
2995:
2983:
2978:
2976:
2975:
2957:
2951:
2946:
2930:
2925:
2909:
2893:
2889:
2888:
2878:
2872:
2861:
2838:
2836:
2835:
2830:
2825:
2824:
2815:
2814:
2804:
2799:
2760:
2758:
2757:
2752:
2747:
2746:
2726:
2724:
2723:
2718:
2706:
2704:
2703:
2698:
2693:
2692:
2672:
2670:
2669:
2664:
2649:
2647:
2646:
2641:
2636:
2635:
2589:
2587:
2586:
2581:
2579:
2577:
2576:
2572:
2571:
2569:
2568:
2559:
2558:
2543:
2523:
2517:
2506:
2484:
2482:
2481:
2476:
2471:
2469:
2467:
2456:
2440:
2429:
2427:
2426:
2417:
2416:
2401:
2386:
2384:
2383:
2378:
2373:
2371:
2360:
2349:
2347:
2346:
2337:
2336:
2321:
2306:
2304:
2303:
2298:
2293:
2292:
2287:
2283:
2282:
2280:
2269:
2255:
2254:
2242:
2241:
2229:
2228:
2196:
2194:
2193:
2188:
2183:
2182:
2167:
2163:
2162:
2154:
2138:
2134:
2129:
2128:
2127:
2105:
2099:
2091:
2086:
2085:
2039:being less than
2022:
2020:
2019:
2014:
1994:
1992:
1991:
1986:
1984:
1976:
1959:
1957:
1956:
1951:
1949:
1947:
1936:
1919:
1906:
1904:
1903:
1898:
1896:
1895:
1878:
1877:
1851:
1849:
1848:
1843:
1841:
1839:
1838:
1837:
1825:
1824:
1814:
1813:
1812:
1803:
1802:
1789:
1783:
1772:
1759:
1748:
1688:is greater than
1671:
1669:
1668:
1663:
1661:
1659:
1636:
1619:
1613:
1602:
1580:
1578:
1577:
1572:
1535:
1533:
1532:
1527:
1519:
1518:
1510:
1476:is equal to the
1451:
1449:
1448:
1443:
1441:
1431:
1423:
1408:
1407:
1361:
1359:
1358:
1353:
1351:
1347:
1339:
1324:
1323:
1301:
1299:
1298:
1293:
1291:
1287:
1279:
1264:
1263:
1213:
1211:
1210:
1205:
1189:
1187:
1186:
1181:
1179:
1178:
1159:
1157:
1156:
1151:
1131:
1129:
1128:
1123:
1121:
1113:
1096:
1094:
1093:
1088:
1086:
1081:
1080:
1076:
1068:
1059:
1055:
1047:
1038:
1034:
1026:
1017:
1013:
1005:
996:
992:
984:
975:
971:
963:
956:
943:
941:
940:
935:
933:
931:
930:
918:
908:
907:
887:
885:
884:
879:
877:
875:
874:
862:
859:
854:
839:
831:
826:
824:
823:
811:
792:non-overlapping
770:
768:
767:
762:
757:
755:
745:
744:
736:
729:
709:
703:
692:
673:
671:
670:
665:
654:and solving for
653:
651:
650:
645:
634:
610:
608:
607:
602:
591:
580:
579:
571:
561:
559:
558:
553:
550:
539:
523:
521:
520:
515:
504:
493:
492:
484:
471:
469:
468:
463:
458:
456:
448:
428:
414:
387:
385:
384:
379:
367:allele frequency
364:
362:
361:
356:
354:
253:coalescence time
177:
175:
174:
169:
147:
145:
144:
139:
119:coalescence time
78:
71:
67:
64:
58:
38:
37:
30:
21:
6573:
6572:
6568:
6567:
6566:
6564:
6563:
6562:
6533:
6532:
6531:
6526:
6517:
6503:Systems ecology
6391:
6362:Extinction debt
6327:Ecological debt
6317:Bioluminescence
6298:
6291:
6260:marine habitats
6235:Ecological trap
6216:
6096:
6089:
6032:
6026:
5982:Rapoport's rule
5977:Priority effect
5918:Endemic species
5886:
5845:Population size
5761:
5754:
5744:
5714:
5709:
5662:
5656:
5642:Trophic cascade
5552:Bioaccumulation
5535:
5462:
5419:
5341:
5308:
5205:
5117:
5078:Ecosystem model
5011:
4997:
4967:
4962:
4949:
4884:
4858:
4820:
4804:
4802:
4795:
4762:
4693:Genetic linkage
4676:
4671:
4641:
4636:
4605:
4569:
4560:
4517:
4515:
4501:
4488:
4485:
4480:
4422:
4421:
4417:
4410:
4397:
4396:
4392:
4359:(6): e1002740.
4346:
4345:
4341:
4300:
4299:
4295:
4259:
4258:
4254:
4218:
4217:
4213:
4183:
4182:
4178:
4132:
4131:
4127:
4081:
4080:
4076:
4066:
4065:
4061:
4023:
4022:
4018:
3974:
3973:
3969:
3941:
3940:
3936:
3921:10.2307/2406998
3906:
3905:
3901:
3847:
3846:
3839:
3783:
3782:
3775:
3728:
3727:
3723:
3699:
3698:
3694:
3650:
3645:
3644:
3640:
3630:
3628:
3618:
3617:
3613:
3609:
3592:
3579:
3547:
3546:
3527:
3526:
3515:
3496:
3494:
3427:
3423:
3388:
3387:
3352:
3351:
3335:
3332:
3329:
3327:
3256:
3252:
3217:
3216:
3181:
3180:
3119:
3115:
3105:
3060:
3059:
3043:
3036:
3024:
2987:
2961:
2894:
2880:
2879:
2848:
2847:
2816:
2806:
2774:
2773:
2767:generation time
2738:
2733:
2732:
2709:
2708:
2684:
2679:
2678:
2655:
2654:
2627:
2622:
2621:
2614:
2602:
2560:
2544:
2535:
2531:
2527:
2493:
2492:
2444:
2418:
2402:
2395:
2394:
2364:
2338:
2322:
2315:
2314:
2273:
2261:
2257:
2256:
2246:
2233:
2220:
2209:
2208:
2168:
2146:
2142:
2113:
2106:
2100:
2077:
2072:
2071:
2063:
2049:
2038:
2002:
2001:
1967:
1966:
1937:
1920:
1910:
1909:
1887:
1882:
1881:
1872:
1863:
1829:
1816:
1815:
1804:
1794:
1790:
1735:
1734:
1720:
1709:
1700:
1687:
1637:
1620:
1589:
1588:
1545:
1544:
1485:
1484:
1458:
1439:
1438:
1399:
1394:
1393:
1384:
1349:
1348:
1315:
1310:
1309:
1289:
1288:
1255:
1250:
1249:
1235:
1223:arithmetic mean
1193:
1192:
1170:
1165:
1164:
1139:
1138:
1104:
1103:
1078:
1077:
1057:
1056:
1036:
1035:
1015:
1014:
994:
993:
973:
972:
957:
947:
946:
922:
912:
911:
866:
815:
805:
804:
786:
777:
730:
710:
679:
678:
656:
655:
627:
613:
612:
584:
564:
563:
526:
525:
497:
477:
476:
449:
429:
407:
393:
392:
370:
369:
347:
342:
341:
330:
311:
293:
278:
274:
259:divided by the
205:
189:in 1931 by the
154:
153:
130:
129:
97:
79:
68:
62:
59:
51:help improve it
48:
39:
35:
28:
23:
22:
15:
12:
11:
5:
6571:
6569:
6561:
6560:
6555:
6550:
6545:
6535:
6534:
6528:
6527:
6522:
6519:
6518:
6516:
6515:
6510:
6505:
6500:
6495:
6490:
6485:
6483:Microecosystem
6480:
6475:
6470:
6465:
6460:
6455:
6450:
6445:
6440:
6435:
6430:
6425:
6420:
6415:
6410:
6405:
6399:
6397:
6393:
6392:
6390:
6389:
6384:
6382:Thorson's rule
6379:
6374:
6369:
6364:
6359:
6354:
6349:
6344:
6339:
6334:
6329:
6324:
6319:
6314:
6309:
6307:Assembly rules
6303:
6301:
6293:
6292:
6290:
6289:
6284:
6279:
6274:
6269:
6264:
6263:
6262:
6252:
6247:
6242:
6237:
6232:
6226:
6224:
6218:
6217:
6215:
6214:
6209:
6204:
6192:
6190:Patch dynamics
6187:
6185:Metapopulation
6182:
6177:
6172:
6167:
6162:
6157:
6152:
6147:
6142:
6137:
6132:
6127:
6122:
6117:
6112:
6107:
6101:
6099:
6091:
6090:
6088:
6087:
6082:
6080:Storage effect
6077:
6072:
6067:
6062:
6057:
6052:
6047:
6042:
6036:
6034:
6028:
6027:
6025:
6024:
6019:
6014:
6009:
6004:
5999:
5994:
5989:
5984:
5979:
5974:
5969:
5964:
5962:Neutral theory
5959:
5954:
5949:
5947:Native species
5940:
5935:
5930:
5925:
5920:
5915:
5910:
5905:
5900:
5894:
5892:
5888:
5887:
5885:
5884:
5879:
5878:
5877:
5872:
5862:
5857:
5852:
5847:
5842:
5837:
5832:
5827:
5822:
5820:Overpopulation
5817:
5812:
5807:
5802:
5797:
5792:
5787:
5782:
5777:
5772:
5766:
5764:
5756:
5755:
5745:
5743:
5742:
5735:
5728:
5720:
5711:
5710:
5708:
5707:
5702:
5697:
5692:
5687:
5682:
5677:
5672:
5666:
5664:
5658:
5657:
5655:
5654:
5649:
5644:
5639:
5634:
5629:
5627:Nutrient cycle
5624:
5619:
5617:Feeding frenzy
5614:
5609:
5604:
5599:
5597:Energy quality
5594:
5589:
5584:
5579:
5574:
5569:
5564:
5559:
5557:Cascade effect
5554:
5549:
5543:
5541:
5537:
5536:
5534:
5533:
5532:
5531:
5526:
5521:
5516:
5511:
5506:
5501:
5491:
5486:
5481:
5476:
5470:
5468:
5464:
5463:
5461:
5460:
5455:
5450:
5445:
5440:
5435:
5429:
5427:
5421:
5420:
5418:
5417:
5412:
5407:
5402:
5400:Microbial loop
5397:
5392:
5387:
5382:
5377:
5372:
5367:
5365:Lithoautotroph
5362:
5357:
5351:
5349:
5347:Microorganisms
5343:
5342:
5340:
5339:
5334:
5329:
5324:
5318:
5316:
5310:
5309:
5307:
5306:
5304:Prey switching
5301:
5296:
5291:
5286:
5281:
5276:
5271:
5266:
5261:
5256:
5251:
5246:
5241:
5236:
5231:
5226:
5221:
5215:
5213:
5207:
5206:
5204:
5203:
5198:
5193:
5188:
5183:
5181:Photosynthesis
5178:
5173:
5168:
5163:
5158:
5153:
5148:
5143:
5138:
5136:Chemosynthesis
5133:
5127:
5125:
5119:
5118:
5116:
5115:
5110:
5105:
5100:
5095:
5090:
5085:
5080:
5075:
5070:
5065:
5060:
5055:
5050:
5045:
5040:
5035:
5030:
5028:Abiotic stress
5025:
5019:
5017:
5013:
5012:
4998:
4996:
4995:
4988:
4981:
4973:
4964:
4963:
4961:
4960:
4954:
4951:
4950:
4948:
4947:
4942:
4940:Phylogeography
4937:
4932:
4930:Microevolution
4927:
4918:
4913:
4908:
4903:
4898:
4892:
4890:
4889:Related topics
4886:
4885:
4883:
4882:
4877:
4872:
4866:
4864:
4860:
4859:
4857:
4856:
4851:
4846:
4844:Founder effect
4841:
4836:
4830:
4828:
4822:
4821:
4819:
4818:
4813:
4807:
4805:
4800:
4797:
4796:
4794:
4793:
4788:
4783:
4778:
4772:
4770:
4764:
4763:
4761:
4760:
4755:
4750:
4745:
4740:
4735:
4730:
4725:
4723:Price equation
4720:
4715:
4713:Neutral theory
4710:
4705:
4700:
4695:
4690:
4684:
4682:
4678:
4677:
4672:
4670:
4669:
4662:
4655:
4647:
4638:
4637:
4635:
4634:
4629:
4624:
4619:
4613:
4611:
4610:Related Topics
4607:
4606:
4604:
4603:
4598:
4596:Candidate gene
4593:
4588:
4583:
4577:
4575:
4571:
4570:
4561:
4559:
4558:
4551:
4544:
4536:
4530:
4529:
4523:
4499:
4497:on 2005-05-24.
4484:
4483:External links
4481:
4479:
4478:
4415:
4408:
4390:
4339:
4293:
4272:(2): 255–265.
4252:
4211:
4176:
4147:(3): 581–591.
4125:
4096:(4): 581–597.
4074:
4059:
4038:10.1086/282557
4016:
3967:
3934:
3915:(4): 367–402.
3899:
3860:(4): 975–996.
3837:
3807:10.1.1.135.974
3773:
3744:(3): 609–611.
3721:
3692:
3638:
3610:
3608:
3605:
3604:
3603:
3598:
3591:
3588:
3578:
3575:
3554:
3534:
3514:
3511:
3510:
3509:
3493:
3489:
3484:
3476:
3471:
3467:
3463:
3458:
3451:
3446:
3443:
3440:
3436:
3432:
3426:
3420:
3415:
3410:
3405:
3402:
3399:
3395:
3391:
3384:
3379:
3374:
3369:
3366:
3363:
3359:
3355:
3348:
3344:
3340:
3334:
3331:
3330:
3317:
3313:
3305:
3300:
3296:
3292:
3287:
3280:
3275:
3272:
3269:
3265:
3261:
3255:
3249:
3244:
3239:
3234:
3231:
3228:
3224:
3220:
3213:
3208:
3203:
3198:
3195:
3192:
3188:
3184:
3177:
3173:
3169:
3162:
3157:
3153:
3149:
3144:
3137:
3132:
3128:
3124:
3118:
3111:
3108:
3104:
3099:
3092:
3089:
3086:
3081:
3077:
3073:
3068:
3067:
3041:
3034:
3023:
3020:
3019:
3018:
3007:
3001:
2994:
2990:
2986:
2981:
2974:
2971:
2968:
2964:
2960:
2955:
2950:
2945:
2942:
2939:
2935:
2929:
2924:
2921:
2918:
2914:
2908:
2904:
2900:
2897:
2892:
2887:
2883:
2876:
2871:
2868:
2865:
2860:
2856:
2841:
2840:
2828:
2823:
2819:
2813:
2809:
2803:
2798:
2795:
2792:
2788:
2784:
2781:
2763:
2762:
2750:
2745:
2741:
2729:
2728:
2716:
2696:
2691:
2687:
2675:
2674:
2662:
2639:
2634:
2630:
2613:
2610:
2601:
2595:
2591:
2590:
2575:
2567:
2563:
2557:
2554:
2551:
2547:
2541:
2538:
2534:
2530:
2526:
2521:
2516:
2513:
2510:
2505:
2501:
2486:
2485:
2474:
2466:
2463:
2460:
2455:
2451:
2447:
2443:
2438:
2435:
2432:
2425:
2421:
2415:
2412:
2409:
2405:
2388:
2387:
2376:
2370:
2367:
2363:
2358:
2355:
2352:
2345:
2341:
2335:
2332:
2329:
2325:
2308:
2307:
2296:
2291:
2286:
2279:
2276:
2272:
2267:
2264:
2260:
2253:
2249:
2245:
2240:
2236:
2232:
2227:
2223:
2219:
2216:
2198:
2197:
2186:
2181:
2178:
2175:
2171:
2166:
2160:
2157:
2152:
2149:
2145:
2141:
2137:
2132:
2126:
2123:
2120:
2116:
2112:
2109:
2103:
2097:
2094:
2089:
2084:
2080:
2059:
2048:
2045:
2034:
2028:
2027:
2024:
2023:
2012:
2009:
1999:
1996:
1995:
1982:
1979:
1974:
1964:
1961:
1960:
1946:
1943:
1940:
1935:
1932:
1929:
1926:
1923:
1917:
1907:
1894:
1890:
1868:
1859:
1853:
1852:
1836:
1832:
1828:
1823:
1819:
1811:
1807:
1801:
1797:
1793:
1787:
1782:
1779:
1776:
1771:
1767:
1763:
1758:
1755:
1752:
1747:
1743:
1719:
1716:
1705:
1696:
1683:
1673:
1672:
1658:
1655:
1652:
1649:
1646:
1643:
1640:
1635:
1632:
1629:
1626:
1623:
1617:
1612:
1609:
1606:
1601:
1597:
1582:
1581:
1570:
1567:
1564:
1561:
1558:
1555:
1552:
1537:
1536:
1525:
1522:
1516:
1513:
1507:
1504:
1501:
1498:
1495:
1492:
1457:
1454:
1453:
1452:
1437:
1434:
1429:
1426:
1421:
1420:
1417:
1414:
1411:
1406:
1402:
1380:
1363:
1362:
1345:
1342:
1337:
1336:
1333:
1330:
1327:
1322:
1318:
1303:
1302:
1285:
1282:
1277:
1276:
1273:
1270:
1267:
1262:
1258:
1234:
1231:
1219:
1218:
1215:
1214:
1203:
1200:
1190:
1177:
1173:
1161:
1160:
1149:
1146:
1136:
1133:
1132:
1119:
1116:
1111:
1101:
1098:
1097:
1084:
1074:
1071:
1066:
1065:
1062:
1053:
1050:
1045:
1044:
1041:
1032:
1029:
1024:
1023:
1020:
1011:
1008:
1003:
1002:
999:
990:
987:
982:
981:
978:
969:
966:
961:
960:
954:
944:
929:
925:
921:
889:
888:
873:
869:
865:
858:
853:
850:
847:
843:
837:
834:
829:
822:
818:
814:
785:
782:
776:
773:
772:
771:
760:
754:
751:
748:
742:
739:
733:
728:
725:
722:
719:
716:
713:
707:
702:
699:
696:
691:
687:
663:
643:
640:
637:
633:
630:
626:
623:
620:
600:
597:
594:
590:
587:
583:
577:
574:
549:
546:
543:
538:
534:
513:
510:
507:
503:
500:
496:
490:
487:
473:
472:
461:
455:
452:
447:
444:
441:
438:
435:
432:
426:
423:
420:
417:
413:
410:
406:
403:
400:
377:
353:
350:
329:
326:
307:
292:
289:
276:
272:
204:
201:
167:
164:
161:
150:heterozygosity
137:
93:
81:
80:
42:
40:
33:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
6570:
6559:
6556:
6554:
6551:
6549:
6546:
6544:
6541:
6540:
6538:
6525:
6520:
6514:
6511:
6509:
6508:Urban ecology
6506:
6504:
6501:
6499:
6496:
6494:
6491:
6489:
6486:
6484:
6481:
6479:
6476:
6474:
6471:
6469:
6466:
6464:
6461:
6459:
6456:
6454:
6451:
6449:
6446:
6444:
6441:
6439:
6436:
6434:
6431:
6429:
6426:
6424:
6421:
6419:
6416:
6414:
6411:
6409:
6406:
6404:
6401:
6400:
6398:
6394:
6388:
6385:
6383:
6380:
6378:
6375:
6373:
6370:
6368:
6367:Kleiber's law
6365:
6363:
6360:
6358:
6355:
6353:
6350:
6348:
6345:
6343:
6340:
6338:
6335:
6333:
6330:
6328:
6325:
6323:
6320:
6318:
6315:
6313:
6310:
6308:
6305:
6304:
6302:
6300:
6294:
6288:
6285:
6283:
6280:
6278:
6275:
6273:
6270:
6268:
6265:
6261:
6258:
6257:
6256:
6253:
6251:
6248:
6246:
6243:
6241:
6238:
6236:
6233:
6231:
6228:
6227:
6225:
6223:
6219:
6213:
6210:
6208:
6205:
6203:
6201:
6197:
6193:
6191:
6188:
6186:
6183:
6181:
6178:
6176:
6173:
6171:
6168:
6166:
6163:
6161:
6158:
6156:
6153:
6151:
6148:
6146:
6143:
6141:
6140:Foster's rule
6138:
6136:
6133:
6131:
6128:
6126:
6123:
6121:
6118:
6116:
6113:
6111:
6108:
6106:
6103:
6102:
6100:
6098:
6092:
6086:
6083:
6081:
6078:
6076:
6073:
6071:
6068:
6066:
6063:
6061:
6058:
6056:
6053:
6051:
6048:
6046:
6043:
6041:
6038:
6037:
6035:
6029:
6023:
6020:
6018:
6015:
6013:
6010:
6008:
6005:
6003:
6000:
5998:
5995:
5993:
5990:
5988:
5985:
5983:
5980:
5978:
5975:
5973:
5970:
5968:
5965:
5963:
5960:
5958:
5955:
5953:
5950:
5948:
5944:
5941:
5939:
5936:
5934:
5931:
5929:
5926:
5924:
5921:
5919:
5916:
5914:
5911:
5909:
5906:
5904:
5901:
5899:
5896:
5895:
5893:
5889:
5883:
5880:
5876:
5873:
5871:
5868:
5867:
5866:
5863:
5861:
5858:
5856:
5853:
5851:
5848:
5846:
5843:
5841:
5838:
5836:
5833:
5831:
5828:
5826:
5823:
5821:
5818:
5816:
5813:
5811:
5808:
5806:
5803:
5801:
5798:
5796:
5793:
5791:
5788:
5786:
5783:
5781:
5778:
5776:
5773:
5771:
5768:
5767:
5765:
5763:
5757:
5752:
5748:
5741:
5736:
5734:
5729:
5727:
5722:
5721:
5718:
5706:
5703:
5701:
5698:
5696:
5693:
5691:
5688:
5686:
5683:
5681:
5678:
5676:
5673:
5671:
5668:
5667:
5665:
5659:
5653:
5650:
5648:
5645:
5643:
5640:
5638:
5635:
5633:
5630:
5628:
5625:
5623:
5620:
5618:
5615:
5613:
5610:
5608:
5605:
5603:
5600:
5598:
5595:
5593:
5590:
5588:
5585:
5583:
5580:
5578:
5575:
5573:
5570:
5568:
5565:
5563:
5560:
5558:
5555:
5553:
5550:
5548:
5545:
5544:
5542:
5538:
5530:
5527:
5525:
5522:
5520:
5517:
5515:
5512:
5510:
5507:
5505:
5502:
5500:
5497:
5496:
5495:
5492:
5490:
5487:
5485:
5482:
5480:
5477:
5475:
5472:
5471:
5469:
5465:
5459:
5458:Trophic level
5456:
5454:
5451:
5449:
5446:
5444:
5441:
5439:
5436:
5434:
5431:
5430:
5428:
5426:
5422:
5416:
5415:Phage ecology
5413:
5411:
5408:
5406:
5405:Microbial mat
5403:
5401:
5398:
5396:
5393:
5391:
5388:
5386:
5383:
5381:
5378:
5376:
5373:
5371:
5368:
5366:
5363:
5361:
5360:Bacteriophage
5358:
5356:
5353:
5352:
5350:
5348:
5344:
5338:
5335:
5333:
5330:
5328:
5327:Decomposition
5325:
5323:
5320:
5319:
5317:
5315:
5311:
5305:
5302:
5300:
5297:
5295:
5292:
5290:
5287:
5285:
5282:
5280:
5277:
5275:
5274:Mesopredators
5272:
5270:
5267:
5265:
5262:
5260:
5257:
5255:
5252:
5250:
5247:
5245:
5242:
5240:
5237:
5235:
5232:
5230:
5227:
5225:
5222:
5220:
5219:Apex predator
5217:
5216:
5214:
5212:
5208:
5202:
5199:
5197:
5194:
5192:
5189:
5187:
5184:
5182:
5179:
5177:
5174:
5172:
5169:
5167:
5164:
5162:
5159:
5157:
5154:
5152:
5149:
5147:
5144:
5142:
5139:
5137:
5134:
5132:
5129:
5128:
5126:
5124:
5120:
5114:
5111:
5109:
5106:
5104:
5101:
5099:
5096:
5094:
5091:
5089:
5086:
5084:
5081:
5079:
5076:
5074:
5071:
5069:
5066:
5064:
5061:
5059:
5056:
5054:
5053:Biotic stress
5051:
5049:
5046:
5044:
5041:
5039:
5036:
5034:
5031:
5029:
5026:
5024:
5021:
5020:
5018:
5014:
5009:
5005:
5001:
4994:
4989:
4987:
4982:
4980:
4975:
4974:
4971:
4959:
4956:
4955:
4952:
4946:
4943:
4941:
4938:
4936:
4933:
4931:
4928:
4926:
4922:
4919:
4917:
4914:
4912:
4909:
4907:
4904:
4902:
4899:
4897:
4894:
4893:
4891:
4887:
4881:
4880:Sewall Wright
4878:
4876:
4873:
4871:
4868:
4867:
4865:
4861:
4855:
4852:
4850:
4847:
4845:
4842:
4840:
4837:
4835:
4832:
4831:
4829:
4827:
4826:Genetic drift
4823:
4817:
4814:
4812:
4809:
4808:
4806:
4798:
4792:
4789:
4787:
4784:
4782:
4779:
4777:
4774:
4773:
4771:
4769:
4765:
4759:
4756:
4754:
4751:
4749:
4746:
4744:
4741:
4739:
4736:
4734:
4731:
4729:
4726:
4724:
4721:
4719:
4716:
4714:
4711:
4709:
4706:
4704:
4701:
4699:
4696:
4694:
4691:
4689:
4686:
4685:
4683:
4679:
4675:
4668:
4663:
4661:
4656:
4654:
4649:
4648:
4645:
4633:
4630:
4628:
4625:
4623:
4620:
4618:
4615:
4614:
4612:
4608:
4602:
4599:
4597:
4594:
4592:
4589:
4587:
4584:
4582:
4579:
4578:
4576:
4572:
4568:
4564:
4557:
4552:
4550:
4545:
4543:
4538:
4537:
4534:
4527:
4524:
4514:on 2009-07-23
4513:
4509:
4505:
4500:
4496:
4492:
4487:
4486:
4482:
4474:
4470:
4465:
4460:
4455:
4450:
4446:
4442:
4438:
4434:
4430:
4426:
4419:
4416:
4411:
4405:
4401:
4394:
4391:
4386:
4382:
4377:
4372:
4367:
4362:
4358:
4354:
4353:PLOS Genetics
4350:
4343:
4340:
4335:
4331:
4327:
4323:
4319:
4315:
4312:(8): 701–10.
4311:
4307:
4303:
4302:Masel, Joanna
4297:
4294:
4289:
4285:
4280:
4275:
4271:
4267:
4263:
4256:
4253:
4248:
4244:
4239:
4234:
4230:
4226:
4222:
4215:
4212:
4207:
4203:
4199:
4195:
4192:(1): 86–107.
4191:
4187:
4180:
4177:
4172:
4168:
4163:
4158:
4154:
4150:
4146:
4142:
4141:
4136:
4129:
4126:
4121:
4117:
4112:
4107:
4103:
4099:
4095:
4091:
4090:
4085:
4078:
4075:
4070:
4063:
4060:
4055:
4051:
4047:
4043:
4039:
4035:
4031:
4027:
4020:
4017:
4012:
4008:
4003:
3998:
3994:
3990:
3986:
3982:
3981:Proc Biol Sci
3978:
3971:
3968:
3962:
3957:
3954:(2): 95–107.
3953:
3949:
3945:
3938:
3935:
3930:
3926:
3922:
3918:
3914:
3910:
3903:
3900:
3895:
3891:
3886:
3881:
3877:
3873:
3868:
3863:
3859:
3855:
3851:
3844:
3842:
3838:
3833:
3829:
3825:
3821:
3817:
3813:
3808:
3803:
3799:
3795:
3791:
3787:
3780:
3778:
3774:
3769:
3765:
3760:
3755:
3751:
3747:
3743:
3739:
3735:
3731:
3730:James F. Crow
3725:
3722:
3717:
3713:
3709:
3705:
3704:
3696:
3693:
3688:
3684:
3679:
3674:
3670:
3666:
3663:(2): 97–159.
3662:
3658:
3657:
3649:
3642:
3639:
3627:
3626:
3621:
3615:
3612:
3606:
3602:
3599:
3597:
3594:
3593:
3589:
3587:
3584:
3576:
3574:
3572:
3566:
3552:
3532:
3524:
3520:
3512:
3491:
3487:
3482:
3474:
3469:
3465:
3461:
3456:
3449:
3444:
3441:
3438:
3434:
3430:
3424:
3418:
3413:
3408:
3403:
3400:
3397:
3393:
3389:
3382:
3377:
3372:
3367:
3364:
3361:
3357:
3353:
3346:
3342:
3338:
3311:
3303:
3298:
3294:
3290:
3285:
3278:
3273:
3270:
3267:
3263:
3259:
3253:
3247:
3242:
3237:
3232:
3229:
3226:
3222:
3218:
3211:
3206:
3201:
3196:
3193:
3190:
3186:
3182:
3175:
3171:
3167:
3160:
3155:
3151:
3147:
3142:
3135:
3130:
3126:
3122:
3116:
3109:
3106:
3102:
3097:
3087:
3079:
3075:
3071:
3058:
3057:
3056:
3053:
3051:
3047:
3040:
3033:
3028:
3021:
3005:
2992:
2988:
2984:
2979:
2972:
2969:
2966:
2962:
2958:
2948:
2943:
2940:
2937:
2933:
2927:
2922:
2919:
2916:
2912:
2906:
2902:
2898:
2895:
2890:
2885:
2881:
2874:
2866:
2858:
2854:
2846:
2845:
2844:
2826:
2821:
2817:
2811:
2807:
2796:
2793:
2790:
2786:
2782:
2779:
2772:
2771:
2770:
2768:
2748:
2743:
2739:
2731:
2730:
2714:
2694:
2689:
2685:
2677:
2676:
2660:
2652:
2637:
2632:
2628:
2620:
2619:
2618:
2611:
2609:
2607:
2599:
2596:
2594:
2573:
2565:
2561:
2555:
2552:
2549:
2545:
2539:
2536:
2532:
2528:
2524:
2519:
2511:
2503:
2499:
2491:
2490:
2489:
2472:
2461:
2453:
2449:
2445:
2441:
2436:
2433:
2430:
2423:
2419:
2413:
2410:
2407:
2403:
2393:
2392:
2391:
2374:
2368:
2365:
2361:
2356:
2353:
2350:
2343:
2339:
2333:
2330:
2327:
2323:
2313:
2312:
2311:
2294:
2289:
2284:
2277:
2274:
2270:
2265:
2262:
2258:
2251:
2247:
2243:
2238:
2234:
2230:
2225:
2221:
2217:
2214:
2207:
2206:
2205:
2203:
2184:
2179:
2176:
2173:
2169:
2164:
2158:
2155:
2150:
2147:
2143:
2139:
2135:
2130:
2124:
2121:
2118:
2114:
2110:
2107:
2101:
2095:
2092:
2087:
2082:
2078:
2070:
2069:
2068:
2065:
2062:
2058:
2054:
2046:
2044:
2042:
2037:
2033:
2010:
2007:
2000:
1998:
1997:
1980:
1977:
1972:
1965:
1963:
1962:
1944:
1941:
1938:
1933:
1930:
1927:
1924:
1921:
1915:
1908:
1892:
1888:
1880:
1879:
1876:
1875:
1874:
1871:
1867:
1862:
1858:
1834:
1830:
1826:
1821:
1817:
1809:
1805:
1799:
1795:
1791:
1785:
1777:
1769:
1765:
1761:
1753:
1745:
1741:
1733:
1732:
1731:
1729:
1725:
1717:
1715:
1713:
1708:
1704:
1699:
1695:
1691:
1686:
1682:
1678:
1653:
1647:
1644:
1641:
1638:
1633:
1630:
1627:
1624:
1621:
1615:
1607:
1599:
1595:
1587:
1586:
1585:
1568:
1565:
1559:
1553:
1550:
1543:
1542:
1541:
1523:
1520:
1511:
1505:
1499:
1493:
1490:
1483:
1482:
1481:
1479:
1475:
1471:
1467:
1463:
1455:
1435:
1432:
1427:
1424:
1415:
1412:
1409:
1404:
1400:
1392:
1391:
1390:
1388:
1383:
1379:
1375:
1370:
1368:
1343:
1340:
1331:
1328:
1325:
1320:
1316:
1308:
1307:
1306:
1283:
1280:
1271:
1268:
1265:
1260:
1256:
1248:
1247:
1246:
1244:
1240:
1233:Dioeciousness
1232:
1230:
1228:
1224:
1201:
1198:
1191:
1175:
1171:
1163:
1162:
1147:
1144:
1137:
1135:
1134:
1117:
1114:
1109:
1102:
1100:
1099:
1082:
1072:
1069:
1060:
1051:
1048:
1039:
1030:
1027:
1018:
1009:
1006:
997:
988:
985:
976:
967:
964:
952:
945:
927:
923:
919:
910:
909:
906:
905:
904:
902:
901:harmonic mean
898:
894:
871:
867:
863:
856:
851:
848:
845:
841:
835:
832:
827:
820:
816:
812:
803:
802:
801:
799:
798:harmonic mean
795:
791:
783:
781:
774:
758:
749:
740:
737:
731:
723:
720:
717:
711:
705:
697:
689:
685:
677:
676:
675:
661:
638:
635:
631:
628:
621:
618:
595:
592:
588:
585:
575:
572:
544:
536:
532:
508:
505:
501:
498:
488:
485:
459:
453:
450:
442:
439:
436:
430:
424:
418:
415:
411:
408:
401:
398:
391:
390:
389:
375:
368:
351:
348:
339:
335:
327:
325:
323:
318:
315:
310:
306:
301:
298:
290:
288:
286:
285:genetic drift
282:
270:
266:
263:, yielding a
262:
261:mutation rate
258:
254:
250:
245:
243:
239:
235:
230:
226:
225:genetic drift
223:under random
222:
218:
214:
213:Sewall Wright
210:
209:Ronald Fisher
202:
200:
198:
197:Sewall Wright
195:
192:
188:
183:
181:
165:
162:
159:
152:is equal to
151:
135:
128:
127:mutation rate
124:
120:
116:
110:
108:
105:
101:
96:
92:
88:
77:
74:
66:
63:November 2020
56:
52:
46:
43:This article
41:
32:
31:
19:
6493:Regime shift
6478:Macroecology
6199:
6195:
6135:Edge effects
6105:Biogeography
6050:Commensalism
5898:Biodiversity
5794:
5775:Allee effect
5514:kelp forests
5467:Example webs
5332:Detritivores
5171:Organotrophs
5151:Kinetotrophs
5103:Productivity
4896:Biogeography
4870:R. A. Fisher
4748:Heritability
4681:Key concepts
4600:
4581:Heritability
4516:. Retrieved
4512:the original
4507:
4495:the original
4436:
4432:
4418:
4399:
4393:
4356:
4352:
4342:
4309:
4305:
4296:
4269:
4265:
4255:
4228:
4224:
4214:
4189:
4185:
4179:
4144:
4138:
4128:
4093:
4087:
4077:
4068:
4062:
4029:
4025:
4019:
3984:
3980:
3970:
3951:
3947:
3937:
3912:
3908:
3902:
3857:
3853:
3789:
3785:
3741:
3737:
3724:
3707:
3701:
3695:
3660:
3654:
3641:
3629:. Retrieved
3623:
3614:
3580:
3567:
3516:
3054:
3049:
3045:
3038:
3031:
3029:
3025:
2842:
2764:
2615:
2603:
2592:
2487:
2389:
2309:
2201:
2199:
2066:
2060:
2056:
2050:
2040:
2035:
2031:
2029:
1869:
1865:
1860:
1856:
1854:
1721:
1711:
1706:
1702:
1697:
1693:
1689:
1684:
1680:
1676:
1674:
1583:
1538:
1473:
1468:so that the
1459:
1386:
1381:
1377:
1373:
1371:
1366:
1364:
1304:
1236:
1220:
896:
892:
890:
789:
787:
778:
674:which gives
474:
365:, given the
331:
319:
313:
308:
304:
302:
296:
294:
268:
264:
246:
242:F-statistics
237:
233:
206:
184:
111:
106:
94:
90:
86:
84:
69:
60:
44:
6130:Disturbance
6033:interaction
5855:Recruitment
5785:Depensation
5577:Copiotrophs
5448:Energy flow
5370:Lithotrophy
5314:Decomposers
5294:Planktivore
5269:Insectivore
5259:Heterotroph
5224:Bacterivore
5191:Phototrophs
5141:Chemotrophs
5113:Restoration
5063:Competition
4849:Coalescence
4423:Rajon, E.;
2606:life tables
2598:Theoretical
1148:0.032416667
794:generations
6537:Categories
6498:Sexecology
6075:Parasitism
6040:Antibiosis
5875:Resistance
5870:Resilience
5760:Population
5680:Camouflage
5632:Oligotroph
5547:Ascendency
5509:intertidal
5499:cold seeps
5453:Food chain
5254:Herbivores
5229:Carnivores
5156:Mixotrophs
5131:Autotrophs
5010:components
4791:Ecological
4781:Artificial
4518:2005-02-25
3607:References
3545:Ne, where
3525:equal to 4
1376:is large,
1227:bottleneck
297:Drosophila
229:inbreeding
194:geneticist
6403:Allometry
6357:Emergence
6085:Symbiosis
6070:Mutualism
5865:Stability
5770:Abundance
5582:Dominance
5540:Processes
5529:tide pool
5425:Food webs
5299:Predation
5284:Omnivores
5211:Consumers
5166:Mycotroph
5123:Producers
5068:Ecosystem
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4306:BioEssays
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