352:, then mutations affect only one trait at a time, and adaptation is much less constrained. In a modular gene network, for example, a gene that induces a limited set of other genes that control a specific trait under selection may evolve more readily than one that also induces other gene pathways controlling traits not under selection. Individual genes also exhibit modularity. A mutation in one cis-regulatory element of a gene's promoter region may allow the expression of the gene to be altered only in specific tissues, developmental stages, or environmental conditions rather than changing gene activity in the entire organism simultaneously.
432:. Identifying evolvable proteins and manipulating their evolvability is becoming increasingly necessary in order to achieve ever larger functional modification of enzymes. Proteins are also often studied as part of the basic science of evolvability, because the biophysical properties and chemical functions can be easily changed by a few mutations. More evolvable proteins can tolerate a broader range of amino acid changes and allow them to evolve toward new functions. The study of evolvability has fundamental importance for understanding very long term evolution of
369:. Short-term selection for low variation most of the time is likely to be more powerful than long-term selection for evolvability, making it difficult for natural selection to cause the evolution of evolvability. Other forces of selection also affect the generation of variation; for example, mutation and recombination may in part be byproducts of mechanisms to cope with DNA damage.
258:, increasing evolvability according to the second sense. Even without genetic diversity, some genotypes have higher evolvability than others, and selection for robustness can increase the "neighborhood richness" of phenotypes that can be accessed from the same starting genotype by mutation. For example, one reason many proteins are less robust to mutation is that they have marginal
303:. When this happens, natural selection weeds out the very bad mutations, while leaving the others relatively unaffected. While evolution has no "foresight" to know which environment will be encountered in the future, some mutations cause major disruption to a basic biological process, and will never be adaptive in any environment. Screening these out in advance leads to
399:
the risk that a future environment will be similar or different. Theoretical models also predict the evolution of evolvability via modularity. When the costs of evolvability are sufficiently short-lived, more evolvable lineages may be the most successful in the long-term. However, the hypothesis that
131:
Pigliucci's second definition of evolvability includes
Altenberg's quantitative concept of evolvability, being not a single number, but the entire upper tail of the fitness distribution of the offspring produced by the population. This quantity was considered a "local" property of the instantaneous
113:
in the population. Each allele catalyzes the same reaction, but with a different level of activity. However, even after millions of years of evolution, exploring many sequences with similar function, no mutation might exist that gives this enzyme the ability to catalyze a different reaction. Thus,
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exists, many mutants will persist in a cryptic state. Mutations tend to fall into two categories, having either a very bad effect or very little effect: few mutations fall somewhere in between. Sometimes, these mutations will not be completely invisible, but still have rare effects, with very low
140:
More heritable phenotypic variation means more evolvability. While mutation is the ultimate source of heritable variation, its permutations and combinations also make a big difference. Sexual reproduction generates more variation (and thereby evolvability) relative to asexual reproduction (see
310:
Another way that phenotypes can be explored, prior to strong genetic commitment, is through learning. An organism that learns gets to "sample" several different phenotypes during its early development, and later sticks to whatever worked best. Later in evolution, the optimal phenotype can be
253:
does not increase evolvability in the first sense. In organisms with a high level of robustness, mutations have smaller phenotypic effects than in organisms with a low level of robustness. Thus, robustness reduces the amount of heritable genetic variation on which selection can act. However,
262:, and most mutations reduce this stability further. Proteins that are more thermostable can tolerate a wider range of mutations and are more evolvable. For polygenic traits, neighborhood richness contributes more to evolvability than does genetic diversity or "spread" across genotype space.
66:
cannot occur. Early failed efforts to evolve computer programs by random mutation and selection showed that evolvability is not a given, but depends on the representation of the program as a data structure, because this determines how changes in the program map to changes in its behavior.
145:). Evolvability is further increased by generating more variation when an organism is stressed, and thus likely to be less well adapted, but less variation when an organism is doing well. The amount of variation generated can be adjusted in many different ways, for example via the
412:
we wish to increase evolvability, and in medicine and agriculture we wish to decrease it. Protein evolvability is defined as the ability of the protein to acquire sequence diversity and conformational flexibility which can enable it to evolve toward a new function.
326:
can also increase evolvability even when it has no direction, for example when the flattening is a result of random errors in molecular and/or developmental processes. This increase in evolvability can happen when evolution is faced with crossing a "valley" in an
114:
although the enzyme's activity is evolvable in the first sense, that does not mean that the enzyme's function is evolvable in the second sense. However, every system evolvable in the second sense must also be evolvable in the first.
331:. This means that two mutations exist that are deleterious by themselves, but beneficial in combination. These combinations can evolve more easily when the landscape is first flattened, and the discovered phenotype is then fixed by
463:. Predicting the evolution and evolvability of our pathogens, and devising strategies to slow or circumvent the development of resistance, demands deeper knowledge of the complex forces driving evolution at the molecular level.
343:
If every mutation affected every trait, then a mutation that was an improvement for one trait would be a disadvantage for other traits. This means that almost no mutations would be beneficial overall. But if
360:
While variation yielding high evolvability could be useful in the long term, in the short term most of that variation is likely to be a disadvantage. For example, naively it would seem that increasing the
241:
The relationship between robustness and evolvability depends on whether recombination can be ignored. Recombination can generally be ignored in asexual populations and for traits affected by single genes.
2844:
Pan D, Xue W, Zhang W, Liu H, Yao X (October 2012). "Understanding the drug resistance mechanism of hepatitis C virus NS3/4A to ITMN-191 due to R155K, A156V, D168A/E mutations: a computational study".
132:
state of a population, and its integration over the population's evolutionary trajectory, and over many possible populations, would be necessary to give a more global measure of evolvability.
380:
of microbes. Mutator alleles can also evolve more easily when they only increase mutation rates in nearby DNA sequences, not across the whole genome: this is known as a contingency locus.
75:
are structured in ways that make beneficial changes more likely. This has been taken as evidence that evolution has created fitter populations of organisms that are better able to evolve.
181:
Rather than creating more phenotypic variation, some mechanisms increase the intensity and effectiveness with which selection acts on existing phenotypic variation. For example:
376:
on the success of adaptive mutations that they cause. In this case, selection can take place at the level of the lineage. This may explain why mutators are often seen during
219:, where different genotypes contain different adaptive mutations. Recombination brings the two alleles together, creating a super-genotype in place of two competing lineages.
3259:
Danchin É, Charmantier A, Champagne FA, Mesoudi A, Pujol B, Blanchet S (June 2011). "Beyond DNA: integrating inclusive inheritance into an extended theory of evolution".
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via a mutator allele would increase evolvability. But as an extreme example, if the mutation rate is too high then all individuals will be dead or at least carry a heavy
120:
recognizes three classes of definition, depending on timescale. The first corresponds to Wagner's first, and represents the very short timescales that are described by
2660:
Dellus-Gur E, Toth-Petroczy A, Elias M, Tawfik DS (July 2013). "What makes a protein fold amenable to functional innovation? Fold polarity and stability trade-offs".
2067:"The spontaneous appearance rate of the yeast prion [PSI+] and its implications for the evolution of the evolvability properties of the [PSI+] system"
54:
In order for a biological organism to evolve by natural selection, there must be a certain minimum probability that new, heritable variants are beneficial. Random
749:
274:, may lead to the accumulation of significant quantities of cryptic genetic variation. In a new environment or genetic background, this variation may be
2353:
Bommarius AS, Blum JK, Abrahamson MJ (April 2011). "Status of protein engineering for biocatalysts: how to design an industrially useful biocatalyst".
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RodrĂguez-Rojas A, RodrĂguez-Beltrán J, Couce A, Blázquez J (August 2013). "Antibiotics and antibiotic resistance: a bitter fight against evolution".
2423:
Wang X, Minasov G, Shoichet BK (June 2002). "Evolution of an antibiotic resistance enzyme constrained by stability and activity trade-offs".
286:
Different genetic codes have the potential to change robustness and evolvability by changing the effect of single-base mutational changes.
2609:
Ranea JA, Sillero A, Thornton JM, Orengo CA (October 2006). "Protein superfamily evolution and the last universal common ancestor (LUCA)".
124:. He divides Wagner's second definition into two categories, one representing the intermediate timescales that can be studied using
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204:
above which selection becomes an important player. This could happen through an increase in the census population size, decreasing
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2732:
2707:
865:
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Vaishnav ED, de Boer CG, Molinet J, Yassour M, Fan L, Adiconis X, Thompson DA, Levine JZ, Cubillos FA, Regev A (March 2022).
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3774:
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approaches aim to create changes rapidly through mutations with large effects. Such mutations, however, commonly destroy
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Fudala A, Korona R (August 2009). "Low frequency of mutations with strongly deleterious but nonlethal fitness effects".
3619:
3347:
17:
3319:
2318:
Carter PJ (May 2011). "Introduction to current and future protein therapeutics: a protein engineering perspective".
1884:"Complex adaptations can drive the evolution of the capacitor [PSI], even with realistic rates of yeast sex"
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3362:
3357:
384:
142:
467:
387:, or via the tendency of variation-generating mechanisms to become more active when an organism is stressed. The
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if it can acquire novel functions through genetic change, functions that help the organism survive and reproduce.
2499:
Salverda ML, Dellus E, Gorter FA, Debets AJ, van der Oost J, Hoekstra RF, Tawfik DS, de Visser JA (March 2011).
400:
evolvability is an adaptation is often rejected in favor of alternative hypotheses, e.g. minimization of costs.
86:
describes two definitions of evolvability. According to the first definition, a biological system is evolvable:
1360:
Whitacre J, Bender A (March 2010). "Degeneracy: a design principle for achieving robustness and evolvability".
3753:
2971:
Neve P (October 2007). "Challenges for herbicide resistance evolution and management: 50 years after Harper".
62:, are expected to be mostly detrimental. Beneficial mutations are always rare, but if they are too rare, then
3828:
3604:
2801:
Merlo LM, Pepper JW, Reid BJ, Maley CC (December 2006). "Cancer as an evolutionary and ecological process".
3571:
395:. An evolutionary capacitor is a switch that turns genetic variation on and off. This is very much like
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3629:
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3456:
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Soskine M, Tawfik DS (August 2010). "Mutational effects and the evolution of new protein functions".
197:
611:
1519:
Eyre-Walker A, Keightley PD (August 2007). "The distribution of fitness effects of new mutations".
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275:
170:
2700:
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3634:
3609:
3492:
3412:
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Tokuriki N, Tawfik DS (October 2009). "Stability effects of mutations and protein evolvability".
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resistance. It is possible that we are facing the end of the effective life of most of available
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3535:
3436:
747:
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Intelligence: Bioengineers' Ultimate Best Friend".
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2022:, Bergman A (July 2003). "The evolution of the evolvability properties of the yeast prion ".
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Labbé P, Berticat C, Berthomieu A, Unal S, Bernard C, Weill M, Lenormand T (November 2007).
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2980:
2618:
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2231:
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Draghi J, Wagner GP (February 2008). "Evolution of evolvability in a developmental model".
2131:
1933:
Michod RE (1986). "On fitness and adaptedness and their role in evolutionary explanation".
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1379:
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995:
809:
552:
332:
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Learning biases phenotypes in a beneficial direction. But an exploratory flattening of the
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2922:"Forty years of erratic insecticide resistance evolution in the mosquito Culex pipiens"
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173:. A large population size increases the influx of novel mutations in each generation.
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2005:
1950:
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Kim Y (August 2007). "Rate of adaptive peak shifts with partial genetic robustness".
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Altenberg L (1995). "Genome growth and the evolution of the genotype–phenotype map".
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so it becomes the default behavior rather than a rare behavior. This is known as the
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1962:
1817:
1741:"The effect of phenotypic plasticity on evolution in multipeaked fitness landscapes"
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1407:
1064:
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Hansen TF, PĂ©labon C, Houle D (September 2011). "Heritability is not evolvability".
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1080:"Evidence that adaptation in Drosophila is not limited by mutation at single sites"
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37:. Evolvability is the ability of a population of organisms to not merely generate
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3482:
3110:
3059:
2938:
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1900:
1096:
954:
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223:
212:, or through changes in the probability distribution of the numbers of offspring.
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2220:"Second-order selection for evolvability in a large Escherichia coli population"
2082:
1625:
1328:
3297:
3019:
2695:
2556:
Proceedings of the
National Academy of Sciences of the United States of America
2366:
2218:
Woods RJ, Barrick JE, Cooper TF, Shrestha U, Kauth MR, Lenski RE (March 2011).
1667:
Proceedings of the
National Academy of Sciences of the United States of America
1391:
1254:
Proceedings of the
National Academy of Sciences of the United States of America
821:
545:
Proceedings of the
National Academy of Sciences of the United States of America
316:
83:
59:
2673:
2630:
2401:
2143:
1216:
1144:"Adaptive evolution: evaluating empirical support for theoretical predictions"
1007:
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169:, and via access to previously cryptic variants through the switching of an
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2019:
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915:
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499:
366:
1663:"Evolution of molecular error rates and the consequences for evolvability"
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972:
584:
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2725:
Arthropod biology and evolution : molecules, development, morphology
1438:
460:
383:
The evolution of evolvability is less controversial if it occurs via the
158:
3741:
3441:
3394:
3043:"Quantifying the adaptive potential of an antibiotic resistance enzyme"
1997:
1946:
1830:
717:
657:
349:
345:
300:
162:
128:, and one representing exceedingly rare long-term innovations of form.
63:
43:
34:
1031:"The evolution of stress-induced hypermutation in asexual populations"
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According to the second definition, a biological system is evolvable:
3692:
3660:
3427:
3333:
456:
452:
3272:
2814:
2771:
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1532:
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907:
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A better understanding of evolvability is proposed to be part of an
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3258:
2659:
1374:
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1976:
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1720:
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1610:"Cryptic genetic variation is enriched for potential adaptations"
1423:"The genetic code constrains yet facilitates Darwinian evolution"
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440:
110:
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391:
may also be an example of the evolution of evolvability through
281:
3367:
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860:. Princeton Studies in Complexity. Princeton University Press.
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When recombination is low, mutator alleles may still sometimes
106:
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2457:
1142:
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795:
443:, bacteria, fungi and cancers evolve to be resistant to host
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1137:
1135:
1133:
1131:
1129:
1127:
1125:
208:, through an increase in the recombination rate, decreasing
67:
Analogously, the evolvability of organisms depends on their
3665:
2919:
2550:
Bloom JD, Labthavikul ST, Otey CR, Arnold FH (April 2006).
1248:
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1122:
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Evolvability phenomena have practical applications. For
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2352:
1470:
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if its properties show heritable genetic variation, and
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3155:
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2177:
Evolution; International
Journal of Organic Evolution
2024:
Evolution; International Journal of Organic Evolution
1790:
Evolution; International Journal of Organic Evolution
1564:
Evolution; International Journal of Organic Evolution
1469:
1035:
Evolution; International Journal of Organic Evolution
696:
Evolution; International Journal of Organic Evolution
254:
robustness may allow exploration of large regions of
2881:"The emergence of antibiotic resistance by mutation"
2800:
2722:
1518:
2422:
1353:
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2878:
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2268:
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538:
497:
451:. These same problems occur in agriculture with
3091:
1833:"The look-ahead effect of phenotypic mutations"
689:
215:Recombination decreasing the importance of the
2843:
2387:
1359:
874:
439:Many human diseases are capable of evolution.
230:
47:genetic diversity, and thereby evolve through
3313:
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2274:
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1191:
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1187:
1078:Karasov T, Messer PW, Petrov DA (June 2010).
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78:
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2879:Woodford N, Ellington MJ (January 2007).
2747:
2585:
2575:
2552:"Protein stability promotes evolvability"
2526:
2516:
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1446:
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1273:
1250:"Protein stability promotes evolvability"
1224:
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1028:
962:
883:
829:
707:
670:
643:
610:
600:
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1421:Firnberg E, Ostermeier M (August 2013).
855:
245:
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14:
3821:
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2694:
2317:
1932:
746:
200:increasing the threshold value of the
3301:
2390:Current Opinion in Structural Biology
1604:
936:
307:stocks of cryptic genetic variation.
265:
2970:
690:Wagner GP, Altenberg L (June 1996).
539:Kirschner M, Gerhart J (July 1998).
319:, and it can increase evolvability.
2885:Clinical Microbiology and Infection
2355:Current Opinion in Chemical Biology
1787:
498:Colegrave N, Collins S (May 2008).
348:is restricted to within functional
24:
3620:Evolutionary developmental biology
2036:10.1111/j.0014-3820.2003.tb00358.x
1882:Griswold CK, Masel J (June 2009).
709:10.1111/j.1558-5646.1996.tb02339.x
251:Robustness in the face of mutation
189:on "good genes", and so intensify
25:
3845:
1935:Journal of the History of Biology
3577:Evolution of sexual reproduction
3230:10.1111/j.1749-6632.2009.04578.x
3168:10.1111/j.1558-5646.2007.00246.x
2993:10.1111/j.1365-3180.2007.00581.x
2898:10.1111/j.1469-0691.2006.01492.x
2189:10.1111/j.1558-5646.2007.00303.x
1802:10.1111/j.1558-5646.2007.00166.x
1758:10.1111/j.1420-9101.2006.01125.x
1576:10.1111/j.1558-5646.2009.00713.x
1048:10.1111/j.1558-5646.2012.01576.x
385:evolution of sexual reproduction
143:evolution of sexual reproduction
3834:Extended evolutionary synthesis
3252:
3191:
3136:
3085:
3034:
2999:
2964:
2913:
2872:
2837:
2794:
2741:
2716:
2688:
2653:
2602:
2451:
2416:
2381:
2346:
2311:
2211:
2168:
2107:
2058:
2012:
1969:
1926:
1875:
1824:
1781:
1745:Journal of Evolutionary Biology
1732:
1713:
1650:
1598:
1555:
1512:
1463:
1414:
1300:
1241:
1199:, Trotter MV (September 2010).
1071:
1022:
979:
930:
677:Advances in Genetic Programming
468:Extended Evolutionary Synthesis
403:
3348:Genotype–phenotype distinction
2611:Journal of Molecular Evolution
2124:Theoretical Population Biology
1978:Journal of Applied Probability
1362:Journal of Theoretical Biology
789:
683:
664:
637:
532:
491:
430:tolerance to further mutations
60:DNA sequences with no function
13:
1:
3605:Regulation of gene expression
2437:10.1016/s0022-2836(02)00400-x
1201:"Robustness and evolvability"
1029:Ram Y, Hadany L (July 2012).
767:10.1089/genbio.2022.29027.sbi
485:
338:
3775:Endless Forms Most Beautiful
3555:Evolution of genetic systems
3363:Gene–environment correlation
3358:Gene–environment interaction
3111:10.1371/journal.pbio.1000058
3060:10.1371/journal.pgen.1002783
2939:10.1371/journal.pgen.0030205
2858:10.1016/j.bbagen.2012.06.001
2753:"Is evolvability evolvable?"
2662:Journal of Molecular Biology
2518:10.1371/journal.pgen.1001321
2425:Journal of Molecular Biology
1901:10.1371/journal.pgen.1000517
1097:10.1371/journal.pgen.1000924
889:"Is evolvability evolvable?"
603:Evolution and Biocomputation
7:
3754:Christiane NĂĽsslein-Volhard
2702:. Oxford University Press.
2332:10.1016/j.yexcr.2011.02.013
2083:10.1534/genetics.109.110213
1626:10.1534/genetics.105.051649
1329:10.1534/genetics.112.148627
473:
420:, both rational design and
278:and sometimes be adaptive.
231:Robustness and evolvability
10:
3850:
3630:Hedgehog signaling pathway
3507:Developmental architecture
3020:10.1016/j.ijmm.2013.02.004
2367:10.1016/j.cbpa.2010.11.011
2320:Experimental Cell Research
1392:10.1016/j.jtbi.2009.11.008
955:10.1093/genetics/130.1.195
822:10.1038/s41586-022-04506-6
234:
185:Mating rituals that allow
3807:
3786:
3715:
3643:
3597:
3590:
3554:
3506:
3470:
3457:Transgressive segregation
3403:
3340:
2674:10.1016/j.jmb.2013.03.033
2631:10.1007/s00239-005-0289-7
2402:10.1016/j.sbi.2009.08.003
2144:10.1016/j.tpb.2007.08.006
1217:10.1016/j.tig.2010.06.002
1008:10.1007/s11692-011-9127-6
356:Evolution of evolvability
290:Exploration ahead of time
270:Temporary robustness, or
198:effective population size
157:, via the probability of
149:, via the probability of
136:Generating more variation
105:For example, consider an
937:Houle D (January 1992).
621:10.1007/3-540-59046-3_11
393:evolutionary capacitance
177:Enhancement of selection
3635:Notch signaling pathway
3610:Gene regulatory network
3493:Dual inheritance theory
3261:Nature Reviews Genetics
2760:Nature Reviews Genetics
2577:10.1073/pnas.0510098103
2277:Nature Reviews Genetics
2244:10.1126/science.1198914
1688:10.1073/pnas.1012918108
1521:Nature Reviews Genetics
1275:10.1073/pnas.0510098103
1148:Nature Reviews Genetics
896:Nature Reviews Genetics
566:10.1073/pnas.95.15.8420
313:genetically assimilated
260:thermodynamic stability
79:Alternative definitions
58:, unless they occur in
3683:cis-regulatory element
3591:Control of development
3471:Non-genetic influences
3437:evolutionary landscape
2803:Nature Reviews. Cancer
2477:10.1093/protein/gzl029
1850:10.1186/1745-6150-3-18
1427:Nucleic Acids Research
517:10.1038/sj.hdy.6801095
480:Evolutionary trade-off
378:experimental evolution
171:evolutionary capacitor
3794:Nature versus nurture
3698:Cell surface receptor
3615:Evo-devo gene toolkit
3514:Developmental biology
3452:Polygenic inheritance
3378:Quantitative genetics
1488:10.1128/mBio.01654-17
434:protein superfamilies
296:mutational robustness
246:Without recombination
237:Mutational robustness
235:Further information:
217:Hill-Robertson effect
202:selection coefficient
122:quantitative genetics
71:map. This means that
3829:Evolutionary biology
3703:Transcription factor
3418:Genetic assimilation
3405:Genetic architecture
988:Evolutionary Biology
449:pharmaceutical drugs
333:genetic assimilation
155:asexual reproduction
3799:Morphogenetic field
3716:Influential figures
3222:2009NYASA1168..218P
2985:2007WeedR..47..365N
2623:2006JMolE..63..513R
2568:2006PNAS..103.5869B
2236:2011Sci...331.1433W
2136:2007TPBio..72..560K
1679:2011PNAS..108.1082R
1384:2010JThBi.263..143W
1266:2006PNAS..103.5869B
1000:2011EvBio..38..258H
814:2022Natur.603..455V
557:1998PNAS...95.8420K
428:or at least reduce
418:protein engineering
410:protein engineering
126:population genetics
3488:Genomic imprinting
1947:10.1007/bf00138880
1439:10.1093/nar/gkt536
1205:Trends in Genetics
658:10.1147/rd.21.0002
422:directed evolution
329:adaptive landscape
266:With recombination
69:genotype–phenotype
41:, but to generate
35:adaptive evolution
18:Adaptive potential
3816:
3815:
3749:Eric F. Wieschaus
3711:
3710:
3529:Pattern formation
3433:Fitness landscape
3146:(December 2007).
2734:978-3-642-36159-3
2709:978-0-19-969259-0
2698:(July 14, 2011).
2118:(December 2007).
867:978-0-691-12240-3
856:Wagner A (2005).
808:(7901): 455–463.
750:GEN Biotechnology
630:978-3-540-59046-0
324:fitness landscape
191:natural selection
118:Massimo Pigliucci
49:natural selection
39:genetic diversity
16:(Redirected from
3841:
3759:William McGinnis
3728:Richard Lewontin
3723:C. H. Waddington
3595:
3594:
3572:Neutral networks
3322:
3315:
3308:
3299:
3298:
3293:
3292:
3256:
3250:
3249:
3207:
3195:
3189:
3188:
3170:
3152:
3140:
3134:
3133:
3123:
3113:
3089:
3083:
3082:
3072:
3062:
3038:
3032:
3031:
3003:
2997:
2996:
2968:
2962:
2961:
2951:
2941:
2917:
2911:
2910:
2900:
2876:
2870:
2869:
2841:
2835:
2834:
2798:
2792:
2791:
2757:
2751:(January 2008).
2745:
2739:
2738:
2720:
2714:
2713:
2692:
2686:
2685:
2657:
2651:
2650:
2606:
2600:
2599:
2589:
2579:
2547:
2541:
2540:
2530:
2520:
2496:
2490:
2489:
2479:
2455:
2449:
2448:
2420:
2414:
2413:
2385:
2379:
2378:
2350:
2344:
2343:
2315:
2309:
2308:
2272:
2266:
2265:
2255:
2230:(6023): 1433–6.
2215:
2209:
2208:
2172:
2166:
2165:
2155:
2111:
2105:
2104:
2094:
2062:
2056:
2055:
2016:
2010:
2009:
1973:
1967:
1966:
1930:
1924:
1923:
1913:
1903:
1879:
1873:
1872:
1862:
1852:
1828:
1822:
1821:
1785:
1779:
1778:
1760:
1736:
1730:
1729:
1717:
1711:
1710:
1700:
1690:
1661:(January 2011).
1654:
1648:
1647:
1637:
1602:
1596:
1595:
1559:
1553:
1552:
1516:
1510:
1509:
1499:
1467:
1461:
1460:
1450:
1418:
1412:
1411:
1377:
1357:
1351:
1350:
1340:
1304:
1298:
1297:
1287:
1277:
1245:
1239:
1238:
1228:
1193:
1182:
1181:
1171:
1139:
1120:
1119:
1109:
1099:
1075:
1069:
1068:
1050:
1026:
1020:
1019:
983:
977:
976:
966:
934:
928:
927:
893:
887:(January 2008).
881:
872:
871:
853:
844:
843:
833:
793:
787:
786:
759:Mary Ann Liebert
744:
738:
737:
711:
687:
681:
680:
668:
662:
661:
641:
635:
634:
614:
598:
589:
588:
578:
568:
536:
530:
529:
519:
495:
187:sexual selection
21:
3849:
3848:
3844:
3843:
3842:
3840:
3839:
3838:
3819:
3818:
3817:
3812:
3803:
3782:
3769:Sean B. Carroll
3707:
3639:
3586:
3550:
3502:
3483:Maternal effect
3466:
3399:
3336:
3326:
3296:
3273:10.1038/nrg3028
3257:
3253:
3205:
3196:
3192:
3150:
3141:
3137:
3104:(4): e1000058.
3090:
3086:
3053:(6): e1002783.
3039:
3035:
3004:
3000:
2969:
2965:
2918:
2914:
2877:
2873:
2852:(10): 1526–34.
2842:
2838:
2815:10.1038/nrc2013
2799:
2795:
2772:10.1038/nrg2278
2755:
2746:
2742:
2735:
2721:
2717:
2710:
2693:
2689:
2668:(14): 2609–21.
2658:
2654:
2607:
2603:
2562:(15): 5869–74.
2548:
2544:
2511:(3): e1001321.
2497:
2493:
2456:
2452:
2421:
2417:
2386:
2382:
2351:
2347:
2316:
2312:
2289:10.1038/nrg2808
2273:
2269:
2216:
2212:
2173:
2169:
2112:
2108:
2063:
2059:
2030:(7): 1498–512.
2017:
2013:
1990:10.2307/3212376
1974:
1970:
1931:
1927:
1894:(6): e1000517.
1880:
1876:
1829:
1825:
1786:
1782:
1737:
1733:
1722:Complex Systems
1718:
1714:
1655:
1651:
1603:
1599:
1560:
1556:
1533:10.1038/nrg2146
1517:
1513:
1468:
1464:
1419:
1415:
1358:
1354:
1305:
1301:
1260:(15): 5869–74.
1246:
1242:
1194:
1185:
1160:10.1038/nrg3322
1140:
1123:
1090:(6): e1000924.
1076:
1072:
1027:
1023:
984:
980:
935:
931:
908:10.1038/nrg2278
891:
882:
875:
868:
854:
847:
794:
790:
745:
741:
688:
684:
669:
665:
642:
638:
631:
612:10.1.1.493.6534
599:
592:
537:
533:
496:
492:
488:
476:
445:immune defences
426:enzyme function
406:
358:
341:
292:
284:
268:
248:
239:
233:
224:generation time
179:
138:
81:
28:
23:
22:
15:
12:
11:
5:
3847:
3837:
3836:
3831:
3814:
3813:
3808:
3805:
3804:
3802:
3801:
3796:
3790:
3788:
3784:
3783:
3781:
3780:
3779:
3778:
3766:
3761:
3756:
3751:
3746:
3745:
3744:
3733:François Jacob
3730:
3725:
3719:
3717:
3713:
3712:
3709:
3708:
3706:
3705:
3700:
3695:
3690:
3685:
3680:
3675:
3670:
3669:
3668:
3658:
3653:
3647:
3645:
3641:
3640:
3638:
3637:
3632:
3627:
3622:
3617:
3612:
3607:
3601:
3599:
3592:
3588:
3587:
3585:
3584:
3579:
3574:
3569:
3564:
3558:
3556:
3552:
3551:
3549:
3548:
3543:
3538:
3533:
3532:
3531:
3526:
3516:
3510:
3508:
3504:
3503:
3501:
3500:
3495:
3490:
3485:
3480:
3474:
3472:
3468:
3467:
3465:
3464:
3462:Sequence space
3459:
3454:
3449:
3444:
3439:
3430:
3425:
3420:
3415:
3409:
3407:
3401:
3400:
3398:
3397:
3392:
3391:
3390:
3380:
3375:
3370:
3365:
3360:
3355:
3350:
3344:
3342:
3338:
3337:
3325:
3324:
3317:
3310:
3302:
3295:
3294:
3251:
3190:
3161:(12): 2743–9.
3135:
3084:
3033:
3014:(6–7): 293–7.
2998:
2979:(5): 365–369.
2963:
2912:
2871:
2836:
2809:(12): 924–35.
2793:
2740:
2733:
2715:
2708:
2687:
2652:
2601:
2542:
2491:
2470:(10): 439–42.
2450:
2415:
2396:(5): 596–604.
2380:
2361:(2): 194–200.
2345:
2310:
2267:
2210:
2167:
2106:
2077:(2): 393–400.
2057:
2011:
1984:(4): 728–738.
1968:
1941:(2): 289–302.
1925:
1874:
1837:Biology Direct
1823:
1796:(8): 1847–56.
1780:
1751:(5): 1555–70.
1731:
1712:
1649:
1620:(3): 1985–91.
1608:(March 2006).
1597:
1570:(8): 2164–71.
1554:
1511:
1462:
1433:(15): 7420–8.
1413:
1352:
1323:(4): 1209–20.
1311:(April 2013).
1299:
1240:
1183:
1154:(12): 867–77.
1121:
1070:
1041:(7): 2315–28.
1021:
994:(3): 258–277.
978:
949:(1): 195–204.
929:
873:
866:
845:
788:
739:
702:(3): 967–976.
682:
663:
636:
629:
590:
551:(15): 8420–7.
541:"Evolvability"
531:
489:
487:
484:
483:
482:
475:
472:
405:
402:
357:
354:
340:
337:
317:Baldwin effect
291:
288:
283:
280:
267:
264:
256:genotype space
247:
244:
232:
229:
228:
227:
220:
213:
194:
178:
175:
137:
134:
109:with multiple
103:
102:
95:
94:
91:
84:Andreas Wagner
80:
77:
26:
9:
6:
4:
3:
2:
3846:
3835:
3832:
3830:
3827:
3826:
3824:
3811:
3806:
3800:
3797:
3795:
3792:
3791:
3789:
3785:
3777:
3776:
3772:
3771:
3770:
3767:
3765:
3762:
3760:
3757:
3755:
3752:
3750:
3747:
3743:
3740:
3739:
3738:
3737:Jacques Monod
3734:
3731:
3729:
3726:
3724:
3721:
3720:
3718:
3714:
3704:
3701:
3699:
3696:
3694:
3691:
3689:
3686:
3684:
3681:
3679:
3676:
3674:
3671:
3667:
3664:
3663:
3662:
3659:
3657:
3654:
3652:
3651:Homeotic gene
3649:
3648:
3646:
3642:
3636:
3633:
3631:
3628:
3626:
3623:
3621:
3618:
3616:
3613:
3611:
3608:
3606:
3603:
3602:
3600:
3596:
3593:
3589:
3583:
3580:
3578:
3575:
3573:
3570:
3568:
3565:
3563:
3560:
3559:
3557:
3553:
3547:
3544:
3542:
3539:
3537:
3534:
3530:
3527:
3525:
3522:
3521:
3520:
3519:Morphogenesis
3517:
3515:
3512:
3511:
3509:
3505:
3499:
3496:
3494:
3491:
3489:
3486:
3484:
3481:
3479:
3476:
3475:
3473:
3469:
3463:
3460:
3458:
3455:
3453:
3450:
3448:
3445:
3443:
3440:
3438:
3434:
3431:
3429:
3426:
3424:
3421:
3419:
3416:
3414:
3411:
3410:
3408:
3406:
3402:
3396:
3393:
3389:
3386:
3385:
3384:
3381:
3379:
3376:
3374:
3371:
3369:
3366:
3364:
3361:
3359:
3356:
3354:
3353:Reaction norm
3351:
3349:
3346:
3345:
3343:
3339:
3335:
3331:
3323:
3318:
3316:
3311:
3309:
3304:
3303:
3300:
3290:
3286:
3282:
3278:
3274:
3270:
3267:(7): 475–86.
3266:
3262:
3255:
3247:
3243:
3239:
3235:
3231:
3227:
3223:
3219:
3216:(1): 218–28.
3215:
3211:
3204:
3201:(June 2009).
3200:
3194:
3186:
3182:
3178:
3174:
3169:
3164:
3160:
3156:
3149:
3145:
3139:
3131:
3127:
3122:
3117:
3112:
3107:
3103:
3099:
3095:
3088:
3080:
3076:
3071:
3066:
3061:
3056:
3052:
3048:
3047:PLOS Genetics
3044:
3037:
3029:
3025:
3021:
3017:
3013:
3009:
3002:
2994:
2990:
2986:
2982:
2978:
2974:
2973:Weed Research
2967:
2959:
2955:
2950:
2945:
2940:
2935:
2931:
2927:
2926:PLOS Genetics
2923:
2916:
2908:
2904:
2899:
2894:
2890:
2886:
2882:
2875:
2867:
2863:
2859:
2855:
2851:
2847:
2840:
2832:
2828:
2824:
2820:
2816:
2812:
2808:
2804:
2797:
2789:
2785:
2781:
2777:
2773:
2769:
2765:
2761:
2754:
2750:
2744:
2736:
2730:
2726:
2719:
2711:
2705:
2701:
2697:
2691:
2683:
2679:
2675:
2671:
2667:
2663:
2656:
2648:
2644:
2640:
2636:
2632:
2628:
2624:
2620:
2617:(4): 513–25.
2616:
2612:
2605:
2597:
2593:
2588:
2583:
2578:
2573:
2569:
2565:
2561:
2557:
2553:
2546:
2538:
2534:
2529:
2524:
2519:
2514:
2510:
2506:
2505:PLOS Genetics
2502:
2495:
2487:
2483:
2478:
2473:
2469:
2465:
2461:
2454:
2446:
2442:
2438:
2434:
2430:
2426:
2419:
2411:
2407:
2403:
2399:
2395:
2391:
2384:
2376:
2372:
2368:
2364:
2360:
2356:
2349:
2341:
2337:
2333:
2329:
2326:(9): 1261–9.
2325:
2321:
2314:
2306:
2302:
2298:
2294:
2290:
2286:
2283:(8): 572–82.
2282:
2278:
2271:
2263:
2259:
2254:
2249:
2245:
2241:
2237:
2233:
2229:
2225:
2221:
2214:
2206:
2202:
2198:
2194:
2190:
2186:
2183:(2): 301–15.
2182:
2178:
2171:
2163:
2159:
2154:
2149:
2145:
2141:
2137:
2133:
2130:(4): 560–75.
2129:
2125:
2121:
2117:
2110:
2102:
2098:
2093:
2088:
2084:
2080:
2076:
2072:
2068:
2061:
2053:
2049:
2045:
2041:
2037:
2033:
2029:
2025:
2021:
2015:
2007:
2003:
1999:
1995:
1991:
1987:
1983:
1979:
1972:
1964:
1960:
1956:
1952:
1948:
1944:
1940:
1936:
1929:
1921:
1917:
1912:
1907:
1902:
1897:
1893:
1889:
1888:PLOS Genetics
1885:
1878:
1870:
1866:
1861:
1856:
1851:
1846:
1842:
1838:
1834:
1827:
1819:
1815:
1811:
1807:
1803:
1799:
1795:
1791:
1784:
1776:
1772:
1768:
1764:
1759:
1754:
1750:
1746:
1742:
1735:
1727:
1723:
1716:
1708:
1704:
1699:
1694:
1689:
1684:
1680:
1676:
1673:(3): 1082–7.
1672:
1668:
1664:
1660:
1653:
1645:
1641:
1636:
1631:
1627:
1623:
1619:
1615:
1611:
1607:
1601:
1593:
1589:
1585:
1581:
1577:
1573:
1569:
1565:
1558:
1550:
1546:
1542:
1538:
1534:
1530:
1526:
1522:
1515:
1507:
1503:
1498:
1493:
1489:
1485:
1481:
1477:
1473:
1466:
1458:
1454:
1449:
1444:
1440:
1436:
1432:
1428:
1424:
1417:
1409:
1405:
1401:
1397:
1393:
1389:
1385:
1381:
1376:
1371:
1368:(1): 143–53.
1367:
1363:
1356:
1348:
1344:
1339:
1334:
1330:
1326:
1322:
1318:
1314:
1310:
1303:
1295:
1291:
1286:
1281:
1276:
1271:
1267:
1263:
1259:
1255:
1251:
1244:
1236:
1232:
1227:
1222:
1218:
1214:
1211:(9): 406–14.
1210:
1206:
1202:
1198:
1192:
1190:
1188:
1179:
1175:
1170:
1165:
1161:
1157:
1153:
1149:
1145:
1138:
1136:
1134:
1132:
1130:
1128:
1126:
1117:
1113:
1108:
1103:
1098:
1093:
1089:
1085:
1084:PLOS Genetics
1081:
1074:
1066:
1062:
1058:
1054:
1049:
1044:
1040:
1036:
1032:
1025:
1017:
1013:
1009:
1005:
1001:
997:
993:
989:
982:
974:
970:
965:
960:
956:
952:
948:
944:
940:
933:
925:
921:
917:
913:
909:
905:
901:
897:
890:
886:
880:
878:
869:
863:
859:
852:
850:
841:
837:
832:
827:
823:
819:
815:
811:
807:
803:
799:
792:
784:
780:
776:
772:
768:
764:
760:
756:
752:
751:
743:
735:
731:
727:
723:
719:
715:
710:
705:
701:
697:
693:
686:
678:
674:
667:
659:
655:
651:
647:
640:
632:
626:
622:
618:
613:
608:
604:
597:
595:
586:
582:
577:
572:
567:
562:
558:
554:
550:
546:
542:
535:
527:
523:
518:
513:
510:(5): 464–70.
509:
505:
501:
494:
490:
481:
478:
477:
471:
469:
464:
462:
458:
454:
450:
447:, as well as
446:
442:
437:
435:
431:
427:
423:
419:
414:
411:
401:
398:
394:
390:
386:
381:
379:
375:
370:
368:
367:mutation load
364:
363:mutation rate
353:
351:
347:
336:
334:
330:
325:
320:
318:
314:
308:
306:
302:
297:
287:
279:
277:
273:
263:
261:
257:
252:
243:
238:
225:
221:
218:
214:
211:
210:genetic draft
207:
206:genetic drift
203:
199:
195:
192:
188:
184:
183:
182:
174:
172:
168:
164:
160:
156:
152:
148:
147:mutation rate
144:
133:
129:
127:
123:
119:
115:
112:
108:
100:
99:
98:
92:
89:
88:
87:
85:
76:
74:
70:
65:
61:
57:
52:
50:
46:
45:
40:
36:
32:
19:
3773:
3666:eyeless gene
3562:Evolvability
3561:
3536:Segmentation
3413:Canalisation
3383:Heterochrony
3373:Heritability
3341:Key concepts
3264:
3260:
3254:
3213:
3209:
3193:
3158:
3154:
3138:
3101:
3098:PLOS Biology
3097:
3087:
3050:
3046:
3036:
3011:
3007:
3001:
2976:
2972:
2966:
2932:(11): e205.
2929:
2925:
2915:
2888:
2884:
2874:
2849:
2845:
2839:
2806:
2802:
2796:
2766:(1): 75–82.
2763:
2759:
2743:
2727:. Springer.
2724:
2718:
2699:
2690:
2665:
2661:
2655:
2614:
2610:
2604:
2559:
2555:
2545:
2508:
2504:
2494:
2467:
2463:
2453:
2431:(1): 85–95.
2428:
2424:
2418:
2393:
2389:
2383:
2358:
2354:
2348:
2323:
2319:
2313:
2280:
2276:
2270:
2227:
2223:
2213:
2180:
2176:
2170:
2127:
2123:
2109:
2074:
2070:
2060:
2027:
2023:
2014:
1981:
1977:
1971:
1938:
1934:
1928:
1891:
1887:
1877:
1840:
1836:
1826:
1793:
1789:
1783:
1748:
1744:
1734:
1725:
1721:
1715:
1670:
1666:
1652:
1617:
1613:
1600:
1567:
1563:
1557:
1527:(8): 610–8.
1524:
1520:
1514:
1479:
1475:
1465:
1430:
1426:
1416:
1365:
1361:
1355:
1320:
1316:
1302:
1257:
1253:
1243:
1208:
1204:
1151:
1147:
1087:
1083:
1073:
1038:
1034:
1024:
991:
987:
981:
946:
942:
932:
902:(1): 75–82.
899:
895:
857:
805:
801:
791:
754:
748:
742:
699:
695:
685:
676:
666:
649:
645:
639:
602:
548:
544:
534:
507:
503:
493:
465:
438:
415:
407:
404:Applications
382:
371:
359:
342:
321:
309:
293:
285:
272:canalisation
269:
249:
240:
180:
139:
130:
116:
104:
96:
82:
53:
42:
31:Evolvability
30:
29:
3764:Mike Levine
3673:Distal-less
3498:Polyphenism
3478:Epigenetics
3330:development
3199:Pigliucci M
3144:Pigliucci M
2891:(1): 5–18.
2749:Pigliucci M
885:Pigliucci M
761:: 140–141.
652:(1): 2–13.
461:antibiotics
397:bet-hedging
389:yeast prion
159:outcrossing
3823:Categories
3742:Lac operon
3567:Robustness
3546:Modularity
3541:Metamerism
3447:Plasticity
3442:Pleiotropy
3395:Heterotopy
1728:: 495–502.
486:References
346:pleiotropy
339:Modularity
305:preadapted
301:penetrance
163:inbreeding
64:adaptation
3693:Morphogen
3678:Engrailed
3661:Pax genes
3582:Tinkering
3428:Epistasis
3423:Dominance
3334:phenotype
2114:King OD,
2006:123907349
1843:(1): 18.
1657:Rajon E,
1375:0907.0510
1307:Rajon E,
783:248313305
775:2768-1572
607:CiteSeerX
457:herbicide
453:pesticide
374:hitchhike
167:dispersal
56:mutations
3656:Hox gene
3644:Elements
3625:Homeobox
3281:21681209
3238:19566710
3177:17924956
3130:19355786
3079:22761587
3028:23517688
2958:18020711
2907:17184282
2866:22698669
2823:17109012
2780:18059367
2696:Wagner A
2682:23542341
2647:25258028
2639:17021929
2596:16581913
2537:21408208
2486:16868005
2445:12079336
2410:19765975
2375:21115265
2340:21371474
2297:20634811
2262:21415350
2205:11560256
2197:18031304
2162:17915273
2101:19917766
2071:Genetics
2052:30954684
2044:12940355
1963:42288730
1955:11611993
1920:19521499
1869:18479505
1818:13150906
1810:17683428
1767:16910985
1707:21199946
1644:16387877
1614:Genetics
1592:12103318
1584:19473394
1549:10868777
1541:17637733
1506:29138304
1457:23754851
1408:11511132
1400:19925810
1347:23335336
1317:Genetics
1294:16581913
1235:20598394
1178:23154809
1116:20585551
1065:35770307
1057:22759304
1016:11359207
943:Genetics
916:18059367
840:35264797
734:21040413
726:28565291
679:: 47–74.
526:18212804
504:Heredity
474:See also
276:revealed
222:Shorter
44:adaptive
3787:Debates
3598:Systems
3524:Eyespot
3388:Neoteny
3289:8837202
3246:5710484
3218:Bibcode
3185:2703146
3121:3279047
3070:3386231
2981:Bibcode
2949:2077897
2831:8040576
2788:3164124
2619:Bibcode
2587:1458665
2564:Bibcode
2528:3048372
2305:8951755
2253:3176658
2232:Bibcode
2224:Science
2153:2118055
2132:Bibcode
2116:Masel J
2092:2828720
2020:Masel J
1998:3212376
1911:2686163
1860:2423361
1775:6964065
1698:3024668
1675:Bibcode
1659:Masel J
1635:1456269
1606:Masel J
1497:5686537
1448:3753648
1380:Bibcode
1338:3606098
1309:Masel J
1285:1458665
1262:Bibcode
1226:3198833
1197:Masel J
1169:3748133
1107:2887467
996:Bibcode
973:1732160
964:1204793
924:3164124
831:8934302
810:Bibcode
718:2410639
585:9671692
553:Bibcode
441:Viruses
350:modules
111:alleles
73:genomes
3688:Ligand
3368:Operon
3287:
3279:
3244:
3236:
3183:
3175:
3128:
3118:
3077:
3067:
3026:
2956:
2946:
2905:
2864:
2829:
2821:
2786:
2778:
2731:
2706:
2680:
2645:
2637:
2594:
2584:
2535:
2525:
2484:
2443:
2408:
2373:
2338:
2303:
2295:
2260:
2250:
2203:
2195:
2160:
2150:
2099:
2089:
2050:
2042:
2004:
1996:
1961:
1953:
1918:
1908:
1867:
1857:
1816:
1808:
1773:
1765:
1705:
1695:
1642:
1632:
1590:
1582:
1547:
1539:
1504:
1494:
1455:
1445:
1406:
1398:
1345:
1335:
1292:
1282:
1233:
1223:
1176:
1166:
1114:
1104:
1063:
1055:
1014:
971:
961:
922:
914:
864:
838:
828:
802:Nature
781:
773:
732:
724:
716:
627:
609:
583:
573:
524:
196:Large
165:, via
151:sexual
107:enzyme
3285:S2CID
3242:S2CID
3206:(PDF)
3181:S2CID
3151:(PDF)
2827:S2CID
2784:S2CID
2756:(PDF)
2643:S2CID
2301:S2CID
2201:S2CID
2048:S2CID
2002:S2CID
1994:JSTOR
1959:S2CID
1814:S2CID
1771:S2CID
1588:S2CID
1545:S2CID
1482:(6).
1404:S2CID
1370:arXiv
1061:S2CID
1012:S2CID
920:S2CID
892:(PDF)
779:S2CID
757:(2).
730:S2CID
714:JSTOR
576:33871
294:When
3328:The
3277:PMID
3234:PMID
3214:1168
3173:PMID
3126:PMID
3075:PMID
3024:PMID
2954:PMID
2903:PMID
2862:PMID
2850:1820
2819:PMID
2776:PMID
2729:ISBN
2704:ISBN
2678:PMID
2635:PMID
2592:PMID
2533:PMID
2482:PMID
2441:PMID
2406:PMID
2371:PMID
2336:PMID
2293:PMID
2258:PMID
2193:PMID
2158:PMID
2097:PMID
2040:PMID
1951:PMID
1916:PMID
1865:PMID
1806:PMID
1763:PMID
1703:PMID
1640:PMID
1580:PMID
1537:PMID
1502:PMID
1476:mBio
1453:PMID
1396:PMID
1343:PMID
1290:PMID
1231:PMID
1174:PMID
1112:PMID
1053:PMID
969:PMID
912:PMID
862:ISBN
836:PMID
771:ISSN
722:PMID
625:ISBN
581:PMID
522:PMID
455:and
161:vs.
153:vs.
3332:of
3269:doi
3226:doi
3163:doi
3116:PMC
3106:doi
3065:PMC
3055:doi
3016:doi
3012:303
2989:doi
2944:PMC
2934:doi
2893:doi
2854:doi
2811:doi
2768:doi
2670:doi
2666:425
2627:doi
2582:PMC
2572:doi
2560:103
2523:PMC
2513:doi
2472:doi
2433:doi
2429:320
2398:doi
2363:doi
2328:doi
2324:317
2285:doi
2248:PMC
2240:doi
2228:331
2185:doi
2148:PMC
2140:doi
2087:PMC
2079:doi
2075:184
2032:doi
1986:doi
1943:doi
1906:PMC
1896:doi
1855:PMC
1845:doi
1798:doi
1753:doi
1693:PMC
1683:doi
1671:108
1630:PMC
1622:doi
1618:172
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1443:PMC
1435:doi
1388:doi
1366:263
1333:PMC
1325:doi
1321:193
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1156:doi
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1092:doi
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826:PMC
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806:603
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