3479:. This selective pressure resulted in the development of processes allowing the viruses to spread horizontally throughout the population. The outcome of this selection was cell-to-cell fusion. (This is distinct from the conjugation methods used by bacterial plasmids under evolutionary pressure, with important consequences.) The possibility of this kind of fusion is supported by the presence of fusion proteins in the envelopes of the pox viruses that allow them to fuse with host membranes. These proteins could have been transferred to the cell membrane during viral reproduction, enabling cell-to-cell fusion between the virus host and an uninfected cell. The theory proposes meiosis originated from the fusion between two cells infected with related but different viruses which recognised each other as uninfected. After the fusion of the two cells, incompatibilities between the two viruses result in a meiotic-like cell division.
1898:
male partition is suggested to be an "experimental" part of the species that allows the species to expand their ecological niche, and to have alternative configurations. This theory underlines the higher variability and higher mortality in males, in comparison to females. This functional partitioning also explains the higher susceptibility to disease in males, in comparison to females and therefore includes the idea of "protection against parasites" as another functionality of male sex. Geodakyan's evolutionary theory of sex was developed in Russia in 1960–1980 and was not known to the West till the era of the
Internet. Trofimova, who analysed psychological sex differences, hypothesised that the male sex might also provide a "redundancy pruning" function.
3483:
centromeres would incite the grouping of both sets into tetrads. It is speculated that this grouping may be the origin of crossing over, characteristic of the first division in modern meiosis. The partitioning apparatus of the mitotic-like cell cycle the cells used to replicate independently would then pull each set of chromosomes to one side of the cell, still bound by centromeres. These centromeres would prevent their replication in subsequent division, resulting in four daughter cells with one copy of one of the two original pox-like viruses. The process resulting from combination of two similar pox viruses within the same host closely mimics meiosis.
1809:
648:
1434:
select against sex. They concluded that, although the Red Queen hypothesis favors sex under certain circumstances, it alone does not account for the ubiquity of sex. Otto and
Gerstein further stated that "it seems doubtful to us that strong selection per gene is sufficiently commonplace for the Red Queen hypothesis to explain the ubiquity of sex". Parker reviewed numerous genetic studies on plant disease resistance and failed to uncover a single example consistent with the assumptions of the Red Queen hypothesis.
637:
35:
3453:. This model suggests that the nucleus originated when the lysogenic virus incorporated genetic material from the archaean and the bacterium and took over the role of information storage for the amalgam. The archaeal host transferred much of its functional genome to the virus during the evolution of cytoplasm, but retained the function of gene translation and general metabolism. The bacterium transferred most of its functional genome to the virus as it transitioned into a
9574:
8993:
7708:
590:
1669:). Recombinational repair is prevalent from the simplest viruses to the most complex multicellular eukaryotes. It is effective against many different types of genomic damage, and in particular is highly efficient at overcoming double-strand damages. Studies of the mechanism of meiotic recombination indicate that meiosis is an adaptation for repairing DNA. These considerations form the basis for the first part of the repair and complementation hypothesis.
3503:. Cavalier-Smith's theory of the Neomuran revolution has implications for the evolutionary history of the cellular machinery for recombination and sex. It suggests that this machinery evolved in two distinct bouts separated by a long period of stasis; first the appearance of recombination machinery in a bacterial ancestor which was maintained for 3 Gy(billion years), until the neomuran revolution when the mechanics were adapted to the presence of
1768:. Each mutation that arises in asexually reproducing organisms turns the ratchet once. The ratchet is unable to be rotated backwards, only forwards. The next mutation that occurs turns the ratchet once more. Additional mutations in a population continually turn the ratchet and the mutations, mostly deleterious, continually accumulate without recombination. These mutations are passed onto the next generation because the offspring are exact genetic
9003:
9550:
2040:
1129:
974:
1381:). The number of sexuals, the number asexuals, and the rates of parasite infection for both were monitored. It was found that clones that were plentiful at the beginning of the study became more susceptible to parasites over time. As parasite infections increased, the once plentiful clones dwindled dramatically in number. Some clonal types disappeared entirely. Meanwhile, sexual snail populations remained much more stable over time.
1081:
from the population (i.e. sex aids in the removal of deleterious genes). However, in organisms containing only one set of chromosomes, deleterious mutations would be eliminated immediately, and therefore removal of harmful mutations is an unlikely benefit for sexual reproduction. Lastly, sex creates new gene combinations that may be more fit than previously existing ones, or may simply lead to reduced competition among relatives.
3356:
1524:
899:
9562:
1467:
1226:
577:
3507:. The archaeal products of the revolution maintained recombination machinery that was essentially bacterial, whereas the eukaryotic products broke with this bacterial continuity. They introduced cell fusion and ploidy cycles into cell life histories. Cavalier-Smith argues that both bouts of mechanical evolution were motivated by similar selective forces: the need for accurate DNA replication without loss of viability.
1880:, the mutation rate per effective genome per sexual generation is 0.036. Secondly, there should be strong interactions among loci (synergistic epistasis), a mutation-fitness relation for which there is only limited evidence. Conversely, there is also the same amount of evidence that mutations show no epistasis (purely additive model) or antagonistic interactions (each additional mutation has a disproportionally
1254:. However, should the same two alleles arise in different members of an asexual population, the only way that one chromosome can develop the other allele is to independently gain the same mutation, which would take much longer. Several studies have addressed counterarguments, and the question of whether this model is sufficiently robust to explain the predominance of sexual versus asexual reproduction remains.
1653:, are adaptive responses to the two major sources of "noise" in transmitting genetic information. Genetic noise can occur as either physical damage to the genome (e.g. chemically altered bases of DNA or breaks in the chromosome) or replication errors (mutations). This alternative view is referred to as the repair and complementation hypothesis, to distinguish it from the traditional variation hypothesis.
960:
advantages given by sexual differentiation, i.e. the benefits of two separate sexes compared to hermaphrodites rather than to explain benefits of sexual forms (hermaphrodite + dioecious) over asexual ones. It has already been understood that since sexual reproduction is not associated with any clear reproductive advantages over asexual reproduction, there should be some important advantages in evolution.
3822:
generation; … " "But from the sexual, or amatorial, generation of plants new varieties, or improvements, are frequently obtained; as many of the young plants from seeds are dissimilar to the parent, and some of them supererior to the parent in the qualities we wish to possess; … " " … another advantage occurs from sexual generation, which is the production of new species of plants, or mules, … "
3499:. The designation "Neomuran revolution" refers to the appearances of the common ancestors of eukaryotes and archaea. Cavalier-Smith proposes that the first neomurans emerged 850 million years ago. Other molecular biologists assume that this group appeared much earlier, but Cavalier-Smith dismisses these claims because they are based on the "theoretically and empirically" unsound model of
1506:) engage in sexual reproduction but all members of the species are capable of bearing offspring. The two-fold reproductive disadvantage assumes that males contribute only genes to their offspring and sexual females spend half their reproductive potential on sons. Thus, in this formulation, the principal cost of sex is that males and females must successfully
1700:, who concluded that the adaptive advantage of sex is hybrid vigor; or as he put it, "the offspring of two individuals, especially if their progenitors have been subjected to very different conditions, have a great advantage in height, weight, constitutional vigor and fertility over the self fertilised offspring from either one of the same parents."
1828:. It assumes that the majority of deleterious mutations are only slightly deleterious, and affect the individual such that the introduction of each additional mutation has an increasingly large effect on the fitness of the organism. This relationship between number of mutations and fitness is known as
3482:
The two viruses established in the cell would initiate replication in response to signals from the host cell. A mitosis-like cell cycle would proceed until the viral membranes dissolved, at which point linear chromosomes would be bound together with centromeres. The homologous nature of the two viral
1991:
bubbles. The contact among the pre-biotic bubbles could, through simple food or parasitic reactions, promote the transfer of genetic material from one bubble to another. That interactions between two organisms be in balance appear to be a sufficient condition to make these interactions evolutionarily
1859:
There has been much criticism of
Kondrashov's theory, since it relies on two key restrictive conditions. The first requires that the rate of deleterious mutation should exceed one per genome per generation in order to provide a substantial advantage for sex. While there is some empirical evidence for
1462:
generation). Contrast this outcome with an asexual species, in which each and every member of an equally sized 100-organism population is capable of bearing young. If all capable members of this asexual population procreated once, a total of 100 offspring would be produced – twice as many as produced
1114:
In contrast to the view that sex promotes genetic variation, Heng, and
Gorelick and Heng reviewed evidence that sex actually acts as a constraint on genetic variation. They consider that sex acts as a coarse filter, weeding out major genetic changes, such as chromosomal rearrangements, but permitting
1080:
in the same individual (i.e. sex aids in the spread of advantageous traits) without the mutations having to have occurred one after another in a single line of descendants. Second, sex acts to bring together currently deleterious mutations to create severely unfit individuals that are then eliminated
3287:
Exposure to conditions that cause RNA damage could have led to blockage of replication and death of these early RNA life forms. Sex would have allowed re-assortment of segments between two individuals with damaged RNA, permitting undamaged combinations of RNA segments to come together, thus allowing
3262:
in eukaryotic microorganisms are induced by stressful circumstances such as overcrowding, resource depletion, and DNA damaging conditions. This suggests that these sexual processes are adaptations for dealing with stress, particularly stress that causes DNA damage. In bacteria, these stresses induce
1790:
suggest that this unicellular initial stage is important for resisting mutations due to the importance of high relatedness. Highly related individuals are more closely related, and more clonal, whereas less related individuals are less so, increasing the likelihood that an individual in a population
1707:
In the view of the repair and complementation hypothesis, the removal of DNA damage by recombinational repair produces a new, less deleterious form of informational noise, allelic recombination, as a by-product. This lesser informational noise generates genetic variation, viewed by some as the major
1703:
However, outcrossing may be abandoned in favor of parthenogenesis or selfing (which retain the advantage of meiotic recombinational repair) under conditions in which the costs of mating are very high. For instance, costs of mating are high when individuals are rare in a geographic area, such as when
1585:
are genes that distort meiosis to produce gametes containing themselves more than the 50% of the time expected by chance. A selfish cytoplasmic gene is a gene located in an organelle, plasmid or intracellular parasite that modifies reproduction to cause its own increase at the expense of the cell or
1294:
When an environment changes, previously neutral or deleterious alleles can become favourable. If the environment changed sufficiently rapidly (i.e. between generations), these changes in the environment can make sex advantageous for the individual. Such rapid changes in environment are caused by the
1897:
Geodakyan suggested that sexual dimorphism provides a partitioning of a species' phenotypes into at least two functional partitions: a female partition that secures beneficial features of the species and a male partition that emerged in species with more variable and unpredictable environments. The
1580:
Sexual reproduction implies that chromosomes and alleles segregate and recombine in every generation, but not all genes are transmitted together to the offspring. There is a chance of spreading mutants that cause unfair transmission at the expense of their non-mutant colleagues. These mutations are
1433:
Critics of the Red Queen hypothesis question whether the constantly changing environment of hosts and parasites is sufficiently common to explain the evolution of sex. In particular, Otto and
Nuismer presented results showing that species interactions (e.g. host vs parasite interactions) typically
955:
The concept of sex includes two fundamental phenomena: the sexual process (fusion of genetic information of two individuals) and sexual differentiation (separation of this information into two parts). Depending on the presence or absence of these phenomena, all of the existing forms of reproduction
3460:
For these transformations to lead to the eukaryotic cell cycle, the VE hypothesis specifies a pox-like virus as the lysogenic virus. A pox-like virus is a likely ancestor because of its fundamental similarities with eukaryotic nuclei. These include a double stranded DNA genome, a linear chromosome
1442:
The paradox of the existence of sexual reproduction is that though it is ubiquitous in multicellular organisms, there are ostensibly many inherent disadvantages to reproducing sexually when weighed against the relative advantages of alternative forms of reproduction, such as asexual reproduction.
1341:
In reality, there will be several genes involved in the relationship between hosts and parasites. In an asexual population of hosts, offspring will only have the different parasitic resistance if a mutation arises. In a sexual population of hosts, however, offspring will have a new combination of
959:
Reproductive advantages of the asexual forms are in quantity of the progeny, and the advantages of the hermaphrodite forms are in maximal diversity. Transition from the hermaphrodite to dioecious state leads to a loss of at least half of the diversity. So, the primary challenge is to explain the
1910:
breaks up favourable gene combinations more often than it creates them, and sex is maintained because it ensures selection is longer-term than in asexual populations – so the population is less affected by short-term changes. This explanation is not widely accepted, as its assumptions are very
1719:
can have many different effects upon an organism. It is generally believed that the majority of non-neutral mutations are deleterious, which means that they will cause a decrease in the organism's overall fitness. If a mutation has a deleterious effect, it will then usually be removed from the
1637:
As discussed in the earlier part of this article, sexual reproduction is conventionally explained as an adaptation for producing genetic variation through allelic recombination. As acknowledged above, however, serious problems with this explanation have led many biologists to conclude that the
1454:
For example, assume that the entire population of some theoretical species has 100 total organisms consisting of two sexes (i.e. males and females), with 50:50 male-to-female representation, and that only the females of this species can bear offspring. If all capable members of this population
3821:
From p. 104: "As the progeny by lateral generation so exactly resembles the parent stock, it follows, that though any new variety, or improvement, may be thus continued for a century or two, as in grafted fruit-trees, yet that no new variety or improvements can be obtained by this mode of
3283:
that preceded DNA cellular life forms. One proposed origin of sex in the RNA world was based on the type of sexual interaction that is known to occur in extant single-stranded segmented RNA viruses, such as influenza virus, and in extant double-stranded segmented RNA viruses such as reovirus.
1103:), and different hypotheses may apply in different species. However, a research framework based on creation of variation has yet to be found that allows one to determine whether the reason for sex is universal for all sexual species, and, if not, which mechanisms are acting in each species.
1664:
it is an adaptation for repairing the genomic DNA which is passed on to progeny. Recombinational repair is the only repair process known which can accurately remove double-strand damages in DNA, and such damages are both common in nature and ordinarily lethal if not repaired. For instance,
3270:
If environmental stresses leading to DNA damage were a persistent challenge to the survival of early microorganisms, then selection would likely have been continuous through the prokaryote to eukaryote transition, and adaptative adjustments would have followed a course in which bacterial
1245:
Sex could be a method by which novel genotypes are created. Because sex combines genes from two individuals, sexually reproducing populations can more easily combine advantageous genes than can asexual populations. If, in a sexual population, two different advantageous
1855:
these genotypes, creating some individuals with fewer deleterious mutations, and some with more. Because there is a major selective disadvantage to individuals with more mutations, these individuals die out. In essence, sex compartmentalises the deleterious mutations.
3416:(fusion sex) arose from prokaryan unilateral sex-as-infection, when infected hosts began swapping nuclearised genomes containing coevolved, vertically transmitted symbionts that provided protection against horizontal superinfection by other, more virulent symbionts.
1791:
of low relatedness may have a detrimental mutation. Highly related populations also tend to thrive better than lowly related because the cost of sacrificing an individual is greatly offset by the benefit gained by its relatives and in turn, its genes, according to
956:
can be classified as asexual, hermaphrodite or dioecious. The sexual process and sexual differentiation are different phenomena, and, in essence, are diametrically opposed. The first creates (increases) diversity of genotypes, and the second decreases it by half.
2020:. Therefore, current sexual species could be descendants of primitive organisms that practiced more stable exchanges in the long term, while asexual species have emerged, much more recently in evolutionary history, from the conflict of interest resulting from
1923:
An information theoretic analysis using a simplified but useful model shows that in asexual reproduction, the information gain per generation of a species is limited to 1 bit per generation, while in sexual reproduction, the information gain is bounded by
3300:
that exchange genetic material (that is: copies of their own genome) for their transmission and propagation. In some organisms, sexual reproduction has been shown to enhance the spread of parasitic genetic elements (e.g. yeast, filamentous fungi).
3169:
organisms likely derive from a single-celled common ancestor. It is probable that the evolution of sex was an integral part of the evolution of the first eukaryotic cell. There are a few species which have secondarily lost this feature, such as
1784:, which is unicellular. This passage through a single cell is beneficial in that it lowers the chance of mutations from being passed on through multiple individuals. Instead, the mutation is only passed onto one individual. Further studies using
3210:
mechanism would have no source from which to recover the original undamaged sequence. The most primitive form of sex may have been one organism with damaged DNA replicating an undamaged strand from a similar organism in order to repair itself.
1775:
For sexually reproducing populations, studies have shown that single-celled bottlenecks are beneficial for resisting mutation build-up. Passaging a population through a single-celled bottleneck involves the fertilization event occurring with
799:
advantages because, despite the two-fold cost of sex (see below), it dominates among multicellular forms of life, implying that the fitness of offspring produced by sexual processes outweighs the costs. Sexual reproduction derives from
1753:, DNA is also susceptible to mutations within the sequence that can affect an organism in a negative manner. Asexual organisms do not have the ability to recombine their genetic information to form new and differing alleles. Once a
1263:
Supporters of these theories respond to the balance argument that the individuals produced by sexual and asexual reproduction may differ in other respects too – which may influence the persistence of sexuality. For example, in the
2108:
accumulation can lead to an increased advantage to sex under conditions not otherwise predicted (Nature, 1994). Using computer simulations, they showed that when the two mechanisms act simultaneously the advantage to sex over
1757:
occurs in the DNA or other genetic carrying sequence, there is no way for the mutation to be removed from the population until another mutation occurs that ultimately deletes the primary mutation. This is rare among organisms.
787:
of reproduction compared to sexual reproduction, where 50% of offspring are males and unable to produce offspring themselves. In hermaphroditic reproduction, each of the two parent organisms required for the formation of a
5012:
1598:. Again, however, this is not applicable to all sexual organisms. There are numerous species which are sexual but do not have a genetic-loss problem because they do not produce males or females. Yeast, for example, are
2011:
formation and genetic exchanges. So that, rather than providing selective advantages through reproduction, sex could be thought of as a series of separate events which combines step-by-step some very weak benefits of
1498:
and allows eggs to develop into offspring genetically identical to the mother by mitotic division. The mutant-asexual lineage would double its representation in the population each generation, all else being equal.
885:, Harris Bernstein, Carol Bernstein, Michael M. Cox, Frederic A. Hopf and Richard E. Michod – have suggested competing explanations for how a vast array of different living species maintain sexual reproduction.
1407:
to generate a host-parasite coevolutionary system in a controlled environment, allowing them to conduct more than 70 evolution experiments testing the Red Queen
Hypothesis. They genetically manipulated the
1799:
showed that conditions of high relatedness resisted mutant individuals more effectively than those of low relatedness, suggesting the importance of high relatedness to resist mutations from proliferating.
760:), but it is unclear if these processes are evolutionarily related to sexual reproduction in Eukaryotes. In eukaryotes, true sexual reproduction by meiosis and cell fusion is thought to have arisen in the
3186:
Organisms need to replicate their genetic material in an efficient and reliable manner. The necessity to repair genetic damage is one of the leading theories explaining the origin of sexual reproduction.
1443:
Thus, because sexual reproduction abounds in complex multicellular life, there must be some significant benefit(s) to sex and sexual reproduction that compensates for these fundamental disadvantages.
1424:
were rapidly driven extinct by the coevolving parasites while sex allowed populations to keep pace with their parasites, a result consistent with the Red Queen
Hypothesis. In natural populations of
869:(complementation) "is amply sufficient to account for the … genesis of the two sexes". This is consistent with the repair and complementation hypothesis, described below. Since the emergence of the
3427:
While theories positing fitness benefits that led to the origin of sex are often problematic, several theories addressing the emergence of the mechanisms of sexual reproduction have been proposed.
1772:
of their parents. The genetic load of organisms and their populations will increase due to the addition of multiple deleterious mutations and decrease the overall reproductive success and fitness.
1451:
Among the most limiting disadvantages to the evolution of sexual reproduction by natural selection is that an asexual population can grow much more rapidly than a sexual one with each generation.
1606:
which fuse and recombine their haploid genomes. Both sexes reproduce during the haploid and diploid stages of their life cycle and have a 100% chance of passing their genes into their offspring.
5191:
Kuzdzal-Fick, Jennie J.; Fox, Sara A.; Strassmann, Joan E.; Queller, David C. (16 December 2011). "High
Relatedness Is Necessary and Sufficient to Maintain Multicellularity in Dictyostelium".
1260:
suggested that sex might facilitate the spread of advantageous genes by allowing them to better escape their genetic surroundings, if they should arise on a chromosome with deleterious genes.
1581:
referred to as "selfish" because they promote their own spread at the cost of alternative alleles or of the host organism; they include nuclear meiotic drivers and selfish cytoplasmic genes.
3986:
Bernstein H, Byerly HC, Hopf FA, Michod RE (September 1985). "Genetic damage, mutation, and the evolution of sex". Science. 229 (4719): 1277–81 doi:10.1126/science.3898363. PMID 3898363
1594:
A sexually reproducing organism only passes on ~50% of its own genetic material to each L2 offspring. This is a consequence of the fact that gametes from sexually reproducing species are
4270:"Functional constraints against variations on molecules from the tissue-level - slowly evolving brain-specific genes demonstrated by protein-kinase and immunoglobulin supergene families"
1851:
Kondrashov argues that the slightly deleterious nature of mutations means that the population will tend to be composed of individuals with a small number of mutations. Sex will act to
1096:
to each sexual partner is avoidance of the bad effects of their deleterious recessive genes in progeny by the masking effect of normal dominant genes contributed by the other partner.
783:– possess apparent advantages over it. Asexual reproduction can proceed by budding, fission, or spore formation and does not involve the union of gametes, which accordingly results in
1992:
efficient, i.e. to select bubbles that tolerate these interactions ("libertine" bubbles) through a blind evolutionary process of self-reinforcing gene correlations and compatibility.
1704:
there has been a forest fire and the individuals entering the burned area are the initial ones to arrive. At such times mates are hard to find, and this favors parthenogenic species.
6305:
Bernstein, H., Bernstein, C. Evolutionary origin and adaptive function of meiosis. In "Meiosis", Intech Publ (Carol
Bernstein and Harris Bernstein editors), Chapter 3: 41-75 (2013).
4126:
Gorelick R, Viillablanca, FX. Meiosis decreases recombination load: Mitosis increases recombination load. Ideas in
Ecology and Evolution 11:19-28, 2018 doi:10.4033/iee.2018.11.3.n
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arise at different loci on a chromosome in different members of the population, a chromosome containing the two advantageous alleles can be produced within a few generations by
1237:
occur at random. The two alleles are recombined rapidly in a sexual population (top), but in an asexual population (bottom) the two alleles must independently arise because of
1375:
Further evidence for the Red Queen hypothesis was provided by observing long-term dynamics and parasite coevolution in a "mixed" (sexual and asexual) population of snails (
1780:
sets of DNA, forming one fertilized cell. For example, humans undergo a single-celled bottleneck in that the haploid sperm fertilizes the haploid egg, forming the diploid
1947:
1502:
Technically the problem above is not one of sexual reproduction but of having a subset of organisms incapable of bearing offspring. Indeed, some multicellular organisms (
1099:
The classes of hypotheses based on the creation of variation are further broken down below. Any number of these hypotheses may be true in any given species (they are not
3263:
an altered physiologic state, termed competence, that allows active take-up of DNA from a donor bacterium and the integration of this DNA into the recipient genome (see
3419:
Consequently, sex-as-meiosis (fission sex) would evolve as a host strategy for uncoupling from (and thereby render impotent) the acquired symbiotic/parasitic genes.
1845:
with several minor faults. Each is not sufficient alone to prevent the car from running, but in combination, the faults combine to prevent the car from functioning.
1076:
For the advantage due to genetic variation, there are three possible reasons this might happen. First, sexual reproduction can combine the effects of two beneficial
7297:
3998:
Bernstein H, Hopf FA, Michod RE (1987). "The molecular basis of the evolution of sex". Adv Genet. 24: 323–70. doi:10.1016/s0065-2660(08)60012-7. PMID 3324702
1416:, causing populations to mate either sexually, by self-fertilization, or a mixture of both within the same population. Then they exposed those populations to the
775:
form) by natural selection in a highly competitive world has long been one of the major mysteries of biology, since both other known mechanisms of reproduction –
8725:
5986:"Bangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes"
306:
1969:
1510:, which almost always involves expending energy to come together through time and space. Asexual organisms need not expend the energy necessary to find a mate.
771:), most current work has focused on the persistence of sexual reproduction over evolutionary time. The maintenance of sexual reproduction (specifically, of its
5426:
Sohail, M; Vakhrusheva, OA; Sul, JH; Pulit, SL; Francioli, LC; van den Berg, LH; Veldink, JH; de Bakker, PIW; Bazykin, GA; Kondrashov, AS; Sunyaev, SR (2017).
3342:: One primitive organism ate another one, but instead of completely digesting it, some of the eaten organism's DNA was incorporated into the DNA of the eater.
1764:
introduced the idea that mutations build up in asexual reproducing organisms. Muller described this occurrence by comparing the mutations that accumulate as a
1392:, they found that the prevalence, abundance and mean intensity of mites in sexual geckos was significantly higher than in asexuals sharing the same habitat.
3465:
repeats, a complex membrane bound capsid, the ability to produce capped mRNA, and the ability to export the capped mRNA across the viral membrane into the
3537:? Grouping itself offers a survival advantage. A binary recognition based system is the most simple and effective method in maintaining species grouping.
1092:(assuming the initial mutation rate is higher than optimal), since this removal allows greater survival of progeny with undamaged DNA. The advantage of
3475:
division in the VE hypothesis arose because of the evolutionary pressures placed on the lysogenic virus as a result of its inability to enter into the
3412:
Sex may also be derived from another prokaryotic process. A comprehensive theory called "origin of sex as vaccination" proposes that eukaryan sex-as-
2061:
1146:
991:
716:. Sexual reproduction is widespread in eukaryotes, though a few eukaryotic species have secondarily lost the ability to reproduce sexually, such as
3469:. The presence of a lysogenic pox-like virus ancestor explains the development of meiotic division, an essential component of sexual reproduction.
4826:
4213:
2101:
1283:
One of the most widely discussed theories to explain the persistence of sex is that it is maintained to assist sexual individuals in resisting
1265:
1193:
1038:
621:
3307:
is a form of genetic exchange that some sources describe as "sex", but technically is not a form of reproduction, even though it is a form of
1165:
1010:
8474:
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2125:
229:
4559:
Barrière A, Félix MA (July 2005). "High local genetic diversity and low outcrossing rate in Caenorhabditis elegans natural populations".
1848:
Similarly, an organism may be able to cope with a few defects, but the presence of many mutations could overwhelm its backup mechanisms.
1172:
1017:
540:
5277:
3519:
Why does sexual reproduction exist, if in many organisms it has a 50% cost (fitness disadvantage) in relation to asexual reproduction?
8978:
8433:
7629:
4094:
Birdsell, JA; Wills, C (2003). "The Evolutionary Origin and Maintenance of Sexual Recombination: A Review of Contemporary Models".
348:
1816:, which is represented by the red line – each subsequent mutation has a disproportionately large effect on the organism's fitness.
1494:. In his manuscript, Smith further speculated on the impact of an asexual mutant arising in a sexual population, which suppresses
1179:
1024:
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7329:
5985:
2155:
9529:
7784:
3366:
1534:
909:
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can provide either the male or the female gamete, which leads to advantages in both size and genetic variance of a population.
54:
7293:
5898:
1724:. Sexual reproduction is believed to be more efficient than asexual reproduction in removing those mutations from the genome.
1430:, self-fertilization is the predominant mode of reproduction, but infrequent out-crossing events occur at a rate of about 1%.
8741:
8501:
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1115:
minor variation, such as changes at the nucleotide or gene level (that are often neutral) to pass through the sexual sieve.
8098:
4362:"The Maintenance of Sex, Clonal Dynamics, and Host-Parasite Coevolution in a Mixed Population of Sexual and Asexual Snails"
676:
482:
3288:
survival. Such a regeneration phenomenon, known as multiplicity reactivation, occurs in the influenza virus and reovirus.
9494:
8824:
7744:
6505:
Barry RD (1961). "The multiplication of influenza virus. II. Multiplicity reactivation of ultraviolet irradiated virus".
4008:
Heng HH; Heng, Henry H.Q. (2007). "Elimination of altered karyotypes by sexual reproduction preserves species identity".
804:, where parent genotypes are reorganized and shared with the offspring. This stands in contrast to single-parent asexual
4437:
Hanley KA; Fisher RN; Case TJ (1995). "Lower mite infestations in an asexual gecko compared with its sexual ancestors".
1106:
On the other hand, the maintenance of sex based on DNA repair and complementation applies widely to all sexual species.
9339:
9067:
8486:
8146:
7209:
6122:
4799:
4156:
614:
343:
174:
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6783:
Bell, PJ (2006). "Sex and the eukaryotic cell cycle is consistent with a viral ancestry for the eukaryotic nucleus".
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4361:
4309:
Wolfe KH, Sharp PM; Sharp (1993). "Mammalian gene evolution - nucleotide-sequence divergence between mouse and rat".
3399:
2087:
1567:
1212:
1057:
942:
8553:
2069:
1688:, the second fundamental aspect of sex, is maintained by the advantage of masking mutations and the disadvantage of
9082:
9077:
8156:
7684:
5278:"Factors affecting the genetic load in Drosophila: synergistic epistasis and correlations among fitness components"
3196:
1812:
Diagram illustrating different relationships between numbers of mutations and fitness. Kondrashov's model requires
1666:
1275:, sexual offspring form eggs which are better able to survive the winter versus those the fleas produce asexually.
76:
8803:
8793:
8717:
7992:
7109:
7004:
3228:
1876:(fungus), the mutation rate per genome per replication are 0.0027 and 0.0030 respectively. For the nematode worm
761:
333:
301:
6011:
5326:
Elena, S. F.; Lenski, R. E. (1997). "Test of synergistic interactions among deleterious mutations in bacteria".
5296:
4671:
Otto SP, Gerstein AC; Gerstein (August 2006). "Why have sex? The population genetics of sex and recombination".
8931:
8798:
8578:
8322:
8120:
8024:
7913:
7607:
7250:
7002:
Hurst, L.D.; J.R. Peck (1996). "Recent advances in the understanding of the evolution and maintenance of sex".
4164:
3165:, although a later date, 1.2 billion years ago, has also been presented. Nonetheless, all sexually reproducing
2922:
2720:
2065:
1150:
995:
882:
555:
338:
9473:
6318:
Bernstein, H (2017). "Sexual communication in archaea, the precursor to meiosis". In Witzany, Guenther (ed.).
1186:
1031:
9594:
9324:
8996:
8583:
8327:
8066:
7664:
7110:"Scientists put sex origin mystery to bed, Wild strawberry research provides evidence on when gender emerges"
5009:
Darwin CR (1876). The effects of cross and self fertilisation in the vegetable kingdom. London: John Murray.
870:
607:
594:
8528:
6041:
9599:
8936:
8438:
7679:
7479:
7462:
3311:. However, it does support the "selfish gene" part theory, since the gene itself is propagated through the
2780:
1349:'s Red Queen, sexual hosts are continually "running" (adapting) to "stay in one place" (resist parasites).
525:
3377:
3373:
1676:
stage of the sexual cycle, which was at first transient, became the predominant stage, because it allowed
1545:
1541:
920:
916:
9566:
9291:
9006:
8600:
7408:
7403:
7398:
7393:
7322:
5787:
4480:
Morran, Levi T.; Schmidt, Olivia G.; Gelarden, Ian A.; Parrish Rc, Raymond C.; Lively, Curtis M. (2011).
3161:. In the eukaryotic fossil record, sexual reproduction first appeared about 2.0 billion years ago in the
2396:
2148:
581:
808:, where the offspring is always identical to the parents (barring mutation). Recombination supplies two
9540:
9483:
9349:
9181:
8702:
7876:
7617:
7457:
5540:
Trofimova, I. (2015). "Do psychological sex differences reflect evolutionary bi-sexual partitioning?".
1677:
1093:
829:
81:
2484:
9604:
9176:
8496:
8293:
8103:
7871:
7764:
7447:
3232:
3023:
2740:
1229:
This diagram illustrates how sex might create novel genotypes more rapidly. Two advantageous alleles
1100:
1071:
753:
447:
422:
402:
382:
59:
8625:
6144:
4538:"Sex -- As We Know It -- Works Thanks to Ever-Evolving Host-Parasite Relationships, Biologists Find"
3877:
1692:(mating with a close relative) which allows expression of recessive mutations (commonly observed as
8919:
8783:
8755:
8730:
8687:
8590:
8523:
8398:
8271:
8239:
8212:
8202:
7388:
5485:"Background Selection From Unlinked Sites Causes Nonindependent Evolution of Deleterious Mutations"
3845:
3592:
3381:
3308:
3192:
2700:
2364:
2050:
1786:
1549:
1377:
924:
768:
437:
432:
407:
362:
328:
322:
311:
8403:
3441:
The viral eukaryogenesis (VE) theory proposes that eukaryotic cells arose from a combination of a
3271:
transformation or archaeal DNA transfer naturally gave rise to sexual reproduction in eukaryotes.
1609:
Some species avoid the 50% cost of sexual reproduction, although they have "sex" (in the sense of
1088:, there is an immediate large benefit of removing DNA damage by recombinational DNA repair during
9417:
9354:
9329:
9212:
9132:
8924:
8854:
8763:
8491:
8373:
8141:
7737:
7669:
7157:
7135:
6961:
4537:
4415:
3858:
3231:. In extant organisms, proteins with central functions in meiosis are similar to key proteins in
2054:
1928:
1388:
gecko species and its two related sexual ancestral species. Contrary to expectation based on the
1139:
984:
757:
550:
457:
452:
397:
358:
159:
6246:
Dacks J, Roger AJ (June 1999). "The first sexual lineage and the relevance of facultative sex".
5148:
Niklas, Karl J. (1 January 2014). "The evolutionary-developmental origins of multicellularity".
1352:
Evidence for this explanation for the evolution of sex is provided by comparison of the rate of
9402:
9301:
9286:
9255:
9156:
8963:
8612:
8481:
8443:
8344:
8315:
8288:
8283:
7881:
7585:
7580:
7376:
7371:
7315:
5990:
4046:
Gorelick R, Heng HH; Heng (2011). "Sex reduces genetic variation: a multidisciplinary review".
3638:
3134:
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2428:
2141:
1426:
1397:
462:
417:
239:
134:
8469:
6222:
6216:
5929:
3834:
3814:
3585:"Interactive Tree of Life (iTOL): An online tool for phylogenetic tree display and annotation"
963:
9513:
9407:
9334:
9233:
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9166:
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8778:
8697:
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8378:
8334:
8298:
8266:
8207:
8185:
8166:
8029:
7982:
7933:
7928:
7886:
7689:
6892:
5788:"The origin of sex was interaction, not reproduction (what's sex really all about), Big Idea"
4830:
3492:
3304:
3109:
2821:
2013:
1988:
1907:
1852:
1761:
1693:
1657:
1646:
1610:
1251:
824:
801:
560:
442:
387:
353:
266:
4890:
3631:"Interactive Tree of Life v2: Online annotation and display of phylogenetic trees made easy"
1808:
9422:
9265:
9260:
9142:
9137:
9124:
8878:
8773:
8707:
8508:
8423:
8339:
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7753:
7624:
7013:
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6428:
6255:
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5627:
5584:
5439:
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5114:
5063:
4964:
4917:
4715:
4629:
4578:
4493:
4318:
4222:
3833:
Weismann, August (1889). Poulton, Edward B.; Schönland, Selmar; Shipley, Arthur E. (eds.).
3436:
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2110:
1507:
1389:
1288:
776:
721:
412:
184:
69:
26:
8513:
8383:
4286:
4269:
3841:
Ch. 5: The Significance of Sexual Reproduction in the Theory of Natural Selection (1886)
3206:
individual, on the other hand, is more likely to become fixed (i.e. permanent), since any
767:
Since hypotheses for the origin of sex are difficult to verify experimentally (outside of
8:
9518:
8941:
8899:
8849:
8768:
8616:
8608:
8538:
8518:
8464:
8308:
8061:
8004:
7866:
7849:
7827:
7521:
7489:
7278:
7201:
2882:
2841:
1987:'s "libertine bubble theory", sex originated from an archaic gene transfer process among
1765:
1740:
1403:
1353:
1238:
502:
492:
427:
392:
276:
179:
104:
39:
8219:
7230:
7017:
6796:
6738:
6688:
6475:
6432:
6259:
6071:
Bernstein H, Bernstein C (2010). "Evolutionary origin of recombination during meiosis".
6003:
5631:
5588:
5517:
5484:
5443:
5339:
5204:
5118:
5067:
4968:
4921:
4719:
4633:
4582:
4497:
4322:
4226:
1868:), there is also strong evidence against it. Thus, for instance, for the sexual species
1645:" approach to this problem has led to the view that the two fundamental aspects of sex,
9207:
8914:
8866:
8859:
8456:
8368:
8229:
8190:
8014:
7958:
7948:
7908:
7822:
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7730:
7634:
7612:
7597:
7205:
7089:
7064:
7043:
6852:
6827:
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6625:
6600:
6385:
6360:
6341:
6279:
6187:
6162:
6132:
6088:
6015:
5959:
5724:
5661:
5460:
5427:
5403:
5378:
5359:
5308:
5232:
5087:
4749:
4731:
4653:
4602:
4568:
4514:
4481:
4454:
4397:
4381:
4342:
4071:
3961:
3934:
3790:
3757:
3738:
3660:
3264:
3244:
3003:
2902:
2800:
1954:
1642:
1491:
749:
670:
545:
477:
261:
189:
154:
7288:
6935:
6908:
6576:
6551:
6483:
6440:
5803:
4929:
4620:
Otto SP, Nuismer SL; Nuismer (2004). "Species interactions and the evolution of sex".
9468:
9248:
9243:
9152:
9002:
8956:
8682:
8630:
8408:
8224:
8125:
8088:
8083:
8039:
8034:
7987:
7953:
7639:
7425:
7354:
7258:
7236:
7215:
7187:
7165:
7143:
7095:
7074:
7068:
7050:
7029:
7025:
6988:
6969:
6940:
6857:
6808:
6752:
6700:
6630:
6581:
6552:"Multiplicity reactivation of reovirus particles after exposure to ultraviolet light"
6532:
6518:
6487:
6444:
6390:
6331:
6271:
6226:
6192:
6118:
5951:
5934:
5879:
5768:
5716:
5680:
5653:
5600:
5596:
5557:
5522:
5504:
5465:
5408:
5363:
5351:
5312:
5300:
5236:
5224:
5216:
5173:
5165:
5130:
5126:
5079:
5049:
4992:
4987:
4952:
4933:
4866:
4838:
4805:
4795:
4761:
4754:
4688:
4645:
4594:
4519:
4462:
4389:
4334:
4291:
4250:
4245:
4208:
4168:
4160:
4152:
4107:
4063:
4059:
4025:
3966:
3913:
3903:
3795:
3777:
3730:
3665:
3611:
1821:
1721:
878:
858:
796:
736:. The evolution of sexual reproduction contains two related yet distinct themes: its
246:
124:
114:
109:
9452:
6764:
6345:
6092:
6019:
5963:
5728:
4735:
4346:
3892:
Crow J.F. (1994). Advantages of Sexual Reproduction, Dev. Gen., vol.15, pp. 205-213.
3606:
1665:
double-strand breaks in DNA occur about 50 times per cell cycle in human cells (see
9478:
9447:
9442:
9397:
9025:
8548:
8115:
8093:
7901:
7592:
7474:
7301:
7021:
6930:
6920:
6847:
6839:
6800:
6742:
6692:
6657:
6620:
6616:
6612:
6571:
6567:
6563:
6522:
6514:
6479:
6436:
6380:
6372:
6361:"DNA repair and the evolution of transformation in the bacterium Bacillus subtilis"
6323:
6283:
6263:
6182:
6174:
6080:
6007:
5943:
5903:
5869:
5833:
5799:
5758:
5708:
5665:
5643:
5635:
5592:
5549:
5512:
5496:
5455:
5447:
5398:
5390:
5343:
5292:
5208:
5157:
5122:
5091:
5071:
5054:
4982:
4972:
4925:
4723:
4680:
4657:
4637:
4606:
4586:
4509:
4501:
4446:
4401:
4373:
4326:
4281:
4240:
4230:
4099:
4075:
4055:
4017:
3956:
3946:
3785:
3769:
3742:
3722:
3655:
3647:
3601:
805:
647:
281:
8393:
5575:
Eshel, I.; Feldman, MW (May 1970). "On the evolutionary effect of recombination".
5394:
4482:"Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex"
3247:
search and strand exchange in the bacterial sexual process of transformation, has
1727:
There are two main hypotheses which explain how sex may act to remove deleterious
1306:
conferring two types of parasitic ability, and one gene in hosts with two alleles
845:
Reproduction, including modes of sexual reproduction, features in the writings of
9488:
9392:
9202:
8909:
8533:
8360:
8349:
8303:
8246:
8195:
7975:
7779:
7674:
7656:
7602:
7546:
6416:
5553:
5016:
3951:
3500:
2983:
1385:
854:
809:
780:
764:, possibly via several processes of varying success, and then to have persisted.
729:
234:
224:
86:
6376:
4103:
3773:
1984:
636:
9578:
9554:
9017:
8749:
8621:
8563:
7923:
7918:
7856:
7834:
7430:
6804:
6412:
4204:
3909:
3318:
A similar origin of sexual reproduction is proposed to have evolved in ancient
3175:
1697:
1622:
1474:. If each individual were to contribute to the same number of offspring (two),
1361:
1314:, conferring two types of parasite resistance, such that parasites with allele
874:
862:
850:
706:
316:
219:
6327:
4953:"Historical overview: searching for replication help in all of the rec places"
4590:
4385:
3850:
Tageblatt der 58. Versammlung Deutscher Naturforscher und Aerzte in Strassburg
849:; modern philosophical-scientific thinking on the problem dates from at least
34:
9588:
9456:
9370:
9238:
9072:
9049:
8951:
8829:
8788:
8634:
8388:
8056:
8051:
7644:
7575:
7541:
7452:
7418:
6408:
6150:
5947:
5508:
5428:"Negative selection in humans and fruit flies involves synergistic epistasis"
5220:
5169:
3781:
3534:
3413:
3323:
3064:
1916:
1792:
1582:
1409:
1365:
1346:
1257:
467:
139:
7283:
6723:"Viral eukaryogenesis: Was the ancestor of the nucleus a complex DNA virus?"
6462:
Bernstein H, Byerly HC, Hopf FA, Michod RE (October 1984). "Origin of sex".
6084:
5451:
5212:
5105:
Muller, H.J. (1964). "The Relation of Recombination to Mutational Advance".
4809:
4641:
4505:
861:, as detailed in the majority of the explanations below. On the other hand,
9281:
9102:
9092:
8946:
8894:
8839:
8672:
8667:
8261:
8019:
7711:
7563:
7440:
7435:
7381:
7033:
6944:
6925:
6895:
6861:
6843:
6812:
6756:
6704:
6696:
6536:
6275:
6196:
6178:
5955:
5883:
5772:
5763:
5746:
5720:
5712:
5657:
5561:
5526:
5469:
5376:
5304:
5228:
5177:
5134:
4996:
4977:
4893:
4692:
4649:
4598:
4523:
4466:
4393:
4235:
4067:
4029:
3970:
3799:
3734:
3669:
3615:
3454:
2942:
2680:
964:
Advantages due to genetic variation, DNA repair and genetic complementation
487:
472:
256:
251:
169:
6747:
6722:
6634:
6585:
6491:
6448:
6394:
5604:
5500:
5412:
5355:
5083:
5052:(1988). "Deleterious mutations and the evolution of sexual reproduction".
4937:
4338:
4295:
4254:
3526:(types of gametes, according to their compatibility) arise as a result of
1372:. The genes coding for immune system proteins evolve considerably faster.
1298:
Imagine, for example that there is one gene in parasites with two alleles
9217:
9197:
8973:
8844:
8573:
8108:
7938:
7844:
7802:
7179:
5925:
5699:
Lesbarrères D (2011). "Sex or no sex, reproduction is not the question".
4135:
3651:
3523:
3476:
3339:
3319:
3297:
3162:
2626:
2588:
1906:
Ilan Eshel suggested that sex prevents rapid evolution. He suggests that
1685:
1650:
1603:
497:
214:
164:
8413:
6661:
5639:
5161:
3584:
3195:, since there are two copies of the gene in the cell and if one copy is
2003:
to solve a problem that bacteria did not have, namely a large amount of
1330:. Such a situation will lead to cyclic changes in allele frequency – as
9461:
9161:
9114:
8834:
8735:
8657:
8644:
8278:
8078:
7965:
7943:
7896:
7891:
7839:
7807:
7722:
6527:
6407:
6267:
5838:
5821:
4727:
4684:
4458:
4330:
3504:
3207:
3171:
3166:
2962:
2008:
2000:
1861:
1842:
1689:
1478:
the sexual population remains the same size each generation, where the
1268:
1153: in this section. Unsourced material may be challenged and removed.
1085:
998: in this section. Unsourced material may be challenged and removed.
815:
745:
717:
655:
271:
194:
149:
129:
43:
9573:
6832:
Philosophical Transactions of the Royal Society B: Biological Sciences
5874:
5857:
4794:. Hoekstra, Rolf F. (2nd ed.). Oxford : Oxford University Press.
4573:
3846:"Die Bedeutung der sexuellen Fortpflanzung für die Selektions-Theorie"
3710:
3630:
3251:
in eukaryotes that perform similar functions in meiotic recombination
2104:
were the first to suggest that the combined effects of parasitism and
1660:
is fundamentally a DNA repair process, and that when it occurs during
9412:
9380:
9147:
9053:
8968:
8819:
8428:
8046:
7794:
7551:
7526:
7516:
7504:
7119:
6117:] (in Estonian). Vol. XXV. Tartu: Sulemees. pp. 51–56.
5747:"Sex is not a solution for reproduction: the libertine bubble theory"
5075:
3726:
3527:
3466:
3450:
3442:
3312:
3280:
3248:
3220:
2472:
2412:
2021:
1829:
1681:
866:
846:
834:
772:
710:
702:
530:
144:
64:
5648:
4908:
Bernstein H; Byerly HC; Hopf FA; Michod RE (1984). "Origin of sex".
4907:
4450:
4021:
3879:
The Effects of Cross and Self Fertilisation in the Vegetable Kingdom
3860:
Die Bedeutung der sexuellen Fortpflanzung für die Selektions-Theorie
2039:
1638:
benefit of sex is a major unsolved problem in evolutionary biology.
1128:
973:
9549:
9375:
9344:
8871:
8073:
7568:
7469:
7114:
6046:
5618:
Colegrave, N. (2002). "Sex releases the speed limit on evolution".
4377:
3462:
3279:
Sex might also have been present even earlier, in the hypothesized
3044:
2493:
2475:
2105:
1754:
1716:
1618:
1614:
1284:
1077:
725:
661:
520:
119:
6966:
The masterpiece of nature: the evolution and genetics of sexuality
5347:
4151:
MacIntyre, Ross J.; Clegg, Michael, T (Eds.), Springer. Hardcover
3267:) allowing recombinational repair of the recipients' damaged DNA.
1708:
effect of sex, as discussed in the earlier parts of this article.
857:
picked up the thread in 1885, arguing that sex serves to generate
9107:
8256:
7970:
7536:
7509:
7232:
The prehistory of sex: four million years of human sexual culture
3530:(gamete dimorphism), or did mating types evolve before anisogamy?
3496:
3472:
3446:
3327:
3259:
3255:
3236:
3224:
3203:
3188:
3146:
2560:
2527:
2017:
1996:
1983:
exchange that is independent from reproduction. According to the
1979:
The evolution of sex can alternatively be described as a kind of
1865:
1838:
1777:
1769:
1750:
1661:
1599:
1595:
1503:
1495:
1490:
of sexual reproduction. It was first described mathematically by
1369:
1247:
1089:
820:
748:) have processes that can transfer DNA from one cell to another (
713:
698:
640:
5377:
Drake JW; Charlesworth B; Charlesworth D; Crow JF (April 1998).
5190:
1466:
1455:
procreated once, a total of 50 offspring would be produced (the
1420:
parasite. It was found that the self-fertilizing populations of
1225:
9561:
9087:
7558:
7531:
7364:
4479:
3154:
2537:
2509:
1781:
1673:
1626:
1437:
1357:
789:
694:
686:
651:
576:
8177:
3199:, the other copy is unlikely to be damaged at the same site.
3158:
3150:
3089:
2444:
690:
5036:
Modern Genetic Analysis, New York, W.H. Freeman and Company.
4670:
3515:
Some questions biologists have attempted to answer include:
3422:
888:
673:
led to the evolution and maintenance of sexual reproduction?
9385:
7359:
7200:
6012:
10.1666/0094-8373(2000)026<0386:BPNGNS>2.0.CO;2
5297:
10.1554/0014-3820(2000)054[1654:fatgli]2.0.co;2
3296:
Another theory is that sexual reproduction originated from
3240:
2518:
1980:
1728:
6358:
5688:. Cambridge: Cambridge University Press. pp. 269–280.
4619:
4209:"Sexual reproduction as an adaptation to resist parasites"
3191:
individuals can repair a damaged section of their DNA via
1672:
In some lines of descent from the earliest organisms, the
654:
production is an essential step in sexual reproduction of
7338:
6982:
6828:"Cell evolution and Earth history: Stasis and revolution"
6675:
Sterrer W (2002). "On the origin of sex as vaccination".
6461:
5425:
4436:
3852:(in German) Strassburg, Germany: G. Fischbach pp. 42-56.
3219:
Sexual reproduction appears to have arisen very early in
3126:
3085:
2004:
1892:
1746:
1109:
733:
6878:. Cambridge University Press, Cambridge; 2011, p. 22-29.
6070:
3905:
36 Arguments for the Existence of God: A Work of Fiction
1920:
algae that sex can remove the speed limit on evolution.
7162:
Mendel's demon: gene justice and the complexity of life
4856:
4854:
4308:
4045:
3561:
1680:— the masking of deleterious recessive mutations (i.e.
1656:
The repair and complementation hypothesis assumes that
7307:
7040:
6825:
6601:"Selfish DNA: a sexually-transmitted nuclear parasite"
5682:
Information Theory, Inference, and Learning Algorithms
3555:
3553:
1384:
However, Hanley et al. studied mite infestations of a
9538:
7045:
The Evolution of sex: an examination of current ideas
4360:
Jokela, Jukka; Dybdahl, Mark; Lively, Curtis (2009).
4203:
3839:. Oxford, England: Clarendon Press. pp. 252–332.
3243:
recombinase, that catalyses the key functions of DNA
1957:
1931:
795:
Sexual reproduction therefore must offer significant
7184:
The Red Queen: sex and the evolution of human nature
5574:
4851:
4140:
The Red Queen: Sex and the Evolution of Human Nature
3836:
Essays Upon Heredity and Kindred Biological Problems
3254:
Natural transformation in bacteria, DNA transfer in
1734:
1395:
In 2011, researchers used the microscopic roundworm
1278:
6419:(April 1981). "The origin of genetic information".
6313:
6311:
4359:
3550:
3223:evolution, implying that the essential features of
2120:
1589:
1482:
asexual population doubles in size each generation.
1446:
873:in the 20th century, numerous biologists including
7042:
6301:
6299:
6297:
6295:
6293:
4753:
2862:
2113:is larger than for either factor operating alone.
1963:
1941:
720:, and some plants and animals routinely reproduce
6648:DasSarma, Shiladitya (2007). "Extreme Microbes".
4706:Parker MA (1994). "Pathogens and sex in plants".
4416:"Parasites May Have Had Role In Evolution Of Sex"
3755:
1711:
1632:
1463:by the sexual population in a single generation.
9586:
9047:
6549:
6359:Michod RE, Wojciechowski MF, Hoelzer MA (1988).
6308:
6066:
6064:
5483:Matheson, Joseph; Masel, Joanna (2 March 2024).
5275:
4944:
4185:Van Valen, L. (1973). "A New Evolutionary Law".
3994:
3992:
3982:
3980:
3338:A third theory is that sex evolved as a form of
2116:
7091:Eros and evolution: a natural philosophy of sex
6290:
6104:
6102:
4267:
4214:Proceedings of the National Academy of Sciences
4041:
4039:
3322:as a combination of two independent processes:
1914:It has recently been shown in experiments with
1803:
1318:can attach themselves to hosts with the allele
7257:. Princeton, N.J: Princeton University Press.
7001:
6906:
6160:
5899:"Sexual Reproduction and the Evolution of Sex"
5858:"Have sex or not ? Lessons from bacteria"
5815:
5813:
3685:"Sexual Reproduction and the Evolution of Sex"
3274:
9033:
7738:
7323:
7073:. Cambridge, UK: Cambridge University Press.
7063:
6214:
6061:
5851:
5849:
5698:
5482:
5044:
5042:
4903:
4901:
4785:
4783:
4781:
4779:
4777:
4558:
4093:
3989:
3977:
3540:Why do some organisms have gamete dimorphism?
3149:reproduce sexually, as do many multicellular
2149:
615:
7294:Sexual Reproduction and the Evolution of Sex
6968:. Berkeley: University of California Press.
6778:
6776:
6774:
6716:
6714:
6154:
6108:
6099:
6033:
5918:
5678:
4552:
4036:
3882:. London, England: John Murray. p. 462.
3756:Goodenough, U.; Heitman, J. (1 March 2014).
1995:The "libertine bubble theory" proposes that
1629:), "sex" and reproduction occur separately.
1513:
1438:Disadvantages of sex and sexual reproduction
7041:Levin, Bruce R.; Richard E. Michod (1988).
6983:Bernstein, Carol; Harris Bernstein (1991).
5983:
5810:
5740:
5738:
5325:
4821:
4819:
4089:
4087:
4085:
4007:
3863:(in German). Jena, Germany: Gustav Fischer.
3758:"Origins of Eukaryotic Sexual Reproduction"
3628:
3582:
3559:
3345:
2068:. Unsourced material may be challenged and
663:
9040:
9026:
7745:
7731:
7330:
7316:
6245:
6210:
6208:
6206:
5846:
5269:
5048:
5039:
4898:
4774:
3762:Cold Spring Harbor Perspectives in Biology
3533:Why do most sexual organisms use a binary
2156:
2142:
2007:material, occurring in an archaic step of
1974:
1486:This idea is sometimes referred to as the
1295:co-evolution between hosts and parasites.
622:
608:
7094:. Reading, Mass: Addison-Wesley Pub. Co.
6934:
6924:
6851:
6771:
6746:
6711:
6674:
6624:
6575:
6526:
6384:
6317:
6221:. New York: Metropolitan Books. pp.
6186:
6115:Why the eukaryotic cell memory was needed
5873:
5837:
5822:"Sex and the origin of genetic exchanges"
5762:
5647:
5617:
5539:
5533:
5516:
5459:
5402:
5022:
4986:
4976:
4835:The Evolution of Sex and its Consequences
4705:
4572:
4513:
4285:
4244:
4234:
4184:
3960:
3950:
3901:
3819:. Dublin, Ireland: P. Byrne. p. 104.
3789:
3659:
3605:
3423:Mechanistic origin of sexual reproduction
3400:Learn how and when to remove this message
3291:
2088:Learn how and when to remove this message
1568:Learn how and when to remove this message
1213:Learn how and when to remove this message
1058:Learn how and when to remove this message
943:Learn how and when to remove this message
889:Advantages of sex and sexual reproduction
865:(1809–1882) concluded that the effect of
840:
837:, hybrid vigor or masking of mutations).
8979:Transgenerational epigenetic inheritance
7752:
7249:
7049:. Sunderland, Mass: Sinauer Associates.
6782:
6720:
6647:
6598:
6504:
6218:Dr. Tatiana's sex advice to all creation
6163:"How oxygen gave rise to eukaryotic sex"
5890:
5735:
4825:
4816:
4268:Kuma, K.; Iwabe, N.; Miyata, T. (1995).
4197:
4082:
3856:
3832:
3708:
3372:Relevant discussion may be found on the
1807:
1540:Relevant discussion may be found on the
1465:
1224:
915:Relevant discussion may be found on the
646:
635:
6887:Beukeboom, L. & Perrin, N. (2014).
6819:
6668:
6203:
6039:
5977:
5819:
5785:
5744:
5319:
4885:Beukeboom, L. & Perrin, N. (2014).
4789:
4742:
3430:
9587:
9530:Index of evolutionary biology articles
7228:
7178:
7156:
7134:
7087:
6592:
6161:Hörandl E, Speijer D (February 2018).
6149:Abstract in English available online:
5930:"On The Origin Of Sexual Reproduction"
5924:
5276:Whitlock, M. C.; Bourguet, D. (2000).
5262:Charlesworth B, Charlesworth D (2010)
5147:
5104:
4950:
4860:
4129:
3875:
3812:
3486:
3384:to additional sources at this section.
3122:
1893:Geodakyan's evolutionary theory of sex
1552:to additional sources at this section.
1401:as a host and the pathogenic bacteria
1110:Protection from major genetic mutation
927:to additional sources at this section.
9021:
8742:Dialogues Concerning Natural Religion
7726:
7311:
7289:The Evolutionary Theory of Sex (2012)
7164:. London: Weidenfeld & Nicolson.
6907:Czárán, T.L.; Hoekstra, R.F. (2006).
5611:
4748:
4302:
4287:10.1093/oxfordjournals.molbev.a040181
3709:Redfield, Rosemary J. (August 2001).
2134:
1901:
1887:
6960:
5896:
5855:
5253:, 3rd edition. Blackwell Publishing.
4865:(3rd ed.). Wiley. p. 314.
4261:
3932:
3682:
3349:
3333:
3098:
3074:
3053:
3033:
3012:
2992:
2972:
2951:
2931:
2911:
2891:
2871:
2851:
2830:
2810:
2789:
2769:
2749:
2729:
2709:
2689:
2669:
2123:
2066:adding citations to reliable sources
2033:
1517:
1151:adding citations to reliable sources
1122:
996:adding citations to reliable sources
967:
892:
7426:Heterogametic sex / Homogametic sex
4178:
3491:An alternative theory, proposed by
1971:is the size of the genome in bits.
812:mechanisms at the molecular level:
685:describes how sexually reproducing
677:(more unsolved problems in biology)
13:
9340:Evolutionary developmental biology
8147:Evolutionary developmental biology
7211:The Major Transitions in Evolution
6954:
6889:The Evolution of Sex Determination
6042:"The Real Reasons Why We Have Sex"
4889:. Oxford University Press, p. 5–6
4887:The Evolution of Sex Determination
4207:; Axelrod, R.; Tanese, R. (1990).
3365:relies largely or entirely upon a
3298:selfish parasitic genetic elements
1533:relies largely or entirely upon a
1162:"Evolution of sexual reproduction"
1118:
1007:"Evolution of sexual reproduction"
908:relies largely or entirely upon a
14:
9616:
7272:
7214:. Oxford: W.H. Freeman Spektrum.
6891:. Oxford University Press, p. 25
6550:McClain ME, Spendlove RS (1966).
6441:10.1038/scientificamerican0481-88
5984:Butterfield, Nicholas J. (2000).
5266:. Roberts and Company Publishers.
5264:Elements of Evolutionary Genetics
4098:. Vol. 33. pp. 27–137.
1826:deterministic mutation hypothesis
1735:Evading harmful mutation build-up
1279:Increased resistance to parasites
853:(1731–1802) in the 18th century.
9572:
9560:
9548:
9297:Evolution of sexual reproduction
9001:
8992:
8991:
7707:
7706:
7500:Evolution of sexual reproduction
7142:. Oxford: Blackwell Scientific.
5032:1999. Gene mutations, p197-234,
4833:. In Stearns, Stephen C. (ed.).
4792:Evolution : an introduction
4673:Biochemical Society Transactions
4060:10.1111/j.1558-5646.2010.01173.x
3354:
3235:in bacteria and DNA transfer in
2038:
1824:, and is sometimes known as the
1820:This hypothesis was proposed by
1602:sexual organisms which have two
1590:Genetic heritability cost of sex
1522:
1447:Population expansion cost of sex
1127:
972:
897:
683:Evolution of sexual reproduction
589:
588:
575:
33:
8804:Extended evolutionary synthesis
7993:Gene-centered view of evolution
7005:Trends in Ecology and Evolution
6909:"Evolution of sexual asymmetry"
6900:
6881:
6868:
6826:Cavalier-Smith, Thomas (2006).
6641:
6543:
6498:
6455:
6401:
6352:
6239:
6040:Cumming, Vivian (4 July 2016).
5779:
5692:
5672:
5568:
5476:
5419:
5379:"Rates of spontaneous mutation"
5370:
5256:
5243:
5184:
5141:
5098:
5003:
4879:
4699:
4664:
4613:
4530:
4473:
4430:
4408:
4353:
4274:Molecular Biology and Evolution
4145:
4120:
4001:
3926:
3895:
3886:
3229:last eukaryotic common ancestor
1749:is able to recombine to modify
1138:needs additional citations for
983:needs additional citations for
762:last eukaryotic common ancestor
732:) without entirely having lost
582:Evolutionary biology portal
9068:Genotype–phenotype distinction
8932:Hologenome theory of evolution
8799:History of molecular evolution
8025:Evolutionarily stable strategy
7914:Last universal common ancestor
7608:Sexual reproduction in animals
6785:Journal of Theoretical Biology
6677:Journal of Theoretical Biology
6568:10.1128/JB.92.5.1422-1429.1966
6464:Journal of Theoretical Biology
6248:Journal of Molecular Evolution
5577:Theoretical Population Biology
5542:American Journal of Psychology
4760:. Cambridge University Press.
4311:Journal of Molecular Evolution
3869:
3825:
3806:
3749:
3702:
3676:
3622:
3575:
2668:
2029:Parasites and Muller's ratchet
1712:Deleterious mutation clearance
1667:naturally occurring DNA damage
1633:DNA repair and complementation
1342:parasitic resistance alleles.
541:Creation–evolution controversy
295:History of evolutionary theory
1:
9325:Regulation of gene expression
8726:Renaissance and Enlightenment
6484:10.1016/S0022-5193(84)80178-2
5804:10.1016/S0262-4079(11)62719-X
4930:10.1016/S0022-5193(84)80178-2
4418:. Science Daily. 31 July 2009
3607:10.1093/bioinformatics/btl529
3544:
2117:Origin of sexual reproduction
2016:, meiosis, gametogenesis and
1720:population by the process of
1470:This diagram illustrates the
871:modern evolutionary synthesis
9495:Endless Forms Most Beautiful
9275:Evolution of genetic systems
9083:Gene–environment correlation
9078:Gene–environment interaction
8937:Missing heritability problem
8564:Gamete differentiation/sexes
7480:Sex as a biological variable
7463:Simultaneous hermaphroditism
7284:The Evolution of Sexs (2003)
7026:10.1016/0169-5347(96)81041-X
6727:Journal of Molecular Biology
6617:10.1093/genetics/101.3-4.519
6519:10.1016/0042-6822(61)90330-0
5597:10.1016/0040-5809(70)90043-2
5554:10.5406/amerjpsyc.128.4.0485
5489:Genome Biology and Evolution
5127:10.1016/0027-5107(64)90047-8
4957:Proc. Natl. Acad. Sci. U.S.A
4540:. Science Daily. 9 July 2011
3952:10.1371/journal.pbio.0060052
3646:(Web Server issue): W475–8.
3629:Letunic, I; Bork, P (2011).
3583:Letunic, I; Bork, P (2007).
3560:Letunic, I; Bork, P (2006).
3510:
3227:were already present in the
1804:Removal of deleterious genes
526:Evolution as fact and theory
7:
9474:Christiane Nüsslein-Volhard
7186:. New York: Penguin Books.
7088:Michod, Richard E. (1995).
6427:(4): 88–92, 96, et passim.
6320:Biocommunication of Archaea
5395:10.1093/genetics/148.4.1667
4104:10.1007/978-1-4757-5190-1_2
3774:10.1101/cshperspect.a016154
3275:Virus-like RNA-based origin
3181:
3123:
2883:Earliest multicellular life
2353:
1942:{\displaystyle {\sqrt {G}}}
1613:). In these species (e.g.,
784:
664:Unsolved problem in biology
10:
9621:
9350:Hedgehog signaling pathway
9227:Developmental architecture
8569:Life cycles/nuclear phases
8121:Trivers–Willard hypothesis
7618:Penile-vaginal intercourse
7458:Sequential hermaphroditism
7235:. New York: Bantam Books.
6987:. Boston: Academic Press.
6985:Aging, sex, and DNA repair
6876:The Evolution of Anisogamy
6874:T. Togashi, P. Cox (Eds.)
6805:10.1016/j.jtbi.2006.05.015
6111:Biosüsteemide mälu teooria
5150:American Journal of Botany
3855:Revised and expanded in:
3562:"Interactive Tree of Life"
3434:
3214:
1738:
1696:). This is in accord with
1586:organism that carries it.
1069:
561:Nature-nurture controversy
9527:
9506:
9435:
9363:
9317:
9310:
9274:
9226:
9190:
9177:Transgressive segregation
9123:
9060:
8987:
8887:
8812:
8716:
8643:
8599:
8454:
8358:
8175:
8134:
8067:Parent–offspring conflict
8003:
7872:Earliest known life forms
7793:
7760:
7702:
7655:
7488:
7448:Testis-determining factor
7345:
6377:10.1093/genetics/118.1.31
6328:10.1007/978-3-319-65536-9
4591:10.1016/j.cub.2005.06.022
3857:Weismann, August (1886).
1682:hybrid vigor or heterosis
1514:Selfish cytoplasmic genes
1084:For the advantage due to
709:that was a single-celled
448:Evolutionary neuroscience
423:Evolutionary epistemology
403:Evolutionary anthropology
383:Applications of evolution
8920:Cultural group selection
8784:The eclipse of Darwinism
8756:On the Origin of Species
8731:Transmutation of species
7389:Sex-determination system
7229:Taylor, Timothy (1996).
6913:BMC Evolutionary Biology
5948:10.1126/science.324_1254
5015:19 February 2016 at the
4831:"The Evolution of Sexes"
3876:Darwin, Charles (1876).
3813:Darwin, Erasmus (1800).
3346:Vaccination-like process
3309:horizontal gene transfer
3193:homologous recombination
2309:−1000 —
2289:−1500 —
2269:−2000 —
2249:−2500 —
2229:−3000 —
2209:−3500 —
2189:−4000 —
2169:−4500 —
2166:
1870:Saccharomyces cerevisiae
1787:Dictyostelium discoideum
1378:Potamopyrgus antipodarum
1368:with genes coding other
1334:increases in frequency,
769:evolutionary computation
438:Evolutionary linguistics
433:Evolutionary game theory
408:Evolutionary computation
9355:Notch signaling pathway
9330:Gene regulatory network
9213:Dual inheritance theory
8925:Dual inheritance theory
8764:History of paleontology
7337:
6085:10.1525/bio.2010.60.7.5
5897:Otto, Sarah P. (2008).
5452:10.1126/science.aah5238
5213:10.1126/science.1213272
4790:Stearns, S. C. (2005).
4642:10.1126/science.1094072
4506:10.1126/science.1206360
4366:The American Naturalist
3902:Goldstein, R N (2010).
3831:English translation:
3715:Nature Reviews Genetics
3711:"Do bacteria have sex?"
2329:−500 —
1975:Libertine bubble theory
827:pair at that time) and
551:Objections to evolution
458:Evolutionary psychology
453:Evolutionary physiology
398:Evolutionary aesthetics
377:Fields and applications
359:History of paleontology
9403:cis-regulatory element
9311:Control of development
9191:Non-genetic influences
9157:evolutionary landscape
8613:Punctuated equilibrium
7934:Non-adaptive radiation
7882:Evolutionary arms race
7372:Sexual differentiation
7279:Why Sex is Good (2005)
6926:10.1186/1471-2148-4-34
6844:10.1098/rstb.2006.1842
6697:10.1006/jtbi.2002.3008
6215:Olivia Judson (2002).
6179:10.1098/rspb.2017.2706
5764:10.1002/bies.201000125
5713:10.1002/bies.201100105
4978:10.1073/pnas.131004998
4236:10.1073/pnas.87.9.3566
3844:Weismann, Aug. (1885)
3639:Nucleic Acids Research
3292:Parasitic DNA elements
3233:natural transformation
3202:A harmful damage in a
2100:R. Stephen Howard and
1965:
1943:
1878:Caenorhabditis elegans
1817:
1483:
1398:Caenorhabditis elegans
1242:
841:Historical perspective
825:homologous chromosomes
744:Bacteria and Archaea (
659:
644:
483:Speciation experiments
463:Experimental evolution
418:Evolutionary economics
240:Recent human evolution
98:Processes and outcomes
9514:Nature versus nurture
9418:Cell surface receptor
9335:Evo-devo gene toolkit
9234:Developmental biology
9172:Polygenic inheritance
9098:Quantitative genetics
8905:Evolutionary medicine
8779:Mendelian inheritance
8487:Biological complexity
8475:Programmed cell death
8167:Phenotypic plasticity
7887:Evolutionary pressure
7877:Evidence of evolution
7775:Timeline of evolution
7414:Temperature-dependent
6748:10.1007/s002390010215
6109:Ploompuu, T. (1999).
5679:David MacKay (2003).
4861:Ridley, Mark (2003).
4837:. Springer Basel AG.
3493:Thomas Cavalier-Smith
3305:Bacterial conjugation
1999:sex evolved in proto-
1966:
1944:
1831:synergistic epistasis
1814:synergistic epistasis
1811:
1762:Hermann Joseph Muller
1694:inbreeding depression
1658:genetic recombination
1647:genetic recombination
1611:genetic recombination
1469:
1345:In other words, like
1338:will be disfavoured.
1228:
1072:Hill–Robertson effect
650:
639:
443:Evolutionary medicine
388:Biosocial criminology
354:History of speciation
267:Evolutionary taxonomy
230:Timeline of evolution
9595:Evolutionary biology
9567:Evolutionary biology
9423:Transcription factor
9138:Genetic assimilation
9125:Genetic architecture
8879:Teleology in biology
8774:Blending inheritance
8152:Genetic assimilation
8015:Artificial selection
7754:Evolutionary biology
7070:The evolution of sex
6894:. Online resources,
6417:Winkler-Oswatitsch R
6322:. pp. 103–117.
4892:. Online resources,
4756:The Evolution of Sex
4708:Evolutionary Ecology
4096:Evolutionary Biology
3933:Baer, C. F. (2008).
3683:Otto, Sarah (2014).
3437:Viral eukaryogenesis
3431:Viral eukaryogenesis
3378:improve this article
2111:asexual reproduction
2062:improve this section
1955:
1929:
1546:improve this article
1472:two-fold cost of sex
1390:Red Queen hypothesis
1289:Red Queen Hypothesis
1287:, also known as the
1147:improve this article
992:improve this article
921:improve this article
777:asexual reproduction
413:Evolutionary ecology
27:Evolutionary biology
9600:Sexual reproduction
9519:Morphogenetic field
9436:Influential figures
8942:Molecular evolution
8900:Ecological genetics
8769:Transitional fossil
8559:Sexual reproduction
8399:endomembrane system
8328:pollinator-mediated
8284:dolphins and whales
8062:Parental investment
7630:Hormonal motivation
7603:Fungal reproduction
7522:Reproductive system
7122:on 19 December 2015
7065:Maynard Smith, John
7018:1996TEcoE..11...46H
6797:2006JThBi.243...54B
6739:2001JMolE..53..251L
6689:2002JThBi.216..387S
6662:10.1511/2007.65.224
6476:1984JThBi.110..323B
6433:1981SciAm.244d..88E
6421:Scientific American
6260:1999JMolE..48..779D
6004:2000Pbio...26..386B
5942:(5932): 1254–1256.
5640:10.1038/nature01191
5632:2002Natur.420..664C
5589:1970TPBio...1...88E
5501:10.1093/gbe/evae050
5444:2017Sci...356..539S
5340:1997Natur.390..395E
5205:2011Sci...334.1548K
5199:(6062): 1548–1551.
5162:10.3732/ajb.1300314
5119:1964MRFMM...1....2M
5068:1988Natur.336..435K
4969:2001PNAS...98.8173C
4922:1984JThBi.110..323B
4720:1994EvEco...8..560P
4634:2004Sci...304.1018O
4628:(5673): 1018–1020.
4583:2005CBio...15.1176B
4498:2011Sci...333..216M
4323:1993JMolE..37..441W
4227:1990PNAS...87.3566H
4187:Evolutionary Theory
3848:Stilling, J. (ed.)
3497:Neomuran revolution
3487:Neomuran revolution
2863:Sexual reproduction
2842:Huronian glaciation
1860:it (for example in
1795:. The studies with
1404:Serratia marcescens
1354:molecular evolution
1239:clonal interference
671:selection pressures
515:Social implications
503:Universal Darwinism
493:Island biogeography
428:Evolutionary ethics
393:Ecological genetics
339:Molecular evolution
277:Transitional fossil
105:Population genetics
21:Part of a series on
9208:Genomic imprinting
8915:Cultural evolution
8030:Fisher's principle
7959:Handicap principle
7949:Parallel evolution
7813:Adaptive radiation
7635:Human reproduction
7613:Sexual intercourse
7598:Plant reproduction
7206:John Maynard Smith
6838:(1470): 969–1006.
6650:American Scientist
6268:10.1007/PL00013156
6173:(1872): 20172706.
5839:10.4081/eb.2012.e1
4728:10.1007/BF01238258
4685:10.1042/BST0340519
4331:10.1007/BF00178874
3652:10.1093/nar/gkr201
3495:, was labeled the
3265:Natural competence
3110:Quaternary ice age
3045:Earliest tetrapods
3004:Cambrian explosion
2963:Cryogenian ice age
2822:Atmospheric oxygen
2801:Pongola glaciation
2429:Multicellular life
2381:Single-celled life
1961:
1939:
1902:Speed of evolution
1888:Other explanations
1818:
1492:John Maynard Smith
1484:
1243:
1101:mutually exclusive
883:George C. Williams
785:a much faster rate
660:
645:
546:Theistic evolution
478:Selective breeding
190:Parallel evolution
155:Adaptive radiation
9536:
9535:
9469:Eric F. Wieschaus
9431:
9430:
9249:Pattern formation
9153:Fitness landscape
9015:
9014:
8631:Uniformitarianism
8584:Sex-determination
8089:Sexual dimorphism
8084:Natural selection
7988:Unit of selection
7954:Signalling theory
7720:
7719:
7640:Lordosis behavior
7355:Sexual dimorphism
7264:978-0-691-08147-2
7255:Sex and evolution
7242:978-0-553-09694-1
7221:978-0-7167-4525-9
7193:978-0-14-024548-6
7171:978-0-297-64634-1
7149:978-0-632-03481-9
7101:978-0-201-40754-9
7080:978-0-521-21887-0
7056:978-0-87893-459-1
6994:978-0-12-092860-6
6975:978-0-520-04583-5
6721:Bell, PJ (2001).
6599:Hickey D (1982).
6337:978-3-319-65535-2
6232:978-0-8050-6331-8
5875:10.1159/000342879
5626:(6916): 664–666.
5438:(6337): 539–542.
5334:(6658): 395–398.
5107:Mutation Research
5062:(6198): 435–440.
5050:Kondrashov, A. S.
4827:Hoekstra, Rolf F.
4750:Smith, J. Maynard
4492:(6039): 216–218.
4173:978-1-4419-3385-0
4113:978-1-4419-3385-0
3919:978-0-307-37818-7
3410:
3409:
3402:
3334:Partial predation
3143:
3142:
3127:million years ago
3118:
3117:
3097:
3096:
3073:
3072:
3052:
3051:
3032:
3031:
3024:Andean glaciation
3011:
3010:
2991:
2990:
2971:
2970:
2950:
2949:
2930:
2929:
2910:
2909:
2890:
2889:
2870:
2869:
2850:
2849:
2829:
2828:
2809:
2808:
2788:
2787:
2768:
2767:
2748:
2747:
2728:
2727:
2708:
2707:
2688:
2687:
2098:
2097:
2090:
1964:{\displaystyle G}
1937:
1874:Neurospora crassa
1822:Alexey Kondrashov
1731:from the genome.
1722:natural selection
1578:
1577:
1570:
1223:
1222:
1215:
1197:
1068:
1067:
1060:
1042:
953:
952:
945:
879:Alexey Kondrashov
859:genetic variation
819:(promoted during
632:
631:
323:Origin of Species
125:Natural selection
9612:
9605:Sexual selection
9577:
9576:
9565:
9564:
9553:
9552:
9544:
9479:William McGinnis
9448:Richard Lewontin
9443:C. H. Waddington
9315:
9314:
9292:Neutral networks
9042:
9035:
9028:
9019:
9018:
9005:
8995:
8994:
8794:Modern synthesis
8554:Multicellularity
8549:Mosaic evolution
8434:auditory ossicle
8116:Social selection
8099:Flowering plants
8094:Sexual selection
7747:
7740:
7733:
7724:
7723:
7710:
7709:
7670:Animal sexuality
7593:Sexual selection
7332:
7325:
7318:
7309:
7308:
7268:
7251:Williams, George
7246:
7225:
7197:
7175:
7153:
7131:
7129:
7127:
7118:. Archived from
7105:
7084:
7060:
7048:
7037:
6998:
6979:
6949:
6948:
6938:
6928:
6904:
6898:
6885:
6879:
6872:
6866:
6865:
6855:
6823:
6817:
6816:
6780:
6769:
6768:
6750:
6718:
6709:
6708:
6672:
6666:
6665:
6645:
6639:
6638:
6628:
6611:(3–4): 519–531.
6596:
6590:
6589:
6579:
6562:(5): 1422–1429.
6547:
6541:
6540:
6530:
6502:
6496:
6495:
6459:
6453:
6452:
6405:
6399:
6398:
6388:
6356:
6350:
6349:
6315:
6306:
6303:
6288:
6287:
6243:
6237:
6236:
6212:
6201:
6200:
6190:
6158:
6152:
6148:
6142:
6138:
6136:
6128:
6106:
6097:
6096:
6068:
6059:
6058:
6056:
6054:
6037:
6031:
6030:
6028:
6026:
5981:
5975:
5974:
5972:
5970:
5922:
5916:
5915:
5913:
5911:
5894:
5888:
5887:
5877:
5856:Lodé, T (2012).
5853:
5844:
5843:
5841:
5826:Trends Evol Biol
5820:Lodé, T (2012).
5817:
5808:
5807:
5786:Lodé, T (2011).
5783:
5777:
5776:
5766:
5745:Lodé, T (2011).
5742:
5733:
5732:
5696:
5690:
5689:
5687:
5676:
5670:
5669:
5651:
5615:
5609:
5608:
5572:
5566:
5565:
5537:
5531:
5530:
5520:
5480:
5474:
5473:
5463:
5423:
5417:
5416:
5406:
5374:
5368:
5367:
5323:
5317:
5316:
5291:(5): 1654–1660.
5282:
5273:
5267:
5260:
5254:
5249:Ridley M (2004)
5247:
5241:
5240:
5188:
5182:
5181:
5145:
5139:
5138:
5102:
5096:
5095:
5076:10.1038/336435a0
5046:
5037:
5026:
5020:
5007:
5001:
5000:
4990:
4980:
4948:
4942:
4941:
4905:
4896:
4883:
4877:
4876:
4858:
4849:
4848:
4823:
4814:
4813:
4787:
4772:
4771:
4759:
4746:
4740:
4739:
4703:
4697:
4696:
4679:(Pt 4): 519–22.
4668:
4662:
4661:
4617:
4611:
4610:
4576:
4556:
4550:
4549:
4547:
4545:
4534:
4528:
4527:
4517:
4477:
4471:
4470:
4434:
4428:
4427:
4425:
4423:
4412:
4406:
4405:
4357:
4351:
4350:
4306:
4300:
4299:
4289:
4265:
4259:
4258:
4248:
4238:
4221:(9): 3566–3573.
4201:
4195:
4194:
4182:
4176:
4149:
4143:
4133:
4127:
4124:
4118:
4117:
4091:
4080:
4079:
4054:(4): 1088–1098.
4043:
4034:
4033:
4005:
3999:
3996:
3987:
3984:
3975:
3974:
3964:
3954:
3930:
3924:
3923:
3899:
3893:
3890:
3884:
3883:
3873:
3867:
3864:
3840:
3829:
3823:
3820:
3810:
3804:
3803:
3793:
3753:
3747:
3746:
3727:10.1038/35084593
3706:
3700:
3699:
3697:
3695:
3680:
3674:
3673:
3663:
3635:
3626:
3620:
3619:
3609:
3589:
3579:
3573:
3572:
3570:
3568:
3557:
3501:molecular clocks
3405:
3398:
3394:
3391:
3385:
3358:
3357:
3350:
3104:
3099:
3080:
3075:
3059:
3054:
3039:
3034:
3018:
3013:
2998:
2993:
2978:
2973:
2957:
2952:
2943:Earliest animals
2937:
2932:
2917:
2912:
2897:
2892:
2877:
2872:
2857:
2852:
2836:
2831:
2816:
2811:
2795:
2790:
2775:
2770:
2755:
2750:
2741:Earliest fossils
2735:
2730:
2715:
2710:
2695:
2690:
2675:
2670:
2649:
2618:
2580:
2578:
2552:
2550:
2457:
2350:
2345:
2340:
2335:
2330:
2325:
2320:
2315:
2310:
2305:
2300:
2295:
2290:
2285:
2280:
2275:
2270:
2265:
2260:
2255:
2250:
2245:
2240:
2235:
2230:
2225:
2220:
2215:
2210:
2205:
2200:
2195:
2190:
2185:
2180:
2175:
2170:
2158:
2151:
2144:
2138:
2128:
2121:
2093:
2086:
2082:
2079:
2073:
2042:
2034:
1970:
1968:
1967:
1962:
1950:
1948:
1946:
1945:
1940:
1938:
1933:
1741:Muller's ratchet
1641:An alternative "
1573:
1566:
1562:
1559:
1553:
1526:
1525:
1518:
1218:
1211:
1207:
1204:
1198:
1196:
1155:
1131:
1123:
1063:
1056:
1052:
1049:
1043:
1041:
1000:
976:
968:
948:
941:
937:
934:
928:
901:
900:
893:
814:recombinational
665:
624:
617:
610:
597:
592:
591:
584:
580:
579:
556:Level of support
349:Current research
334:Modern synthesis
329:Before synthesis
282:Extinction event
40:Darwin's finches
37:
18:
17:
9620:
9619:
9615:
9614:
9613:
9611:
9610:
9609:
9585:
9584:
9583:
9571:
9559:
9547:
9539:
9537:
9532:
9523:
9502:
9489:Sean B. Carroll
9427:
9359:
9306:
9270:
9222:
9203:Maternal effect
9186:
9119:
9056:
9046:
9016:
9011:
8983:
8910:Group selection
8883:
8808:
8712:
8639:
8601:Tempo and modes
8595:
8450:
8354:
8171:
8130:
8006:
7999:
7976:Species complex
7789:
7780:History of life
7756:
7751:
7721:
7716:
7698:
7685:Differentiation
7675:Human sexuality
7665:Plant sexuality
7651:
7547:Spermatogenesis
7491:
7484:
7347:
7341:
7336:
7298:Archived (2023)
7275:
7265:
7243:
7222:
7202:Szathmáry, Eörs
7194:
7172:
7150:
7125:
7123:
7108:
7102:
7081:
7057:
6995:
6976:
6957:
6955:Further reading
6952:
6905:
6901:
6886:
6882:
6873:
6869:
6824:
6820:
6781:
6772:
6719:
6712:
6673:
6669:
6646:
6642:
6597:
6593:
6548:
6544:
6503:
6499:
6460:
6456:
6406:
6402:
6357:
6353:
6338:
6316:
6309:
6304:
6291:
6244:
6240:
6233:
6213:
6204:
6167:Proc. Biol. Sci
6159:
6155:
6140:
6139:
6130:
6129:
6125:
6107:
6100:
6069:
6062:
6052:
6050:
6038:
6034:
6024:
6022:
5982:
5978:
5968:
5966:
5928:(5 June 2009).
5923:
5919:
5909:
5907:
5895:
5891:
5854:
5847:
5818:
5811:
5798:(2837): 30–31.
5784:
5780:
5743:
5736:
5697:
5693:
5685:
5677:
5673:
5616:
5612:
5573:
5569:
5538:
5534:
5481:
5477:
5424:
5420:
5375:
5371:
5324:
5320:
5280:
5274:
5270:
5261:
5257:
5248:
5244:
5189:
5185:
5146:
5142:
5103:
5099:
5047:
5040:
5027:
5023:
5017:Wayback Machine
5008:
5004:
4963:(15): 8173–80.
4951:Cox MM (2001).
4949:
4945:
4906:
4899:
4884:
4880:
4873:
4859:
4852:
4845:
4824:
4817:
4802:
4788:
4775:
4768:
4747:
4743:
4704:
4700:
4669:
4665:
4618:
4614:
4567:(13): 1176–84.
4557:
4553:
4543:
4541:
4536:
4535:
4531:
4478:
4474:
4451:10.2307/2410266
4435:
4431:
4421:
4419:
4414:
4413:
4409:
4358:
4354:
4307:
4303:
4266:
4262:
4205:Hamilton, W. D.
4202:
4198:
4183:
4179:
4150:
4146:
4134:
4130:
4125:
4121:
4114:
4092:
4083:
4044:
4037:
4022:10.1139/g07-039
4006:
4002:
3997:
3990:
3985:
3978:
3931:
3927:
3920:
3900:
3896:
3891:
3887:
3874:
3870:
3830:
3826:
3811:
3807:
3754:
3750:
3707:
3703:
3693:
3691:
3681:
3677:
3633:
3627:
3623:
3587:
3580:
3576:
3566:
3564:
3558:
3551:
3547:
3513:
3489:
3439:
3433:
3425:
3406:
3395:
3389:
3386:
3380:by introducing
3371:
3359:
3355:
3348:
3336:
3294:
3277:
3239:. For example,
3217:
3184:
3163:Proterozoic Eon
3139:
3138:
3130:
3114:
3113:
3102:
3093:
3092:
3078:
3069:
3068:
3057:
3048:
3047:
3037:
3028:
3027:
3016:
3007:
3006:
2996:
2987:
2986:
2984:Ediacaran biota
2976:
2967:
2966:
2955:
2946:
2945:
2935:
2926:
2925:
2923:Earliest plants
2915:
2906:
2905:
2895:
2886:
2885:
2875:
2866:
2865:
2855:
2846:
2845:
2834:
2825:
2824:
2814:
2805:
2804:
2793:
2784:
2783:
2781:Earliest oxygen
2773:
2764:
2763:
2753:
2744:
2743:
2733:
2724:
2723:
2713:
2704:
2703:
2693:
2684:
2683:
2673:
2666:
2665:
2664:
2659:
2658:
2657:
2652:
2651:
2650:
2646:
2644:
2642:
2640:
2638:
2636:
2634:
2632:
2630:
2628:
2625:
2621:
2620:
2619:
2615:
2613:
2611:
2609:
2607:
2605:
2603:
2601:
2599:
2597:
2595:
2594:
2593:
2592:
2591:
2590:
2589:
2587:
2583:
2582:
2581:
2576:
2574:
2572:
2570:
2568:
2566:
2564:
2563:
2562:
2561:
2559:
2555:
2554:
2553:
2548:
2546:
2544:
2542:
2540:
2538:
2536:
2532:
2531:
2530:
2523:
2522:
2521:
2514:
2513:
2512:
2505:
2504:
2503:
2498:
2497:
2496:
2489:
2488:
2487:
2480:
2479:
2478:
2468:
2467:
2466:
2461:
2460:
2459:
2455:
2453:
2451:
2449:
2447:
2445:
2440:
2439:
2438:
2433:
2432:
2431:
2424:
2423:
2422:
2417:
2416:
2415:
2408:
2407:
2406:
2401:
2400:
2399:
2392:
2391:
2390:
2385:
2384:
2383:
2376:
2375:
2374:
2369:
2368:
2367:
2360:
2359:
2358:
2351:
2348:
2346:
2343:
2341:
2338:
2336:
2333:
2331:
2328:
2326:
2323:
2321:
2318:
2316:
2313:
2311:
2308:
2306:
2303:
2301:
2298:
2296:
2293:
2291:
2288:
2286:
2283:
2281:
2278:
2276:
2273:
2271:
2268:
2266:
2263:
2261:
2258:
2256:
2253:
2251:
2248:
2246:
2243:
2241:
2238:
2236:
2233:
2231:
2228:
2226:
2223:
2221:
2218:
2216:
2213:
2211:
2208:
2206:
2203:
2201:
2198:
2196:
2193:
2191:
2188:
2186:
2183:
2181:
2178:
2176:
2173:
2171:
2168:
2162:
2136:
2130:
2126:
2119:
2094:
2083:
2077:
2074:
2059:
2043:
2030:
1977:
1956:
1953:
1952:
1932:
1930:
1927:
1926:
1925:
1904:
1895:
1890:
1806:
1743:
1737:
1714:
1678:complementation
1635:
1623:dinoflagellates
1592:
1583:Meiotic drivers
1574:
1563:
1557:
1554:
1548:by introducing
1539:
1527:
1523:
1516:
1460:
1449:
1440:
1386:parthenogenetic
1362:immunoglobulins
1281:
1219:
1208:
1202:
1199:
1156:
1154:
1144:
1132:
1121:
1119:Novel genotypes
1112:
1094:complementation
1074:
1064:
1053:
1047:
1044:
1001:
999:
989:
977:
966:
949:
938:
932:
929:
923:by introducing
914:
902:
898:
891:
855:August Weismann
843:
833:(also known as
830:complementation
810:fault-tolerance
781:hermaphroditism
730:parthenogenesis
707:common ancestor
680:
679:
674:
667:
628:
587:
574:
573:
566:
565:
516:
508:
507:
378:
370:
369:
368:
296:
288:
287:
286:
235:Human evolution
225:History of life
209:
208:Natural history
201:
200:
199:
99:
91:
46:
12:
11:
5:
9618:
9608:
9607:
9602:
9597:
9582:
9581:
9569:
9557:
9534:
9533:
9528:
9525:
9524:
9522:
9521:
9516:
9510:
9508:
9504:
9503:
9501:
9500:
9499:
9498:
9486:
9481:
9476:
9471:
9466:
9465:
9464:
9453:François Jacob
9450:
9445:
9439:
9437:
9433:
9432:
9429:
9428:
9426:
9425:
9420:
9415:
9410:
9405:
9400:
9395:
9390:
9389:
9388:
9378:
9373:
9367:
9365:
9361:
9360:
9358:
9357:
9352:
9347:
9342:
9337:
9332:
9327:
9321:
9319:
9312:
9308:
9307:
9305:
9304:
9299:
9294:
9289:
9284:
9278:
9276:
9272:
9271:
9269:
9268:
9263:
9258:
9253:
9252:
9251:
9246:
9236:
9230:
9228:
9224:
9223:
9221:
9220:
9215:
9210:
9205:
9200:
9194:
9192:
9188:
9187:
9185:
9184:
9182:Sequence space
9179:
9174:
9169:
9164:
9159:
9150:
9145:
9140:
9135:
9129:
9127:
9121:
9120:
9118:
9117:
9112:
9111:
9110:
9100:
9095:
9090:
9085:
9080:
9075:
9070:
9064:
9062:
9058:
9057:
9045:
9044:
9037:
9030:
9022:
9013:
9012:
9010:
9009:
8999:
8988:
8985:
8984:
8982:
8981:
8976:
8971:
8966:
8961:
8960:
8959:
8949:
8944:
8939:
8934:
8929:
8928:
8927:
8922:
8917:
8907:
8902:
8897:
8891:
8889:
8885:
8884:
8882:
8881:
8876:
8875:
8874:
8869:
8864:
8863:
8862:
8852:
8847:
8842:
8837:
8832:
8822:
8816:
8814:
8810:
8809:
8807:
8806:
8801:
8796:
8791:
8786:
8781:
8776:
8771:
8766:
8761:
8760:
8759:
8750:Charles Darwin
8747:
8746:
8745:
8733:
8728:
8722:
8720:
8714:
8713:
8711:
8710:
8705:
8700:
8695:
8690:
8688:Non-ecological
8685:
8680:
8675:
8670:
8665:
8660:
8655:
8649:
8647:
8641:
8640:
8638:
8637:
8628:
8619:
8605:
8603:
8597:
8596:
8594:
8593:
8588:
8587:
8586:
8581:
8576:
8571:
8566:
8556:
8551:
8546:
8541:
8536:
8531:
8526:
8521:
8516:
8511:
8506:
8505:
8504:
8494:
8489:
8484:
8479:
8478:
8477:
8472:
8461:
8459:
8452:
8451:
8449:
8448:
8447:
8446:
8441:
8439:nervous system
8436:
8431:
8426:
8418:
8417:
8416:
8411:
8406:
8401:
8396:
8391:
8381:
8376:
8371:
8365:
8363:
8356:
8355:
8353:
8352:
8347:
8342:
8337:
8332:
8331:
8330:
8320:
8319:
8318:
8313:
8312:
8311:
8306:
8296:
8291:
8286:
8281:
8276:
8275:
8274:
8269:
8259:
8249:
8244:
8243:
8242:
8232:
8227:
8222:
8217:
8216:
8215:
8205:
8200:
8199:
8198:
8188:
8182:
8180:
8173:
8172:
8170:
8169:
8164:
8159:
8154:
8149:
8144:
8138:
8136:
8132:
8131:
8129:
8128:
8123:
8118:
8113:
8112:
8111:
8106:
8101:
8091:
8086:
8081:
8076:
8071:
8070:
8069:
8064:
8054:
8049:
8044:
8043:
8042:
8032:
8027:
8022:
8017:
8011:
8009:
8001:
8000:
7998:
7997:
7996:
7995:
7985:
7980:
7979:
7978:
7973:
7963:
7962:
7961:
7951:
7946:
7941:
7939:Origin of life
7936:
7931:
7926:
7924:Microevolution
7921:
7919:Macroevolution
7916:
7911:
7906:
7905:
7904:
7894:
7889:
7884:
7879:
7874:
7869:
7864:
7859:
7857:Common descent
7854:
7853:
7852:
7842:
7837:
7835:Baldwin effect
7832:
7831:
7830:
7825:
7815:
7810:
7805:
7799:
7797:
7791:
7790:
7788:
7787:
7782:
7777:
7772:
7767:
7761:
7758:
7757:
7750:
7749:
7742:
7735:
7727:
7718:
7717:
7715:
7714:
7703:
7700:
7699:
7697:
7696:
7695:
7694:
7693:
7692:
7687:
7682:
7667:
7661:
7659:
7653:
7652:
7650:
7649:
7648:
7647:
7642:
7637:
7632:
7627:
7622:
7621:
7620:
7605:
7600:
7595:
7590:
7589:
7588:
7583:
7573:
7572:
7571:
7566:
7556:
7555:
7554:
7549:
7539:
7534:
7529:
7524:
7519:
7514:
7513:
7512:
7507:
7496:
7494:
7486:
7485:
7483:
7482:
7477:
7472:
7467:
7466:
7465:
7460:
7450:
7445:
7444:
7443:
7438:
7431:Sex chromosome
7428:
7423:
7422:
7421:
7416:
7411:
7406:
7401:
7396:
7386:
7385:
7384:
7379:
7369:
7368:
7367:
7362:
7351:
7349:
7343:
7342:
7335:
7334:
7327:
7320:
7312:
7306:
7305:
7302:Nature journal
7291:
7286:
7281:
7274:
7273:External links
7271:
7270:
7269:
7263:
7247:
7241:
7226:
7220:
7198:
7192:
7176:
7170:
7154:
7148:
7132:
7106:
7100:
7085:
7079:
7061:
7055:
7038:
6999:
6993:
6980:
6974:
6956:
6953:
6951:
6950:
6899:
6880:
6867:
6818:
6770:
6733:(3): 251–256.
6710:
6683:(4): 387–396.
6667:
6656:(3): 224–231.
6640:
6591:
6542:
6513:(4): 398–405.
6497:
6470:(3): 323–351.
6454:
6411:, Gardiner W,
6400:
6351:
6336:
6307:
6289:
6254:(6): 779–783.
6238:
6231:
6202:
6153:
6141:|journal=
6124:978-9985908150
6123:
6098:
6079:(7): 498–505.
6060:
6032:
5976:
5917:
5889:
5868:(6): 325–328.
5845:
5809:
5778:
5757:(6): 419–422.
5734:
5691:
5671:
5610:
5567:
5548:(4): 485–514.
5532:
5495:(3): evae050.
5475:
5418:
5389:(4): 1667–86.
5369:
5318:
5268:
5255:
5242:
5183:
5140:
5097:
5038:
5021:
5002:
4943:
4910:J. Theor. Biol
4897:
4878:
4871:
4850:
4843:
4815:
4801:978-0199255634
4800:
4773:
4766:
4741:
4714:(5): 560–584.
4698:
4663:
4612:
4551:
4529:
4472:
4445:(3): 418–426.
4429:
4407:
4386:10.1086/599080
4378:10.1086/599080
4372:(s1): S43–53.
4352:
4317:(4): 441–456.
4301:
4280:(1): 123–130.
4260:
4196:
4177:
4157:978-0306472619
4144:
4128:
4119:
4112:
4081:
4035:
4016:(5): 517–524.
4000:
3988:
3976:
3925:
3918:
3894:
3885:
3868:
3866:
3865:
3853:
3824:
3805:
3768:(3): a016154.
3748:
3721:(8): 634–639.
3701:
3675:
3621:
3593:Bioinformatics
3574:
3548:
3546:
3543:
3542:
3541:
3538:
3531:
3520:
3512:
3509:
3488:
3485:
3435:Main article:
3432:
3429:
3424:
3421:
3408:
3407:
3376:. Please help
3362:
3360:
3353:
3347:
3344:
3335:
3332:
3293:
3290:
3276:
3273:
3216:
3213:
3183:
3180:
3176:parthenocarpic
3141:
3140:
3131:
3124:
3120:
3119:
3116:
3115:
3108:
3107:
3105:
3095:
3094:
3084:
3083:
3081:
3071:
3070:
3063:
3062:
3060:
3050:
3049:
3043:
3042:
3040:
3030:
3029:
3022:
3021:
3019:
3009:
3008:
3002:
3001:
2999:
2989:
2988:
2982:
2981:
2979:
2969:
2968:
2961:
2960:
2958:
2948:
2947:
2941:
2940:
2938:
2928:
2927:
2921:
2920:
2918:
2908:
2907:
2903:Earliest fungi
2901:
2900:
2898:
2888:
2887:
2881:
2880:
2878:
2868:
2867:
2861:
2860:
2858:
2848:
2847:
2840:
2839:
2837:
2827:
2826:
2820:
2819:
2817:
2807:
2806:
2799:
2798:
2796:
2786:
2785:
2779:
2778:
2776:
2766:
2765:
2761:LHB meteorites
2759:
2758:
2756:
2746:
2745:
2739:
2738:
2736:
2726:
2725:
2719:
2718:
2716:
2706:
2705:
2701:Earliest water
2699:
2698:
2696:
2686:
2685:
2679:
2678:
2676:
2667:
2662:
2661:
2660:
2655:
2654:
2653:
2624:
2623:
2622:
2586:
2585:
2584:
2558:
2557:
2556:
2535:
2534:
2533:
2526:
2525:
2524:
2517:
2516:
2515:
2508:
2507:
2506:
2501:
2500:
2499:
2492:
2491:
2490:
2483:
2482:
2481:
2471:
2470:
2469:
2464:
2463:
2462:
2443:
2442:
2441:
2436:
2435:
2434:
2427:
2426:
2425:
2420:
2419:
2418:
2411:
2410:
2409:
2404:
2403:
2402:
2397:Photosynthesis
2395:
2394:
2393:
2388:
2387:
2386:
2379:
2378:
2377:
2372:
2371:
2370:
2363:
2362:
2361:
2356:
2355:
2354:
2352:
2349:0 —
2347:
2342:
2337:
2332:
2327:
2322:
2317:
2312:
2307:
2302:
2297:
2292:
2287:
2282:
2277:
2272:
2267:
2262:
2257:
2252:
2247:
2242:
2237:
2232:
2227:
2222:
2217:
2212:
2207:
2202:
2197:
2192:
2187:
2182:
2177:
2172:
2167:
2164:
2163:
2161:
2160:
2153:
2146:
2135:
2132:
2131:
2124:
2118:
2115:
2096:
2095:
2046:
2044:
2037:
2028:
1976:
1973:
1960:
1936:
1903:
1900:
1894:
1891:
1889:
1886:
1805:
1802:
1739:Main article:
1736:
1733:
1713:
1710:
1698:Charles Darwin
1634:
1631:
1591:
1588:
1576:
1575:
1544:. Please help
1530:
1528:
1521:
1515:
1512:
1458:
1448:
1445:
1439:
1436:
1280:
1277:
1221:
1220:
1135:
1133:
1126:
1120:
1117:
1111:
1108:
1066:
1065:
980:
978:
971:
965:
962:
951:
950:
919:. Please help
905:
903:
896:
890:
887:
875:W. D. Hamilton
863:Charles Darwin
851:Erasmus Darwin
842:
839:
754:transformation
675:
668:
662:
630:
629:
627:
626:
619:
612:
604:
601:
600:
599:
598:
585:
568:
567:
564:
563:
558:
553:
548:
543:
538:
536:Social effects
533:
528:
523:
517:
514:
513:
510:
509:
506:
505:
500:
495:
490:
485:
480:
475:
470:
465:
460:
455:
450:
445:
440:
435:
430:
425:
420:
415:
410:
405:
400:
395:
390:
385:
379:
376:
375:
372:
371:
367:
366:
356:
351:
346:
341:
336:
331:
326:
319:
314:
309:
304:
298:
297:
294:
293:
290:
289:
285:
284:
279:
274:
269:
264:
262:Classification
259:
254:
249:
244:
243:
242:
232:
227:
222:
220:Common descent
217:
215:Origin of life
211:
210:
207:
206:
203:
202:
198:
197:
192:
187:
182:
177:
172:
167:
162:
157:
152:
147:
142:
137:
132:
127:
122:
117:
112:
107:
101:
100:
97:
96:
93:
92:
90:
89:
84:
79:
73:
72:
67:
62:
57:
51:
48:
47:
38:
30:
29:
23:
22:
9:
6:
4:
3:
2:
9617:
9606:
9603:
9601:
9598:
9596:
9593:
9592:
9590:
9580:
9575:
9570:
9568:
9563:
9558:
9556:
9551:
9546:
9545:
9542:
9531:
9526:
9520:
9517:
9515:
9512:
9511:
9509:
9505:
9497:
9496:
9492:
9491:
9490:
9487:
9485:
9482:
9480:
9477:
9475:
9472:
9470:
9467:
9463:
9460:
9459:
9458:
9457:Jacques Monod
9454:
9451:
9449:
9446:
9444:
9441:
9440:
9438:
9434:
9424:
9421:
9419:
9416:
9414:
9411:
9409:
9406:
9404:
9401:
9399:
9396:
9394:
9391:
9387:
9384:
9383:
9382:
9379:
9377:
9374:
9372:
9371:Homeotic gene
9369:
9368:
9366:
9362:
9356:
9353:
9351:
9348:
9346:
9343:
9341:
9338:
9336:
9333:
9331:
9328:
9326:
9323:
9322:
9320:
9316:
9313:
9309:
9303:
9300:
9298:
9295:
9293:
9290:
9288:
9285:
9283:
9280:
9279:
9277:
9273:
9267:
9264:
9262:
9259:
9257:
9254:
9250:
9247:
9245:
9242:
9241:
9240:
9239:Morphogenesis
9237:
9235:
9232:
9231:
9229:
9225:
9219:
9216:
9214:
9211:
9209:
9206:
9204:
9201:
9199:
9196:
9195:
9193:
9189:
9183:
9180:
9178:
9175:
9173:
9170:
9168:
9165:
9163:
9160:
9158:
9154:
9151:
9149:
9146:
9144:
9141:
9139:
9136:
9134:
9131:
9130:
9128:
9126:
9122:
9116:
9113:
9109:
9106:
9105:
9104:
9101:
9099:
9096:
9094:
9091:
9089:
9086:
9084:
9081:
9079:
9076:
9074:
9073:Reaction norm
9071:
9069:
9066:
9065:
9063:
9059:
9055:
9051:
9043:
9038:
9036:
9031:
9029:
9024:
9023:
9020:
9008:
9004:
9000:
8998:
8990:
8989:
8986:
8980:
8977:
8975:
8972:
8970:
8967:
8965:
8962:
8958:
8955:
8954:
8953:
8952:Phylogenetics
8950:
8948:
8945:
8943:
8940:
8938:
8935:
8933:
8930:
8926:
8923:
8921:
8918:
8916:
8913:
8912:
8911:
8908:
8906:
8903:
8901:
8898:
8896:
8893:
8892:
8890:
8886:
8880:
8877:
8873:
8870:
8868:
8865:
8861:
8858:
8857:
8856:
8855:Structuralism
8853:
8851:
8848:
8846:
8843:
8841:
8838:
8836:
8833:
8831:
8830:Catastrophism
8828:
8827:
8826:
8823:
8821:
8818:
8817:
8815:
8811:
8805:
8802:
8800:
8797:
8795:
8792:
8790:
8789:Neo-Darwinism
8787:
8785:
8782:
8780:
8777:
8775:
8772:
8770:
8767:
8765:
8762:
8758:
8757:
8753:
8752:
8751:
8748:
8744:
8743:
8739:
8738:
8737:
8734:
8732:
8729:
8727:
8724:
8723:
8721:
8719:
8715:
8709:
8706:
8704:
8703:Reinforcement
8701:
8699:
8696:
8694:
8691:
8689:
8686:
8684:
8681:
8679:
8676:
8674:
8671:
8669:
8666:
8664:
8661:
8659:
8656:
8654:
8651:
8650:
8648:
8646:
8642:
8636:
8635:Catastrophism
8632:
8629:
8627:
8626:Macromutation
8623:
8622:Micromutation
8620:
8618:
8614:
8610:
8607:
8606:
8604:
8602:
8598:
8592:
8589:
8585:
8582:
8580:
8577:
8575:
8572:
8570:
8567:
8565:
8562:
8561:
8560:
8557:
8555:
8552:
8550:
8547:
8545:
8542:
8540:
8537:
8535:
8532:
8530:
8529:Immune system
8527:
8525:
8522:
8520:
8517:
8515:
8512:
8510:
8507:
8503:
8500:
8499:
8498:
8495:
8493:
8490:
8488:
8485:
8483:
8480:
8476:
8473:
8471:
8468:
8467:
8466:
8463:
8462:
8460:
8458:
8453:
8445:
8442:
8440:
8437:
8435:
8432:
8430:
8427:
8425:
8422:
8421:
8419:
8415:
8412:
8410:
8407:
8405:
8402:
8400:
8397:
8395:
8392:
8390:
8389:symbiogenesis
8387:
8386:
8385:
8382:
8380:
8377:
8375:
8372:
8370:
8367:
8366:
8364:
8362:
8357:
8351:
8348:
8346:
8343:
8341:
8338:
8336:
8333:
8329:
8326:
8325:
8324:
8321:
8317:
8314:
8310:
8307:
8305:
8302:
8301:
8300:
8297:
8295:
8292:
8290:
8287:
8285:
8282:
8280:
8277:
8273:
8270:
8268:
8265:
8264:
8263:
8260:
8258:
8255:
8254:
8253:
8250:
8248:
8245:
8241:
8238:
8237:
8236:
8233:
8231:
8228:
8226:
8223:
8221:
8218:
8214:
8211:
8210:
8209:
8206:
8204:
8201:
8197:
8194:
8193:
8192:
8189:
8187:
8184:
8183:
8181:
8179:
8174:
8168:
8165:
8163:
8160:
8158:
8155:
8153:
8150:
8148:
8145:
8143:
8140:
8139:
8137:
8133:
8127:
8124:
8122:
8119:
8117:
8114:
8110:
8107:
8105:
8102:
8100:
8097:
8096:
8095:
8092:
8090:
8087:
8085:
8082:
8080:
8077:
8075:
8072:
8068:
8065:
8063:
8060:
8059:
8058:
8057:Kin selection
8055:
8053:
8052:Genetic drift
8050:
8048:
8045:
8041:
8038:
8037:
8036:
8033:
8031:
8028:
8026:
8023:
8021:
8018:
8016:
8013:
8012:
8010:
8008:
8002:
7994:
7991:
7990:
7989:
7986:
7984:
7981:
7977:
7974:
7972:
7969:
7968:
7967:
7964:
7960:
7957:
7956:
7955:
7952:
7950:
7947:
7945:
7942:
7940:
7937:
7935:
7932:
7930:
7927:
7925:
7922:
7920:
7917:
7915:
7912:
7910:
7907:
7903:
7900:
7899:
7898:
7895:
7893:
7890:
7888:
7885:
7883:
7880:
7878:
7875:
7873:
7870:
7868:
7865:
7863:
7860:
7858:
7855:
7851:
7848:
7847:
7846:
7843:
7841:
7838:
7836:
7833:
7829:
7826:
7824:
7821:
7820:
7819:
7816:
7814:
7811:
7809:
7806:
7804:
7801:
7800:
7798:
7796:
7792:
7786:
7783:
7781:
7778:
7776:
7773:
7771:
7768:
7766:
7763:
7762:
7759:
7755:
7748:
7743:
7741:
7736:
7734:
7729:
7728:
7725:
7713:
7705:
7704:
7701:
7691:
7688:
7686:
7683:
7681:
7678:
7677:
7676:
7673:
7672:
7671:
7668:
7666:
7663:
7662:
7660:
7658:
7654:
7646:
7645:Pelvic thrust
7643:
7641:
7638:
7636:
7633:
7631:
7628:
7626:
7623:
7619:
7616:
7615:
7614:
7611:
7610:
7609:
7606:
7604:
7601:
7599:
7596:
7594:
7591:
7587:
7584:
7582:
7579:
7578:
7577:
7576:Fertilization
7574:
7570:
7567:
7565:
7562:
7561:
7560:
7557:
7553:
7550:
7548:
7545:
7544:
7543:
7542:Gametogenesis
7540:
7538:
7535:
7533:
7530:
7528:
7525:
7523:
7520:
7518:
7515:
7511:
7508:
7506:
7503:
7502:
7501:
7498:
7497:
7495:
7493:
7487:
7481:
7478:
7476:
7475:parasexuality
7473:
7471:
7468:
7464:
7461:
7459:
7456:
7455:
7454:
7453:Hermaphrodite
7451:
7449:
7446:
7442:
7439:
7437:
7434:
7433:
7432:
7429:
7427:
7424:
7420:
7419:Haplodiploidy
7417:
7415:
7412:
7410:
7407:
7405:
7402:
7400:
7397:
7395:
7392:
7391:
7390:
7387:
7383:
7380:
7378:
7375:
7374:
7373:
7370:
7366:
7363:
7361:
7358:
7357:
7356:
7353:
7352:
7350:
7344:
7340:
7333:
7328:
7326:
7321:
7319:
7314:
7313:
7310:
7303:
7299:
7295:
7292:
7290:
7287:
7285:
7282:
7280:
7277:
7276:
7266:
7260:
7256:
7252:
7248:
7244:
7238:
7234:
7233:
7227:
7223:
7217:
7213:
7212:
7207:
7203:
7199:
7195:
7189:
7185:
7181:
7177:
7173:
7167:
7163:
7159:
7155:
7151:
7145:
7141:
7137:
7133:
7121:
7117:
7116:
7111:
7107:
7103:
7097:
7093:
7092:
7086:
7082:
7076:
7072:
7071:
7066:
7062:
7058:
7052:
7047:
7046:
7039:
7035:
7031:
7027:
7023:
7019:
7015:
7011:
7007:
7006:
7000:
6996:
6990:
6986:
6981:
6977:
6971:
6967:
6963:
6959:
6958:
6946:
6942:
6937:
6932:
6927:
6922:
6918:
6914:
6910:
6903:
6896:
6893:
6890:
6884:
6877:
6871:
6863:
6859:
6854:
6849:
6845:
6841:
6837:
6833:
6829:
6822:
6814:
6810:
6806:
6802:
6798:
6794:
6790:
6786:
6779:
6777:
6775:
6766:
6762:
6758:
6754:
6749:
6744:
6740:
6736:
6732:
6728:
6724:
6717:
6715:
6706:
6702:
6698:
6694:
6690:
6686:
6682:
6678:
6671:
6663:
6659:
6655:
6651:
6644:
6636:
6632:
6627:
6622:
6618:
6614:
6610:
6606:
6602:
6595:
6587:
6583:
6578:
6573:
6569:
6565:
6561:
6557:
6553:
6546:
6538:
6534:
6529:
6524:
6520:
6516:
6512:
6508:
6501:
6493:
6489:
6485:
6481:
6477:
6473:
6469:
6465:
6458:
6450:
6446:
6442:
6438:
6434:
6430:
6426:
6422:
6418:
6414:
6410:
6404:
6396:
6392:
6387:
6382:
6378:
6374:
6370:
6366:
6362:
6355:
6347:
6343:
6339:
6333:
6329:
6325:
6321:
6314:
6312:
6302:
6300:
6298:
6296:
6294:
6285:
6281:
6277:
6273:
6269:
6265:
6261:
6257:
6253:
6249:
6242:
6234:
6228:
6224:
6220:
6219:
6211:
6209:
6207:
6198:
6194:
6189:
6184:
6180:
6176:
6172:
6168:
6164:
6157:
6151:
6146:
6134:
6126:
6120:
6116:
6112:
6105:
6103:
6094:
6090:
6086:
6082:
6078:
6074:
6067:
6065:
6049:
6048:
6043:
6036:
6021:
6017:
6013:
6009:
6005:
6001:
5997:
5993:
5992:
5987:
5980:
5965:
5961:
5957:
5953:
5949:
5945:
5941:
5937:
5936:
5931:
5927:
5921:
5906:
5905:
5900:
5893:
5885:
5881:
5876:
5871:
5867:
5863:
5859:
5852:
5850:
5840:
5835:
5831:
5827:
5823:
5816:
5814:
5805:
5801:
5797:
5793:
5792:New Scientist
5789:
5782:
5774:
5770:
5765:
5760:
5756:
5752:
5748:
5741:
5739:
5730:
5726:
5722:
5718:
5714:
5710:
5706:
5702:
5695:
5684:
5683:
5675:
5667:
5663:
5659:
5655:
5650:
5645:
5641:
5637:
5633:
5629:
5625:
5621:
5614:
5606:
5602:
5598:
5594:
5590:
5586:
5583:(1): 88–100.
5582:
5578:
5571:
5563:
5559:
5555:
5551:
5547:
5543:
5536:
5528:
5524:
5519:
5514:
5510:
5506:
5502:
5498:
5494:
5490:
5486:
5479:
5471:
5467:
5462:
5457:
5453:
5449:
5445:
5441:
5437:
5433:
5429:
5422:
5414:
5410:
5405:
5400:
5396:
5392:
5388:
5384:
5380:
5373:
5365:
5361:
5357:
5353:
5349:
5348:10.1038/37108
5345:
5341:
5337:
5333:
5329:
5322:
5314:
5310:
5306:
5302:
5298:
5294:
5290:
5286:
5279:
5272:
5265:
5259:
5252:
5246:
5238:
5234:
5230:
5226:
5222:
5218:
5214:
5210:
5206:
5202:
5198:
5194:
5187:
5179:
5175:
5171:
5167:
5163:
5159:
5155:
5151:
5144:
5136:
5132:
5128:
5124:
5120:
5116:
5112:
5108:
5101:
5093:
5089:
5085:
5081:
5077:
5073:
5069:
5065:
5061:
5057:
5056:
5051:
5045:
5043:
5035:
5031:
5025:
5018:
5014:
5011:
5006:
4998:
4994:
4989:
4984:
4979:
4974:
4970:
4966:
4962:
4958:
4954:
4947:
4939:
4935:
4931:
4927:
4923:
4919:
4916:(3): 323–51.
4915:
4911:
4904:
4902:
4894:
4891:
4888:
4882:
4874:
4872:9781405103459
4868:
4864:
4857:
4855:
4846:
4844:9783034862738
4840:
4836:
4832:
4828:
4822:
4820:
4811:
4807:
4803:
4797:
4793:
4786:
4784:
4782:
4780:
4778:
4769:
4767:9780521293020
4763:
4758:
4757:
4751:
4745:
4737:
4733:
4729:
4725:
4721:
4717:
4713:
4709:
4702:
4694:
4690:
4686:
4682:
4678:
4674:
4667:
4659:
4655:
4651:
4647:
4643:
4639:
4635:
4631:
4627:
4623:
4616:
4608:
4604:
4600:
4596:
4592:
4588:
4584:
4580:
4575:
4574:q-bio/0508003
4570:
4566:
4562:
4555:
4539:
4533:
4525:
4521:
4516:
4511:
4507:
4503:
4499:
4495:
4491:
4487:
4483:
4476:
4468:
4464:
4460:
4456:
4452:
4448:
4444:
4440:
4433:
4417:
4411:
4403:
4399:
4395:
4391:
4387:
4383:
4379:
4375:
4371:
4367:
4363:
4356:
4348:
4344:
4340:
4336:
4332:
4328:
4324:
4320:
4316:
4312:
4305:
4297:
4293:
4288:
4283:
4279:
4275:
4271:
4264:
4256:
4252:
4247:
4242:
4237:
4232:
4228:
4224:
4220:
4216:
4215:
4210:
4206:
4200:
4192:
4188:
4181:
4174:
4170:
4166:
4162:
4158:
4154:
4148:
4142:1995 Penguin.
4141:
4137:
4132:
4123:
4115:
4109:
4105:
4101:
4097:
4090:
4088:
4086:
4077:
4073:
4069:
4065:
4061:
4057:
4053:
4049:
4042:
4040:
4031:
4027:
4023:
4019:
4015:
4011:
4004:
3995:
3993:
3983:
3981:
3972:
3968:
3963:
3958:
3953:
3948:
3944:
3940:
3936:
3929:
3921:
3915:
3911:
3907:
3906:
3898:
3889:
3881:
3880:
3872:
3862:
3861:
3854:
3851:
3847:
3843:
3842:
3838:
3837:
3828:
3818:
3817:
3809:
3801:
3797:
3792:
3787:
3783:
3779:
3775:
3771:
3767:
3763:
3759:
3752:
3744:
3740:
3736:
3732:
3728:
3724:
3720:
3716:
3712:
3705:
3690:
3686:
3679:
3671:
3667:
3662:
3657:
3653:
3649:
3645:
3641:
3640:
3632:
3625:
3617:
3613:
3608:
3603:
3599:
3595:
3594:
3586:
3578:
3563:
3556:
3554:
3549:
3539:
3536:
3535:mating system
3532:
3529:
3525:
3521:
3518:
3517:
3516:
3508:
3506:
3502:
3498:
3494:
3484:
3480:
3478:
3474:
3470:
3468:
3464:
3458:
3456:
3455:mitochondrion
3452:
3448:
3444:
3438:
3428:
3420:
3417:
3415:
3404:
3401:
3393:
3383:
3379:
3375:
3369:
3368:
3367:single source
3363:This section
3361:
3352:
3351:
3343:
3341:
3331:
3329:
3325:
3324:jumping genes
3321:
3316:
3314:
3310:
3306:
3302:
3299:
3289:
3285:
3282:
3272:
3268:
3266:
3261:
3257:
3252:
3250:
3246:
3242:
3238:
3234:
3230:
3226:
3222:
3212:
3209:
3205:
3200:
3198:
3194:
3190:
3179:
3177:
3173:
3168:
3164:
3160:
3156:
3152:
3148:
3137:
3136:
3128:
3121:
3111:
3106:
3101:
3100:
3091:
3087:
3086:Earliest apes
3082:
3077:
3076:
3066:
3065:Karoo ice age
3061:
3056:
3055:
3046:
3041:
3036:
3035:
3025:
3020:
3015:
3014:
3005:
3000:
2995:
2994:
2985:
2980:
2975:
2974:
2964:
2959:
2954:
2953:
2944:
2939:
2934:
2933:
2924:
2919:
2914:
2913:
2904:
2899:
2894:
2893:
2884:
2879:
2874:
2873:
2864:
2859:
2854:
2853:
2843:
2838:
2833:
2832:
2823:
2818:
2813:
2812:
2802:
2797:
2792:
2791:
2782:
2777:
2772:
2771:
2762:
2757:
2752:
2751:
2742:
2737:
2732:
2731:
2722:
2717:
2712:
2711:
2702:
2697:
2692:
2691:
2682:
2677:
2672:
2671:
2648:
2617:
2579:
2551:
2529:
2520:
2511:
2495:
2486:
2477:
2474:
2458:
2430:
2414:
2398:
2382:
2366:
2165:
2159:
2154:
2152:
2147:
2145:
2140:
2139:
2133:
2129:
2127:Life timeline
2122:
2114:
2112:
2107:
2103:
2102:Curtis Lively
2092:
2089:
2081:
2071:
2067:
2063:
2057:
2056:
2052:
2047:This section
2045:
2041:
2036:
2035:
2032:
2031:
2025:
2023:
2019:
2015:
2014:recombination
2010:
2006:
2002:
1998:
1993:
1990:
1986:
1982:
1972:
1958:
1934:
1921:
1919:
1918:
1917:Chlamydomonas
1912:
1911:restrictive.
1909:
1908:recombination
1899:
1885:
1883:
1879:
1875:
1871:
1867:
1863:
1857:
1854:
1849:
1846:
1844:
1841:, think of a
1840:
1835:
1833:
1832:
1827:
1823:
1815:
1810:
1801:
1798:
1797:D. discoideum
1794:
1793:kin selection
1789:
1788:
1783:
1779:
1773:
1771:
1767:
1763:
1759:
1756:
1752:
1748:
1742:
1732:
1730:
1725:
1723:
1718:
1709:
1705:
1701:
1699:
1695:
1691:
1687:
1683:
1679:
1675:
1670:
1668:
1663:
1659:
1654:
1652:
1648:
1644:
1643:informational
1639:
1630:
1628:
1624:
1620:
1616:
1612:
1607:
1605:
1601:
1597:
1587:
1584:
1572:
1569:
1561:
1551:
1547:
1543:
1537:
1536:
1535:single source
1531:This section
1529:
1520:
1519:
1511:
1509:
1505:
1500:
1497:
1493:
1489:
1488:two-fold cost
1481:
1477:
1473:
1468:
1464:
1461:
1452:
1444:
1435:
1431:
1429:
1428:
1423:
1419:
1418:S. marcescens
1415:
1411:
1410:mating system
1406:
1405:
1400:
1399:
1393:
1391:
1387:
1382:
1380:
1379:
1373:
1371:
1367:
1366:immune system
1363:
1359:
1356:of genes for
1355:
1350:
1348:
1347:Lewis Carroll
1343:
1339:
1337:
1333:
1329:
1325:
1321:
1317:
1313:
1309:
1305:
1301:
1296:
1292:
1290:
1286:
1276:
1274:
1271:of the genus
1270:
1267:
1261:
1259:
1258:Ronald Fisher
1255:
1253:
1252:recombination
1249:
1240:
1236:
1232:
1227:
1217:
1214:
1206:
1195:
1192:
1188:
1185:
1181:
1178:
1174:
1171:
1167:
1164: –
1163:
1159:
1158:Find sources:
1152:
1148:
1142:
1141:
1136:This section
1134:
1130:
1125:
1124:
1116:
1107:
1104:
1102:
1097:
1095:
1091:
1087:
1082:
1079:
1073:
1062:
1059:
1051:
1040:
1037:
1033:
1030:
1026:
1023:
1019:
1016:
1012:
1009: –
1008:
1004:
1003:Find sources:
997:
993:
987:
986:
981:This section
979:
975:
970:
969:
961:
957:
947:
944:
936:
926:
922:
918:
912:
911:
910:single source
906:This section
904:
895:
894:
886:
884:
880:
876:
872:
868:
864:
860:
856:
852:
848:
838:
836:
832:
831:
826:
822:
818:
817:
811:
807:
803:
802:recombination
798:
793:
791:
786:
782:
778:
774:
770:
765:
763:
759:
755:
751:
747:
743:
739:
735:
731:
727:
723:
719:
715:
712:
708:
704:
700:
696:
692:
688:
684:
678:
672:
657:
653:
649:
642:
638:
634:
625:
620:
618:
613:
611:
606:
605:
603:
602:
596:
586:
583:
578:
572:
571:
570:
569:
562:
559:
557:
554:
552:
549:
547:
544:
542:
539:
537:
534:
532:
529:
527:
524:
522:
519:
518:
512:
511:
504:
501:
499:
496:
494:
491:
489:
486:
484:
481:
479:
476:
474:
471:
469:
468:Phylogenetics
466:
464:
461:
459:
456:
454:
451:
449:
446:
444:
441:
439:
436:
434:
431:
429:
426:
424:
421:
419:
416:
414:
411:
409:
406:
404:
401:
399:
396:
394:
391:
389:
386:
384:
381:
380:
374:
373:
364:
360:
357:
355:
352:
350:
347:
345:
342:
340:
337:
335:
332:
330:
327:
325:
324:
320:
318:
315:
313:
312:Before Darwin
310:
308:
305:
303:
300:
299:
292:
291:
283:
280:
278:
275:
273:
270:
268:
265:
263:
260:
258:
255:
253:
250:
248:
245:
241:
238:
237:
236:
233:
231:
228:
226:
223:
221:
218:
216:
213:
212:
205:
204:
196:
193:
191:
188:
186:
183:
181:
178:
176:
173:
171:
168:
166:
163:
161:
158:
156:
153:
151:
148:
146:
143:
141:
140:Genetic drift
138:
136:
133:
131:
128:
126:
123:
121:
118:
116:
113:
111:
108:
106:
103:
102:
95:
94:
88:
85:
83:
80:
78:
75:
74:
71:
68:
66:
63:
61:
58:
56:
53:
52:
50:
49:
45:
41:
36:
32:
31:
28:
25:
24:
20:
19:
16:
9493:
9386:eyeless gene
9296:
9282:Evolvability
9256:Segmentation
9133:Canalisation
9103:Heterochrony
9093:Heritability
9061:Key concepts
8964:Polymorphism
8947:Astrobiology
8895:Biogeography
8850:Saltationism
8840:Orthogenesis
8825:Alternatives
8754:
8740:
8673:Cospeciation
8668:Cladogenesis
8617:Saltationism
8574:Mating types
8558:
8497:Color vision
8482:Avian flight
8404:mitochondria
8142:Canalisation
8020:Biodiversity
7765:Introduction
7564:spermatozoon
7499:
7492:reproduction
7441:Y chromosome
7436:X chromosome
7382:Virilization
7377:Feminization
7254:
7231:
7210:
7183:
7180:Ridley, Matt
7161:
7158:Ridley, Mark
7139:
7136:Ridley, Mark
7124:. Retrieved
7120:the original
7113:
7090:
7069:
7044:
7012:(2): 46–52.
7009:
7003:
6984:
6965:
6962:Bell, Graham
6916:
6912:
6902:
6888:
6883:
6875:
6870:
6835:
6831:
6821:
6791:(1): 54–63.
6788:
6784:
6730:
6726:
6680:
6676:
6670:
6653:
6649:
6643:
6608:
6604:
6594:
6559:
6555:
6545:
6510:
6506:
6500:
6467:
6463:
6457:
6424:
6420:
6403:
6371:(1): 31–39.
6368:
6364:
6354:
6319:
6251:
6247:
6241:
6217:
6170:
6166:
6156:
6114:
6110:
6076:
6072:
6051:. Retrieved
6045:
6035:
6023:. Retrieved
5995:
5991:Paleobiology
5989:
5979:
5967:. Retrieved
5939:
5933:
5926:Zimmer, Carl
5920:
5908:. Retrieved
5902:
5892:
5865:
5861:
5829:
5825:
5795:
5791:
5781:
5754:
5750:
5704:
5700:
5694:
5681:
5674:
5623:
5619:
5613:
5580:
5576:
5570:
5545:
5541:
5535:
5492:
5488:
5478:
5435:
5431:
5421:
5386:
5382:
5372:
5331:
5327:
5321:
5288:
5284:
5271:
5263:
5258:
5250:
5245:
5196:
5192:
5186:
5153:
5149:
5143:
5110:
5106:
5100:
5059:
5053:
5033:
5029:
5024:
5019:see page 462
5005:
4960:
4956:
4946:
4913:
4909:
4886:
4881:
4862:
4834:
4791:
4755:
4744:
4711:
4707:
4701:
4676:
4672:
4666:
4625:
4621:
4615:
4564:
4560:
4554:
4544:19 September
4542:. Retrieved
4532:
4489:
4485:
4475:
4442:
4438:
4432:
4422:19 September
4420:. Retrieved
4410:
4369:
4365:
4355:
4314:
4310:
4304:
4277:
4273:
4263:
4218:
4212:
4199:
4190:
4186:
4180:
4147:
4139:
4131:
4122:
4095:
4051:
4047:
4013:
4009:
4003:
3942:
3939:PLOS Biology
3938:
3928:
3904:
3897:
3888:
3878:
3871:
3859:
3849:
3835:
3827:
3816:Phytologia …
3815:
3808:
3765:
3761:
3751:
3718:
3714:
3704:
3692:. Retrieved
3688:
3678:
3643:
3637:
3624:
3600:(1): 127–8.
3597:
3591:
3577:
3565:. Retrieved
3524:mating types
3514:
3490:
3481:
3471:
3459:
3440:
3426:
3418:
3411:
3396:
3387:
3364:
3337:
3317:
3303:
3295:
3286:
3278:
3269:
3253:
3218:
3201:
3185:
3144:
3133:
2681:Earth formed
2099:
2084:
2075:
2060:Please help
2048:
2027:
2026:
1994:
1985:Thierry Lodé
1978:
1922:
1915:
1913:
1905:
1896:
1881:
1877:
1873:
1872:(yeast) and
1869:
1858:
1850:
1847:
1836:
1830:
1825:
1819:
1813:
1796:
1785:
1774:
1760:
1744:
1726:
1715:
1706:
1702:
1671:
1655:
1640:
1636:
1608:
1604:mating types
1593:
1579:
1564:
1555:
1532:
1501:
1487:
1485:
1479:
1475:
1471:
1456:
1453:
1450:
1441:
1432:
1425:
1421:
1417:
1413:
1402:
1396:
1394:
1383:
1376:
1374:
1351:
1344:
1340:
1335:
1331:
1327:
1323:
1319:
1315:
1311:
1307:
1303:
1299:
1297:
1293:
1282:
1272:
1266:heterogamous
1262:
1256:
1244:
1234:
1230:
1209:
1200:
1190:
1183:
1176:
1169:
1157:
1145:Please help
1140:verification
1137:
1113:
1105:
1098:
1083:
1075:
1054:
1045:
1035:
1028:
1021:
1014:
1002:
990:Please help
985:verification
982:
958:
954:
939:
930:
907:
867:hybrid vigor
844:
828:
813:
794:
766:
758:transduction
742:maintenance.
741:
737:
682:
681:
633:
488:Sociobiology
473:Paleontology
321:
257:Biogeography
252:Biodiversity
170:Coextinction
160:Co-operation
135:Polymorphism
60:Introduction
15:
9484:Mike Levine
9393:Distal-less
9218:Polyphenism
9198:Epigenetics
9050:development
8974:Systematics
8845:Mutationism
8663:Catagenesis
8591:Snake venom
8524:Eusociality
8502:in primates
8492:Cooperation
8420:In animals
8240:butterflies
8213:Cephalopods
8203:Brachiopods
8135:Development
8109:Mate choice
7862:Convergence
7845:Coevolution
7803:Abiogenesis
7126:25 November
6556:J Bacteriol
6528:1885/109240
5707:(11): 818.
5156:(1): 6–25.
4136:Matt Ridley
3694:28 February
3505:nucleosomes
3477:lytic cycle
3461:with short
3390:August 2021
3340:cannibalism
3320:haloarchaea
2137:This box:
2078:August 2021
1686:Outcrossing
1651:outcrossing
1558:August 2021
1269:water fleas
933:August 2021
806:replication
750:conjugation
746:prokaryotes
701:could have
656:seed plants
498:Systematics
307:Renaissance
185:Convergence
175:Contingency
165:Coevolution
9589:Categories
9462:Lac operon
9287:Robustness
9266:Modularity
9261:Metamerism
9167:Plasticity
9162:Pleiotropy
9115:Heterotopy
8835:Lamarckism
8813:Philosophy
8736:David Hume
8698:Peripatric
8693:Parapatric
8678:Ecological
8658:Anagenesis
8653:Allopatric
8645:Speciation
8609:Gradualism
8534:Metabolism
8394:chromosome
8384:Eukaryotes
8162:Modularity
8079:Population
8005:Population
7966:Speciation
7944:Panspermia
7897:Extinction
7892:Exaptation
7867:Divergence
7840:Cladistics
7828:Reciprocal
7808:Adaptation
7625:Copulation
7346:Biological
6413:Schuster P
6073:BioScience
5998:(3): 386.
5862:Sexual Dev
5113:(1): 2–9.
5028:Griffiths
4561:Curr. Biol
4167:Softcover
4165:0306472619
3945:(2): e52.
3545:References
3445:virus, an
3330:swapping.
3221:eukaryotic
3208:DNA repair
3172:Bdelloidea
3167:eukaryotic
2473:Arthropods
2413:Eukaryotes
2009:proto-cell
2001:eukaryotes
1862:Drosophila
1837:By way of
1690:inbreeding
1427:C. elegans
1422:C. elegans
1414:C. elegans
1203:April 2023
1173:newspapers
1086:DNA repair
1070:See also:
1048:April 2023
1018:newspapers
816:DNA repair
718:Bdelloidea
711:eukaryotic
272:Cladistics
195:Extinction
180:Divergence
150:Speciation
130:Adaptation
44:John Gould
9413:Morphogen
9398:Engrailed
9381:Pax genes
9302:Tinkering
9148:Epistasis
9143:Dominance
9054:phenotype
8969:Protocell
8820:Darwinism
8708:Sympatric
8457:processes
8345:Tetrapods
8294:Kangaroos
8220:Dinosaurs
8157:Inversion
8126:Variation
8047:Gene flow
8040:Inclusive
7850:Mutualism
7795:Evolution
7680:Mechanics
7657:Sexuality
7552:Oogenesis
7527:Sex organ
7517:Germ cell
7505:Anisogamy
7140:Evolution
6919:: 34–46.
6143:ignored (
6133:cite book
5969:1 October
5910:1 October
5751:BioEssays
5701:BioEssays
5509:1759-6653
5364:205025450
5313:198153635
5285:Evolution
5251:Evolution
5237:206537272
5221:0036-8075
5170:0002-9122
4863:Evolution
4439:Evolution
4048:Evolution
3782:1943-0264
3528:anisogamy
3511:Questions
3467:cytoplasm
3463:telomeric
3451:bacterium
3443:lysogenic
3382:citations
3374:talk page
3313:F-plasmid
3281:RNA world
3249:orthologs
3174:and some
2494:Dinosaurs
2049:does not
2022:anisogamy
1989:prebiotic
1884:effect).
1853:recombine
1717:Mutations
1600:isogamous
1550:citations
1542:talk page
1504:isogamous
1285:parasites
1273:Cladocera
1078:mutations
925:citations
917:talk page
847:Aristotle
835:heterosis
773:dioecious
722:asexually
531:Dysgenics
247:Phylogeny
145:Gene flow
115:Diversity
110:Variation
9376:Hox gene
9364:Elements
9345:Homeobox
8997:Category
8872:Vitalism
8867:Theistic
8860:Spandrel
8544:Morality
8539:Monogamy
8414:plastids
8379:Flagella
8335:Reptiles
8316:sea cows
8299:primates
8208:Molluscs
8186:Bacteria
8074:Mutation
8007:genetics
7983:Taxonomy
7929:Mismatch
7909:Homology
7823:Cheating
7818:Altruism
7712:Category
7690:Activity
7586:Internal
7581:External
7470:Intersex
7253:(1975).
7208:(1995).
7182:(1995).
7160:(2000).
7138:(1993).
7115:NBC News
7067:(1978).
7034:21237760
6964:(1982).
6945:15383154
6862:16754610
6813:16846615
6765:20542871
6757:11523012
6705:12151256
6605:Genetics
6537:13687359
6507:Virology
6365:Genetics
6346:26593032
6276:10229582
6197:29436502
6093:86663600
6053:12 April
6047:BBC News
6025:12 April
6020:36648568
5964:39987391
5956:19498143
5884:22986519
5773:21472739
5729:46112804
5721:22009640
5658:12478292
5649:1842/692
5562:26721176
5527:38482769
5518:10972689
5470:28473589
5383:Genetics
5305:11108592
5229:22174251
5178:24363320
5135:14195748
5013:Archived
4997:11459950
4829:(1987).
4810:56964580
4752:(1978).
4736:31756267
4693:16856849
4650:15143283
4599:16005289
4524:21737739
4467:28565091
4394:19441961
4347:10437152
4068:21091466
4030:17612621
3971:18303954
3910:Pantheon
3800:24591519
3735:11483988
3689:Scitable
3670:21470960
3616:17050570
3449:, and a
3447:archaean
3245:homology
3182:Diploidy
3178:plants.
3147:protists
3135:Ice Ages
2528:Primates
2476:Molluscs
2106:mutation
1755:mutation
1619:ciliates
1615:bacteria
1508:copulate
1370:proteins
823:because
740:and its
726:apomixis
699:protists
641:Ladybugs
595:Category
521:Eugenics
363:timeline
344:Evo-devo
302:Overview
120:Mutation
82:Evidence
77:Glossary
9579:Science
9555:Biology
9541:Portals
9507:Debates
9318:Systems
9244:Eyespot
9108:Neoteny
8888:Related
8718:History
8579:Meiosis
8514:Empathy
8509:Emotion
8409:nucleus
8350:Viruses
8340:Spiders
8252:Mammals
8235:Insects
8035:Fitness
7971:Species
7770:Outline
7537:Meiosis
7510:Isogamy
7014:Bibcode
6853:1578732
6793:Bibcode
6735:Bibcode
6685:Bibcode
6635:6293914
6626:1201875
6586:5924273
6492:6209512
6472:Bibcode
6449:6164094
6429:Bibcode
6409:Eigen M
6395:8608929
6386:1203263
6284:9441768
6256:Bibcode
6188:5829205
6000:Bibcode
5935:Science
5666:4382757
5628:Bibcode
5605:5527627
5585:Bibcode
5461:6200135
5440:Bibcode
5432:Science
5413:9560386
5404:1460098
5356:9389477
5336:Bibcode
5201:Bibcode
5193:Science
5115:Bibcode
5092:4233528
5084:3057385
5064:Bibcode
4965:Bibcode
4938:6209512
4918:Bibcode
4716:Bibcode
4658:8599387
4630:Bibcode
4622:Science
4607:2229622
4579:Bibcode
4515:3402160
4494:Bibcode
4486:Science
4459:2410266
4402:6797643
4339:8308912
4319:Bibcode
4296:7877487
4255:2185476
4223:Bibcode
4193:: 1–30.
4076:7714974
3962:2253642
3791:3949356
3743:5465846
3661:3125724
3567:23 July
3473:Meiotic
3414:syngamy
3328:plasmid
3260:meiosis
3256:archaea
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