765:(cM) is a unit that describes a recombination frequency of 1%. In this way we can measure the genetic distance between two loci, based upon their recombination frequency. This is a good estimate of the real distance. Double crossovers would turn into no recombination. In this case we cannot tell if crossovers took place. If the loci we're analysing are very close (less than 7 cM) a double crossover is very unlikely. When distances become higher, the likelihood of a double crossover increases. As the likelihood of a double crossover increases one could systematically underestimate the genetic distance between two loci, unless one used an appropriate mathematical model.
337:
505:, is a statistical test often used for linkage analysis in human, animal, and plant populations. The LOD score compares the likelihood of obtaining the test data if the two loci are indeed linked, to the likelihood of observing the same data purely by chance. Positive LOD scores favour the presence of linkage, whereas negative LOD scores indicate that linkage is less likely. Computerised LOD score analysis is a simple way to analyse complex family pedigrees in order to determine the linkage between
988:
485:
477:
approach, whereby the probability that a gene important for a disease is linked to a genetic marker is studied through the LOD score, which assesses the probability that a given pedigree, where the disease and the marker are cosegregating, is due to the existence of linkage (with a given linkage value) or to chance. Non-parametric linkage analysis, in turn, studies the probability of an allele being
669:
1150:
females. In mammals, females often have a higher rate of recombination compared to males. It is theorised that there are unique selections acting or meiotic drivers which influence the difference in rates. The difference in rates may also reflect the vastly different environments and conditions of meiosis in oogenesis and spermatogenesis.
1227:
copy of the parental chromosome is transmitted, a '1' would be assigned to that meiosis. The two alleles in the parent came, one each, from two grandparents. These indicators are then used to determine identical-by-descent (IBD) states or inheritance states, which are in turn used to identify genes responsible for diseases.
421:. The greater the frequency of recombination (segregation) between two genetic markers, the further apart they are assumed to be. Conversely, the lower the frequency of recombination between the markers, the smaller the physical distance between them. Historically, the markers originally used were detectable
1050:). However, after crossover, some progeny could have received one parental chromosome with a dominant allele for one trait (e.g. Purple) linked to a recessive allele for a second trait (e.g. round) with the opposite being true for the other parental chromosome (e.g. red and Long). This is referred to as
541:
1062:
here would still be purple and long but a test cross of this individual with the recessive parent would produce progeny with much greater proportion of the two crossover phenotypes. While such a problem may not seem likely from this example, unfavourable repulsion linkages do appear when breeding for
768:
Double linkage is more of a historical concern for plants. In animals, double crossover happens rarely. In humans, for example, one chromosome has two crossovers on average during meiosis. Furthermore, modern geneticists have enough genes that only nearby genes need to be linkage-analyzed, unlike the
674:
NR denotes the number of non-recombinant offspring, and R denotes the number of recombinant offspring. The reason 0.5 is used in the denominator is that any alleles that are completely unlinked (e.g. alleles on separate chromosomes) have a 50% chance of recombination, due to independent assortment.
308:
frequency is more difficult to compute in an F2 cross than a backcross, but the lack of fit between observed and expected numbers of progeny in the above table indicate it is less than 50%. This indicated that two factors interacted in some way to create this difference by masking the appearance of
1092:
However, it is important to note that recombination frequency tends to underestimate the distance between two linked genes. This is because as the two genes are located farther apart, the chance of double or even number of crossovers between them also increases. Double or even number of crossovers
1226:
in a pedigree. The indicator indicates which copy of the parental chromosome contributes to the transmitted gamete at that position. For example, if the allele from the 'first' copy of the parental chromosome is transmitted, a '0' might be assigned to that meiosis. If the allele from the 'second'
697:
By convention, a LOD score greater than 3.0 is considered evidence for linkage, as it indicates 1000 to 1 odds that the linkage being observed did not occur by chance. On the other hand, a LOD score less than −2.0 is considered evidence to exclude linkage. Although it is very unlikely that a LOD
856:
When two genes are close together on the same chromosome, they do not assort independently and are said to be linked. Whereas genes located on different chromosomes assort independently and have a recombination frequency of 50%, linked genes have a recombination frequency that is less than 50%.
1149:
While recombination of chromosomes is an essential process during meiosis, there is a large range of frequency of cross overs across organisms and within species. Sexually dimorphic rates of recombination are termed heterochiasmy, and are observed more often than a common rate between male and
1136:
Edgar et al. performed mapping experiments with r mutants of bacteriophage T4 showing that recombination frequencies between rII mutants are not strictly additive. The recombination frequency from a cross of two rII mutants (a x d) is usually less than the sum of recombination frequencies for
476:
with the ailment phenotype through families. It can be used to map genes for both binary and quantitative traits. Linkage analysis may be either parametric (if we know the relationship between phenotypic and genetic similarity) or non-parametric. Parametric linkage analysis is the traditional
1221:
With very large pedigrees or with very dense genetic marker data, such as from whole-genome sequencing, it is possible to precisely locate recombinations. With this type of genetic analysis, a meiosis indicator is assigned to each position of the genome for each
1715:
Gusella, James F.; Frontali, Marina; Wasmuth, John J.; Collins, Francis S.; Lehrach, Hans; Myers, Richard; Altherr, Michael; Allitto, Bernice; Taylor, Sherry (1992-05-01). "The
Huntington's disease candidate region exhibits many different haplotypes".
449:, a set of genes which are known to be linked. As knowledge advances, more markers can be added to a group, until the group covers an entire chromosome. For well-studied organisms the linkage groups correspond one-to-one with the chromosomes.
100:. The first experiment to demonstrate linkage was carried out in 1905. At the time, the reason why certain traits tend to be inherited together was unknown. Later work revealed that genes are physical structures related by physical distance.
664:{\displaystyle {\text{LOD}}=Z=\log _{10}{\frac {\text{probability of birth sequence with a given linkage value}}{\text{probability of birth sequence with no linkage}}}=\log _{10}{\frac {(1-\theta )^{NR}\times \theta ^{R}}{0.5^{NR+R}}}}
88:
of potentially deleterious alleles may be influenced by the presence of other alleles, and these other alleles may be located on other chromosomes than that on which a particular potentially deleterious allele is located.
1181:
errors by the defective DNA polymerase that are themselves recombination events such as template switches, i.e. copy choice recombination events. Recombination is also increased by mutations that reduce the expression of
2117:
Fujisawa H, Yonesaki T, Minagawa T. Sequence of the T4 recombination gene, uvsX, and its comparison with that of the recA gene of
Escherichia coli. Nucleic Acids Res. 1985;13(20):7473-7481. doi:10.1093/nar/13.20.7473
444:
Linkage maps help researchers to locate other markers, such as other genes by testing for genetic linkage of the already known markers. In the early stages of developing a linkage map, the data are used to assemble
1039:. The recombination frequency is more difficult to compute in an F2 cross than a backcross, but the lack of fit between observed and expected numbers of progeny in the above table indicate it is less than 50%.
698:
score of 3 would be obtained from a single pedigree, the mathematical properties of the test allow data from a number of pedigrees to be combined by summing their LOD scores. A LOD score of 3 translates to a
679: is the recombinant fraction, i.e. the fraction of births in which recombination has happened between the studied genetic marker and the putative gene associated with the disease. Thus, it is equal to
1137:
adjacent internal sub-intervals (a x b) + (b x c) + (c x d). Although not strictly additive, a systematic relationship was observed that likely reflects the underlying molecular mechanism of
725:
rate and reduce the power to map human quantitative trait loci (QTL). While linkage analysis was successfully used to identify genetic variants that contribute to rare disorders such as
788:. The law of independent assortment always holds true for genes that are located on different chromosomes, but for genes that are on the same chromosome, it does not always hold true.
904:
would occur in a 9:3:3:1 ratio of PL:Pl:pL:pl. To their surprise, they observed an increased frequency of PL and pl and a decreased frequency of Pl and pL (see table below).
1093:
between the two genes results in them being cosegregated to the same gamete, yielding a parental progeny instead of the expected recombinant progeny. As mentioned above, the
107:(cM). A distance of 1 cM between two markers means that the markers are separated to different chromosomes on average once per 100 meiotic product, thus once per 50 meioses.
142:
cross-bred pea plants in experiments similar to Mendel's. They were interested in trait inheritance in the sweet pea and were studying two genes—the gene for flower colour (
2075:
Berger H, Warren AJ, Fry KE. Variations in genetic recombination due to amber mutations in T4D bacteriophage. J Virol. 1969;3(2):171-175. doi:10.1128/JVI.3.2.171-175.1969
2099:
Bernstein H. Repair and recombination in phage T4. I. Genes affecting recombination. Cold Spring Harb Symp Quant Biol. 1968;33:325-331. doi:10.1101/sqb.1968.033.01.037
2108:
Hamlett NV, Berger H. Mutations altering genetic recombination and repair of DNA in bacteriophage T4. Virology. 1975;63(2):539-567. doi:10.1016/0042-6822(75)90326-8
2057:
Bernstein H. The effect on recombination of mutational defects in the DNA-polymerase and deoxycytidylate hydroxymethylase of phage T4D. Genetics. 1967;56(4):755-769
806:. A and a and B and b represent the alleles of genes A and B. Crossing these homozygous parental strains will result in F1 generation offspring that are double
1081:
producing recombination between the genes is related to the distance between the two genes. Thus, the use of recombination frequencies has been used to develop
398:
relative to each other in terms of recombination frequency, rather than a specific physical distance along each chromosome. Linkage maps were first developed by
309:
the other two phenotypes. This led to the conclusion that some traits are related to each other because of their near proximity to each other on a chromosome.
826:
with equal frequencies (25%) because the alleles of gene A assort independently of the alleles for gene B during meiosis. Note that 2 of the 4 gametes (50%)—
181:
would occur in a 9:3:3:1 ratio of PL:Pl:pL:pl. To their surprise, they observed an increased frequency of PL and pl and a decreased frequency of Pl and pL:
2084:
Bernstein H. On the mechanism of intragenic recombination. I. The rII region of bacteriophage T4. (1962) Journal of
Theoretical Biology. 1962; 3, 335-353.
2266:
1294:"Where genotype is not predictive of phenotype: towards an understanding of the molecular basis of reduced penetrance in human inherited disease"
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could be mapped in a linear order. This result provided evidence for the key idea that the gene has a linear structure equivalent to a length of
1959:
Edgar RS, Feynman RP, Klein S, Lielausis I, Steinberg CM. Mapping experiments with r mutants of bacteriophage T4D. Genetics. 1962;47:179–186.
2223:
1201:(gp32) Mutation in the bacteriophage uvsX gene also substantially reduces recombination. The uvsX gene is analogous to the well studied
737:. An explanation for this is that the genetic mechanisms affecting common disorders are different from those causing some rare disorders.
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1941:
Benzer S. Fine structure of a genetic region in bacteriophage. Proc Natl Acad Sci U S A. 1955;41(6):344-354. doi:10.1073/pnas.41.6.344
745:
Recombination frequency is a measure of genetic linkage and is used in the creation of a genetic linkage map. Recombination frequency (
1950:
Benzer S. On the topology of the genetic fine structure. Proc Natl Acad Sci U S A. 1959;45(11):1607-1620. doi:10.1073/pnas.45.11.1607
1385:
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2374:
1664:
Ferreira, Manuel A. R. (2004-10-01). "Linkage
Analysis: Principles and Methods for the Analysis of Human Quantitative Traits".
1932:
Graph of mapping function from compared to idealised 1-1 equivalence of recombination frequency percentage (RF%) to map units.
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2192:
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1925:
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Mark J. Daly; Hirschhorn, Joel N. (2005-02-01). "Genome-wide association studies for common diseases and complex traits".
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between linked genes differs led to the idea that crossover frequency might indicate the distance separating genes on the
1983:
Fisher KM, Bernstein H. The additivity of intervals in the RIIA cistron of phage T4D. Genetics. 1965;52 (6):1127–1136.
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868:. They were interested in trait inheritance in the sweet pea and were studying two genes—the gene for flower colour (
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between them, and the more likely they are to be inherited together. Markers on different chromosomes are perfectly
353:
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Linkage analysis has a number of methodological and theoretical limitations that can significantly increase the
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cell. In this example, the recombination frequency is 50% since 2 of the 4 gametes were recombinant gametes.
123:
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119:
97:
17:
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Cantor, Rita M. (2013), "Analysis of
Genetic Linkage", in Rimoin, David; Pyeritz, Reed; Korf, Bruce (eds.),
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or when they are widely separated on the same chromosome. This is a consequence of independent assortment.
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increase recombination (decrease linkage) several fold. The increase in recombination may be due to
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The two possible arrangements, cis and trans, of alleles in a double heterozygote are referred to as
722:
434:
1415:; Balmukand, B (July 1928). "The estimation of linkage from the offspring of selfed heterozygotes".
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The progeny in this case received two dominant alleles linked on one chromosome (referred to as
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Cooper, DN; Krawczak, M; Polychronakos, C; Tyler-Smith, C; Kehrer-Sawatzki, H (October 2013).
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Recombination is reduced (linkage increased) by mutations in genes that encode proteins with
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390:) is a table for a species or experimental population that shows the position of its known
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states that every trait is inherited independently of every other trait. But shortly after
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Tendency of DNA sequences that are close together on a chromosome to be inherited together
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Meneely, Philip Mark; Dawes Hoang, Rachel; Okeke, Iruka N.; Heston, Katherine (2017).
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that are physically near to each other are unlikely to be separated onto different
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is the process of determining which of the two is present in a given individual.
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Linkage analysis is a genetic method that searches for chromosomal segments that
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131:
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The recombination frequency will be 50% when two genes are located on different
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As an example of independent assortment, consider the crossing of the pure-bred
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729:, it did not perform that well when applied to more common disorders such as
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41:
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and arranged like beads on a string. During 1955 to 1959, Benzer performed
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The estimate with the highest LOD score will be considered the best estimate
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2009:
1972:
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365:
349:
2153:
1996:
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of one gene is independent of alleles of another gene. This is stated in
328:, which expresses the frequency of crossing over, is named in his honour.
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850:
762:
325:
104:
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834:—were not present in the parental generation. These gametes represent
792:
321:
85:
61:
45:
2201:
2154:
Griffiths AJF; Miller JH; Suzuki DT; Lewontin RC; et al. (1993).
1729:
1291:
838:. Recombinant gametes are those gametes that differ from both of the
2680:
2459:
1059:
901:
810:
with genotype AaBb. The F1 offspring AaBb produces gametes that are
422:
178:
2527:
1780:
1165:
that encode proteins involved in the processing of DNA often affect
1125:. He found that, on the basis of recombination tests, the sites of
2014:
Biochimica et
Biophysica Acta (BBA) - Gene Structure and Expression
2010:"Homologous pairing and chromosome dynamics in meiosis and mitosis"
1589:
1194:
1158:
1126:
1077:
When two genes are located on the same chromosome, the chance of a
796:
461:
377:
31:
2572:
1223:
1186:(gp30) and dCMP hydroxymethylase (gp42), two enzymes employed in
843:
839:
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699:
357:
49:
72:
than markers that are far apart. In other words, the nearer two
1105:
In the early 1950s the prevailing view was that the genes in a
777:
773:
734:
512:
The method is described in greater detail by
Strachan and Read.
437:
or those generating restriction fragment length polymorphisms (
1499:
Emery and Rimoin's
Principles and Practice of Medical Genetics
1817:
1714:
860:
As an example of linkage, consider the classic experiment by
391:
73:
1623:"A Note on Multiple Testing Procedures in Linkage Analysis"
1203:
1162:
754:
438:
1153:
876:, red) and the gene affecting the shape of pollen grains (
509:
traits (or between a trait and a marker, or two markers).
150:, red) and the gene affecting the shape of pollen grains (
1910:"Accurate calculation of large map distances, Figure 6-4"
1130:
1100:
312:
The understanding of linkage was expanded by the work of
575:
probability of birth sequence with a given linkage value
1387:
Reports to the
Evolution committee of the Royal Society
1144:
1766:
1372:
1173:, mutations that reduce expression of the replicative
30:"Genetic map" redirects here. Not to be confused with
1907:
544:
525:
Make a number of estimates of recombination frequency
2182:
2168:(5th ed.). New York: W.H. Freeman and Company.
409:A linkage map is a map based on the frequencies of
348:genetic linkage map. This was the first successful
126:, exceptions to this rule were found. In 1905, the
92:Genetic linkage is the most prominent exception to
2161:
2155:
663:
772:During meiosis, chromosomes assort randomly into
488:Pedigree illustrating Parametric Linkage Analysis
2729:
2095:
2093:
2053:
2051:
1824:. Oxford: Oxford University Press. p. 361.
802:with a different pure-bred strain with genotype
1908:Griffiths, AJF; Miller, JH; Suzuki, DT (2000).
1531:"Sequential tests for the detection of linkage"
1411:
1133:with many sites that can independently mutate.
1852:
1501:(6th ed.), Academic Press, pp. 1–9,
492:
264:Their experiment revealed linkage between the
2543:
2217:
2090:
2086:https://doi.org/10.1016/S0022-5193(62)80030-7
2071:
2069:
2067:
2065:
2063:
2048:
769:early days when only a few genes were known.
578:probability of birth sequence with no linkage
425:(enzyme production, eye colour) derived from
364:chromosome. The distances between the genes (
352:work and provides important evidence for the
76:are on a chromosome, the lower the chance of
2183:Poehlman JM; Sleper DA (1995). "Chapter 3".
1492:
1490:
1213:that plays a central role in recombination.
1097:attempt to correct for multiple crossovers.
501:(logarithm (base 10) of odds), developed by
429:sequences; eventually, confirmed or assumed
372:events that occur between different alleles.
103:The typical unit of genetic linkage is the
2550:
2536:
2224:
2210:
2060:
1853:Punnett, R. C.; Bateson, W. (1908-05-15).
740:
356:. The map shows the relative positions of
316:. Morgan's observation that the amount of
1640:
1597:
1546:
1528:
1487:
1476:
1319:
186:Bateson, Saunders, and Punnett experiment
2187:(4th ed.). Iowa: Iowa State Press.
1663:
1407:
1405:
1345:"Discovery and Types of Genetic Linkage"
1031:is greater than that of the recombinant
986:
535:The LOD score is calculated as follows:
483:
335:
296:is greater than that of the recombinant
2231:
1821:Genetics: genes, genomes, and evolution
1154:Genes affecting recombination frequency
884:, round). They crossed the pure lines
749:) is the frequency with which a single
528:Calculate a LOD score for each estimate
14:
2730:
2517:Index of evolutionary biology articles
1569:
1496:
1456:. New York: McGraw-Hill. p. 209.
1109:are discrete entities, indivisible by
1101:Linkage of genetic sites within a gene
158:, round). They crossed the pure lines
2557:
2531:
2205:
2007:
1620:
1451:
1402:
1390:. London: Harrison and Sons, Printers
1285:
1216:
1342:
1145:Variation of recombination frequency
892:and then self-crossed the resulting
166:and then self-crossed the resulting
68:, and are therefore said to be more
48:to be inherited together during the
2164:An Introduction to Genetic Analysis
1914:An Introduction to Genetic Analysis
1480:An Introduction to Genetic Analysis
1095:Kosambi and Haldane transformations
467:
24:
1628:American Journal of Human Genetics
1577:American Journal of Human Genetics
1535:American Journal of Human Genetics
1507:10.1016/b978-0-12-383834-6.00010-0
1063:disease resistance in some crops.
842:gametes that made up the original
368:) are equal to the percentages of
25:
2754:
786:the law of independent assortment
1666:Twin Research and Human Genetics
1572:"All LODs Are Not Created Equal"
1197:functions (gp46 and gp47) and a
354:chromosome theory of inheritance
2614:Single-nucleotide polymorphisms
2111:
2102:
2078:
2001:
1977:
1953:
1944:
1935:
1901:
1846:
1811:
1760:
1708:
1657:
1614:
991:Unlinked Genes vs. Linked Genes
776:, such that the segregation of
2717:Human Genome Diversity Project
2317:Constructive neutral evolution
2008:McKee, Bruce D. (2004-03-15).
1570:Nyholt, Dale R (August 2000).
1563:
1522:
1483:(7th ed.). W. H. Freeman.
1470:
1445:
1366:
1336:
909:Bateson and Punnett experiment
716:
705:of approximately 0.05, and no
614:
601:
515:Briefly, it works as follows:
331:
124:Mendel's work was rediscovered
13:
1:
2676:Genome-wide association study
1279:
1253:Genome-wide association study
120:Law of Independent Assortment
98:Law of Independent Assortment
44:that are close together on a
2707:International HapMap Project
2267:Fisher's fundamental theorem
2144:Resources in other libraries
2026:10.1016/j.bbaexp.2003.11.017
921:Expected from 9:3:3:1 ratio
753:will take place between two
198:Expected from 9:3:3:1 ratio
110:
7:
2292:Coefficient of relationship
1343:Lobo, Ingrid; Shaw, Kenna.
1230:
1015:alleles. The frequency of
707:multiple testing correction
493:Parametric linkage analysis
10:
2759:
1916:(7th ed.). New York:
1879:10.1126/science.27.698.785
995:Their experiment revealed
456:a physical map (such as a
375:
280:alleles. The frequency of
29:
2694:
2663:
2632:
2586:
2565:
2512:
2447:
2421:
2383:
2358:
2325:
2287:Coefficient of inbreeding
2239:
2139:Resources in your library
1621:Risch, Neil (June 1991).
1312:10.1007/s00439-013-1331-2
2465:Evolutionary game theory
2247:Hardy–Weinberg principle
1027:occurring together with
1019:occurring together with
458:radiation reduced hybrid
292:occurring together with
284:occurring together with
2671:Whole genome sequencing
2604:Human genetic variation
2277:Shifting balance theory
1769:Nature Reviews Genetics
1477:Griffiths, AJF (2000).
741:Recombination frequency
413:between markers during
345:Drosophila melanogaster
2262:Linkage disequilibrium
1452:Mader, Sylvia (2007).
1268:Linkage disequilibrium
1263:Lander–Green algorithm
992:
915:Phenotype and genotype
733:or different forms of
665:
489:
419:homologous chromosomes
373:
192:Phenotype and genotype
136:Edith Rebecca Saunders
2640:Personalized medicine
2504:Quantitative genetics
2413:Balding–Nichols model
2398:Population bottleneck
2393:Small population size
2297:Selection coefficient
1855:"The Heredity of Sex"
1454:Biology Ninth Edition
1139:genetic recombination
1115:genetic recombination
1111:genetic recombination
990:
896:lines. According to
795:parental strain with
751:chromosomal crossover
711:Bonferroni correction
666:
487:
370:chromosomal crossover
339:
66:chromosomal crossover
2712:1000 Genomes Project
2702:Human Genome Project
2650:Genetic epidemiology
2375:Background selection
2362:on genomic variation
2360:Effects of selection
2312:Population structure
2185:Breeding Field Crops
1678:10.1375/twin.7.5.513
1248:Genetic epidemiology
542:
479:identical by descent
341:Thomas Hunt Morgan's
2743:Population genetics
2664:Analysis techniques
2645:Predictive medicine
2619:Identity by descent
2594:Biological specimen
2578:Biological database
2494:Population genomics
2370:Genetic hitchhiking
2257:Identity by descent
2233:Population genetics
1871:1908Sci....27..785P
1418:Journal of Genetics
1258:Identity by descent
1243:Genetic association
1199:DNA-binding protein
911:
836:recombinant gametes
782:Mendel's Second Law
188:
54:sexual reproduction
40:is the tendency of
2738:Classical genetics
2480:Landscape genetics
1529:Morton NE (1955).
1431:10.1007/BF02983317
1217:Meiosis indicators
1117:experiments using
993:
907:
898:Mendelian genetics
727:Huntington disease
661:
490:
441:) have been used.
433:sequences such as
404:Thomas Hunt Morgan
374:
314:Thomas Hunt Morgan
184:
175:Mendelian genetics
2725:
2724:
2599:De-identification
2559:Personal genomics
2525:
2524:
2475:Genetic genealogy
2470:Fitness landscape
2194:978-0-8138-2427-7
2175:978-0-7167-2285-4
2125:Library resources
1927:978-0-7167-3520-5
1831:978-0-19-879536-0
1730:10.1038/ng0592-99
1463:978-0-07-325839-3
1056:trans arrangement
985:
984:
659:
580:
579:
576:
548:
452:A linkage map is
400:Alfred Sturtevant
386:(also known as a
262:
261:
16:(Redirected from
2750:
2655:Pharmacogenomics
2624:Genetic disorder
2552:
2545:
2538:
2529:
2528:
2434:J. B. S. Haldane
2226:
2219:
2212:
2203:
2202:
2198:
2179:
2167:
2159:
2118:
2115:
2109:
2106:
2100:
2097:
2088:
2082:
2076:
2073:
2058:
2055:
2046:
2045:
2020:(1–3): 165–180.
2005:
1999:
1981:
1975:
1957:
1951:
1948:
1942:
1939:
1933:
1931:
1905:
1899:
1898:
1865:(698): 785–787.
1850:
1844:
1843:
1815:
1809:
1808:
1764:
1758:
1757:
1712:
1706:
1705:
1661:
1655:
1654:
1644:
1635:(6): 1058–1064.
1618:
1612:
1611:
1601:
1567:
1561:
1560:
1550:
1526:
1520:
1519:
1494:
1485:
1484:
1474:
1468:
1467:
1449:
1443:
1442:
1409:
1400:
1399:
1397:
1395:
1370:
1364:
1363:
1361:
1359:
1353:Nature Education
1340:
1334:
1333:
1323:
1306:(10): 1077–130.
1289:
1273:Structural motif
1210:Escherichia coli
1171:bacteriophage T4
1123:bacteriophage T4
1007:alleles and the
912:
906:
866:Reginald Punnett
784:and is known as
693:
670:
668:
667:
662:
660:
658:
657:
639:
638:
637:
625:
624:
599:
594:
593:
581:
577:
574:
573:
568:
567:
549:
546:
468:Linkage analysis
272:alleles and the
189:
183:
140:Reginald Punnett
21:
2758:
2757:
2753:
2752:
2751:
2749:
2748:
2747:
2728:
2727:
2726:
2721:
2690:
2686:Genetic testing
2659:
2628:
2609:Genetic linkage
2582:
2566:Data collection
2561:
2556:
2526:
2521:
2508:
2443:
2417:
2379:
2363:
2361:
2354:
2321:
2252:Genetic linkage
2235:
2230:
2195:
2176:
2150:
2149:
2148:
2133:
2132:
2128:
2121:
2116:
2112:
2107:
2103:
2098:
2091:
2083:
2079:
2074:
2061:
2056:
2049:
2006:
2002:
1982:
1978:
1958:
1954:
1949:
1945:
1940:
1936:
1928:
1906:
1902:
1851:
1847:
1832:
1816:
1812:
1781:10.1038/nrg1521
1765:
1761:
1718:Nature Genetics
1713:
1709:
1662:
1658:
1619:
1615:
1568:
1564:
1527:
1523:
1517:
1495:
1488:
1475:
1471:
1464:
1450:
1446:
1410:
1403:
1393:
1391:
1384:(18 May 1904).
1371:
1367:
1357:
1355:
1341:
1337:
1290:
1286:
1282:
1277:
1233:
1219:
1169:frequency. In
1156:
1147:
1103:
1048:cis arrangement
941:Purple, round (
900:, the expected
862:William Bateson
743:
719:
713:) is required.
680:
644:
640:
633:
629:
617:
613:
600:
598:
589:
585:
572:
563:
559:
545:
543:
540:
539:
495:
470:
435:microsatellites
402:, a student of
396:genetic markers
380:
334:
218:Purple, round (
177:, the expected
132:William Bateson
113:
84:, although the
58:genetic markers
38:Genetic linkage
35:
28:
23:
22:
15:
12:
11:
5:
2756:
2746:
2745:
2740:
2723:
2722:
2720:
2719:
2714:
2709:
2704:
2698:
2696:
2695:Major projects
2692:
2691:
2689:
2688:
2683:
2678:
2673:
2667:
2665:
2661:
2660:
2658:
2657:
2652:
2647:
2642:
2636:
2634:
2630:
2629:
2627:
2626:
2621:
2616:
2611:
2606:
2601:
2596:
2590:
2588:
2587:Field concepts
2584:
2583:
2581:
2580:
2575:
2569:
2567:
2563:
2562:
2555:
2554:
2547:
2540:
2532:
2523:
2522:
2520:
2519:
2513:
2510:
2509:
2507:
2506:
2501:
2499:Phylogeography
2496:
2491:
2489:Microevolution
2486:
2477:
2472:
2467:
2462:
2457:
2451:
2449:
2448:Related topics
2445:
2444:
2442:
2441:
2436:
2431:
2425:
2423:
2419:
2418:
2416:
2415:
2410:
2405:
2403:Founder effect
2400:
2395:
2389:
2387:
2381:
2380:
2378:
2377:
2372:
2366:
2364:
2359:
2356:
2355:
2353:
2352:
2347:
2342:
2337:
2331:
2329:
2323:
2322:
2320:
2319:
2314:
2309:
2304:
2299:
2294:
2289:
2284:
2282:Price equation
2279:
2274:
2272:Neutral theory
2269:
2264:
2259:
2254:
2249:
2243:
2241:
2237:
2236:
2229:
2228:
2221:
2214:
2206:
2200:
2199:
2193:
2180:
2174:
2147:
2146:
2141:
2135:
2134:
2123:
2122:
2120:
2119:
2110:
2101:
2089:
2077:
2059:
2047:
2000:
1976:
1952:
1943:
1934:
1926:
1900:
1845:
1830:
1810:
1759:
1707:
1672:(5): 513–530.
1656:
1613:
1590:10.1086/303029
1584:(2): 282–288.
1562:
1541:(3): 277–318.
1521:
1515:
1486:
1469:
1462:
1444:
1401:
1365:
1335:
1299:Human genetics
1283:
1281:
1278:
1276:
1275:
1270:
1265:
1260:
1255:
1250:
1245:
1240:
1234:
1232:
1229:
1218:
1215:
1175:DNA polymerase
1155:
1152:
1146:
1143:
1102:
1099:
1068:gametic phases
983:
982:
979:
976:
968:
967:
964:
961:
953:
952:
949:
946:
938:
937:
934:
931:
926:Purple, long (
923:
922:
919:
916:
872:, purple, and
742:
739:
718:
715:
672:
671:
656:
653:
650:
647:
643:
636:
632:
628:
623:
620:
616:
612:
609:
606:
603:
597:
592:
588:
584:
571:
566:
562:
558:
555:
552:
533:
532:
529:
526:
523:
494:
491:
469:
466:
447:linkage groups
360:on the second
333:
330:
260:
259:
256:
253:
245:
244:
241:
238:
230:
229:
226:
223:
215:
214:
211:
208:
203:Purple, long (
200:
199:
196:
193:
146:, purple, and
112:
109:
26:
9:
6:
4:
3:
2:
2755:
2744:
2741:
2739:
2736:
2735:
2733:
2718:
2715:
2713:
2710:
2708:
2705:
2703:
2700:
2699:
2697:
2693:
2687:
2684:
2682:
2679:
2677:
2674:
2672:
2669:
2668:
2666:
2662:
2656:
2653:
2651:
2648:
2646:
2643:
2641:
2638:
2637:
2635:
2631:
2625:
2622:
2620:
2617:
2615:
2612:
2610:
2607:
2605:
2602:
2600:
2597:
2595:
2592:
2591:
2589:
2585:
2579:
2576:
2574:
2571:
2570:
2568:
2564:
2560:
2553:
2548:
2546:
2541:
2539:
2534:
2533:
2530:
2518:
2515:
2514:
2511:
2505:
2502:
2500:
2497:
2495:
2492:
2490:
2487:
2485:
2481:
2478:
2476:
2473:
2471:
2468:
2466:
2463:
2461:
2458:
2456:
2453:
2452:
2450:
2446:
2440:
2439:Sewall Wright
2437:
2435:
2432:
2430:
2427:
2426:
2424:
2420:
2414:
2411:
2409:
2406:
2404:
2401:
2399:
2396:
2394:
2391:
2390:
2388:
2386:
2385:Genetic drift
2382:
2376:
2373:
2371:
2368:
2367:
2365:
2357:
2351:
2348:
2346:
2343:
2341:
2338:
2336:
2333:
2332:
2330:
2328:
2324:
2318:
2315:
2313:
2310:
2308:
2305:
2303:
2300:
2298:
2295:
2293:
2290:
2288:
2285:
2283:
2280:
2278:
2275:
2273:
2270:
2268:
2265:
2263:
2260:
2258:
2255:
2253:
2250:
2248:
2245:
2244:
2242:
2238:
2234:
2227:
2222:
2220:
2215:
2213:
2208:
2207:
2204:
2196:
2190:
2186:
2181:
2177:
2171:
2166:
2165:
2158:
2152:
2151:
2145:
2142:
2140:
2137:
2136:
2131:
2126:
2114:
2105:
2096:
2094:
2087:
2081:
2072:
2070:
2068:
2066:
2064:
2054:
2052:
2043:
2039:
2035:
2031:
2027:
2023:
2019:
2015:
2011:
2004:
1998:
1994:
1990:
1986:
1980:
1974:
1970:
1966:
1962:
1956:
1947:
1938:
1929:
1923:
1919:
1918:W. H. Freeman
1915:
1911:
1904:
1896:
1892:
1888:
1884:
1880:
1876:
1872:
1868:
1864:
1860:
1856:
1849:
1841:
1837:
1833:
1827:
1823:
1822:
1814:
1806:
1802:
1798:
1794:
1790:
1786:
1782:
1778:
1775:(2): 95–108.
1774:
1770:
1763:
1755:
1751:
1747:
1743:
1739:
1735:
1731:
1727:
1724:(2): 99–103.
1723:
1719:
1711:
1703:
1699:
1695:
1691:
1687:
1683:
1679:
1675:
1671:
1667:
1660:
1652:
1648:
1643:
1638:
1634:
1630:
1629:
1624:
1617:
1609:
1605:
1600:
1595:
1591:
1587:
1583:
1579:
1578:
1573:
1566:
1558:
1554:
1549:
1544:
1540:
1536:
1532:
1525:
1518:
1516:9780123838346
1512:
1508:
1504:
1500:
1493:
1491:
1482:
1481:
1473:
1465:
1459:
1455:
1448:
1440:
1436:
1432:
1428:
1424:
1420:
1419:
1414:
1408:
1406:
1389:
1388:
1383:
1379:
1375:
1369:
1354:
1350:
1346:
1339:
1331:
1327:
1322:
1317:
1313:
1309:
1305:
1301:
1300:
1295:
1288:
1284:
1274:
1271:
1269:
1266:
1264:
1261:
1259:
1256:
1254:
1251:
1249:
1246:
1244:
1241:
1239:
1236:
1235:
1228:
1225:
1214:
1212:
1211:
1206:
1205:
1200:
1196:
1191:
1189:
1188:DNA synthesis
1185:
1180:
1176:
1172:
1168:
1167:recombination
1164:
1160:
1151:
1142:
1140:
1134:
1132:
1128:
1124:
1120:
1116:
1112:
1108:
1098:
1096:
1090:
1088:
1084:
1080:
1075:
1073:
1069:
1064:
1061:
1057:
1053:
1049:
1045:
1040:
1038:
1034:
1030:
1026:
1022:
1018:
1014:
1010:
1006:
1002:
998:
989:
980:
977:
974:
970:
969:
965:
962:
959:
955:
954:
950:
947:
944:
940:
939:
935:
932:
929:
925:
924:
920:
917:
914:
913:
910:
905:
903:
899:
895:
891:
887:
883:
879:
875:
871:
867:
863:
858:
854:
852:
847:
845:
841:
837:
833:
829:
825:
821:
817:
813:
809:
808:heterozygotes
805:
801:
798:
794:
789:
787:
783:
779:
775:
770:
766:
764:
760:
756:
752:
748:
738:
736:
732:
731:heart disease
728:
724:
714:
712:
708:
704:
702:
695:
691:
687:
683:
678:
654:
651:
648:
645:
641:
634:
630:
626:
621:
618:
610:
607:
604:
595:
590:
586:
582:
569:
564:
560:
556:
553:
550:
538:
537:
536:
530:
527:
524:
522:
518:
517:
516:
514:
510:
508:
504:
503:Newton Morton
500:
486:
482:
481:with itself.
480:
475:
465:
463:
459:
455:
450:
448:
442:
440:
436:
432:
431:noncoding DNA
428:
424:
420:
416:
412:
411:recombination
407:
405:
401:
397:
393:
389:
385:
379:
371:
367:
363:
359:
355:
351:
347:
346:
342:
338:
329:
327:
323:
319:
318:crossing over
315:
310:
307:
306:recombination
303:
299:
295:
291:
287:
283:
279:
275:
271:
267:
257:
254:
251:
247:
246:
242:
239:
236:
232:
231:
227:
224:
221:
217:
216:
212:
209:
206:
202:
201:
197:
194:
191:
190:
187:
182:
180:
176:
173:According to
171:
169:
165:
161:
157:
153:
149:
145:
141:
137:
133:
129:
125:
121:
117:
116:Gregor Mendel
108:
106:
101:
99:
95:
94:Gregor Mendel
90:
87:
83:
79:
78:recombination
75:
71:
67:
63:
59:
55:
51:
47:
43:
42:DNA sequences
39:
33:
19:
18:Linkage group
2633:Applications
2608:
2455:Biogeography
2429:R. A. Fisher
2307:Heritability
2251:
2240:Key concepts
2184:
2163:
2130:Gene mapping
2129:
2113:
2104:
2080:
2017:
2013:
2003:
1979:
1955:
1946:
1937:
1913:
1903:
1862:
1858:
1848:
1820:
1813:
1772:
1768:
1762:
1721:
1717:
1710:
1669:
1665:
1659:
1632:
1626:
1616:
1581:
1575:
1565:
1538:
1534:
1524:
1498:
1479:
1472:
1453:
1447:
1425:(1): 79–92.
1422:
1416:
1392:. Retrieved
1386:
1378:Saunders, ER
1368:
1356:. Retrieved
1348:
1338:
1303:
1297:
1287:
1220:
1208:
1202:
1192:
1157:
1148:
1135:
1104:
1091:
1087:genetic maps
1086:
1083:linkage maps
1082:
1076:
1071:
1065:
1055:
1051:
1047:
1043:
1041:
1036:
1032:
1028:
1024:
1020:
1016:
1012:
1008:
1004:
1000:
999:between the
996:
994:
972:
971:Red, round (
957:
942:
927:
908:
893:
889:
885:
881:
880:, long, and
877:
873:
869:
859:
855:
848:
835:
831:
827:
823:
819:
815:
811:
803:
799:
790:
785:
771:
767:
746:
744:
723:type-1 error
720:
700:
696:
689:
685:
681:
676:
673:
534:
519:Establish a
511:
498:
496:
471:
453:
451:
446:
443:
408:
387:
383:
381:
366:centimorgans
361:
350:gene mapping
343:
311:
301:
297:
293:
289:
285:
281:
277:
273:
269:
265:
263:
249:
248:Red, round (
234:
219:
204:
185:
172:
167:
163:
159:
155:
154:, long, and
151:
147:
143:
130:geneticists
114:
102:
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81:
69:
37:
36:
2408:Coalescence
2157:"Chapter 5"
1382:Punnett, RC
1238:Centimorgan
1179:replication
1121:mutants of
956:Red, long (
851:chromosomes
763:centimorgan
717:Limitations
474:cosegregate
388:genetic map
384:linkage map
332:Linkage map
326:centimorgan
233:Red, long (
105:centimorgan
2732:Categories
2350:Ecological
2340:Artificial
1413:Fisher, RA
1394:21 January
1374:Bateson, W
1358:21 January
1280:References
1184:DNA ligase
1107:chromosome
902:phenotypes
793:homozygote
427:coding DNA
423:phenotypes
376:See also:
362:Drosophila
322:chromosome
179:phenotypes
86:penetrance
62:chromatids
46:chromosome
2681:SNP array
2460:Evolution
2327:Selection
2034:0006-3002
1887:0036-8075
1840:951645141
1789:1471-0064
1738:1546-1718
1702:199001341
1686:2053-6003
1159:Mutations
1079:crossover
1060:phenotype
1052:repulsion
1023:and with
631:θ
627:×
611:θ
608:−
596:
570:
507:Mendelian
499:LOD score
415:crossover
111:Discovery
52:phase of
2484:genomics
2422:Founders
2042:15020057
1973:13889186
1895:17791047
1797:15716906
1754:25472459
1694:15527667
1608:10884360
1557:13258560
1439:27688031
1349:Scitable
1330:23820649
1231:See also
1207:gene of
1195:nuclease
1127:mutation
1044:coupling
918:Observed
797:genotype
521:pedigree
462:gene map
460:map) or
378:Gene map
195:Observed
82:unlinked
32:Gene map
2573:Biobank
2335:Natural
2302:Fitness
1997:5882191
1989:1210971
1965:1210321
1867:Bibcode
1859:Science
1805:2813666
1746:1302016
1651:2035526
1642:1683115
1599:1287176
1548:1716611
1321:3778950
1224:meiosis
1072:phasing
997:linkage
844:diploid
840:haploid
778:alleles
774:gametes
759:meiosis
757:during
358:alleles
170:lines.
128:British
64:during
50:meiosis
2345:Sexual
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1058:. The
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735:cancer
709:(e.g.
703:-value
324:. The
304:. The
70:linked
56:. Two
1801:S2CID
1750:S2CID
1698:S2CID
1435:S2CID
1163:genes
1054:or a
755:genes
439:RFLPs
392:genes
74:genes
2482:and
2189:ISBN
2170:ISBN
2038:PMID
2030:ISSN
2018:1677
1993:PMID
1969:PMID
1922:ISBN
1891:PMID
1883:ISSN
1836:OCLC
1826:ISBN
1793:PMID
1785:ISSN
1742:PMID
1734:ISSN
1690:PMID
1682:ISSN
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1604:PMID
1553:PMID
1511:ISBN
1458:ISBN
1396:2017
1360:2017
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1985:PMC
1961:PMC
1875:doi
1777:doi
1726:doi
1674:doi
1637:PMC
1594:PMC
1586:doi
1543:PMC
1503:doi
1427:doi
1316:PMC
1308:doi
1304:132
1161:in
1131:DNA
1119:rII
1085:or
1046:or
981:24
966:72
951:72
933:284
684:/ (
642:0.5
587:log
561:log
547:LOD
454:not
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