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Mendelian inheritance

Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularized by William Bateson.[1] These principles were initially controversial. When Mendel's theories were integrated with the Boveri–Sutton chromosome theory of inheritance by Thomas Hunt Morgan in 1915, they became the core of classical genetics. Ronald Fisher combined these ideas with the theory of natural selection in his 1930 book The Genetical Theory of Natural Selection, putting evolution onto a mathematical footing and forming the basis for population genetics within the modern evolutionary synthesis.[2]

Gregor Mendel, the Moravian Augustinian monk who founded the modern science of genetics

History

The principles of Mendelian inheritance were named for and first derived by Gregor Johann Mendel,[3] a nineteenth-century Moravian monk who formulated his ideas after conducting simple hybridisation experiments with pea plants (Pisum sativum) he had planted in the garden of his monastery.[4] Between 1856 and 1863, Mendel cultivated and tested some 5,000 pea plants. From these experiments, he induced two generalizations which later became known as Mendel's Principles of Heredity or Mendelian inheritance. He described his experiments in a two-part paper, Versuche über Pflanzen-Hybriden (Experiments on Plant Hybridization),[5] that he presented to the Natural History Society of Brno on 8 February and 8 March 1865, and which was published in 1866.[6][7][8][9]

Mendel's results were at first largely ignored. Although they were not completely unknown to biologists of the time, they were not seen as generally applicable, even by Mendel himself, who thought they only applied to certain categories of species or traits. A major roadblock to understanding their significance was the importance attached by 19th-century biologists to the apparent blending of many inherited traits in the overall appearance of the progeny, now known to be due to multi-gene interactions, in contrast to the organ-specific binary characters studied by Mendel.[4] In 1900, however, his work was "re-discovered" by three European scientists, Hugo de Vries, Carl Correns, and Erich von Tschermak. The exact nature of the "re-discovery" has been debated: De Vries published first on the subject, mentioning Mendel in a footnote, while Correns pointed out Mendel's priority after having read De Vries' paper and realizing that he himself did not have priority. De Vries may not have acknowledged truthfully how much of his knowledge of the laws came from his own work and how much came only after reading Mendel's paper. Later scholars have accused Von Tschermak of not truly understanding the results at all.[4][10][11][12]

Regardless, the "re-discovery" made Mendelism an important but controversial theory. Its most vigorous promoter in Europe was William Bateson, who coined the terms "genetics" and "allele" to describe many of its tenets. The model of heredity was contested by other biologists because it implied that heredity was discontinuous, in opposition to the apparently continuous variation observable for many traits. Many biologists also dismissed the theory because they were not sure it would apply to all species. However, later work by biologists and statisticians such as Ronald Fisher showed that if multiple Mendelian factors were involved in the expression of an individual trait, they could produce the diverse results observed, thus demonstrating that Mendelian genetics is compatible with natural selection. Thomas Hunt Morgan and his assistants later integrated Mendel's theoretical model with the chromosome theory of inheritance, in which the chromosomes of cells were thought to hold the actual hereditary material, and created what is now known as classical genetics, a highly successful foundation which eventually cemented Mendel's place in history.

Mendel's findings allowed scientists such as Fisher and J.B.S. Haldane to predict the expression of traits on the basis of mathematical probabilities. An important aspect of Mendel's success can be traced to his decision to start his crosses only with plants he demonstrated were true-breeding. He only measured discrete (binary) characteristics, such as color, shape, and position of the seeds, rather than quantitatively variable characteristics. He expressed his results numerically and subjected them to statistical analysis. His method of data analysis and his large sample size gave credibility to his data. He had the foresight to follow several successive generations (P, F1, F2, F3) of pea plants and record their variations. Finally, he performed "test crosses" (backcrossing descendants of the initial hybridization to the initial true-breeding lines) to reveal the presence and proportions of recessive characters.

Mendel's genetic discoveries

Five parts of Mendel's discoveries were an important divergence from the common theories at the time and were the prerequisite for the establishment of his rules.

  1. Characters are unitary, that is, they are discrete e.g.: purple vs. white, tall vs. dwarf. There is no medium-sized plant or light purple flower.
  2. Genetic characteristics have alternate forms, each inherited from one of two parents. Today these are called alleles.
  3. One allele is dominant over the other. The phenotype reflects the dominant allele.
  4. Gametes are created by random segregation. Heterozygotic individuals produce gametes with an equal frequency of the two alleles.
  5. Different traits have independent assortment. In modern terms, genes are unlinked.

According to customary terminology, the principles of inheritance discovered by Gregor Mendel are here referred to as Mendelian laws, although today's geneticists also speak of Mendelian rules or Mendelian principles,[13][14] as there are many exceptions summarized under the collective term Non-Mendelian inheritance.

 
Characteristics Mendel used in his experiments[15]
 
P-Generation and F1-Generation: The dominant allele for purple-red flower hides the phenotypic effect of the recessive allele for white flowers. F2-Generation: The recessive trait from the P-Generation phenotypically reappears in the individuals that are homozygous with the recessive genetic trait.
 
Myosotis: Colour and distribution of colours are inherited independently.[16]

Mendel selected for the experiment the following characters of pea plants:

  • Form of the ripe seeds (round or roundish, surface shallow or wrinkled)
  • Colour of the seed–coat (white, gray, or brown, with or without violet spotting)
  • Colour of the seeds and cotyledons (yellow or green)
  • Flower colour (white or violet-red)
  • Form of the ripe pods (simply inflated, not contracted, or constricted between the seeds and wrinkled)
  • Colour of the unripe pods (yellow or green)
  • Position of the flowers (axial or terminal)
  • Length of the stem [17]

When he crossed purebred white flower and purple flower pea plants (the parental or P generation) by artificial pollination, the resulting flower colour was not a blend. Rather than being a mix of the two, the offspring in the first generation (F1-generation) were all purple-flowered. Therefore, he called this biological trait dominant. When he allowed self-fertilization in the uniform looking F1-generation, he obtained both colours in the F2 generation with a purple flower to white flower ratio of 3 : 1. In some of the other characters also one of the traits was dominant.

He then conceived the idea of heredity units, which he called hereditary "factors". Mendel found that there are alternative forms of factors—now called genes—that account for variations in inherited characteristics. For example, the gene for flower color in pea plants exists in two forms, one for purple and the other for white. The alternative "forms" are now called alleles. For each trait, an organism inherits two alleles, one from each parent. These alleles may be the same or different. An organism that has two identical alleles for a gene is said to be homozygous for that gene (and is called a homozygote). An organism that has two different alleles for a gene is said be heterozygous for that gene (and is called a heterozygote).

Mendel hypothesized that allele pairs separate randomly, or segregate, from each other during the production of the gametes in the seed plant (egg cell) and the pollen plant (sperm). Because allele pairs separate during gamete production, a sperm or egg carries only one allele for each inherited trait. When sperm and egg unite at fertilization, each contributes its allele, restoring the paired condition in the offspring. Mendel also found that each pair of alleles segregates independently of the other pairs of alleles during gamete formation.

The genotype of an individual is made up of the many alleles it possesses. The phenotype is the result of the expression of all characteristics that are genetically determined by its alleles as well as by its environment. The presence of an allele does not mean that the trait will be expressed in the individual that possesses it. If the two alleles of an inherited pair differ (the heterozygous condition), then one determines the organism's appearance and is called the dominant allele; the other has no noticeable effect on the organism's appearance and is called the recessive allele.

Mendel's laws of inheritance
Law Definition
Law of dominance and uniformity Some alleles are dominant while others are recessive; an organism with at least one dominant allele will display the effect of the dominant allele.[18]
Law of segregation During gamete formation, the alleles for each gene segregate from each other so that each gamete carries only one allele for each gene.
Law of independent assortment Genes of different traits can segregate independently during the formation of gametes.

Law of Dominance and Uniformity

 
F1 generation: All individuals have the same genotype and same phenotype expressing the dominant trait (red).
F2 generation: The phenotypes in the second generation show a 3 : 1 ratio.
In the genotype 25 % are homozygous with the dominant trait, 50 % are heterozygous genetic carriers of the recessive trait, 25 % are homozygous with the recessive genetic trait and expressing the recessive character.
 
In Mirabilis jalapa and Antirrhinum majus are examples for intermediate inheritance.[19][20] As seen in the F1-generation, heterozygous plants have "light pink" flowers—a mix of "red" and "white". The F2-generation shows a 1:2:1 ratio of red : light pink : white

If two parents are mated with each other who differ in one genetic characteristic for which they are both homozygous (each pure-bred), all offspring in the first generation (F1) are equal to the examined characteristic in genotype and phenotype showing the dominant trait. This uniformity rule or reciprocity rule applies to all individuals of the F1-generation.[21]

The principle of dominant inheritance discovered by Mendel states that in a heterozygote the dominant allele will cause the recessive allele to be "masked": that is, not expressed in the phenotype. Only if an individual is homozygous with respect to the recessive allele will the recessive trait be expressed. Therefore, a cross between a homozygous dominant and a homozygous recessive organism yields a heterozygous organism whose phenotype displays only the dominant trait.

The F1 offspring of Mendel's pea crosses always looked like one of the two parental varieties. In this situation of "complete dominance," the dominant allele had the same phenotypic effect whether present in one or two copies.

But for some characteristics, the F1 hybrids have an appearance in between the phenotypes of the two parental varieties. A cross between two four o'clock (Mirabilis jalapa) plants shows an exception to Mendel's principle, called incomplete dominance. Flowers of heterozygous plants have a phenotype somewhere between the two homozygous genotypes. In cases of intermediate inheritance (incomplete dominance) in the F1-generation Mendel's principle of uniformity in genotype and phenotype applies as well. Research about intermediate inheritance was done by other scientists. The first was Carl Correns with his studies about Mirabilis jalapa.[19][22][23][24][25]

Law of Segregation of genes

 
A Punnett square for one of Mendel's pea plant experiments – self-fertilization of the F1 generation

The Law of Segregation of genes applies when two individuals, both heterozygous for a certain trait are crossed, for example hybrids of the F1-generation. The offspring in the F2-generation differ in genotype and phenotype, so that the characteristics of the grandparents (P-generation) regularly occur again. In a dominant-recessive inheritance an average of 25% are homozygous with the dominant trait, 50% are heterozygous showing the dominant trait in the phenotype (genetic carriers), 25% are homozygous with the recessive trait and therefore express the recessive trait in the phenotype. The genotypic ratio is 1 : 2 : 1, the phenotypic ratio is 3 : 1.

In the pea plant example, the capital "B" represents the dominant allele for purple blossom and lowercase "b" represents the recessive allele for white blossom. The pistil plant and the pollen plant are both F1-hybrids with genotype "B b". Each has one allele for purple and one allele for white. In the offspring, in the F2-plants in the Punnett-square, three combinations are possible. The genotypic ratio is 1 BB : 2 Bb : 1 bb. But the phenotypic ratio of plants with purple blossoms to those with white blossoms is 3 : 1 due to the dominance of the allele for purple. Plants with homozygous "b b" are white flowered like one of the grandparents in the P-generation.

In cases of incomplete dominance the same segregation of alleles takes place in the F2-generation, but here also the phenotypes show a ratio of 1 : 2 : 1, as the heterozygous are different in phenotype from the homozygous because the genetic expression of one allele compensates the missing expression of the other allele only partially. This results in an intermediate inheritance which was later described by other scientists.

In some literature sources the principle of segregation is cited as "first law". Nevertheless, Mendel did his crossing experiments with heterozygous plants after obtaining these hybrids by crossing two purebred plants, discovering the principle of dominance and uniformity first.[26][18]

Molecular proof of segregation of genes was subsequently found through observation of meiosis by two scientists independently, the German botanist Oscar Hertwig in 1876, and the Belgian zoologist Edouard Van Beneden in 1883. Most alleles are located in chromosomes in the cell nucleus. Paternal and maternal chromosomes get separated in meiosis, because during spermatogenesis the chromosomes are segregated on the four sperm cells that arise from one mother sperm cell, and during oogenesis the chromosomes are distributed between the polar bodies and the egg cell. Every individual organism contains two alleles for each trait. They segregate (separate) during meiosis such that each gamete contains only one of the alleles.[27] When the gametes unite in the zygote the alleles—one from the mother one from the father—get passed on to the offspring. An offspring thus receives a pair of alleles for a trait by inheriting homologous chromosomes from the parent organisms: one allele for each trait from each parent.[27] Heterozygous individuals with the dominant trait in the phenotype are genetic carriers of the recessive trait.

Law of Independent Assortment

 
Segregation and independent assortment are consistent with the chromosome theory of inheritance.
 
When the parents are homozygous for two different genetic traits (llSS and LL sP sP), their children in the F1 generation are heterozygous at both loci and only show the dominant phenotypes (Ll S sP). P-Generation: Each parent possesses one dominant and one recessive trait purebred (homozygous). In this example, solid coat color is indicated by S (dominant), Piebald spotting by sP (recessive), while fur length is indicated by L (short, dominant) or l (long, recessive). All individuals are equal in genotype and phenotype. In the F2 generation all combinations of coat color and fur length occur: 9 are short haired with solid colour, 3 are short haired with spotting, 3 are long haired with solid colour and 1 is long haired with spotting. The traits are inherited independently, so that new combinations can occur. Average number ratio of phenotypes 9:3:3:1[28]
 
For example 3 pairs of homologous chromosomes allow 8 possible combinations, all equally likely to move into the gamete during meiosis. This is the main reason for independent assortment. The equation to determine the number of possible combinations given the number of homologous pairs = 2x (x = number of homologous pairs)

The Law of Independent Assortment states that alleles for separate traits are passed independently of one another.[29][26] That is, the biological selection of an allele for one trait has nothing to do with the selection of an allele for any other trait. Mendel found support for this law in his dihybrid cross experiments. In his monohybrid crosses, an idealized 3:1 ratio between dominant and recessive phenotypes resulted. In dihybrid crosses, however, he found a 9:3:3:1 ratios. This shows that each of the two alleles is inherited independently from the other, with a 3:1 phenotypic ratio for each.

Independent assortment occurs in eukaryotic organisms during meiotic metaphase I, and produces a gamete with a mixture of the organism's chromosomes. The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent chromosome along the metaphase plate with respect to the other bivalent chromosomes. Along with crossing over, independent assortment increases genetic diversity by producing novel genetic combinations.

There are many deviations from the principle of independent assortment due to genetic linkage.

Of the 46 chromosomes in a normal diploid human cell, half are maternally derived (from the mother's egg) and half are paternally derived (from the father's sperm). This occurs as sexual reproduction involves the fusion of two haploid gametes (the egg and sperm) to produce a zygote and a new organism, in which every cell has two sets of chromosomes (diploid). During gametogenesis the normal complement of 46 chromosomes needs to be halved to 23 to ensure that the resulting haploid gamete can join with another haploid gamete to produce a diploid organism.

In independent assortment, the chromosomes that result are randomly sorted from all possible maternal and paternal chromosomes. Because zygotes end up with a mix instead of a pre-defined "set" from either parent, chromosomes are therefore considered assorted independently. As such, the zygote can end up with any combination of paternal or maternal chromosomes. For human gametes, with 23 chromosomes, the number of possibilities is 223 or 8,388,608 possible combinations.[30] This contributes to the genetic variability of progeny. Generally, the recombination of genes has important implications for many evolutionary processes.[31][32][33]

Mendelian trait

A Mendelian trait is one whose inheritance follows Mendel’s principles—namely, the trait depends only on a single locus, whose alleles are either dominant or recessive.

Many traits are inherited in a non-Mendelian fashion.[34]

Non-Mendelian inheritance

Mendel himself warned that care was needed in extrapolating his patterns to other organisms or traits. Indeed, many organisms have traits whose inheritance works differently from the principles he described; these traits are called non-Mendelian.[35][36]

For example, Mendel focused on traits whose genes have only two alleles, such as "A" and "a". However, many genes have more than two alleles. He also focused on traits determined by a single gene. But some traits, such as height, depend on many genes rather than just one. Traits dependent on multiple genes are called polygenic traits.

See also

References

  1. ^ William Bateson: Mendel's Principles of Heredity - A Defence, with a Translation of Mendel's Original Papers on Hybridisation Cambridge University Press 2009, ISBN 978-1-108-00613-2
  2. ^ Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way We Think. New York, New York: Oxford University Press. p. 69. ISBN 978-0-19-929116-8.
  3. ^ Ilona Miko: Gregor Mendel and the Principles of Inheritance Nature Education 2008
  4. ^ a b c Henig, Robin Marantz (2009). The Monk in the Garden : The Lost and Found Genius of Gregor Mendel, the Father of Modern Genetics. Houghton Mifflin. ISBN 978-0-395-97765-1. The article, written by an Austrian monk named Gregor Johann Mendel...
  5. ^ Gregor Mendel: Experiments in Plant Hybridization 1965
  6. ^ National Center for Biotechnology Information: Mendel's experiments U.S. National Library of Medicine, 2000
  7. ^ Mendel's paper in English: Gregor Mendel (1865). "Experiments in Plant Hybridization".
  8. ^ J. G. Mendel: Versuche über Pflanzenhybriden. In: Verhandlungen des Naturforschenden Vereins Brünn. Band 4, 1865, S. 3–47.
  9. ^ Uwe Hoßfeld, Michael V. Simunek: 150 Jahre Mendels Vortrag „Versuche über Pflanzen-Hybriden“. 2015, S. 238
  10. ^ Ernst Mayr: The Growth of Biological Thought, Belknap Press, S. 730 (1982)
  11. ^ Floyd Monaghan, Alain Corcos: Journal of Heredity 77: 468f (1986) und 78: 208-210 (1987
  12. ^ Michal Simunek, Uwe Hoßfeld, Florian Thümmler, Olaf Breidbach (Hg.): The Mendelian Dioskuri – Correspondence of Armin with Erich von Tschermak-Seysenegg, 1898–1951; „Studies in the History of Sciences and Humanities“, Band Nr. 27; Prag 2011; ISBN 978-80-87378-67-0.
  13. ^ Science Learning Hub: Mendel's principles of inheritance
  14. ^ Noel Clarke: Mendelian Genetics - An overview
  15. ^ Gregor Mendel: Versuche über Pflanzenhybriden Verhandlungen des Naturforschenden Vereines in Brünn. Bd. IV. 1866, page 8
  16. ^ Write Work: Mendel's Impact
  17. ^ Gregor Mendel: Experiments in Plant Hybridization 1965, page 5
  18. ^ a b Rutgers: Mendelian Principles
  19. ^ a b Biology University of Hamburg: Mendelian Genetics
  20. ^ Neil A. Campbell, Jane B. Reece: Biologie. Spektrum-Verlag Heidelberg-Berlin 2003, ISBN 3-8274-1352-4, page 302-303.
  21. ^ Ulrich Weber: Biologie Gesamtband Oberstufe, 1st edition, Cornelsen Verlag Berlin 2001, ISBN 3-464-04279-0, page 170 - 171.
  22. ^ Biologie Schule - kompaktes Wissen: Uniformitätsregel (1. Mendelsche Regel)
  23. ^ Frustfrei Lernen: Uniformitätsregel (1. Mendelsche Regel)
  24. ^ Spektrum Biologie: Unvollständige Dominanz
  25. ^ Spektrum Biologie: Intermediärer Erbgang
  26. ^ a b Neil A. Campbell, Jane B. Reece: Biologie. Spektrum-Verlag 2003, page 293-315. ISBN 3-8274-1352-4
  27. ^ a b Bailey, Regina (5 November 2015). "Mendel's Law of Segregation". about education. About.com. Retrieved 2 February 2016.
  28. ^ Spectrum Dictionary of Biology Mendel Rules
  29. ^ Bailey, Regina. "Independent Assortment". Thoughtco. About.com. Retrieved 24 February 2016.
  30. ^ Perez, Nancy. "Meiosis". Retrieved 15 February 2007.
  31. ^ Stapley, J.; Feulner, P. G.; Johnston, S. E.; Santure, A. W.; Smadja, C. M. (2017). "Recombination: The good, the bad and the variable". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 372 (1736). doi:10.1098/rstb.2017.0279. PMC 5698631. PMID 29109232.
  32. ^ Reeve, James; Ortiz-Barrientos, Daniel; Engelstädter, Jan (2016). "The evolution of recombination rates in finite populations during ecological speciation". Proceedings of the Royal Society B: Biological Sciences. 283 (1841). doi:10.1098/rspb.2016.1243. PMC 5095376. PMID 27798297.
  33. ^ Hickey, Donal A.; Golding, G. Brian (2018). "The advantage of recombination when selection is acting at many genetic Loci". Journal of Theoretical Biology. 442: 123–128. Bibcode:2018JThBi.442..123H. doi:10.1016/j.jtbi.2018.01.018. PMID 29355539.
  34. ^ "Genetic Disorders". National Human Genome Research Institute. 18 May 2018.
  35. ^ Schacherer, Joseph (2016). "Beyond the simplicity of Mendelian inheritance". Comptes Rendus Biologies. 339 (7–8): 284–288. doi:10.1016/j.crvi.2016.04.006. PMID 27344551.
  36. ^ Khan Academy: Variations on Mendel's laws (overview)

Notes

  • Bowler, Peter J. (1989). The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. Johns Hopkins University Press.
  • Atics, Jean. Genetics: The life of DNA. ANDRNA press.
  • Reece, Jane B.; Campbell, Neil A. (2011). Mendel and the Gene Idea. Campbell Biology (9th ed.). Benjamin Cummings / Pearson Education. p. 265.

External links

  • Khan Academy, video lecture
  • Probability of Inheritance
  • Mendel's principles of Inheritance
  • Mendelian genetics

mendelian, inheritance, technical, introduction, topic, introduction, genetics, this, article, needs, additional, citations, verification, please, help, improve, this, article, adding, citations, reliable, sources, unsourced, material, challenged, removed, fin. For a non technical introduction to the topic see Introduction to genetics This article needs additional citations for verification Please help improve this article by adding citations to reliable sources Unsourced material may be challenged and removed Find sources Mendelian inheritance news newspapers books scholar JSTOR December 2017 Learn how and when to remove this template message Mendelian inheritance also known as Mendelism is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866 re discovered in 1900 by Hugo de Vries and Carl Correns and later popularized by William Bateson 1 These principles were initially controversial When Mendel s theories were integrated with the Boveri Sutton chromosome theory of inheritance by Thomas Hunt Morgan in 1915 they became the core of classical genetics Ronald Fisher combined these ideas with the theory of natural selection in his 1930 book The Genetical Theory of Natural Selection putting evolution onto a mathematical footing and forming the basis for population genetics within the modern evolutionary synthesis 2 Gregor Mendel the Moravian Augustinian monk who founded the modern science of genetics Contents 1 History 2 Mendel s genetic discoveries 2 1 Law of Dominance and Uniformity 2 2 Law of Segregation of genes 2 3 Law of Independent Assortment 3 Mendelian trait 4 Non Mendelian inheritance 5 See also 6 References 7 Notes 8 External linksHistory EditMain article History of genetics The principles of Mendelian inheritance were named for and first derived by Gregor Johann Mendel 3 a nineteenth century Moravian monk who formulated his ideas after conducting simple hybridisation experiments with pea plants Pisum sativum he had planted in the garden of his monastery 4 Between 1856 and 1863 Mendel cultivated and tested some 5 000 pea plants From these experiments he induced two generalizations which later became known as Mendel s Principles of Heredity or Mendelian inheritance He described his experiments in a two part paper Versuche uber Pflanzen Hybriden Experiments on Plant Hybridization 5 that he presented to the Natural History Society of Brno on 8 February and 8 March 1865 and which was published in 1866 6 7 8 9 Mendel s results were at first largely ignored Although they were not completely unknown to biologists of the time they were not seen as generally applicable even by Mendel himself who thought they only applied to certain categories of species or traits A major roadblock to understanding their significance was the importance attached by 19th century biologists to the apparent blending of many inherited traits in the overall appearance of the progeny now known to be due to multi gene interactions in contrast to the organ specific binary characters studied by Mendel 4 In 1900 however his work was re discovered by three European scientists Hugo de Vries Carl Correns and Erich von Tschermak The exact nature of the re discovery has been debated De Vries published first on the subject mentioning Mendel in a footnote while Correns pointed out Mendel s priority after having read De Vries paper and realizing that he himself did not have priority De Vries may not have acknowledged truthfully how much of his knowledge of the laws came from his own work and how much came only after reading Mendel s paper Later scholars have accused Von Tschermak of not truly understanding the results at all 4 10 11 12 Regardless the re discovery made Mendelism an important but controversial theory Its most vigorous promoter in Europe was William Bateson who coined the terms genetics and allele to describe many of its tenets The model of heredity was contested by other biologists because it implied that heredity was discontinuous in opposition to the apparently continuous variation observable for many traits Many biologists also dismissed the theory because they were not sure it would apply to all species However later work by biologists and statisticians such as Ronald Fisher showed that if multiple Mendelian factors were involved in the expression of an individual trait they could produce the diverse results observed thus demonstrating that Mendelian genetics is compatible with natural selection Thomas Hunt Morgan and his assistants later integrated Mendel s theoretical model with the chromosome theory of inheritance in which the chromosomes of cells were thought to hold the actual hereditary material and created what is now known as classical genetics a highly successful foundation which eventually cemented Mendel s place in history Mendel s findings allowed scientists such as Fisher and J B S Haldane to predict the expression of traits on the basis of mathematical probabilities An important aspect of Mendel s success can be traced to his decision to start his crosses only with plants he demonstrated were true breeding He only measured discrete binary characteristics such as color shape and position of the seeds rather than quantitatively variable characteristics He expressed his results numerically and subjected them to statistical analysis His method of data analysis and his large sample size gave credibility to his data He had the foresight to follow several successive generations P F1 F2 F3 of pea plants and record their variations Finally he performed test crosses backcrossing descendants of the initial hybridization to the initial true breeding lines to reveal the presence and proportions of recessive characters Mendel s genetic discoveries EditFive parts of Mendel s discoveries were an important divergence from the common theories at the time and were the prerequisite for the establishment of his rules Characters are unitary that is they are discrete e g purple vs white tall vs dwarf There is no medium sized plant or light purple flower Genetic characteristics have alternate forms each inherited from one of two parents Today these are called alleles One allele is dominant over the other The phenotype reflects the dominant allele Gametes are created by random segregation Heterozygotic individuals produce gametes with an equal frequency of the two alleles Different traits have independent assortment In modern terms genes are unlinked According to customary terminology the principles of inheritance discovered by Gregor Mendel are here referred to as Mendelian laws although today s geneticists also speak of Mendelian rules or Mendelian principles 13 14 as there are many exceptions summarized under the collective term Non Mendelian inheritance Characteristics Mendel used in his experiments 15 P Generation and F1 Generation The dominant allele for purple red flower hides the phenotypic effect of the recessive allele for white flowers F2 Generation The recessive trait from the P Generation phenotypically reappears in the individuals that are homozygous with the recessive genetic trait Myosotis Colour and distribution of colours are inherited independently 16 Mendel selected for the experiment the following characters of pea plants Form of the ripe seeds round or roundish surface shallow or wrinkled Colour of the seed coat white gray or brown with or without violet spotting Colour of the seeds and cotyledons yellow or green Flower colour white or violet red Form of the ripe pods simply inflated not contracted or constricted between the seeds and wrinkled Colour of the unripe pods yellow or green Position of the flowers axial or terminal Length of the stem 17 When he crossed purebred white flower and purple flower pea plants the parental or P generation by artificial pollination the resulting flower colour was not a blend Rather than being a mix of the two the offspring in the first generation F1 generation were all purple flowered Therefore he called this biological trait dominant When he allowed self fertilization in the uniform looking F1 generation he obtained both colours in the F2 generation with a purple flower to white flower ratio of 3 1 In some of the other characters also one of the traits was dominant He then conceived the idea of heredity units which he called hereditary factors Mendel found that there are alternative forms of factors now called genes that account for variations in inherited characteristics For example the gene for flower color in pea plants exists in two forms one for purple and the other for white The alternative forms are now called alleles For each trait an organism inherits two alleles one from each parent These alleles may be the same or different An organism that has two identical alleles for a gene is said to be homozygous for that gene and is called a homozygote An organism that has two different alleles for a gene is said be heterozygous for that gene and is called a heterozygote Mendel hypothesized that allele pairs separate randomly or segregate from each other during the production of the gametes in the seed plant egg cell and the pollen plant sperm Because allele pairs separate during gamete production a sperm or egg carries only one allele for each inherited trait When sperm and egg unite at fertilization each contributes its allele restoring the paired condition in the offspring Mendel also found that each pair of alleles segregates independently of the other pairs of alleles during gamete formation The genotype of an individual is made up of the many alleles it possesses The phenotype is the result of the expression of all characteristics that are genetically determined by its alleles as well as by its environment The presence of an allele does not mean that the trait will be expressed in the individual that possesses it If the two alleles of an inherited pair differ the heterozygous condition then one determines the organism s appearance and is called the dominant allele the other has no noticeable effect on the organism s appearance and is called the recessive allele Mendel s laws of inheritance Law DefinitionLaw of dominance and uniformity Some alleles are dominant while others are recessive an organism with at least one dominant allele will display the effect of the dominant allele 18 Law of segregation During gamete formation the alleles for each gene segregate from each other so that each gamete carries only one allele for each gene Law of independent assortment Genes of different traits can segregate independently during the formation of gametes Law of Dominance and Uniformity Edit F1 generation All individuals have the same genotype and same phenotype expressing the dominant trait red F2 generation The phenotypes in the second generation show a 3 1 ratio In the genotype 25 are homozygous with the dominant trait 50 are heterozygous genetic carriers of the recessive trait 25 are homozygous with the recessive genetic trait and expressing the recessive character In Mirabilis jalapa and Antirrhinum majus are examples for intermediate inheritance 19 20 As seen in the F1 generation heterozygous plants have light pink flowers a mix of red and white The F2 generation shows a 1 2 1 ratio of red light pink white If two parents are mated with each other who differ in one genetic characteristic for which they are both homozygous each pure bred all offspring in the first generation F1 are equal to the examined characteristic in genotype and phenotype showing the dominant trait This uniformity rule or reciprocity rule applies to all individuals of the F1 generation 21 The principle of dominant inheritance discovered by Mendel states that in a heterozygote the dominant allele will cause the recessive allele to be masked that is not expressed in the phenotype Only if an individual is homozygous with respect to the recessive allele will the recessive trait be expressed Therefore a cross between a homozygous dominant and a homozygous recessive organism yields a heterozygous organism whose phenotype displays only the dominant trait The F1 offspring of Mendel s pea crosses always looked like one of the two parental varieties In this situation of complete dominance the dominant allele had the same phenotypic effect whether present in one or two copies But for some characteristics the F1 hybrids have an appearance in between the phenotypes of the two parental varieties A cross between two four o clock Mirabilis jalapa plants shows an exception to Mendel s principle called incomplete dominance Flowers of heterozygous plants have a phenotype somewhere between the two homozygous genotypes In cases of intermediate inheritance incomplete dominance in the F1 generation Mendel s principle of uniformity in genotype and phenotype applies as well Research about intermediate inheritance was done by other scientists The first was Carl Correns with his studies about Mirabilis jalapa 19 22 23 24 25 Law of Segregation of genes Edit A Punnett square for one of Mendel s pea plant experiments self fertilization of the F1 generation The Law of Segregation of genes applies when two individuals both heterozygous for a certain trait are crossed for example hybrids of the F1 generation The offspring in the F2 generation differ in genotype and phenotype so that the characteristics of the grandparents P generation regularly occur again In a dominant recessive inheritance an average of 25 are homozygous with the dominant trait 50 are heterozygous showing the dominant trait in the phenotype genetic carriers 25 are homozygous with the recessive trait and therefore express the recessive trait in the phenotype The genotypic ratio is 1 2 1 the phenotypic ratio is 3 1 In the pea plant example the capital B represents the dominant allele for purple blossom and lowercase b represents the recessive allele for white blossom The pistil plant and the pollen plant are both F1 hybrids with genotype B b Each has one allele for purple and one allele for white In the offspring in the F2 plants in the Punnett square three combinations are possible The genotypic ratio is 1 BB 2 Bb 1 bb But the phenotypic ratio of plants with purple blossoms to those with white blossoms is 3 1 due to the dominance of the allele for purple Plants with homozygous b b are white flowered like one of the grandparents in the P generation In cases of incomplete dominance the same segregation of alleles takes place in the F2 generation but here also the phenotypes show a ratio of 1 2 1 as the heterozygous are different in phenotype from the homozygous because the genetic expression of one allele compensates the missing expression of the other allele only partially This results in an intermediate inheritance which was later described by other scientists In some literature sources the principle of segregation is cited as first law Nevertheless Mendel did his crossing experiments with heterozygous plants after obtaining these hybrids by crossing two purebred plants discovering the principle of dominance and uniformity first 26 18 Molecular proof of segregation of genes was subsequently found through observation of meiosis by two scientists independently the German botanist Oscar Hertwig in 1876 and the Belgian zoologist Edouard Van Beneden in 1883 Most alleles are located in chromosomes in the cell nucleus Paternal and maternal chromosomes get separated in meiosis because during spermatogenesis the chromosomes are segregated on the four sperm cells that arise from one mother sperm cell and during oogenesis the chromosomes are distributed between the polar bodies and the egg cell Every individual organism contains two alleles for each trait They segregate separate during meiosis such that each gamete contains only one of the alleles 27 When the gametes unite in the zygote the alleles one from the mother one from the father get passed on to the offspring An offspring thus receives a pair of alleles for a trait by inheriting homologous chromosomes from the parent organisms one allele for each trait from each parent 27 Heterozygous individuals with the dominant trait in the phenotype are genetic carriers of the recessive trait Law of Independent Assortment Edit Segregation and independent assortment are consistent with the chromosome theory of inheritance When the parents are homozygous for two different genetic traits llSS and LL sP sP their children in the F1 generation are heterozygous at both loci and only show the dominant phenotypes Ll S sP P Generation Each parent possesses one dominant and one recessive trait purebred homozygous In this example solid coat color is indicated by S dominant Piebald spotting by sP recessive while fur length is indicated by L short dominant or l long recessive All individuals are equal in genotype and phenotype In the F2 generation all combinations of coat color and fur length occur 9 are short haired with solid colour 3 are short haired with spotting 3 are long haired with solid colour and 1 is long haired with spotting The traits are inherited independently so that new combinations can occur Average number ratio of phenotypes 9 3 3 1 28 For example 3 pairs of homologous chromosomes allow 8 possible combinations all equally likely to move into the gamete during meiosis This is the main reason for independent assortment The equation to determine the number of possible combinations given the number of homologous pairs 2x x number of homologous pairs The Law of Independent Assortment states that alleles for separate traits are passed independently of one another 29 26 That is the biological selection of an allele for one trait has nothing to do with the selection of an allele for any other trait Mendel found support for this law in his dihybrid cross experiments In his monohybrid crosses an idealized 3 1 ratio between dominant and recessive phenotypes resulted In dihybrid crosses however he found a 9 3 3 1 ratios This shows that each of the two alleles is inherited independently from the other with a 3 1 phenotypic ratio for each Independent assortment occurs in eukaryotic organisms during meiotic metaphase I and produces a gamete with a mixture of the organism s chromosomes The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent chromosome along the metaphase plate with respect to the other bivalent chromosomes Along with crossing over independent assortment increases genetic diversity by producing novel genetic combinations There are many deviations from the principle of independent assortment due to genetic linkage Of the 46 chromosomes in a normal diploid human cell half are maternally derived from the mother s egg and half are paternally derived from the father s sperm This occurs as sexual reproduction involves the fusion of two haploid gametes the egg and sperm to produce a zygote and a new organism in which every cell has two sets of chromosomes diploid During gametogenesis the normal complement of 46 chromosomes needs to be halved to 23 to ensure that the resulting haploid gamete can join with another haploid gamete to produce a diploid organism In independent assortment the chromosomes that result are randomly sorted from all possible maternal and paternal chromosomes Because zygotes end up with a mix instead of a pre defined set from either parent chromosomes are therefore considered assorted independently As such the zygote can end up with any combination of paternal or maternal chromosomes For human gametes with 23 chromosomes the number of possibilities is 223 or 8 388 608 possible combinations 30 This contributes to the genetic variability of progeny Generally the recombination of genes has important implications for many evolutionary processes 31 32 33 Mendelian trait EditA Mendelian trait is one whose inheritance follows Mendel s principles namely the trait depends only on a single locus whose alleles are either dominant or recessive Many traits are inherited in a non Mendelian fashion 34 Non Mendelian inheritance EditMain article Non Mendelian inheritance Mendel himself warned that care was needed in extrapolating his patterns to other organisms or traits Indeed many organisms have traits whose inheritance works differently from the principles he described these traits are called non Mendelian 35 36 For example Mendel focused on traits whose genes have only two alleles such as A and a However many genes have more than two alleles He also focused on traits determined by a single gene But some traits such as height depend on many genes rather than just one Traits dependent on multiple genes are called polygenic traits See also Edit History of Science portal Biology portalList of Mendelian traits in humans Mendelian diseases monogenic disease Mendelian error Particulate inheritance Punnett squareReferences Edit William Bateson Mendel s Principles of Heredity A Defence with a Translation of Mendel s Original Papers on Hybridisation Cambridge University Press 2009 ISBN 978 1 108 00613 2 Grafen Alan Ridley Mark 2006 Richard Dawkins How A Scientist Changed the Way We Think New York New York Oxford University Press p 69 ISBN 978 0 19 929116 8 Ilona Miko Gregor Mendel and the Principles of Inheritance Nature Education 2008 a b c Henig Robin Marantz 2009 The Monk in the Garden The Lost and Found Genius of Gregor Mendel the Father of Modern Genetics Houghton Mifflin ISBN 978 0 395 97765 1 The article written by an Austrian monk named Gregor Johann Mendel Gregor Mendel Experiments in Plant Hybridization 1965 National Center for Biotechnology Information Mendel s experiments U S National Library of Medicine 2000 Mendel s paper in English Gregor Mendel 1865 Experiments in Plant Hybridization J G Mendel Versuche uber Pflanzenhybriden In Verhandlungen des Naturforschenden Vereins Brunn Band 4 1865 S 3 47 Uwe Hossfeld Michael V Simunek 150 Jahre Mendels Vortrag Versuche uber Pflanzen Hybriden 2015 S 238 Ernst Mayr The Growth of Biological Thought Belknap Press S 730 1982 Floyd Monaghan Alain Corcos Abstract Tschermak A non discoverer of MendelismJournal of Heredity 77 468f 1986 und 78 208 210 1987 Michal Simunek Uwe Hossfeld Florian Thummler Olaf Breidbach Hg The Mendelian Dioskuri Correspondence of Armin with Erich von Tschermak Seysenegg 1898 1951 Studies in the History of Sciences and Humanities Band Nr 27 Prag 2011 ISBN 978 80 87378 67 0 Science Learning Hub Mendel s principles of inheritance Noel Clarke Mendelian Genetics An overview Gregor Mendel Versuche uber Pflanzenhybriden Verhandlungen des Naturforschenden Vereines in Brunn Bd IV 1866 page 8 Write Work Mendel s Impact Gregor Mendel Experiments in Plant Hybridization 1965 page 5 a b Rutgers Mendelian Principles a b Biology University of Hamburg Mendelian Genetics Neil A Campbell Jane B Reece Biologie Spektrum Verlag Heidelberg Berlin 2003 ISBN 3 8274 1352 4 page 302 303 Ulrich Weber Biologie Gesamtband Oberstufe 1st edition Cornelsen Verlag Berlin 2001 ISBN 3 464 04279 0 page 170 171 Biologie Schule kompaktes Wissen Uniformitatsregel 1 Mendelsche Regel Frustfrei Lernen Uniformitatsregel 1 Mendelsche Regel Spektrum Biologie Unvollstandige Dominanz Spektrum Biologie Intermediarer Erbgang a b Neil A Campbell Jane B Reece Biologie Spektrum Verlag 2003 page 293 315 ISBN 3 8274 1352 4 a b Bailey Regina 5 November 2015 Mendel s Law of Segregation about education About com Retrieved 2 February 2016 Spectrum Dictionary of Biology Mendel Rules Bailey Regina Independent Assortment Thoughtco About com Retrieved 24 February 2016 Perez Nancy Meiosis Retrieved 15 February 2007 Stapley J Feulner P G Johnston S E Santure A W Smadja C M 2017 Recombination The good the bad and the variable Philosophical Transactions of the Royal Society of London Series B Biological Sciences 372 1736 doi 10 1098 rstb 2017 0279 PMC 5698631 PMID 29109232 Reeve James Ortiz Barrientos Daniel Engelstadter Jan 2016 The evolution of recombination rates in finite populations during ecological speciation Proceedings of the Royal Society B Biological Sciences 283 1841 doi 10 1098 rspb 2016 1243 PMC 5095376 PMID 27798297 Hickey Donal A Golding G Brian 2018 The advantage of recombination when selection is acting at many genetic Loci Journal of Theoretical Biology 442 123 128 Bibcode 2018JThBi 442 123H doi 10 1016 j jtbi 2018 01 018 PMID 29355539 Genetic Disorders National Human Genome Research Institute 18 May 2018 Schacherer Joseph 2016 Beyond the simplicity of Mendelian inheritance Comptes Rendus Biologies 339 7 8 284 288 doi 10 1016 j crvi 2016 04 006 PMID 27344551 Khan Academy Variations on Mendel s laws overview Notes EditBowler Peter J 1989 The Mendelian Revolution The Emergence of Hereditarian Concepts in Modern Science and Society Johns Hopkins University Press Atics Jean Genetics The life of DNA ANDRNA press Reece Jane B Campbell Neil A 2011 Mendel and the Gene Idea Campbell Biology 9th ed Benjamin Cummings Pearson Education p 265 External links EditKhan Academy video lecture Probability of Inheritance Mendel s principles of Inheritance Mendelian genetics Retrieved from https en wikipedia org w index php title Mendelian inheritance amp oldid 1124627870 Law of Independent Assortment, wikipedia, wiki, book, books, library,

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