Editing Experiments on Plant Hybridization

{{Short description|1865 article by Gregor Mendel}}
{{Use dmy dates|date=March 2019}}
 
"'''Experiments on Plant Hybridization'''" ({{Langx|de|Versuche über Pflanzen-Hybriden}}) is a seminal paper written in 1865 and published in 1866<ref>Mendel, J. G. (1866). "Versuche über Pflanzenhybriden", ''Verhandlungen des naturforschenden Vereines in Brünn'', Bd. IV für das Jahr, 1865, ''Abhandlungen'': 3–47, [https://www.biodiversitylibrary.org/item/124139#page/133/mode/1up]. For the English translation, see: {{cite journal |last1=Druery |first1=C. T. |last2=Bateson |first2=William |year=1901 |title=Experiments in plant hybridization |url=http://www.esp.org/foundations/genetics/classical/gm-65.pdf |journal=Journal of the Royal Horticultural Society |volume=26 |pages=1–32 |access-date=9 October 2009}}</ref><ref>"Mendel's paper "Versuche über Pflanzenhybriden", was read at meetings of the Brunn Natural History Society on 8th February and 8th March 1865, and was published in the ''Verhandlungen des naturforschenden Vereines in Brünn'', '''4''', 1865, which appeared in 1866" (Gregor Mendel, ''Experiments in Plant Hybridisation'', Cosimo, Inc., 2008, p. 7).</ref> by [[Gregor Mendel]], an Augustinian friar considered to be the founder of modern [[genetics]]. The paper was the result after years spent studying genetic traits in ''[[Pea|Pisum sativum]]'', the pea plant.

==Content==
In his paper, Mendel compared 7 pairs of discrete traits found in a pea plant:

{| class="wikitable" style="text-align: center; "
|-
! '''Characteristics''' !! Contrasting traits !! Offspring traits
|-
| Flower Color || Violet and White || Violet
|-
| Flower Position || Axial and Terminal || Axial
|-
| Plant Height || Tall and Dwarf || Tall
|-
| Seed Texture || Round and Wrinkled || Round
|-
| Seed Color || Green and Yellow || Yellow
|-
| Pea Pod Texture || Inflated and Constricted || Inflated
|-
| Pea Pod Color || Green and Yellow || Green
|}

Through experimentation, Mendel discovered that one inheritable trait would invariably be dominant to its recessive alternative. Mendel laid out the genetic model later known as '''[[Mendelian inheritance]]''' or Mendelian genetics. This model provided an alternative to blending inheritance, which was the prevailing theory at the time.

==History==
Mendel read his paper to the Natural History Society of [[Brno|Brünn]]. It was published in the ''[[Proceedings of the Natural History Society of Brünn]]'' the following year.

Mendel's work received little attention from the scientific community and was largely forgotten. It was not until the early 20th century that Mendel's work was rediscovered and his ideas used to help form the [[Modern synthesis (20th century)|modern synthesis]].

Mendel had read a 1863 German translation of Darwin's ''[[On the Origin of Species]]'' (translated by [[Heinrich Georg Bronn|H.G. Bronn]]), and used certain terms from the translation in the final two (10th and 11th) sections of his paper.<ref name=":1" />

There were three main English translations. The first was done in 1901, commissioned by [[William Bateson]] to the [[Royal Horticultural Society|Royal Horticultural Society of London]]. It was mainly done by Charles Thomas Druery, a British poet, author, and botanist. The second was done in 1966 by Curt Stern and Eva Sherwood. The third was done in 2016, with a careful adherence to the Darwinian terminologies that had been used by Mendel, and released to the public domain.<ref name=":1">{{Cite journal |last=Fairbanks |first=Daniel J. |last2=Abbott |first2=Scott |date=October 2016 |title=Darwin's Influence on Mendel: Evidence from a New Translation of Mendel's Paper |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC5068835/ |journal=Genetics |volume=204 |issue=2 |pages=401–405 |doi=10.1534/genetics.116.194613 |issn=1943-2631 |pmc=5068835 |pmid=27729491}}</ref><ref>{{Cite journal |last=Abbott |first=Scott |last2=Fairbanks |first2=Daniel J |date=2016-10-01 |title=Experiments on Plant Hybrids by Gregor Mendel |url=https://academic.oup.com/genetics/article/204/2/407/6072056 |journal=Genetics |volume=204 |issue=2 |pages=407–422 |doi=10.1534/genetics.116.195198 |issn=1943-2631|pmc=5068836 }}</ref>

==Analysis==
In 1936, the statistician [[Ronald Fisher]] used a [[Pearson's chi-squared test]] to analyze Mendel's data and concluded that Mendel's results with the predicted ratios were far too perfect, suggesting that adjustments (intentional or unconscious) had been made to the data to make the observations fit the hypothesis.<ref>{{Cite journal |first1=R. A. |title=Has Mendel's work been rediscovered? |journal=Annals of Science |last1=Fisher |volume=1 |issue=2 |pages=115–126 |year=1936 | doi=10.1080/00033793600200111|hdl=2440/15123 |hdl-access=free }}</ref>

Later authors have suggested Fisher's analysis was flawed, proposing various statistical and botanical explanations for Mendel's numbers.<ref>{{cite book |last=Sturtevant |first=A. H. |title=A History of Genetics |year=2001 |publisher=Cold Springs Harbor Laboratory Press |location=Cold Springs Harbor, New York |isbn=0-87969-607-9 |pages=[https://archive.org/details/historyofgenetic00stur/page/13 13–16] |url-access=registration |url=https://archive.org/details/historyofgenetic00stur/page/13 }}</ref> It is also possible that Mendel's results are "too good" merely because he reported the best subset of his data&mdash;Mendel mentioned in his paper that the data were from a subset of his experiments.

Modern geneticists have inferred the 7 genes studied by Mendel. It is impossible to know for certain, but the identification is possible to a high degree of confidence based on Mendel's description, and the pea varieties grown in central Europe in the 1850s.<ref name=":0">{{Cite journal |last1=Reid |first1=James B. |last2=Ross |first2=John J. |date=September 2011 |title=Mendel's Genes: Toward a Full Molecular Characterization |journal=Genetics |volume=189 |issue=1 |pages=3–10 |doi=10.1534/genetics.111.132118 |issn=0016-6731 |pmc=3176118 |pmid=21908742}}</ref> The table shows that the 7 genes appeared on 5 chromosomes. Of these, the only pair with significant linkage are ''V'' and ''LE'', who are 12.6 [[Centimorgan|map units]] apart. The other pair, ''R'' and ''GP'', are very weakly linked. The effect is that Mendel was unlikely to have encountered genetic linkage. In any case, he did not report dihybrid experiments on either of these pairs, and only reported on the unlinked pairs, and he always found the ratio to be 9:3:3:1.<ref>{{Cite book |last1=Meneely |first1=Philip Mark |url=https://www.worldcat.org/title/951645141 |title=Genetics: genes, genomes, and evolution |last2=Dawes Hoang |first2=Rachel |last3=Okeke |first3=Iruka N. |last4=Heston |first4=Katherine |date=2017 |publisher=Oxford University Press |isbn=978-0-19-879536-0 |location=Oxford |pages=349 |oclc=951645141}}</ref>

{| class="wikitable"
|+ Genes involved in seven pea traits studied by Mendel<ref name=":0" />
! Trait !! Dominant phenotype !! Recessive phenotype !! Symbol group !! Linkage group !! Cloned? !!           Gene function !! Molecular nature of mutation
|-
| Seed shape             ||        Round       ||       Wrinkled      ||      ''R''     ||       V       ||   Yes  ||     Starch branching enzyme 1     ||      | 0.8-kb insertion       
|-
| Stem length            ||        Tall        ||        Dwarf        ||     ''LE''     ||      III      ||   Yes  ||           GA 3-oxidase1           ||    |  G-to-A substitution     
|-
| Cotyledon color        ||       Yellow       ||        Green        ||      ''I''     ||       I       ||   Yes  ||          Stay-green gene          ||     |   6-bp insertion        
|-
| Seed coat/flower color ||       Purple       ||        White        ||      ''A''     ||       II      ||   Yes  ||     bHLH transcription factor     ||  |   G-to-A at splice site    
|-
| Pod color              ||        Green       ||        Yellow       ||     ''GP''     ||       V       ||   No   || Chloroplast structure in pod wall ||     |       Unknown           
|-
| Pod form               ||      Inflated      ||     Constricted     ||     ''V''     ||      III      ||   No   ||   Sclerenchyma formation in pods  ||       |     Unknown           
|-
| Position of flowers    ||        Axial       ||       Terminal      ||     ''FA''     ||       IV      ||   No   ||         Meristem function         ||       |     Unknown           
|}

==Translations==
* The 1901 translation: {{Wikisource-inline|1=Experiments in Plant-Hybridisation|single=true}}
* The 2016 translation: {{Cite journal |last=Abbott |first=Scott |last2=Fairbanks |first2=Daniel J |date=2016-10-01 |title=Experiments on Plant Hybrids by Gregor Mendel |url=https://academic.oup.com/genetics/article/204/2/407/6072056 |journal=Genetics |volume=204 |issue=2 |pages=407–422 |doi=10.1534/genetics.116.195198 |issn=1943-2631|pmc=5068836 }}

==References==
{{reflist}}

[[Category:Gregor Mendel]]
[[Category:Botanical literature]]
[[Category:1865 documents]]
[[Category:1866 documents]]
[[Category:1865 in biology]]
[[Category:1866 in biology]]