{{Short description|Austrian biologist and friar (1822–1884)}}
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{{Infobox scientist
| image             = Gregor Mendel 2.jpg
| caption           = Mendel, {{Circa|1862}}<ref>{{Cite book |last1=Bateson |first1=William |author-link1=William Bateson |url=https://archive.org/details/mendelsprinciple00bate/ |title=Mendel's principles of heredity |last2=Mendel |first2=Gregor |publisher=[[Cambridge University Press]] |year=1909 |location=Cambridge |page=[https://archive.org/details/mendelsprinciple00bate/page/n355/mode/2up 308] |oclc=1048814686 |via=[[Internet Archive]]}}</ref>
| image_size        = 
| honorific_prefix  = [[The Right Reverend]]
| honorific_suffix  = [[Order of Saint Augustine|OSA]]
| birth_name        = Johann Mendel
| birth_date        = {{birth date|df=y|1822|7|20}}
| birth_place       = [[Heinzendorf bei Odrau]], [[Austrian Silesia|Silesia]], Austrian Empire
| death_date        = {{death date and age|df=y|1884|1|6|1822|7|20}}
| death_place       = [[Brno]], Moravia, Austria-Hungary
| education         = [[University of Olmütz]]<br />[[University of Vienna]]
| doctoral_advisor  = <!--Please insert-->
| doctoral_students = <!--Please insert-->
| known_for         = Founder of the modern science of [[genetics]]
| field             = [[Genetics]]
| workplaces        = [[St Thomas's Abbey, Brno]]
| awards            = <!--Please insert-->
| signature         = Gregor Mendel Signature.svg
| module            = {{Infobox clergy
| child    = yes
| ordained = 25 December 1846<ref>{{cite journal|last1=Fr. Richter|first1=Clemens OSA|title=Remembering Johann Gregor Mendel: a human, a Catholic priest, an Augustinian monk, and abbot|journal=Molecular Genetics & Genomic Medicine|year=2015|volume=3|issue=6|pages=483–485|doi=10.1002/mgg3.186|doi-access=free |pmid=26740939|pmc=4694133}}</ref>
| religion = [[Christianity]]
| church   = [[Catholic Church]]
}}
}}
{{Genetics sidebar}}
'''Gregor Johann Mendel''' ({{IPAc-en|ˈ|m|ɛ|n|d|əl}}; {{IPA|de|ˈmɛndl̩|lang}}; {{langx|cs|Řehoř Jan Mendel}};<ref name="card">[http://img.radio.cz/pictures/r/vystavy/mendel_190/umrtni_oznameni.jpg Funeral card in Czech (Brno, 6. January 1884)]</ref> 20 July 1822<!-- Please DO NOT change this date. 20 July is correct; see the talk page. --><ref>20 July is his birthday, often mentioned as 22 July, the date of his baptism. {{cite web |url=https://mendelmuseum.muni.cz/en/g-j-mendel/zivotopis |website=Mendel Museum |title=CV |archive-url=https://web.archive.org/web/20190410150755/https://mendelmuseum.muni.cz/en/g-j-mendel/zivotopis |archive-date=10 April 2019 }}</ref> – 6 January 1884) was an [[Austrian Empire|Austrian]]<ref>{{Cite journal |last=De Castro |first=Mauricio |date=Jan 2016 |title=Johann Gregor Mendel: paragon of experimental science |doi-access=free |journal=Molecular Genetics & Genomic Medicine |language=en |volume=4 |issue=1 |pages=3–8 |doi=10.1002/mgg3.199 |pmc=4707027 |pmid=26788542 }}</ref><ref>{{cite web |url=https://www.genome.gov/genetics-glossary/Gregor-Johann-Mendel |title=Mendel, Johann (Gregor) |website=genome.gov |access-date=22 November 2024 |archive-date=22 November 2024 |archive-url=https://web.archive.org/web/20241122180834/https://www.genome.gov/genetics-glossary/Gregor-Johann-Mendel}}</ref> biologist, meteorologist,<ref>Czech J. Genet. Plant Breed., 50, 2014 (2): 43–51</ref> mathematician, [[Augustinians|Augustinian]] [[friar]] and [[abbot]] of [[St Thomas's Abbey, Brno|St. Thomas' Abbey]] in [[Brno]] (Brünn)<!-- common English name of the city, besides "Brünn" it was the other official name of the city even in Mendel's time, also used by his abbey, see his funeral card from 1884; German names on English Wiki are confusing -->, [[Margraviate of Moravia]]. Mendel was born in a [[Sudeten Germans|German-speaking]] family in the [[Austrian Silesia|Silesian]] part of the [[Austrian Empire]] (today's [[Czech Republic]]) and gained posthumous recognition as the founder of the modern science of [[genetics]].<ref name=":4">{{Cite book|last1=Klein|first1=Jan|title=Solitude of a Humble Genius – Gregor Johann Mendel. Volume 1, Formative years|last2=Klein|first2=Norman|date=2013|publisher=Springer|isbn=978-3-642-35254-6|location=Berlin|pages=91–103|oclc=857364787}}</ref> Though farmers had known for millennia that crossbreeding of animals and plants could favor certain desirable [[trait (biological)|traits]], Mendel's [[pea]] plant experiments conducted between 1856 and 1863 established many of the rules of [[biological inheritance|heredity]], now referred to as the laws of [[Mendelian inheritance]].<ref>{{Cite journal|last=Schacherer|first=Joseph|date=2016|title=Beyond the simplicity of Mendelian inheritance|journal=Comptes Rendus Biologies|volume=339|issue=7–8|pages=284–288|doi=10.1016/j.crvi.2016.04.006|pmid=27344551|doi-access=free}}</ref>

Mendel worked with seven characteristics of [[Pea|pea plants]]: plant height, pod shape and colour, seed shape and colour, and flower position and colour. Taking seed colour as an example, Mendel showed that when a true-breeding yellow pea and a true-breeding green pea were cross-bred, their offspring always produced yellow seeds. However, in the next generation, the green peas reappeared at a ratio of 1 green to 3 yellow. To explain this phenomenon, Mendel coined the terms "[[Dominance (genetics)|recessive]]" and "[[Dominance (genetics)|dominant]]" in reference to certain traits. In the preceding example, the green trait, which seems to have vanished in the first filial generation, is recessive, and the yellow is dominant. He published his work in 1866, demonstrating the actions of invisible "factors"—now called [[gene]]s—in predictably determining the traits of an organism. The actual genes were only discovered in a long process that ended in 2025 when the last three of the seven Mendel genes were identified in the pea [[genome]].<ref name=":6" />

The profound significance of Mendel's work was not recognized until the turn of the 20th century (more than three decades later) with the rediscovery of his laws. [[Erich von Tschermak]], [[Hugo de Vries]] and [[Carl Correns]] independently verified several of Mendel's experimental findings in 1900, ushering in the modern age of genetics.<ref>{{Cite journal|last=Gayon|first=Jean|date=2016|title=From Mendel to epigenetics: History of genetics|journal=Comptes Rendus Biologies|volume=339|issue=7–8|pages=225–230|doi=10.1016/j.crvi.2016.05.009|pmid=27263362|doi-access=free}}</ref><ref>{{Cite journal|last1=Corcos|first1=Alain F.|last2=Monaghan|first2=Floyd V.|date=1990|title=Mendel's work and its rediscovery: A new perspective|url=http://www.tandfonline.com/doi/abs/10.1080/07352689009382287|journal=Critical Reviews in Plant Sciences|volume=9|issue=3|pages=197–212|doi=10.1080/07352689009382287|bibcode=1990CRvPS...9..197C |url-access=subscription}}</ref>

== Early life and education ==
Mendel was born into a [[Sudeten Germans|German-speaking]] family in [[Hynčice (Vražné)|Heinzendorf bei Odrau]],<ref name="card"/> in [[Austrian Silesia|Silesia]], [[Austrian Empire]] (now Hynčice in the [[Czech Republic]]).<ref name=":4" /> He was the son of Anton and Rosine (Schwirtlich) Mendel and had one older sister, Veronika, and one younger, Theresia. They lived and worked on a farm which had been owned by the Mendel family for at least 130 years<ref>Gregor Mendel, Alain F. Corcos, Floyd V. Monaghan, Maria C. Weber "Gregor Mendel's Experiments on Plant Hybrids: A Guided Study", Rutgers University Press, 1993.</ref> (the house where Mendel was born is now a museum devoted to Mendel).<ref>{{cite web |url=http://www.mendel-rodnydum.vrazne.cz/index.php?lang=cs |title=Úvod – Rodný dům Johanna Gregora Mendela}}</ref> During his childhood, Mendel worked as a gardener and studied [[beekeeping]]. As a young man, he attended [[Gymnasium (school)|gymnasium]] in [[Troppau]] ({{langx|cs|Opava}}). Due to illness, he had to take four months off during his gymnasium studies.<ref>{{Cite web |last=Camarena |first=Belia |date=March 20, 2018 |title=Gregor Mendel, the Father of Modern Genetics: Brilliant Scientist or Complete Failure? |url=https://stmuscholars.org/gregor-mendel-the-father-of-modern-genetics-brilliant-scientist-or-complete-failure/ |access-date=10 March 2023 |website=StMU Research Scholars |archive-date=10 March 2023 |archive-url=https://web.archive.org/web/20230310171241/https://stmuscholars.org/gregor-mendel-the-father-of-modern-genetics-brilliant-scientist-or-complete-failure/ |url-status=dead }}</ref> From 1840 to 1843, he studied practical and theoretical philosophy and physics at the Philosophical Institute of the [[Palacký University Olomouc|University of Olomouc]] ({{langx|de|Olmütz}}), taking another year off because of illness. He also struggled financially to pay for his studies, and Theresia gave him her dowry. Later, he helped support her three sons, two of whom became doctors.<ref>{{Cite book|first=Silvia |last=Eckert-Wagner |year=2004 |title=Mendel und seine Erben: Eine Spurensuche |trans-title=Mendel and His Heirs: A search for traces |language=de |location=Norderstedt |publisher=Books on Demand |page=[https://books.google.com/books?id=3sU00yjUKfcC&pg=PA113 113] |isbn=978-3-8334-1706-1 }}</ref>

He became a monk partly because it enabled him to obtain an education without paying for it himself.<ref>{{Cite book|last=Henig|first=Robin Marantz|url=https://archive.org/details/monkingardenlost00heni|title=The Monk in the Garden: The Lost and Found Genius of Gregor Mendel, the Father of Genetics|date=2000|publisher=Houghton Mifflin|isbn=0-395-97765-7|location=Boston|pages=19–21|oclc=43648512}}</ref> As the son of a struggling farmer, the monastic life, in his words, spared him the "perpetual anxiety about a means of livelihood."<ref name=":0">{{Cite journal|last=Iltis|first=Hugo|date=1943|title=Gregor Mendel and His Work|url=https://www.jstor.org/stable/17803|journal=The Scientific Monthly|volume=56|issue=5|pages=414–423|jstor=17803|bibcode=1943SciMo..56..414I}}</ref> Born Johann Mendel, he was given the name "Gregor" ({{lang|cs|Řehoř}} in Czech)<ref name="card" /> when he joined the [[Order of Saint Augustine]].{{sfn|Henig|2000|p=24}}

== Academic career ==
[[File:Zawadzki and Mendel.jpg|left|thumb|Mendel (seated second from right and numbered "2") with other faculty at the Brno Realschule in 1864 ([[Aleksander Zawadzki (naturalist)|Alexander Zawadzki]] is labelled "1".)]]
When Mendel entered the Faculty of Philosophy, the Department of Natural History and Agriculture was headed by [[Johann Karl Nestler]], who conducted extensive research on hereditary traits of plants and animals, especially sheep. Upon recommendation of his [[physics]] teacher [[Friedrich Franz]],<ref name="Hasan">{{Cite book |last=Hasan |first=Heather |title=Mendel and The Laws Of Genetics|publisher=The Rosen Publishing Group|year=2004  |url=https://books.google.com/books?id=lXcRCag4rR8C&q=gregor+mendel+friedrich+franza&pg=PA1842|isbn =978-1-4042-0309-9 }}</ref> Mendel entered the [[Augustinians|Augustinian]] [[St Thomas's Abbey, Brno|St Thomas's Abbey]] in [[Brno]] and began his training as a [[Catholic priest]]. Mendel worked as a substitute high school teacher. In 1850, he failed his exams' oral part, the last of three parts, to become a certified high school teacher. In 1851, he was sent to the [[University of Vienna]] to study under the sponsorship of [[Abbot]] [[Cyril František Napp]] so that he could get a more formal education.{{sfn|Henig|2000|p=24}} At Vienna, his professor of physics was [[Christian Doppler]].<ref name="Mathofinheritance" /> Mendel returned to his abbey in 1853 as a teacher, principally of physics. In 1854, he met [[Aleksander Zawadzki (naturalist)|Aleksander Zawadzki]], who encouraged his research in Brno. In 1856, he took the exam to become a certified teacher and again failed the oral part.{{sfn|Henig|2000|pp=47–62}} In the summer of 1862, he joined an organised group tour to Paris and [[London]], where he visited the [[1862_International_Exhibition|International Exhibition]] and major scientific sites, a trip that may have influenced the final stage of his hybridisation research.<ref>{{Cite journal |last=Van Dijk |first=Peter J. |title=Mendel's journey to Paris and London: context and significance for the origin of genetics |journal=History and Philosophy of the Life Sciences |volume=56 |year=2020 |issue=1–2 |pages=5–47 }}</ref> In 1867, he succeeded Napp as abbot of the monastery.<ref name="MENDELMUSEUM">{{cite web|url=http://www.mendel-museum.com/eng/1online/ |publisher=The Masaryk University Mendel Museum |access-date=20 January 2010 |title=Online Museum Exhibition |url-status=dead |archive-url=https://web.archive.org/web/20141021181034/http://www.mendel-museum.com/eng/1online/ |archive-date=21 October 2014 }}</ref>

After he was elevated as abbot in 1868, his scientific work largely ended, as Mendel became overburdened with administrative responsibilities, especially a dispute with the civil government over its attempt to impose special taxes on religious institutions.<ref>{{cite encyclopedia |last=Windle |first =B.C.A. |others=Looby, John (trans.) |title=Mendel, Mendelism |encyclopedia=Catholic Encyclopedia |year=1911 |url=https://www.newadvent.org/cathen/10180b.htm |access-date=2 April 2007 }}</ref> Mendel died on 6 January 1884, at the age of 61, in Brno,<ref name="card"/> from chronic [[nephritis]]. Czech composer [[Leoš Janáček]] played the organ at his funeral.<ref name=Soudek>{{cite journal |last1=Soudek |first1=Dušan |date=1984 |title=Gregor Mendel and the people around him (commemorative of the centennial of Mendel's death) |journal=American Journal of Human Genetics |volume=36 |issue=3 |pages=495–498 [497]|pmid=6375354 |pmc=1684469 }}</ref> After his death, the succeeding abbot burned all papers in Mendel's collection, to mark an end to the disputes over taxation.<ref name=Carlson>{{cite book|last=Carlson|first=Elof Axel|author-link=Elof Axel Carlson|title=Mendel's Legacy|publisher=[[Cold Spring Harbor Laboratory]] Press|location=Cold Spring Harbor, NY|year=2004|pages=48–49|chapter=Doubts about Mendel's integrity are exaggerated|isbn=978-0-87969-675-7}}</ref> The [[exhumation]] of Mendel's corpse in 2021 delivered some [[physiognomic]] details like body height ({{cvt|168|cm}}). His [[genome analysis|genome was analysed]], revealing that Mendel was predisposed to heart problems.<ref>{{cite web |author=Austria Presse Agentur |url=https://science.apa.at/power-search/5470627088141089095l |language=de |title=Genomanalyse beim ersten Genetiker: Gregor Mendel exhumiert |website=science.apa.at |access-date=2022-07-16 }}{{Dead link|date=June 2024 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

== Contributions ==
=== Experiments on plant hybridization ===
{{Main|Mendelian inheritance}}
[[File:Mendelian inheritance.svg|thumb|Dominant and recessive phenotypes. (1) Parental generation. (2) F1 generation. (3) F2 generation.]]
Mendel, known as the "father of modern genetics," chose to study variation in plants in his monastery's {{convert|2|ha|acre}} experimental garden.<ref>{{cite web |url=https://mendel-museum.com/mendels-experiments-on-peas/ |title=Mendel's Experiments on Peas |publisher=The Masaryk University Mendel Museum |access-date=4 October 2020 |archive-date=9 August 2021 |archive-url=https://web.archive.org/web/20210809014930/https://mendel-museum.com/mendels-experiments-on-peas/ |url-status=dead }}</ref> Mendel was assisted in his experimental design by Aleksander Zawadzki while his superior abbot [[Cyril Napp|Napp]] wrote to discourage him, saying that the Bishop giggled when informed of the detailed genealogies of peas.<ref>{{Cite journal |last=Szybalski |first=W. |date=2010 |title=Professor Alexander Zawadzki of Lviv University – Gregor Mendel's mentor and inspirer |journal=Biopolymers and Cell |volume=26 |issue=2 |pages=83–86 |doi=10.7124/bc.000149|doi-access=free }}</ref>

After initial experiments with pea plants, Mendel settled on studying seven traits that seemed to be inherited independently of other traits: seed shape, flower color, seed coat tint, pod shape, unripe pod color, flower location, and plant height. He first focused on seed shape, which was either angular or round.{{sfn|Henig|2000|pp=78–80}} Between 1856 and 1863 Mendel cultivated and tested some 28,000 plants, the majority of which were [[pea]] plants (''[[Pisum sativum]]'').<ref>{{cite book|last1=Magner|first1=Lois N. |title=History of the Life Sciences|year=2002 |publisher=Marcel Dekker |location=New York|isbn=978-0-203-91100-6|page=380|edition=3, revised|url=https://books.google.com/books?id=YKJ6gVYbrGwC&pg=PA380}}</ref><ref>{{cite book|last1=Gros|first1=Franc̜ois|title=The Gene Civilization|year=1992|publisher=McGraw Hill|location=New York|isbn=978-0-07-024963-9|page=[https://archive.org/details/genecivilization00gros/page/28 28]|edition=English|url=https://archive.org/details/genecivilization00gros|url-access=registration}}</ref><ref name="Moore-2003">{{Cite journal|last=Moore |first=Randy |title=The "Rediscovery" of Mendel's Work |journal=Bioscene |year=2001 |volume=27 |issue=2 |pages=13–24 |url=https://www.cs.uml.edu/~grinstei/91.510/Rediscovery%20of%20Mendel.pdf |archive-url=https://web.archive.org/web/20160216153032/http://courses.pbsci.ucsc.edu/mcdb/bio105/Spring15/Lecture2/Rediscovery%20of%20Mendel.pdf |archive-date=16 February 2016 }}</ref> This study showed that, when true-breeding different varieties were crossed to each other (e.g., tall plants fertilized by short plants), in the second generation, one in four pea plants had [[purebred]] [[recessive]] [[Phenotypic trait|traits]], two out of four were [[hybrid (biology)|hybrids]], and one out of four were purebred [[dominance (genetics)|dominant]]. His experiments led him to make two generalizations, the [[Law of Segregation]] and the [[Law of independent assortment|Law of Independent Assortment]], which later came to be known as Mendel's Laws of Inheritance.<ref>{{cite book|last1=Butler |first1=John M.|title=Fundamentals of Forensic DNA Typing|date=2010|publisher=Elsevier/Academic Press|location=Burlington, MA|isbn=978-0-08-096176-7|pages=34–35|url=https://books.google.com/books?id=-OZeEmqzE4oC&pg=PA34}}</ref>

==== Initial reception of Mendel's work ====
Mendel presented his paper, {{lang|de|Versuche über Pflanzenhybriden}} ("[[Experiments on Plant Hybridization]]"), at two meetings of the Natural History Society of Brno in [[Moravia]] on 8 February and 8 March 1865.{{sfn|Henig|2000|pp=134–138}} It generated a few favorable reports in local newspapers,<ref name="Moore-2003" /> but was ignored by the scientific community. When Mendel's paper was published in 1866 in {{lang|de|[[Proceedings of the Natural History Society of Brünn|Verhandlungen des naturforschenden Vereines in Brünn]]}},<ref>Mendel, J.G. (1866). "[https://www.biodiversitylibrary.org/item/124139#page/133/mode/1up Versuche über Pflanzenhybriden]", ''Verhandlungen des naturforschenden Vereines in Brünn'', Bd. IV für das Jahr, 1865, ''Abhandlungen'': 3–47. 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 |archive-url=https://web.archive.org/web/20000902033224/http://www.esp.org/foundations/genetics/classical/gm-65.pdf |archive-date=2000-09-02 |url-status=live|journal=Journal of the Royal Horticultural Society|volume=26|pages=1–32|access-date=9 October 2009}}</ref> it was seen as essentially about hybridization rather than inheritance, had little impact, and was cited only about three times over the next thirty-five years. His paper was criticized then but is now considered a seminal work.<ref>{{cite journal|last1=Galton|first1=D. J.|title=Did Mendel falsify his data?|journal=QJM|date=2011|volume=105|issue=2|pages=215–16|doi= 10.1093/qjmed/hcr195|pmid=22006558|doi-access=free}}</ref> Notably, [[Charles Darwin]] was not aware of Mendel's paper, and it is envisaged that if he had been aware of it, genetics as it exists now might have taken hold much earlier.<ref>{{cite journal|last1=Lorenzano|first1=P|title=What would have happened if Darwin had known Mendel (or Mendel's work)?|journal=History and Philosophy of the Life Sciences|date=2011|volume=33|issue=1|pages=3–49|pmid=21789954}}</ref><ref>{{cite journal|last1=Liu|first1=Y|title=Darwin and Mendel: who was the pioneer of genetics?|journal=Rivista di Biologia|date=2005|volume=98|issue=2|pages=305–22|pmid=16180199}}</ref> Mendel's scientific biography thus provides an example of the failure of obscure, highly original innovators to receive the attention they deserve.<ref name=":2">{{Cite journal|date=1995|title=The Plight of the Obscure Innovator in Science|journal=Social Studies of Science|volume=25|issue=1|pages=165–83|doi=10.1177/030631295025001008 |last1=Nissani |first1=M.|s2cid=144949936}}</ref>

==== Rediscovery of Mendel's work ====
About forty scientists listened to Mendel's two groundbreaking lectures, but it would appear that they failed to understand the implications of his work. Later, he also carried on a correspondence with [[Carl Nägeli]], one of the leading biologists of the time, but Nägeli also failed to appreciate Mendel's discoveries. At times, Mendel must have entertained doubts about his work, but not always: "My time will come," he reportedly told a friend,<ref name=":0" /> Gustav von Niessl.<ref name=":5">{{Cite journal|last=Gustafsson|first=Åke|author-link=Åke Gustafsson|date=1969|title=The Life of Gregor Johann Mendel — Tragic or Not?|journal=Hereditas|volume=62|issue=1|pages=239–258|doi=10.1111/j.1601-5223.1969.tb02232.x|pmid=4922561|doi-access=free}}</ref>

During Mendel's lifetime, most biologists held the idea that all characteristics were passed to the next generation through [[blending inheritance]] (indeed, many effectively are), in which the traits from each parent are averaged.<ref>{{Cite journal|last=Weldon|first=W. F. R.|date=1902|title=Mendel's Laws of Alternative Inheritance in Peas|url=https://academic.oup.com/biomet/article-lookup/doi/10.1093/biomet/1.2.228|journal=Biometrika|volume=1|issue=2|pages=228–233|doi=10.1093/biomet/1.2.228|url-access=subscription}}</ref><ref>{{Cite journal|last=Bulmer|first=Michael|date=1999|title=The Development of Francis Galton's Ideas on the Mechanism of Heredity|url=http://link.springer.com/10.1023/A:1004608217247|journal=Journal of the History of Biology|volume=32|issue=2|pages=263–292|doi=10.1023/A:1004608217247|pmid=11624207|s2cid=10451997|url-access=subscription}}</ref> Instances of this phenomenon are now explained by the action of multiple genes with [[Quantitative genetics|quantitative effects]]. Charles Darwin tried unsuccessfully to explain inheritance through a theory of [[pangenesis]]. It was not until the early 20th century that the importance of Mendel's ideas was realized.<ref name="Moore-2003" />

By 1900, research aimed at finding a successful theory of discontinuous inheritance rather than [[blending inheritance]] led to independent duplication of his work by [[Hugo de Vries]] and [[Carl Correns]] and the rediscovery of Mendel's writings and laws. Both acknowledged Mendel's priority, and it is thought probable that de Vries did not understand the results he had found until after reading Mendel.<ref name="Moore-2003" /> Though [[Erich von Tschermak]] was originally also credited with rediscovery, this is no longer accepted because he did not understand [[Mendelian inheritance|Mendel's laws]].<ref>{{cite book |author=Mayr, Ernst |author-link=Ernst Mayr |year=1982 |title=The Growth of Biological Thought |location=Cambridge |publisher=The Belknap Press of Harvard University Press |isbn=978-0-674-36446-2 |page=730}}</ref> Though de Vries later lost interest in Mendelism, other biologists started to establish modern genetics as a science. All three of these researchers, each from a different country, published their rediscovery of Mendel's work within a two-month span in the spring of 1900.{{sfn|Henig|2000|pp=1–9}}

Mendel's results were quickly replicated, and genetic linkage was quickly worked out. Biologists flocked to the theory; even though it was not yet applicable to many phenomena, it sought to give a [[genotype|genotypic]] understanding of heredity, which they felt was lacking in previous studies of heredity, which had focused on [[phenotype|phenotypic]] approaches.<ref>{{Cite book|title=Mendel's Legacy: The Origins of Classical Genetics|last=Carlson|first=Elof Axel|publisher=Cold Spring Harbor|year=2004|location=New York}}</ref> Most prominent of these previous approaches was the [[Biostatistics|biometric]] school of [[Karl Pearson]] and [[W. F. R. Weldon]], which was based heavily on statistical studies of phenotype variation. The strongest opposition to this school came from [[William Bateson]], who perhaps did the most in the early days of publicising the benefits of Mendel's theory (the word "[[genetics]]", and much of the discipline's other terminology, originated with Bateson). This debate between the biometricians and the Mendelians was extremely vigorous in the first two decades of the 20th century, with the biometricians claiming statistical and mathematical rigor,<ref>{{cite journal|last1=Deichmann|first1=Ute|title=Early 20th-century research at the interfaces of genetics, development, and evolution: Reflections on progress and dead ends|journal=Developmental Biology|date=2011|volume=357|issue=1|pages=3–12|doi=10.1016/j.ydbio.2011.02.020 |pmid=21392502|doi-access=free}}</ref> whereas the Mendelians claimed a better understanding of biology.<ref>{{cite journal|last1=Elston|first1=RC|last2=Thompson|first2=EA|author2-link= Elizabeth A. Thompson |title=A century of biometrical genetics|journal=Biometrics|date=2000|volume=56|issue=3|pages=659–66|pmid=10985200|doi=10.1111/j.0006-341x.2000.00659.x|s2cid=45142547}}</ref><ref>{{cite journal|last1=Pilpel|first1=Avital|title=Statistics is not enough: revisiting Ronald A. Fisher's critique (1936) of Mendel's experimental results (1866)|journal=Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences|date=September 2007|volume=38|issue=3|pages=618–26|doi=10.1016/j.shpsc.2007.06.009 |pmid=17893069}}</ref> Modern genetics shows that Mendelian heredity is, in fact, an inherently biological process, though not all genes of Mendel's experiments are yet understood.<ref>{{Cite journal|last1=Reid|first1=J. B.|last2=Ross|first2=J. J.|title=Mendel's genes: toward a full molecular characterization|journal=Genetics|date=2011|volume=189|issue=1|pages=3–10|doi=10.1534/genetics.111.132118|pmid=21908742|pmc=3176118}}</ref><ref>{{cite journal|last1=Ellis|first1=T.H. Noel|last2=Hofer|first2=Julie M.I.|last3=Timmerman-Vaughan|first3=Gail M.|last4=Coyne|first4=Clarice J.|last5=Hellens|first5=Roger P.|title=Mendel, 150 years on|journal=Trends in Plant Science|date=2011|volume=16|issue=11|pages=590–96|doi=10.1016/j.tplants.2011.06.006|pmid=21775188|bibcode=2011TPS....16..590E }}</ref>

Ultimately, the two approaches were combined, especially by work conducted by [[R. A. Fisher]] as early as 1918. The combination, in the 1930s and 1940s, of Mendelian genetics with Darwin's theory of [[natural selection]] resulted in the [[Modern synthesis (20th century)|modern synthesis]] of [[evolutionary biology]].<ref>{{cite journal|last1=Kutschera |first1=Ulrich|last2=Niklas|first2=KarlJ.|title=The modern theory of biological evolution: an expanded synthesis|journal=Naturwissenschaften|date=2004|volume=91|issue=6|pages=255–76|doi=10.1007/s00114-004-0515-y|pmid=15241603|bibcode=2004NW.....91..255K|s2cid=10731711}}</ref><ref>{{cite book|last1=Hall|first1=Brian Keith|last2=Hallgrímsson|first2=Benedikt|last3=Strickberger |first3=Monroe W.|title=Strickberger's evolution|date=2014|publisher=Jones & Bartlett Learning|location=Burlington, Mass.|isbn=978-1-4496-1484-3|pages=10–11|edition=5|url=https://books.google.com/books?id=WkcvuVpzjYQC}}</ref>

In the [[Soviet Union]] and the [[People's Republic of China]], Mendelian genetics was rejected in favor of [[Lamarckism]] under the state policy of [[Lysenkoism]], leading to imprisonment and even execution of Mendelian geneticists as well as massive famines in both of those countries.<ref>{{Cite web |date=2021-11-02 |title=Collapse of Reason |url=https://exhibits.library.cornell.edu/cultivating-silence/feature/collapse-of-reason |access-date=2026-04-19 |website=Cultivating Silence - Online exhibitions across Cornell University Library |language=en}}</ref>

=== Modern analysis of the genes causing Mendel's pea phenotypes ===
Mendel postulated that seven "factors" determine the features he studied in peas. These factors are called "[[Gene|genes]]" today, but the nature of these genes remained mysterious for more than a century. The effort to identify these genes lasted until 2025, when the last 3 genes were discovered.<ref name=":6" /> The seven genes are as follows (genes are abbreviated PsXYZ for ''[[Pea|Pisum sativum]]'', the scientific name of the pea): the ''wrinkled'' phenotype of peas (wild-type round) is caused by an [[Insertion (genetics)|insertion]] in the '''PsSBE1''' gene. The ''yellow'' phenotype (wild-type: green) is caused by an insertion or mutation in the '''PsSGR''' gene. The ''white'' phenotype of the flower colour (wild-type: ''purple'') is caused by a [[Deletion (genetics)|deletion]] in the '''PsbHLH''' gene. The ''dwarf'' phenotype is caused by the '''PsGA3ox1''' gene while the pod color phenotype (''yellow'' vs. ''green'') is caused by the '''PsChlG''' gene. Finally, the pod shape is determined by the '''PsCLE41''' gene, which causes the ''constricted'' or ''inflated'' phenotypes, and the '''PsCIK2/3''' gene causes the ''terminal'' and ''axial'' flower position.<ref name=":6">{{Cite journal |last1=Feng |first1=Cong |last2=Chen |first2=Baizhi |last3=Hofer |first3=Julie |last4=Shi |first4=Yan |last5=Jiang |first5=Mei |last6=Song |first6=Bo |last7=Cheng |first7=Hong |last8=Lu |first8=Lu |last9=Wang |first9=Luyao |last10=Howard |first10=Alex |last11=Bendahmane |first11=Abdel |last12=Fouchal |first12=Anissa |last13=Moreau |first13=Carol |last14=Sawada |first14=Chie |last15=LeSignor |first15=Christine |date=2025-04-23 |title=Genomic and genetic insights into Mendel's pea genes |journal=[[Nature (journal)| Nature]] |volume=642 |issue=8069 |language=en |pages=980–989 |doi=10.1038/s41586-025-08891-6 |pmid=40269167 |issn=1476-4687|pmc=12221995 |bibcode=2025Natur.642..980F}}</ref>

=== Other experiments ===
Mendel also experimented with [[hawkweed]] (''Hieracium'').<ref name="Nogler2006">{{cite journal|last1=Nogler|first1=GA|title=The lesser-known Mendel: his experiments on Hieracium.|journal=Genetics|date=2006|volume=172|issue=1|pages=1–6|doi=10.1093/genetics/172.1.1|pmid=16443600|pmc=1456139}}</ref> He published a report on his work with hawkweed,<ref name="Mendel1869">{{cite journal|last1=Mendel|first1=Gregor|title=Ueber einige aus künstlicher Befruchtung gewonnenen Hieracium-Bastarde. (On Hieracium hybrids obtained by artificial fertilisation)|journal=Verh. Naturf. Ver. Brünn|date=1869|volume=8 (Abhandlungen)|pages=26–31}}</ref> a group of plants of great interest to scientists at the time because of their diversity. However, the results of Mendel's inheritance study in hawkweeds were unlike those for peas; the first generation was very variable, and many of their offspring were identical to the maternal parent. In his correspondence with [[Carl Nägeli]] he discussed his results but was unable to explain them.<ref name="Nogler2006" /> It was not appreciated until the end of the nineteenth century that many hawkweed species were [[apomictic]], producing most of their seeds through an asexual process.<ref name=":5" /><ref>{{Cite journal|last1=Koltunow|first1=A. M. G.|last2=Johnson|first2=S. D.|last3=Okada|first3=T.|date=2011|title=Apomixis in hawkweed: Mendel's experimental nemesis|journal=Journal of Experimental Botany|volume=62|issue=5|pages=1699–1707|doi=10.1093/jxb/err011|pmid=21335438|doi-access=free}}</ref>

Mendel appears to have kept animals at the monastery, breeding bees in custom-designed [[bee hive]]s.<ref>{{cite web|url=http://www.mendel-museum.com/eng/1online/room3.htm|title=The Enigma of Generation and the Rise of the Cell|publisher=The Masaryk University Mendel Museum|access-date=20 January 2010|url-status=dead|archive-url=https://web.archive.org/web/20141021181104/http://www.mendel-museum.com/eng/1online/room3.htm|archive-date=21 October 2014}}</ref><ref>{{Cite journal |last=Vecerek |first=O. |date=1965 |title=Johann Gregor Mendel as a Beekeeper |url=https://www.tandfonline.com/doi/full/10.1080/0005772X.1965.11095345 |journal=Bee World|volume=46 |issue=3 |pages=86–96 |doi=10.1080/0005772X.1965.11095345 |issn=0005-772X|url-access=subscription }}</ref>  None of his results on bees survived, except for a passing mention in the reports of the Moravian Apiculture Society.<ref>{{cite book|last1=Orel|first1=Vítězslav|last2=Rozman |first2=Josef|last3=Veselý|first3=Vladimír|title=Mendel as a Beekeeper|date=1965|publisher=Moravian Museum|pages=12–14|url=https://books.google.com/books?id=S9pCAAAAYAAJ}}</ref> All that is known definitely is that he used Cyprian and Carniolan bees,<ref>{{cite book|last1=Demerec|first1=M.|title=Advances in Genetics|date=1956|publisher=Academic Press|location=New York|page=110|url=https://books.google.com/books?id=UuEVFGcJuw4C}}</ref> which were particularly aggressive, to the annoyance of other monks and visitors of the monastery, such that he was asked to get rid of them.<ref>{{cite book|last1=Roberts|first1=Michael|last2=Ingram|first2=Neil|title=Biology|date=2001|publisher=Nelson Thornes|location=Cheltenham|isbn=978-0-7487-6238-5|page=277|edition=2 |url=https://books.google.com/books?id=juiDySqWVYkC}}</ref> Mendel, on the other hand, was fond of his bees and referred to them as "my dearest little animals".<ref>{{cite journal|last1=Matalova|first1=A|last2=Kabelka|first2=A|title=The beehouse of Gregor Mendel|journal=Casopis Moravskeho Musea. Acta Musei Moraviae – Vedy Prirodni. Car Morav Mus Acta Mus Vedy Prir|date=1982|volume=57|pages=207–12|url=https://www.nal.usda.gov/agricola?Search_Arg=IND84032981&DB=local&CNT=25&Search_Code=GKEY&STARTDB=AGRIDB}}</ref>

After his death, Mendel's colleagues remembered that he bred mice, crossing varieties of different sizes, although Mendel had left no record of any such work. A persistent myth has developed that Mendel turned his attention to plants only after Napp declared it unseemly for a celibate priest to closely observe rodent sex. In a 2022 biography, Daniel Fairbanks argued that Napp could hardly have given such a pronouncement, as Napp personally oversaw sheep breeding on the monastery's extensive agricultural estate.<ref>{{cite book|first=Daniel&nbsp;J.|last=Fairbanks|author-link=Daniel Justin Fairbanks|at=Ch.&nbsp;4|title=Gregor Mendel|publisher=Promethean Books|location=Guilford, CT|year=2022|lccn=2021054684|isbn=9781633888395}}</ref> 

Mendel also studied [[astronomy]] and [[meteorology]],<ref name= MENDELMUSEUM /> founding the 'Austrian Meteorological Society' in 1865.<ref name=Mathofinheritance>{{Cite journal|title=The Mathematics of Inheritance |volume=132 |issue=3348 |pages=1012 |publisher=The Masaryk University Mendel Museum |journal=Online Museum Exhibition |bibcode=1933Natur.132.1012F |last1=Fisher |first1=R. A. |year=1933 |doi=10.1038/1321012a0|doi-access=free }}</ref> The majority of his published works were related to meteorology.<ref name=Mathofinheritance />

He also described novel plant [[species]], and these are denoted with the [[List of botanists by author abbreviation|botanical author abbreviation]] "Mendel".<ref>{{cite web|title=Index of Botanists: Mendel, Gregor Johann|url=http://kiki.huh.harvard.edu/databases/botanist_search.php?mode=details&id=68141|website=HUH – Databases – Botanist Search|publisher=Harvard University Herbaria & Libraries|access-date=29 January 2018}}</ref>

== Mendelian paradox<!--'Mendelian paradox' redirects here--> ==
In 1936, [[Ronald Fisher]], a prominent statistician and population geneticist, reconstructed Mendel's experiments, analyzed results from the F<sub>2</sub> (second filial) generation, and found the ratio of dominant to recessive phenotypes (e.g., yellow versus green peas; round versus wrinkled peas) to be implausibly and consistently too close to the expected ratio of 3 to 1.<ref name=fisher36>{{cite journal|last1=Fisher|first1=R.A.|title=Has Mendel's work been rediscovered?|journal=Annals of Science|year=1936|volume=1|issue=2|pages=115–37|doi=10.1080/00033793600200111|url=https://digital.library.adelaide.edu.au/dspace/bitstream/2440/15123/1/144.pdf |archive-url=https://web.archive.org/web/20110413101103/http://digital.library.adelaide.edu.au/dspace/bitstream/2440/15123/1/144.pdf |archive-date=2011-04-13 |url-status=live|hdl=2440/15123|hdl-access=free}}</ref><ref>{{cite journal|last1=Thompson| first1=EA| title=R.A. Fisher's contributions to genetical statistics| journal=Biometrics|year=1990| volume=46|issue=4 |pages=905–14| pmid=2085639|doi=10.2307/2532436| jstor=2532436}}</ref><ref>{{cite journal|last1=Pilgrim|first1=I|title=The too-good-to-be-true paradox and Gregor Mendel|journal=The Journal of Heredity|year=1984|volume=75|issue=6|pages=501–02|pmid=6392413|doi=10.1093/oxfordjournals.jhered.a109998}}</ref> Fisher asserted that "the data of most, if not all, of the experiments have been falsified to agree closely with Mendel's expectations".<ref name=fisher36 /> Mendel's alleged observations, according to Fisher, were "abominable," "shocking," <ref name=hartl /> and "cooked."<ref>{{cite journal|last1=Piegorsch|first1=WW|title=Fisher's contributions to genetics and heredity, with particular emphasis on the Gregor Mendel controversy|journal=Biometrics|year=1990|volume=46|issue=4|pages=915–24|pmid=2085640|doi=10.2307/2532437|jstor=2532437|url=https://zenodo.org/record/1235119}}</ref>

Other scholars agree with Fisher that Mendel's various observations come uncomfortably close to Mendel's expectations. [[A. W. F. Edwards]],<ref name=":3">{{Cite journal|last=Edwards|first=A. W. F.|date=1986|title=More on the too-good-to-be-true paradox and Gregor Mendel|journal=Journal of Heredity|volume=77|issue=2|page=138 |doi=10.1093/oxfordjournals.jhered.a110192}}</ref> for instance, remarks: "One can applaud the lucky gambler; but when he is lucky again tomorrow, and the next day, and the following day, one is entitled to become a little suspicious". Three other lines of evidence likewise lend support to the assertion that Mendel's results are indeed too good to be true.<ref name=":1">{{Cite journal|last=Nissani|first=M.|date=1994|title=Psychological, Historical, and Ethical Reflections on the Mendelian Paradox|journal=Perspectives in Biology and Medicine|volume=37|issue=2|pages=182–96|doi=10.1353/pbm.1994.0027|pmid=11644519|s2cid=33124822}}</ref>

Fisher's analysis gave rise to the '''Mendelian paradox'''<!--boldface per WP:R#PLA-->: Mendel's reported data are, statistically speaking, too good to be true, yet "everything we know about Mendel suggests that he was unlikely to engage in either deliberate fraud or in an unconscious adjustment of his observations".<ref name=":1" /> Several writers have attempted to resolve this paradox.

One attempted explanation invokes [[confirmation bias]].<ref>{{cite journal|last1=Price|first1=Michael|title=Sins against science: Data fabrication and other forms of scientific misconduct may be more prevalent than you think|journal=Monitor on Psychology|year=2010|volume=41|issue=7|page=44|url=http://www.apa.org/monitor/2010/07-08/misconduct.aspx}}</ref> Fisher accused Mendel's experiments as "biased strongly in the direction of agreement with expectation{{nbsp}}[...] to give the theory the benefit of the doubt".<ref name="fisher36" /> In a 2004 article, J.W. Porteous concluded that Mendel's observations were indeed implausible.<ref>{{cite journal|last1=Porteous|first1=JW|title=We still fail to account for Mendel's observations.|journal=Theoretical Biology & Medical Modelling|date=2004|volume=1|article-number=4|pmid=15312231|pmc=516238|doi=10.1186/1742-4682-1-4 |doi-access=free }}</ref> An explanation for Mendel's results based on [[Tetrad (meiosis)|tetrad]] pollen has been proposed, but reproduction of the experiments showed no evidence that the tetrad-pollen model explains any of the bias.<ref>{{Cite journal|last1=Fairbanks|first1=D. J.|last2=Schaalje|first2=G. B.|title=The tetrad-pollen model fails to explain the bias in Mendel's pea (''Pisum sativum'') experiments|journal=Genetics|date=2007|volume=177|issue=4|pages=2531–34|doi=10.1534/genetics.107.079970|pmid=18073445|pmc=2219470}}</ref>

Another attempt<ref name=":1" /> to resolve the Mendelian paradox notes that a conflict may sometimes arise between the moral imperative of a bias-free recounting of one's factual observations and the even more important imperative of advancing scientific knowledge. Mendel might have felt compelled "to simplify his data to meet real, or feared editorial objections."<ref name=":3" /> Such an action could be justified on moral grounds (and hence provide a resolution to the Mendelian paradox) since the alternative—refusing to comply—might have hindered the growth of scientific knowledge. Similarly, like so many other obscure innovators of science,<ref name=":2" /> Mendel, a little-known innovator of working-class background, had to "break through the cognitive paradigms and social prejudices" of his audience.<ref name=":3" /> If such a breakthrough "could be best achieved by deliberately omitting some observations from his report and adjusting others to make them more palatable to his audience, such actions could be justified on moral grounds."<ref name=":1" />

[[Dan Hartl|Daniel L. Hartl]] and [[Daniel J. Fairbanks]] reject Fisher's statistical argument outright, suggesting that Fisher incorrectly interpreted Mendel's experiments. They find it likely that Mendel scored more than ten progeny and that the results matched the expectation. They conclude: "Fisher's allegation of deliberate falsification can finally be put to rest, because on closer analysis it has proved to be unsupported by convincing evidence".<ref name="hartl">{{cite journal |last= Hartl|first=Daniel L. |author2=Fairbanks, Daniel J. |year= 2007|title=Mud sticks: On the alleged falsification of Mendel's Data |journal=Genetics |volume=175 |issue= 3 |pages=975–79 |doi=10.1093/genetics/175.3.975 |pmid= 17384156 |pmc= 1840063}}</ref><ref>{{cite journal|last=Novitski|first=Charles E.|year=2004|title=On Fisher's criticism of Mendel's results with the garden pea|url=http://www.genetics.org/cgi/reprint/166/3/1133|journal=Genetics|volume=166|issue=3|pages=1133–36|doi=10.1534/genetics.166.3.1133|pmc=1470775|pmid=15082533|access-date=20 March 2010|quote=In conclusion, Fisher's criticism of Mendel's data—that Mendel was obtaining data too close to false expectations in the two sets of experiments involving the determination of segregation ratios—is undoubtedly unfounded.}}</ref> In 2008 Hartl and Fairbanks (with Allan Franklin and AWF Edwards) wrote a comprehensive book in which they concluded that there were no reasons to assert Mendel fabricated his results, nor that Fisher deliberately tried to diminish Mendel's legacy.<ref>{{cite book|last1=Franklin|first1=Allan|last2=Edwards|first2=AWF|last3=Fairbanks|first3=Daniel J|last4=Hartl |first4=Daniel L|title=Ending the Mendel-Fisher controversy|year=2008|publisher=University of Pittsburgh Press|location=Pittsburgh, PA|isbn=978-0-8229-4319-8|page=67 |url=https://books.google.com/books?id=C4m6NlmGhjgC}}</ref> Reassessment of Fisher's statistical analysis, according to these authors, also disproves the notion of confirmation bias in Mendel's results.<ref>{{cite journal|last1=Monaghan|first1=F|last2=Corcos|first2=A|title=Chi-square and Mendel's experiments: where's the bias?|journal=The Journal of Heredity|date=1985|volume=76|issue=4|pages=307–09|pmid=4031468|doi=10.1093/oxfordjournals.jhered.a110099}}</ref><ref>{{cite journal|last1=Novitski|first1=C. E.|title=Revision of Fisher's analysis of Mendel's garden pea experiments|journal=Genetics|year=2004|volume=166|issue=3|pages=1139–40|doi=10.1534/genetics.166.3.1139|pmid=15082535|pmc=1470784}}</ref>

==Commemoration==
Mount Mendel in New Zealand's [[Paparoa Range]] was named after him in 1970 by the [[Department of Scientific and Industrial Research (New Zealand)|Department of Scientific and Industrial Research]].<ref>{{LINZ|id=3770|name=Mount Mendel |access-date=21 August 2022}}</ref> In celebration of his 200th birthday, Mendel's body was exhumed and his DNA sequenced.<ref>[https://www.npr.org/sections/health-shots/2022/12/30/1142202365/gregor-mendel-genetics-dna-analyzed Why scientists dug up the father of genetics, Gregor Mendel, and analyzed his DNA]</ref>

== See also ==
* [[List of Roman Catholic cleric–scientists]]
* [[Mendel Museum of Masaryk University|Mendel Museum of Genetics]]
* [[Mendel Polar Station]] in Antarctica
* [[Mendel University in Brno]]
* [[Mendelian error]]
* ''[[The Gardener of God]]'', an Italian docudrama about the life and works of Gregor Mendel

== References ==
{{Reflist}}

== Further reading ==
{{refbegin|colwidth=30em}}
* [[William Bateson]] {{cite book |author1=Mendel, Gregor |author2=Bateson, William |title=Mendel's Principles of Heredity: A Defence, with a Translation of Mendel's Original Papers on Hybridisation (Cambridge Library Collection – Life Sciences) |publisher=Cambridge University Press |location=Cambridge, UK |year=2009 |isbn=978-1-108-00613-2 }} [http://www.esp.org/books/bateson/mendel/facsimile/title3.html On-line Facsimile Edition: Electronic Scholarly Publishing, Prepared by Robert Robbins]
* [[Hugo Iltis]], ''Gregor Johann Mendel. Leben, Werk und Wirkung''. Berlin: J. Springer. 426 pages. (1924)
** Translated by [[Eden Paul|Eden]] and [[Cedar Paul]] as ''Life of Mendel''. New York: W. W. Norton & Co, 1932. 336 pages. New York: Hafner, 1966: London: George Allen & Unwin, 1966.  Ann Arbor: University Microfilms International, 1976.
** Translated by Zhenyao Tan as ''Mên-tê-êrh chuan''. Shanghai: Shang wu yin shu guan, 1924. 2 vols. in 1, 661 pp. Shanghai: Shang wu yin shu guan, Minguo 25 [1936].
** Translated as ''Zasshu shokubutsu no kenkyū. Tsuketari Menderu shōden''. Tōkyō: Iwanami Shoten, Shōwa 3 [1928]. 100 pp. Translated by Yuzuru Nagashima as ''Menderu no shōgai''.  Tōkyō: Sōgensha, Shōwa 17 [1942]. ''Menderu den''. Tōkyō: Tōkyō Sōgensha, 1960.
* {{cite book |author1=Klein, Jan |author2=Klein, Norman |title=Solitude of a Humble Genius – Gregor Johann Mendel: Volume 1 |publisher=Springer |location=Heidelberg |year=2013 |isbn=978-3-642-35253-9}}
* Robert Lock, ''Recent Progress in the Study of Variation, Heredity and Evolution'', London, 1906
* {{cite book |author=Orel, Vítĕzslav |title=Gregor Mendel: the first geneticist |publisher=Oxford University Press |location=Oxford [Oxfordshire] |year=1996 |isbn=978-0-19-854774-7 }}
* {{Cite book |last=Punnett|first=Reginald Crundall|title=Mendelism|url=https://archive.org/stream/mendelism00punn#page/n7/mode/2up |publisher=Macmillan|location=London|year=1922}} (1st Pub. 1905)
* [[Curt Stern (geneticist)|Curt Stern]] and Sherwood ER (1966) ''The Origin of Genetics''.
* {{cite journal|last=Taylor|first=Monica|title=Abbot Mendel|url=https://archive.org/stream/dublinreview171londuoft#page/n7/mode/2up |journal=Dublin Review|date=July–September 1922|publisher=W. Spooner |location=London}}
* {{cite book |author=Tudge, Colin |title=In Mendel's footnotes: an introduction to the science and technologies of genes and genetics from the nineteenth century to the twenty-second |publisher=Vintage |location=London |year=2000 |isbn=978-0-09-928875-6 }}
* {{Cite journal |last1=Waerden |first1=B. L. V. D. |author-link=Bartel Leendert van der Waerden|title=Mendel's Experiments |doi=10.1111/j.1600-0498.1968.tb00098.x |journal=Centaurus |volume=12 |issue=4 |pages=275–88 |year=1968 |pmid=4880928|bibcode=1968Cent...12..275V}} refutes allegations about "data smoothing"
* James Walsh, ''Catholic Churchmen in Science'', Philadelphia: Dolphin Press, 1906
* {{cite book|last=Windle|first=Bertram C. A.|chapter-url=https://archive.org/stream/centuryofscienti00windrich#page/106/mode/2up |chapter=Mendel and His Theory of Heredity |title= A Century of Scientific Thought and Other Essays|publisher=Burns & Oates|year=1915}}
* {{cite journal|last1=Zumkeller |first1=Adolar |last2=Hartmann |first2=Arnulf |date=1971|title=Recently Discovered Sermon Sketches of Gregor Mendel|journal=Folia Mendeliana|volume=6|pages=247–52}}
{{refend}}

== External links ==
{{Commons category|Gregor Mendel}}
{{Wikiquote|Gregor Mendel}}
* {{Gutenberg author |id=40855| name=Gregor Mendel}}
* {{Internet Archive author |sname=Gregor Mendel}}
* {{Librivox author |id=569}}
* [https://www.newadvent.org/cathen/10180b.htm 1913 Catholic Encyclopedia entry, "Mendel, Mendelism"]
* [http://www.opatbrno.cz/ Augustinian Abbey of St. Thomas at Brno] {{Webarchive|url=https://web.archive.org/web/20051122054832/http://www.opatbrno.cz/ |date=22 November 2005 }}
* [http://vlp.mpiwg-berlin.mpg.de/people/data?id=per116 Biography, bibliography and access to digital sources] in the [[Virtual Laboratory]] of the [[Max Planck Institute for the History of Science]]
* [https://web.archive.org/web/20080117233318/http://mendel.imp.ac.at/mendeljsp/biography/biography.jsp Biography of Gregor Mendel]
* [http://www.bbc.co.uk/schools/gcsebitesize/science/add_aqa/celldivision/inheritance3.shtml GCSE student]
* [http://www.accessexcellence.org/RC/AB/BC/Gregor_Mendel.html Gregor Mendel (1822–1884)]
* [https://web.archive.org/web/20170917000119/http://www.thomasmore.edu/library/mendel_collection.cfm?group%20=The%20Mendel%20Collection Gregor Mendel Primary Sources]
* [http://www.weloennig.de/mendel.htm Johann Gregor Mendel: Why his discoveries were ignored for 35 (72) years] {{in lang|de}}
* [https://web.archive.org/web/20090620175253/http://brnonow.com/2009/02/university-to-rebuild-mendels-greenhouse/ Masaryk University to rebuild Mendel's greenhouse | Brno Now]
* [http://www.mendelmuseum.muni.cz/en/ Mendel Museum of Genetics]
* [http://www.mendelweb.org/Mendel.html Mendel's Paper in English]
* [https://www.ncbi.nlm.nih.gov/Omim/ Online Mendelian Inheritance in Man]
* [https://web.archive.org/web/20080419020741/http://biology.clc.uc.edu/Fankhauser/Travel/Berlin/for_web/Mendel_in_Brno.html A photographic tour of St. Thomas' Abbey, Brno, Czech Republic]
* [https://sciencefromthestacks.library.villanova.edu/rediscovering-gregor-mendel-os Villanova University Mendel Collection]

{{History of biology}}
{{Portal bar|Austria|Biology|Catholic Church|Christianity|Czech Republic|Mathematics}}
{{Authority control}}

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[[Category:Gregor Mendel| ]]
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