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Honderd Jaar Mendel / HONDERDJAA R MENDEL Ontwikkeling van de erfelijkheidsleer van 1865 tot 1965 A pudooI 1965 Centrum voor Landbouwpublikaties en Landbouwdocumentatie Wageningen f. r4 o T< r Uitgaven verschenen naar aanleiding van vorige symposia, georganiseerd door de Biologische Raad vand e Koninklijke Nederlandse Akademie van Wetenschappen: Evolutie. De huidige stand van het vraagstuk Utrecht, Het Spectrum, 1960, 208 blz. Voordrachten gehouden tijdens het symposium Evolutie op 20 en 21 november 1959 te Amsterdam Op leven en dood. Problemen rondom de chemische en biologische bestrijding van plagen 2e Druk. Wageningen, Centrum voor Landbouwpublikaties en Landbouwdocumentatie, 1965, 208 blz. Voordrachten gehouden tijdens het symposium Op Leven en Dood op 31 maart en 1 april 1964 te Amsterdam BIBLIOTHEEK OER LANDBOUw HCX;ESCHOOL WAGENINGEN. Druk: N.V. Drukkerij Vada, Wageningen Copyright: Pudoc, Centrum voor Landbouwpublikaties en Landbouwdocumentatie, Wageningen INHOUD Voorwoord Prof. Dr.J . Lever Voorzitter vande Biologische Raad vande Koninklijke Nederlandse Akademie van Wetenschappen 7 Een eeuw erfelijkheidsleer Prof. Dr. S.J . Geerts GenetischLaboratorium van de Katholieke Universiteitte Nijmegen 9 De tijd van Mendel Prof. Dr. H. Engel Zoölogisch Museumvan deUniversiteit van Amsterdam 27 Wie was Mendel? Dr. J. Th. V. Meijknecht o.s.a. Gymnasium Augustinianum te Eindhoven 44 De herontdekking Prof.Dr .J . Heimans Emeritushoogleraar vande Universiteit van Amsterdam 62 Erwten toen en nn Prof.Dr .Ir .S .J . Wellensiek AfdelingTuinboutvplantenteelt van deLandbouwhogeschool te Wageningen 81 Mutatie Prof. Dr. F. H. Sobels Laboratorium voor Stralengenetica van de Rijksuniversiteitte Leiden 93 Mendelisme en micro-organismen Dr. A. Rörsch Medisch Biologisch Laboratorium, Rijksverde- digingsorganisatie TNO teRijswijk 112 Het genbegrip Prof. Dr. G. A. van Arkel Genetisch Instituut vande Rijksuni­ versiteitte Utrecht 128 Erfelijkheid en veredeling Prof. Dr. R. Prakken Afdeling Erfelijkheidsleer van de Land­ bouwhogeschoolte Wageningen 147 Het mendelisme en de mens Prof. Dr. M. Siniscalco Instituut voor Anthropogenetica van de Rijksuniversiteitte Leiden 168 Literatuur 192 Voorwoord PROF. DR. J. LEVER De Biologische Raad is als advies-lichaam verbonden aan deKo ­ ninklijke Nederlandse Akademie van Wetenschappen en functio­ neert voorts als contact-organisatie van alle Nederlandse weten­ schappelijke verenigingen op biologisch gebied. Dit laatste brengt tevensme t zich mede datd eBiologisch e Raad van tijd tot tijd symposia organiseert, gewijd aanonderwerpe n van algemeen biologische betekenis. Tot nu toe vonden een drietal symposia plaats. In ditboe k zijn delezinge n opgenomen vanhe to p2 0e n2 1 april 1965 in het Koninklijk Instituut voor de Tropen te Amsterdam gehouden derde symposium onder detite l 'Honderd jaar Mendel', waarin de aandacht wasgerich t op deontwikkelin g died e erfelijk­ heidsleer heeft doorgemaakt sinds Gregor Mendel in 1865 de re­ sultaten van zijn bekende kruisingsproeven bekend maakte. De Biologische Raad prijst zich gelukkig datoo kd espreker sva n dit symposium bereid waren hunmanuscripte n voor gemeenschap­ pelijke publikatie af te staan. Wiedi t boek doorleest, zal getroffen worden door de rijk geschakeerde deskundigheid van de sprekers, waardoor naast de figuur van Mendel niet alleen de geschiedenis maar ook de huidige problematiek van de erfelijkheidsleer op bij­ zonder heldere en waardevolle wijze wordt belicht. De suggestie tot het houden van dit symposium werd reeds in 1959gedaa n door pater Dr.J . Th.V . Meijknecht enkwa m nader­ hand via de Nederlandse Genetische Vereniging in de Biologische Raad. Deze stelde in 1964ee nspecial e voorbereidingscommissie in onder voorzitterschap vanProf . Dr. C.L .Rümke , hoogleraar ind e erfelijkheidsleer aan de Rijksuniversiteit te Utrecht en bestuurslid van de Biologische Raad. Prof. Rümke heeft zichme tgroo t enthousiasme voor de totstand­ koming en het slagen van dit symposium ingezet. Helaas heeft hij door zijn overlijden op 17 november 1964 de voltooiing niet meer kunnen meemaken. Voor zijn stuwkracht en in het bijzonder ook voor de evenwichtigheid van het programma zijn wij zeer grote dank verschuldigd. De verwachting is gewettigd dat dit boek, evenals de beide vorige, in brede kring de aandacht zal trekken. Eeneeu w erfelijkheidsleer PROF. DR.S .J . GEERTS Het testament van Mendel, in aanwezigheid van een veertig getuigen mondeling verledenop 8 februari en 8 maart 1865 enin 1866 schriftelijk bekrachtigd onder de titel 'Versuche überPflanzen-Hybriden', werdeerst in 1900 doorhet triumviraat De Vries, Correns en Von Tschermak geopenden daarmede aan de vergetelheidontrukt. Voordiencirculeerden wel enkele vage geruchtenover een verborgen schat in boekenalsFocke's 'Die Pflanzenmisch- linge', maar eerstde notariëleopening welke HEIMANS in dit symposium uit­ voerigbespreekt, deed vermoeden welke erfenis ons werd nagelaten. Nu, een eeuw na deze testamentaire beschikking, past het depersoon van de erflaterte belichten (MEIJKNECHT), de geschiedenis van zijn tijd teschetsen (ENGEL) en de verschillende erfgoederen op hun waarde tetaxeren. Deze erfgoederen, de gehele boedel metal het roerend en onroerendgoed, zijn aanzienlijk uitgebreid in de 65jaren, dat de internationale genetische gemeenschap deze in exploitatie heeft genomen. De inboedel bestond in het testament slechts uitPisu m (Erwt) en Phaseolus (Boon) en werdin een later neer­ gelegdebeschikking nog aangevuld met het heelwat ongemakkelijker meubel Hiëracium (Havikskruid). Hoe Pisum tot eenmodern ameublement is ge­ worden, bespreekt WELLENSIEK. Het roerendegoed- machines en apparaten - was bij Mendel wellicht beperkt tot enige pincetten, maar is inmiddels uitgedijd totdoeltreffende uitrustingen ten dienste van landbouw engenees­ kunde, zoals PRAKKEN en SINISCALCO uiteenzullen zetten. Hetonroerende goed, aanvankelijk niet meer dan een stuk veelbelovende grond, telt nu uit­ gestrekte gebieden, zoals het genbegrip ende mutatie, waaroverrespectievelijk VAN ARKEL en SOBELS rapporteren. Vooral met micro-organismen zijn gebieden ontsloten, dieongekende vergezichten opleverden, waar RÖRSCH enkelevan toont. De vraag ligt voor dehand of eendergelijk programma nog ruimte biedt voor een inleiding als deze. Hetantwoord is, dat het inde bedoeling ligt met het testament alsuitgangspunt enkele algemene beschouwingen te geven overde balans en deverlies- en winstrekening 1965. Nagegaan zal worden of wij geheel tevreden kunnen zijn over depopulari­ seringvan de erfenis; daarnaworden enkele doorMendel genoemde be­ schikkingen naderbesproken. {Menzie voor andere overzichten: Iltis, 1923; Blakeslee,1936; Babcock, 1950; Goldschmidt, 1951 en &rkle, 1963). 'Versuche über Pflanzen-Hybriden' Herlezing van het testament maakt duidelijk dat de zgn. 'wetten van Mendel' later naar voren zijn geschoven en dat hijzelf de nadruk legde opd e constantheid en de combinatieleer. De volgende punten in de Versuche vragen de aandacht: 1 Onder de door Mendel gespatieerde passages vindt men even­ min de reciprociteitsregel, als de uniformiteitsregel, de dominan- tieregel, of de splitsingsregel, wel echter de onafhankelijkheids- regel. Hoewel deze empirische regels ook voor Mendel betekenis hadden, legde hij het accent in zijn artikel toch anders. 2 Zo staat er met klem in de paragraaf over 'Die erste Generation der Hybride', dat is dus over de F2: "Uebergangsformen wurden bei keinem Versuche beobachtet". Alle passages, die door hem werden benadrukt, spreken over de constantheid van de kenmer­ ken en over de regels der combinatie. Het slot van de theoretische paragraaf 'Die Befruchtungs-Zellen der Hybriden', waarin hij de combinatieleer nog eens besprak, eindigt met: "Das Gesetz der Combinirung der differirenden Merkmale" (niet gespatieerd). In de passage over zijn Phaseolus(bonen)-kruisinge n - waar­ onder de soortskruising Ph. vulgaris (slaboon), met witte bloemen en zaden, met Ph. multiflorus (pronkboon), met purperrode bloe­ men en zwartgevlekte rose zaden - plaatst hij de voorzichtige opmerking: "Es wird die Aufgabe weiterer Versuche sein, zu ermitteln, ob dasfü r Pisum gefundene Entwicklungsgesetz auch bei den Hybriden anderer Pflanzen Geltung habe". En dan eindigt de paragraaf weer met een spatiëring: "Combinirung mehrerer selbständiger Farbenmerkmale". 3 In de 'Schluss-Bemerkungen' komt duidelijk tot uitdrukking dat Mendel een goed inzicht had in de wezenlijke bijdragen van eicel en pollenkorrel tot het nieuwe embryo en in de grote beteke­ nis die zijn leer van de combinaties van alternatieve kenmerken kon hebben voor de kennis van de cultuurgewassen. 4 Op verschillende plaatsen is het duidelijk dat hij deze ont- 10 wîkkelîngswet - de wet van de combinaties van alternatieve kenmerken - steunt op de hypothese van wat wij thans genen zou­ den noemen: "dass in den Fruchtknoten der Hybriden sovielerle i Keimzellen (Keimbläschen) und in den Antheren so vielerlei Pollenzellen gebildet werden, als constante (spatiëringva n Men­ del) Combinationsformen möglich sind, und dass diese Keim- und Pollenzellen ihrer inneren Beschaffenheit nach den einzelnen Formen entsprechen". Eiders: "Zurückführung des wesentlichen Unterschiedes in der Entwicklung der differirenden Zellelemente kann selbstverständlich nur den Werth einer Hypothese anspre­ chen". Hij denkt dus aan alternatieve celelementen. Hoewel impliciet, is de gedachtengang van Mendel duidelijk gesteund op constante elementen (genen) en alternatieven daarvan (allelen) in de (geslachts)cellen . Zijn accent ligt op de combinatieleer
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