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Stamen Development STAMEN DEVELOPMENT Lfo^ n Promotoren: dr.M.T.M .Willems e hoogleraari nd eplantkund e dr.J.L .va nWen t hoogleraari nd eplantkund e tJfiJO??o\l (0% C.J.Keijzer STAMEN DEVELOPMENT Proefschrift ter verkrijgingva n de graad van doctor in deLandbouwwetenschappen , op gezag van de rector magnificus, dr. C.C.Oosterlee , inhe t openbaar te verdedigen opvrijda g 17oktobe r 1986 des namiddags tevie r uur in de aula van deLandbouwuniversitei tt e Wageningen sW l^lb^l tAiliiisOi: .VKOGESCHOOL WAGENINGIM <• /^N0P2Q\, 10^^ Stellingen 1. Voor het anthere-openingsproces in diverse plantesoorten dienen de 0- vormig verdikte endotheciumcelwanden de loculus niet alleen open, doch in een vroegere fase ook dicht te buigen. Dit proefschrift. 2. Het is onjuist te veronderstellen dat recente wetenschappelijke literatuur de kennis van alle voorafgaande omvat. Bij herhaald refereren blijkt informatie verloren te gaan. Dit proefschrift. 3. De vraag vanuit de veredelingspraktijk naar mannelijke steriliteit in diverse gewassen rechtvaardigt een intensivering van het onderzoek naar de belnvloedbaarheid van het anthere-openingsproces. Dit proefschrift. 4. Ten onrechte zijn de bij micro- en macrogametogenese aan elkaar grenzende genetisch identieke haploide cellen tot voor kort niet met een snelle productie van inteeltlijnen in verband gebracht. C.J. Keijzer (1984). Landbouwkundig Tijdschrift 7/8: 21-26. 5. Mede door het gebruik de ogen te sluiten wanneer men de neus in een bos bloemen steekt, zijn diverse eenvoudig waarneembare fasen van het plantaardige voortplantingsproces slechts bij een relatief klein publiek bekend. 6. De mate waarin de Wageningse promovendus erin slaagt in het kwartier voorafgaand aan de openbare verdediging van haar of zijn proefschrift de inhoud ervan aan een ondeskundig publiek duidelijk te maken, dient mee te wegen in de beoordeling. 7. Wie een Van Leeuwenhoek-microscoop namaakt volgens de reconstructie van Walter en Via, kan onvoldoende nagaan waarom onze Delftenaar een betere kijk op de zaken had dan zijn tijdgenoten. W.G. Walter and H. Via (1968). The American Biology Teacher 8: 537-539. 8.Doo rbi jd etenorbanjo ,naas td ereed sbekend eaanpassingen ,d esnaar -e n velspanning tebenutte no m deklankbode m tegen dekete lt e spannen, ontstaat een instrument dat zelfs de koperblazers van beginnende Dixieland-orkestjeska noverstemmen . 9.Voo rhe toptimalisere nva nd esnelwandeltechnie kka nee nanalys eva n zoweld ebiped eal squadruped evoortbewegin gva nsommig eamfibiee ne n reptielenva nbelan gzijn . R.McNeil lAlexande r (1975).Th eChordates .Cambridg eUniversit yPress . 10.D einvoerin gva nelectronisch ehulpmiddele nvoo rscheidsrechterlijk e beslissingenkom td etennisspor tal sspektake lnie tte ngoede . F.L.M.Vossenaa re nC.J .Keijze r (1983)in :D eideee nligge no pstraat . J.W.Thompson ,Amsterdam ,p.37 . 11.D e gewoonte vanhe tAN P om eenkruispun t tegenwoordigal sknooppun tt e betitelen/ maakthe tverde roverbodi gme et edele nda td ezaa kdaa rwee r eensvas tzit . 12.He ti see nbeleef d gebaarva nhe tIO Co mbelangrijk ebeslissinge npa st e nemennada tee ndaari ngeinteresseer dpromovendu sd everdedigin gva nzij n proefschrifti nall erus theef tkunne nvoorbereiden . Wageningen, 17oktobe r198 6 C.J.Keijze r In herinnering aan mijn vader Dankwoord Gaarne maak ik van de gelegenheid gebruik on eenieder te bedanken die bijgedragen heeft aan het tot stand komen van dit proefschrift. Vrijwel alle medewerkers van de vakgroep Plantencytologie en -morfologie werden meerdere malen geraadpleegd om mij met hun uiteenlopende kundigheden advies of daadwerkeli jke hulp te geven. Ik wil hen alien vanaf deze bladzijde bedanken, slechts een momentopname uit een voortdurende samenwerking die ook in de toekomst de voornaamste pijler van een enthousiaste en productieve vakgroep dient te zijn. Slechts enkelen treft het lot hier bijzonderheden over hun relatie met ondergetekende gepubliceerd te zien. Hooggeleerde Willemse, mijn promotor, ik dank U allereerst voor het mij aanbieden van de betrekking waardoor dit proefschrift mogelijk werd. De verplichte contacten tussen een bioloog en een plantenveredelaar werden, na een gewenningsproces, een uiterst leerzame periode. Dankzij dit fundamentele proefschrift is een meer toegepast vervolg nog slechts een kwestie van fantasie gebruiken en invullen. Hooggeleerde Van Went, mijn co-promotor, ik dank U voor de mogelijkheid die U mij bood om enige tijd in Italie te kunnen werken. Ook waren de vele uren op Uw kamer tijdens de afifonding van dit proefschrift onmisbaar. Carmen Reinders, jouw - nog relatief korte - medewerking betrof de moeilijkste aspecten van ons onderzoek. Dankzij je enthousiasme en doorzettingsvermogen sloeg je je door vele negatieve, bruine resultaten van in-vitro experimenten heen. Het is geen toeval dat onze samenwerking pas in het laatste hoofdstuk naar voren komt. Het geeft daarmee het accent voor de toekomst aan. Professor Cresti, il loro modo di lavorare e organizzare e stato piu istruttivo per me di quello que pensavo possibile in tre mesi. Anche il loro corso "Palio per Olandesi e Americani" era benissimo. Dr. ir. H.J. Wilms, His, onze toegenomen samenwerking in zowel onderzoek als onderwijs heeft ons vakgebied voor de directe oplossing van landbouwkundige problemen toegankelijker gemaakt. Ir. A.H. Bouma ben ik erkentelijk omdat hij zonder aarzeling beaamde dat de aanwezigheid van een deskundig oud-01 tijdens de openbare verdediging van een proefschrift onontbeerlijk is. Contents Introduction 1 1.Hydratio n changes during anther development 3 C.J. Keijzer (1983). In: Pollen, biology and implications for plant breeding,edite d by D.L.Mulcah y and E.Ottaviano ,pp .197 - 201. Elsevier Biomedical,Ne w York, (changed). 2. Theprocesse s of anther dehiscence andpolle n dispersal. 1. The opening mechanism of longitudinally dehiscing anthers 9 C.J. Keijzer (1986). TheNe w Phytologist (inpress) . 3.Th eprocesse s of anther dehiscence andpolle n dispersal. 2. The formation and the transfer mechanism ofpollenkitt , cell wall development inth e locule tissues and a function of the orbicules inpolle n dispersal 31 C.J. Keijzer (1986).Th eNe w Phytologist (in press). 4. A comparison of anther development inmal e sterileAlo e vera and male fertileAlo e ciliaris 49 /, C.J. Keijzer andM . Cresti (1986). Annals of Botany (in press). 5.Functiona l morphological relationships inth e developing locule of Gasteria verrucosa 63 C.J. Keijzer and M.T.M. Willemse (1987). Submitted for publication. 6. The filament development of Gasteria verrucosa 89 C.J. Keijzer, I.H.S.Hoe k andM.T.M . Willemse (1987). Submitted forpublication . 7. Theprocesse s of anther dehiscence andpolle n dispersal. 3. The dehydration of the filament tip and the anther in some monocotylous species 107 C.J. Keijzer, I.H.S. Hoek andM.T.M .Willems e (1987). Submitted forpublication . 8.I n vitro culturean dintraspecifi ctransplantatio n of tetrads intoolde ranther s 123 M.C.Reinder san dC.J .Keijzer . Generalconclusion san ddiscussio n 135 Summary 145 Samenvatting 149 References 153 Curriculumvita e 165 Ifa chapte r ofthi sthesi s isuse d asa referenc e inth e text,th e abbrevation "ch."i s oftenused . Introduction A detailed knowledge of the development and dispersal of angiosperm pollen is important for both biologists and plant breeders. The first reports on this topic appeared as early as in the 17th century in the letters of Van Leeuwenhoek and the work of Grew. During the second half of the last century, the different stages of pollen development were discovered. Besides, the importance for plant breeders became evident after the (re-)discovery of the Mendel laws. Around the turn of the century French and German biologists published many works on the different steps of the anther dehiscence process, with a strong teleological approach. These works concerned light microscopical observations of living tissues together with simple, but ingenious micromanipulations. After 1930 the attention on the dehiscence process faded, whereas the light microscopical investigations on the generative reproduction processes continued. From the early 1960's the electron microscope and modern physiological techniques revealed many of the processes of microsporo- and microgametogenesis in great detail. These works mainly focussed on the pollen and tapetum development and were summarized in some excellent reviews. The mechanical processes in the stamen still received less attention in these years and most of the older works seemed to be forgotten, given the flaws and the call for research on this topic in some botanical standard works. Accordingly, detailed work on the entire stamen did not exist. In this thesis the classical and modern techniques were combined to investigate the stamen as a functional unit for pollen production and dispersal respectively. Chapter 1 HYDRATION CHANGES DURING ANTHER DEVELOPMENT C.J. Keijzer Summary The hydration changes of the anther tissues of Gasteria verrucosa are followed from meiosis to dehiscence. The water percentage of the locule wall is about constant during this development, in spite of remarkable histological changes. The water percentage of the locule contents is about constant during the microspore stages. Shortly before maturation the pollen grains take up water from the surrounding tissues. The water percentage of the locule wall decreases strongly at anthesis, that of the locule contents at dehiscence. Ventilation inside the flower bud leads to desiccation of the anthers by evaporation. Shortly before anthesis
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