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Reproduction in Spinacia Oleracea L

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Reproduction in Spinacia Oleracea L 0000 Reproductioni nSpinaci a oleraceaL . Ultrastructural aspects of pistil development, pollinationan dfertilizatio n • ••-• T a;:: K K. ""• '': rj '^HOGE.SCIIOOl Promotoren: dr. J.L. van Went, hoogleraar in de plantkunde dr. M.T.M. Willemse, hoogleraar in de plantkunde hh (92.01 (9if3 c1 H.J. Wilms REPRODUCTION IN SPINACIA OLERACEA L. Ultrastructural aspects of pistil development, pollination and fertilization. (with a summary in Dutch) PROEFSCHRIFT ter verkrijging vand e graad van doctor ind e 1andbouwwetenschappen op gezag vand e rector magnificus dr. H.C. vande rP Ias , hoogleraar ind eorganisch e scheikunde in hetopenbaa r te verdedigen op vrijdag 15 mei1981 , des namiddags tevie r uur ind e aula van de Landbouwhogeschool teWageningen . >V BIBLIOTHEEKL.H . 0 6 MEI 1981 ONTV. TIJDSCHR. ADM Vakgroep PIantencytologie en -morfologie, Landbouwhogeschool,Wageninge n -1981 . STELLINGEN I. Waargenomen structurele verschillen tussen antipoden, die aanwezig zijn in d e diverse embryozakken binnen de groep van dicotyle planten, berusten vooralo pverschi1le n in snelheid van ontwikkeling en zijn daarom niet soort-specifiek. Dit proefschrift. II. Bij spinazie wordt de kieming van pollenkorrels en de groei van pollenbuizen in stempel en stijl niet gereguleerd door de ennbryozak integenstel1in g tot de groei van de pollenbuis in de nucel1u s. Dit proefschrift. III. De transportweg waarlangs de voeding van zaadknoppen ge- schiedt varieert en is afhankelijk van het ontwikkelings- stadiurn . Dit proefschrift. IV. De 1ichtmicroscopischef1 uorescentietechnie k isnietgeschikt voor het exact bepalen van de weg waarlangs pollenbuizen groeien . V. Voortplantings-en/ofzaadzettingsonderzoeken , waarvoor ge- durende het hele jaar deels met behulp van kassen plant- materiaal moet worden gekweekt,z ij n wetenschappelij k niet verantwoord wanneer de seizoensinvloed daar niet bij be- trokken wordt. VI. De veroudering en het afsterven, die in monocarpe planten optreedt na de bloei en de vorming van vruchten en zaden, kan niet zonder meer verklaard worden door aan te nemen dat zaden een senescence factor produceren. Nooden, L.D., D.C. Rupp en B.D. Derman (1978). Nature 271: 35^-356. VII. "Door bestuiving worden stuifmee lkorre ls bi j d e eieellen ge- brachte nvind t bevruchting piaats".D edoo r deze formulering beoogde vereenvoudiging van de stof weegt niet op tegen de geintroduceerde onjuistheden.Uitdidactisc h oogpunt zou het daarombete rzij n bepaalde onderwerpen uitd ebiologi e in het basisonderwijs niet te behandelen. Meijs, P. (1981). Onderwijstelevisie N.O.T.,De nHaag , p. 41. VIM. Onderwijsmiddelen ter ondersteuning en stimulering van het individuele leerproces zijn lapmiddelen. " IX. Informa ti eove rn ieuw evak literatuu rclien ti nhandbib liotheke n voorhanden te zijn. Indien dergelijke informatie bij voort- during eerst maandenn ahe t verschijnen ter algemene kennis- name komt, is het aan te bevelen de daarvoor verantwoorde- lijke vakgroepen in hun financiele middelen ten behoeve van tijdschriftabonnementen te korten. X. Democratie is gedoemd ten gronde te gaan, enerzijds aan het onvermogen tot besturen, anderzijds aan een intolerante houding van bepaalde groeperingen na genomen besluiten. XI. De voor en tegens van de ''nationale kierenjacht" zijn nog onvoldoende onderzocht om nu reeds paniek te zaaien. Brunekreef, B. en J.S.M. Boleij (1981). Intermediair 17, nrs. 1-2: 1-7. H.J. Wilms Reproduction in Spinaoia oleraceae L. 15 mei 1981. Ter nagedachtenis aanmij n vader aan mijnmoede r aan Anja,Maarte n en Janneke VOORWOORD Het verschijnen van dit proefschrift schenktmijvee l voldoening. Velen hebben bijgedragen aan de tot standkoming ervan. Allen ben ik erkentelijk, enkelen wi1 ik hier noemen. Mijn ouders, die mij in staat hebben gesteld in een goed klimaatee nwetenschappelijk e opleidinn te volgen.Moeder ,s1echt s aan U kan ik mijn dankbaarheid overbrengen. Mijn promotoren, Prof.Dr. J.L. van Went en Prof.Dr. M.T.M. Willemse wil ik dankzeggen voor de inzet en tijd gegeven aan dit onderzoek.Jacque s, jouw heldere en kritischebenaderingvana f het eerste moment heeft mij duidelijk zicht doen houden. Jouw kern- achtige formuleringen hebben de manuscripten duidelijk positief beinvloed. Michiel,jou wkoms t op het laboratorium en je benader- ing van onderzoek hebben een zekere druk gelegd op de afronding van dit onderzoek.D egezamen]ijk e discussies heb ik als zeer be- vruchtend beschouwd. Tijdens zijn verblijfa 1 sgastmedewerke r heeft Prof.Dr. R.N. Kapi1ee nstimulerend e invloed gehad op hetopschrift stellen van de resultaten. Dear Prof. Kapil, I remember our discussions as valuable and pleasant. Ia m much indebted to you, especially for your inspiring way of correcting manuscripts. Alle medewerkers van de vakgroep PIantencytologiee n-morfo - logie dank ik voord ewaardevoll e suggesties en de prettige werk- sfeer. In het bijzonder wil ik de heer A.C. van Aelst bedanken. Adriaan, van jou heb ik de eerste EM-technieken geleerd en daar- naast heb je nog een periode uitstekende assistentie verleend. Ookd estudente n die in het kader van hun studie aanhe tonderzoe k hebben bijgedragen ben ik erkentelijk. De overige mensen hebben mij op verschi11ende manieren dui­ delijk bijgestaan. De heren G. van Geerenstein, J. Verburg en A. van Ommeren voor de verzorging van het piantmateriaa1, de heren T. Zaal en W. van Ooijen voor de hulpvaardigheid en kennis van zakenbijhe t vervaardigen en afwerken van het vele fotomateriaal en de heer E. Rothuisd i e te n alle tijde bereid is geweest assis­ tentie te verlenen.He tomvangrijk e tekenwerk is verricht door de heer A.B. Haasdijk. Allex, de kunstenaar, jouw kwaliteiten zijn aanleidingvoorsteed s meer gecompliceerde tekenopdrachten.Z ooo k deze omslag. De heer J.S. de Block heeft de engelse teksten ge- corrigeerd. Dank gaat ook uit naar Mw. R.J.J.R. Groot-Scholte en Mw. G.G. van de Hoef-van Espelo voor hun bijdrage aan het ver- zorgen van enkele manuscripten. Tenslotte bedank ik Anja voor het opgebrachte geduld. CONTENTS 1. INTRODUCTION !- 9 2. ARTICLES: H 1980 b Development and composition of the spinach ovule. Acta Bot.Neerl . 2^ (h): Ihl - 260 1981 a Ultrastructure of thedevelopin g embryosa c of spinach. Acta Bot.Neerl . 30 (1/2): 75- 9 9 1980a Ultrastructure of thestigm a and style in spinach in relation topolle n germination and pollen tube growth. Acta Bot.Neerl . 29.0) : 33- '• 7 1981 b Pollen tube penetration and fertilization in spinach. Acta Bot.Neerl . 30 (1/2): 101- 12 2 3. PISTIL DEVELOPMENT, POLLINATION AND FERTILIZATION AS PROCESS 123- 12 6 h. SAMENVATTING 127- 12 9 CURRICULUM VITAE INTRODUCTION 1. Regroductjon Plant speciesca nmultipl y asexual1y,orvegetativel yan dsexually , or generatively. Thegenerativ e propagation isaccompanie d bya changeo fth eploid y level,thediploi d nuclear phase is succeeded by thehaploi don ean dexpresse d inth ealternatio no fth egenera ­ tion inwhic h a sporophyte isfollowe d bya gametophyte. There ­ duction of theploid y levelb ymeiosi s implies a recombinationo f genes if fusion of nuclei occurs. Sexual reproduction maintains geneti c viabi1 ity . In angiosperms thehaploi d phase, represented inth egameto ­ phyte, ishighl y reduced. The separation of sexes already isex ­ pressed on thesporophyt e by theformatio n ofmacro-an dm icro - sporophylls. The macrosporophy11s form a stamen with a stigma, style and ovary.I nth eovul e inside theovary ,a megaspor e mother cell develops intoa nembry o saco f 7cells , the megagametophyte. The microsporophy11 is composed bya filament and theanther .I n the anthers many pollen mother cells are formed, each of which gives after meiosis k microspores. Each microspore develops into a three-celled microgametophyte, a vegetative cell and twomal e gametes. Some plant species producethe ir sporophyl1 s togethero n one organism (monoecious) or separated over two organisms(di ­ oecious). The male gametes are transported to thesynergid s andwil l fuse with theeg gcel l and thecentra l cell. Inangiosperm s this double fertilization results in theformatio n ofa zygote and endosperm. Subsequently amatur e seed develops. Seed formation means for practical breeders and growers successful generative multiplication. What happens andwha t cang owron g inth epro ­ cesses of pollination, fertilization and seedsetting hast ob e clarified before these processes can be regulated. This means that it is also necessary to study and analyse these processes morphologically and cytological1y, tocollec t more information on the pistil, pollen-pistil interaction and thegrowt h ofth e pollen tube toan d into theembry osac . 1.2. Pistil structure The pistil constistsofa stigma, style andovary , inwhic hon eo r more ovules with one female gametophyte each arepresent . The morphology of thestigm a shows agrea t diversity. Some comparative light microscopical studies (VASIL &JOHR I 196*0an d several, mainly ultrastructural studies presentdet ai le dfeature s of some plants (KONAR & LINSKENS 1966a). On the basis of a study of the morphology of the stigma surface, the amount of secretion and the nature of the surface, cells of almost 1000 species of about 900 genera of 250 families (HESLOP-HARRISON et al. 1975), HESLOP-HARRISON & SHIVANNA (1977) give a general classification. The main classification dividesth estigma s in types with copious fluidse cr et ion , the wet stigma s and types withlimitedsecretion, the dry stigmas. The morphology and anatomy of the style is very variable. During development many changes take place. In relation to its transmitting function the style has either a canal or a solid core of transmitting tissue. A canal is lined with a glandular epidermis. Sucha styl e is of the open type, which is mostly pre­ sent in monocotyledons. Styles with a solid core of transmitting tissue.thesolid type, are regularindicotyledons . In som eplant s there is a half closed type in which the transmitting tissue is limited only to one side of the stylar canal (VASIL 6 J0HRI 1964; VASIL 197*0• Since the first description of the female gametophyte by STRASBURGER (1884) a large number of species have been studied, mainl yb y 1ightmicroscopica 1 methods.Th eresu1t s of these studies are well reviewed (MAHESHWARI 1950, 1963;RU TISHAUSE R 1969).
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