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Morphological and Physiological Aspects of the Early Phases of Flower Bud Formation of Apple

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Morphological and Physiological Aspects of the Early Phases of Flower Bud Formation of Apple Morphological and physiological aspects of the early phases of flower bud formation of apple CENTRALE LANDBOUWCATALOGUS 0000 0751 3100 Promotor: dr. J. Tromp hoogleraar in de tuinbouwplantenteelt, in het bijzonder de overblijvende gewassen tJtJölZo1 , dO^f- Morphological and physiological aspects of the early phases of flower bud formation of apple F.A. Verheij Proefschrift ter verkrijging van de graad van doctor in de landbouw- en milieuwetenschappen, op gezag van de rector magnificus, dr. C.M. Karssen, in het openbaar te verdedigen op maandag 15 januari 1996 des namiddags te vier uur in de Aula van de Landbouwuniversiteit te Wageningen ï$o-^'^2': E;CLic/ n!:•:•:< : LM MD;» Ü V/UN1 v L-.RSiTEr r WAGKNÏNGEN CIP-DATA KONINKLIJKE BIBLIOTHEEK, DEN HAAG Verheij, F.A. Morphological and physiological aspects of the early phases of flower bud formation of apple / F.A. Verheij. - [S.l. : s.n.]. Fig., Tab. Thesis Wageningen. - With ref. - With summary in Dutch. ISBN 90-5485-484-7 Subject headings: flower bud formation; apple. Stellingen 1. Een korte plastochron en de aanwezigheid van een zeker minimum aantal bladvormingen in de knop zijn geen inducerende factoren in de bloemknopvorming van appel. Dit proefschrift 2. Het antagonisme tussen de groei en bloemknopvorming bemoeilijkt de bestudering van de bloemknopvorming bij appel in hoge mate. Dit proefschrift 3. Tussen het tijdstip van afsluiten van de groei, het begin van de bloemdifferentiatie en de bloemkwalitiet in het volgende voorjaar bestaat geen rechtstreeks verband. Dit proefschrift 4. Het feit dat polyaminen in alle tot nu toe onderzochte organismen zijn aangetroffen, en dat hun betrokkenheid in een groot aantal fysiologische processen is aangetoond, wijst er op dat polyaminen de ontwikkeling van planten beïnvloeden door middel van eenzelfde fundamenteel mechanisme in alle organismen. 5. Bij overblijvende gewassen verdient het dynamische karakter van de relatie tussen vorm en funktie meer aandacht. 6. Binnen populaties onderzoekers is statistische kennis doorgaans niet normaal verdeeld. 7. De in wervingsadvertenties veel gestelde eis van stress-bestendigheid wordt overbodig indien meer waarde wordt gehecht aan het plezier hebben in het werk. 8. Dat een natuurproduct, zoals mest, door toedoen van de mens op grote schaal het milieu bedreigt, toont aan hoezeer het evenwicht tussen mens en natuur is verstoord. 9. Het gebruik van de termen onderwijsinstituten" en onderzoeksscholen" werkt verwarrend. 10. De vrees voor een teloorgang van regionale verschillen bij de éénwording van Europa is ongegrond zolang een snoei-advies aan Nederlandse fruittelers: "verenkelen om verkaling en verdikking tegen te gaan", door Belgische collega's wordt vertaald met: "ontdubbel om ontbloting en aanzetting te voorkomen". Van Kessel, T. 1995. Vuistregels bij de snoei van appelbomen niet klakkeloos toepassen. Fruitteelt, 85: 11-13. 11. Buitensporige mobiliteit blijkt een kwaad dat zichzelf straft: het spoor loopt dood. 12. Ondanks de toename van de gemiddelde levensduur en de verkorting van de gemiddelde arbeidsduur, wordt de roep dat men tijd tekort komt steeds luider. Stellingen behorende bij het proefschrift: 'Morphological and physiological aspects of the early phases of flower bud formation of apple' F.A. Verheij Wageningen, 15januar i 1996 Abstract Verheij, F.A . 1996. Morphological and physiological aspects of the early phases of flower bud formation of apple. Dissertation Wageningen Agricultural Uni­ versity, Wageningen, The Netherlands. 148 pp.; English and Dutch summaries. For consistent yields in apple fruit production, knowledge of the factors affecting flower bud formation isrequired . The aimo fthi sstud y wast ogai nmor e insight in the role of endogenous factors in flower bud formation of apple. The effects oftemperature , appliedgibberelli n (GA4+7), thepresenc e of fruits, defolia­ tion and bending on the plastochron (the time interval between formation of suc­ cessive primordia by the meristem) and the number of appendages per bud at the start of floral differentiation were studied. Although most treatments distinctly affected flower bud formation, the plastochron was only affected by temperature anddefoliation . Considerable variation wasobserve d inth enumbe ro f appendages at the start of floral differentiation, which varied between cultivars, between experiments with thesam ecultiva r andbetwee n budpositions . Iti sconclude d that the length of theplastochro n and the number of appendages are not critical to the occurrence of flower bud formation. Defoliation inhibited flower bud formation and caused animmediat ereduc ­ tion in the soluble sugar content of the shoot. Bending slightly enhanced flower bud formation, or had no effect, and did not affect the soluble sugar level in the shoot, but increased starch content. The poor correlation between the assimilate level of the shoot and the number of flower buds formed suggests that it is not a main regulatory factor in flower bud formation. Itwa sfurthe r investigated whetherammoniu m affectsflower bu d formation of apple through affecting endogenous levels of polyamines (PAs). Ammonium and applied PAs have previously been shown to stimulate flower formation of apple. However, due to alac k of effect of ammonium onflower bu d formation in the present work, no conclusive evidence could be given. PA levels of buds did not greatly respond to treatments affecting flower bud formation (applied GA4+7 and shoot bending) despite the fact that both treatments did affect flower bud formation. PA levels did not show distinct changes during the supposed time of floralinductio n and floral differentiation. It is tentatively suggested that PAs are not major inductive stimuli of flower bud formation of apple. Keywords:ammonium , apple, arginine, assimilates,bu d morphology, defoliation, flower bud formation, gibberellin, Malus domestica Borkh., nitrate, plastochron, polyamines, shoot bending, shoot growth, temperature. Andpluck till time andtimes are done Thesilver apples of the moon Thegolden apples of thesun. (Yeats) Voorwoord Aan de totstandkoming van dit proefschrift hebben veel mensen een bijdrage geleverd en daarvoor wil ik iedereen hartelijk bedanken. Een aantal mensen wil ik in het bijzonder noemen. Allereerst ben ik mijn begeleider en promotor Prof. J. Tromp erkentelijk voor zijn stimulerende en kritische begeleiding, zowel tijdens de uitvoering van hetonderzoe k alsbi j het schrijven vanhe tproefschrift . Zijn begeleiding heeft een grote bijdrage geleverd aan de goede afronding van dit proefschrift. Oege Borsboom wasme tzij njarenlang epraktijkervarin g engrondig ekenni sva nd eteel t vanappelbome nee nwaardevoll ebro nva ninformati e diei kveelvuldi ghe bgeraad ­ pleegd. Oege, je was bovendien een zeer gezellige kamergenoot. Arjen van de Peppel ben ik zeer dankbaar voor zijn expertise en hulp bij de vele chemische analyses. Dankzij jou kennis van de apparatuur is het niet bij malen gebleven. Ruud Lamers, Henk Meurs en het overige kaspersoneel wil ik bedanken voor de goede verzorging van de planten, George Martakis voor zijn hulp bij de statis­ tische verwerking en Jan Peeters van Van Gaal voor de bemestingsadviezen. De studenten Hans de Wild, Marlijn Vos en Nicole van der Werf hebben op enthou­ siaste wijze meegewerkt aan dit onderzoek. Het verrichten van promotie-onderzoek heb ik als zeer leerzaam ervaren, nietallee n ophe tinhoudelijk e vlak, maaroo ko pvee l andereterreinen . Ikhe bme t name geleerd dat een goede werksfeer enorm belangrijk is voor het plezier inj e werk en daarmee de productiviteit. De vakgroepsmedewerkers wil ik bedanken voor de prettige werksfeer. Ik hoop in (een) volgende werkkring(en) weer zulke collega's aan te treffen! Mede door het ontbreken van uitzicht opwer k hebi kd e afgelopen tijd ook als erg zwaar ervaren. Mw. L. Drees van de Geldergroep wil ik van harte bedan­ ken voor het bieden van een luisterend oor, en de professionele hulp bij de (her­ oriëntatie op de arbeidsmarkt. Mijn ouders, Nicole en Alfred bedank ik voor hun betrokkenheid en steun, enoo k mijn vrienden dievoo rd e nodigehul pe n afleiding hebben gezorgd. Tenslotte, Henk, bedankt voor je geduld. Je vormde thuis een waardevol rustpunt. Contents 1. General introduction 1 2. The effect of temperature, applied gibberellin and the presence of fruits on shoot growth and flower bud formation 2.1. The effect on appendage formation of spur buds 17 2.2. The effect on morphological characteristics of the flower cluster 39 3. The effect of defoliation and bending on shoot growth, bud development and on assimilate levels of shoots in relation to flower bud formation 53 4. The effect of ammonium- and nitrate-nutrition on tissue arginine and polyamine content in relation to shoot growth and flowering 79 5. The role of arginine and polyamines in the process of flower bud formation. 95 6. General discussion 111 References 125 Summary 139 Samenvatting 143 Curriculum vitae 147 1. General introduction Flowering, the first step of sexual reproduction of plants, is of paramount importance to agriculture, horticulture, and plant breeding as it is a major factor determining yield. Not only for economic, but also for scientific reasons, the dramatic morphological andphysiologica l changes occurring during the transition of the meristem from vegetative to generative have been the topic of much research. In unravelling the mechanisms involved in the control of generative development, attention has mainly concentrated on herbaceous species, in parti­ cular those
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