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Microspore embryogenesis: reprogramming cell fate from pollen to embryo development
Hui Li, - \ 2014
Wageningen University. Promotor(en): Gerco Angenent, co-promotor(en): Kim Boutilier. - Wageningen : Wageningen University - ISBN 9789462570702 - 224
stuifmeel - embryogenese - embryonale ontwikkeling - biologische ontwikkeling - plantenontwikkeling - in vitro kweek - plantenembryo's - brassica napus - pollen - embryogenesis - embryonic development - biological development - plant development - in vitro culture - plant embryos

Microspore embryogenesis is an expression of plant cell totipotency that leads to the production of haploid embryos. Besides being a widely exploited plant breeding tool, microspore embryogenesis is also a fascinating system that can be used to obtain a deeper mechanistic understanding of plant totipotency. This thesis aims to provide more insight into the process of microspore embryogenesis, from the formation of embryogenic cells to the outgrowth of differentiated embryos.

In Chapter 1 background information is provided on the various aspects of Brassica napus microspore culture and plant development that intersect with the topics that are studied in this thesis. Emphasis is placed on the basic requirements and limitations for successful microspore embryo culture, as well as on the roles of the plant hormone auxin and epigenetic regulation in the development of plant embryos, during both zygotic and in vitro embryo development.

Chapter 2 reviews the recent advances that have been made in understanding the developmental and molecular changes that take place during microspore embryogenesis in model systems. The commonly reported cellular changes associated with the establishment of embryo cell fate are summarized and evaluated. The subsequent differentiation of the embryo is also discussed, specifically, what is known about the establishment of polarity, with emphasis on the importance of exine rupture as a positional clue, and the processes that influence meristem maintenance during culture. Finally, the studies on the molecular changes during microspore embryo induction are put into context of male gametophytic development. Overall, the current perspective on microspore embryo initiation presents a landscape in which several routes can lead to the same final destination.

Stress treatments are widely applied to induce embryogenic growth in microspore culture. Chapter 3 explores the role of histone acetylation status in stress-induced microspore embryogenesis in Brassica napus. Inhibition of histone deactylases (HDACs) using the HDAC inhibitor trichostatin A (TSA), phenocopies the heat stress treatment that is normally used to induce embryogenic cell proliferation in B. napus microspore culture. Arabidopsis is recalcitrant for haploid embryogenesis, yet treatment with TSA also induced embryogenic cell divisions in this model species. Our observations suggest that the totipotency of the male gametophyte is kept in check by an HDAC-dependent mechanism and that the stress treatments used to induce haploid embryo development in culture impinge on this HDAC-dependent pathway. The repression of HDACs or HDAC-mediated pathways by stress and the accompanying changes in histone acetylation status could provide a single, common regulation point for the induction of haploid embryogenesis.

Chapter 4 builds on the knowledge developed in Chapter 3 on the role of HDAC proteins in plant totipotency. A wide variety of chemically distinct HDAC inhibitors was evaluated and additional inhibitors that enhance embryogenic cell induction and/or embryo yield were identified. One surprising observation was made during the course of this study: the initial donor microspore/pollen stage affects the quality of the embryo that is formed. In control cultures, embryos from progressively older stages of donor microspores/pollen became progressively compromised in their basal (axis region) region, characterized by a shift from normal embryos with apical (cotyledons) and basal (root) polarity to abnormal embryos with a reduced basal pole and ball-shaped embryos. These abnormal phenotypes could be partially complemented by treatment with HDAC inhibitors, which promoted growth of the basal region of the embryo. Progressive enhancement of embryo basal identity was accompanied by enhanced and broadened expression of the DR5 auxin response reporter. The embryo phenotypes observed in control and HDAC inhibitor treated microspore cultures are similar to the phenotypes induced by altered expression of the Arabidopsis TOPLESS (TPL)/HDAC19/BODENLOS (BDL) repressor complex, which acts to restrict expression of the AUXIN RESPONSE FACTOR ARF5/MONOPTEROS (MP) to the basal region of the embryo during zygotic embryo development.

To understand why most embryogenic callus failed to develop further, we examined the transcriptome of globular-shaped embryos that have started to histodifferentiate and compared it with embryogenic callus. The transcriptome analysis showed that the expression of many genes that regulate (auxin-related) embryo patterning were downregulated in embryogenic callus compared to globular stage embryos. This result may simply reflect the lack of patterning in these embryos or might indicate a role of auxin-signalling in embryogenic callus formation.

Chapter 5 examines how embryo identity and patterning is established in two B. napus microspore embryo pathways, a zygotic-like pathway, characterized by suspensor and then embryo proper formation, and a pathway characterized by initially unorganized structures that lack a suspensor. We specifically asked the question: how can embryo patterning be established in the absence of an initial asymmetric division and in the absence of a suspensor, two key events in zygotic embryo development. Analysis of embryo fate (GRP) and auxin (PIN1, PIN7 and DR5) markers showed that embryo fate was established prior to cell division, and independent of subsequent division pattern. The suspensorless embryo program was marked by a transient auxin maximum, followed by establishment of the apical and basal poles at the globular stage, coincident with release of the embryo from the pollen exine. Unlike zygotic embryo development, polar auxin transport (PAT) was not required for embryo initiation or polarity establishment in this system. Suspensor embryos developed in a similar fashion as zygotic embryos, PAT was required for specification of the embryo proper from the suspensor. Haploid embryogenesis therefore follows at least two programs, a PAT-dependent program that requires embryo proper specification from the suspensor, and an alternative PAT-independent program marked by an initial auxin maximum.

In the final chapter, Chapter 6, the work presented in this thesis is put in context of the broader plant development field. The epigenetic regulation of developmental transitions that respond to stress and during pollen development are highlighted. A model is provided that histone acetylation levels mediated by HAT and HDAC regulate pollen fate.

Koolzaad - Brassica Napus / Eiwit en oliepad
PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2012
brassica napus - koolzaad - olieleverende planten - brandstofgewassen - planten met oliehoudende zaden - gewassen - akkerbouw - biobased economy - rape - oil plants - fuel crops - oilseed plants - crops - arable farming
Factsheet van het Eiwit en Oliepad met korte informatie over het gewas Koolzaad. Met het project Eiwit & Oliepad wil Innovatief Platteland samen met de gemeente Venray en andere partners het publiek de gelegenheid geven zich een beeld te vormen van de enorme multifunctionaliteit en de nog steeds verder te ontdekken mogelijkheden van in Europa te telen gewassen als grondstof voor de biobased economie
Teelthandleiding groenbemesters : bladkool (Brassica napus)
Timmer, R.D. ; Korthals, G.W. ; Molendijk, L.P.G. - \ 2011 2004 (2011)1 mei.
groenbemesters - brassica napus - teeltsystemen - zaaien - plantenziekten - plagen - onkruiden - nematoda - akkerbouw - teelthandleidingen - green manures - cropping systems - sowing - plant diseases - pests - weeds - arable farming - cultivation manuals
Bladkool is een speciaal op snelheid van beginontwikkeling en bladproductie geselecteerde vorm van winterkoolzaad. Het is vanouds een gewaardeerde groenbemester. Hoewel de beworteling van de bouwvoor niet intensief is geeft het een betere beworteling dan bladrammenas en gele mosterd. Daar staat tegenover dat bladkool niet geschikt is voor late zaai. De gebruiksmogelijkheden zijn beperkt, omdat bladkool een waardplant is voor zowel het bietencysteaaltje als knolvoet. Het is een lang en zeer bladrijk gewas en kan een grote massa vormen. Het bovengrondse deel is erg smakelijk voor dieren en wordt vaak bestemd voor veevoer. Geteeld als groenbemester (stoppelgewas) komt bladkool niet in bloei. Het is op de meeste gronden goed te verbouwen. Veengrond is hierop een uitzondering, terwijl lichte gronden alleen geschikt zijn als er voldoende kalk en magnesium in de bodem aanwezig is. De pH(-KCl) op deze grond moet minimaal 4,5 zijn
Ziektewering van bodemgebonden pathogenen en functionele diversiteit van de bodemmicroflora
Schilder, M.T. ; Postma, J. - \ 2009
Gewasbescherming 40 (2009)1. - ISSN 0166-6495 - p. 23 - 23.
gewasbescherming - bodembiologie - thanatephorus cucumeris - streptomyces - verticillium - streptomyces scabiei - suikerbieten - raphanus - brassica napus - bodemweerbaarheid - bodempathogenen - bodemmicrobiologie - plant protection - soil biology - sugarbeet - soil suppressiveness - soilborne pathogens - soil microbiology
Dit onderzoek richt zich op het vinden van micro-organismen of hun eigenschappen die correleren met ziektewering op verschillende bedrijfstypen en grondsoorten.
Cover crop-based ecological weed management: exploration and optimization
Kruidhof, H.M. - \ 2008
Wageningen University. Promotor(en): Martin Kropff, co-promotor(en): Lammert Bastiaans. - [S.l. : S.n. - ISBN 9789085049357 - 156
gewassen - onkruiden - onkruidbestrijding - gewas onkruid concurrentie - dekgewassen - plantenecologie - biologische landbouw - groenbemesters - allelopathie - secale - brassica napus - medicago sativa - oogstresten - allelochemicaliën - zaailingen - onkruidkunde - agro-ecologie - crops - weeds - weed control - crop weed competition - cover crops - plant ecology - organic farming - green manures - allelopathy - crop residues - allelochemicals - seedlings - weed science - agroecology
Keywords: organic farming, ecologically-based weed management, cover crops, green manure, allelopathy, Secale cereale, Brassica napus, Medicago sativa

Cover crop-based ecological weed management: exploration and optimization. In organic farming systems, weed control is recognized as one of the main production-related bottlenecks. System-oriented approaches for ecological weed management are needed and cover crops may form an important component of such an approach. Inclusion of cover crops in crop rotations introduces two important mechanisms through which the development of weed populations may be hampered. In late summer and autumn the successful introduction of cover crops can prevent growth and seed production of weeds through competition. In springtime, cover crop residues incorporated in the upper layer of the soil may suppress or retard weed development and growth due to, among others, allelopathic effects. The main focus of research was put on the weed suppressive effect of cover crop residue material in spring and particularly on identifying management options to maximize this effect. To better appreciate the potential of cover crop residue material the investigations were focused on three aspects, namely allelochemicals in the cover crop, the residence time of the residue-mediated inhibitory potential in the soil and the variability in inhibitory effects on receptor plants. The study was conducted with representatives of three plant families: Secale cereale L. (winter rye), Brassica napus L. (winter oilseed rape) and Medicago sativa L. (lucerne). Mechanical injuring of field grown cover crops enhanced the allelopathic activity per unit biomass. However, this increase was often just sufficient to compensate for loss of plant material resulting from damaging, implying the limited practical significance of damaging. Different options for pre-treatment and incorporation of cover crop residue material were compared and these were found to influence the size and persistence of an inhibitory effect on seedling emergence. Results were found to be cover crop specific. With regard to species’ sensitivity our results suggest that for inhibition of a receptor plant not just seed size is important. Only if the time course of sensitivity of the receptor plant matches with the time course of residue-mediated inhibitory potential, significant reductions in seedling establishment can be expected. In light of this, it was postulated that variation in synchronicity of receptor species’ sensitivity and potential residue effects may well explain the large degree of variation often noted in field studies of allelopathy.

Breder herbicidenpakket in koolzaad
Zeeland, M.G. van; Hoek, J. - \ 2007
Boerderij/Akkerbouw 93 (2007)1. - ISSN 0169-0116 - p. 1 - 1.
koolzaad - brassica napus - herbiciden - gewasbescherming - rape - herbicides - plant protection
Het middelenaanbod voor de onkruidbestrijding in winterkoolzaad beperkt. Enkele herbiciden blijken perspectiefvol, blijkt uit PPO-onderzoek
Teeltonderzoek aan koolzaad voor biobrandstof 2003-2006 : verslag veldproeven Ebelsheerd en Vredepeel
Geel, W.C.A. van; Borm, G.E.L. ; Beers, T.G. van; Floot, H. ; Meuffels, G.J.H.M. ; Mheen, H.J.C.J. van der; Verstegen, H.A.G. - \ 2007
Wageningen : PPO AGV (Rapport / PPO-AGV ) - 60
koolzaad - brassica napus - bio-energie - brandstofgewassen - rentabiliteit - cultuurmethoden - veldproeven - biobrandstoffen - biobased economy - akkerbouw - rape - bioenergy - fuel crops - profitability - cultural methods - field tests - biofuels - arable farming
In diverse delen van Nederland zijn initiatieven ontwikkeld om een productiekolom op te zetten voor de winning van biobrandstof uit koolzaad. In opdracht van het Hoofdproductschap Akkerbouw heeft PPO van 2003 t/m 2006 teeltonderzoek uitgevoerd in koolzaad. Het onderzoek richtte zich op de mogelijkheden om de rendabiliteit van de teelt te verhogen en op het verkennen van de koolzaadteelt op zandgrond. Naast de traditionele teelt van winterkoolzaad kregen de teelt en opbrengstpotentie van zomerkoolzaad aandacht, door middel van rassenonderzoek.
Peptide signalling in plant development: functional analysis of CLE ligands in Arabidopsis
Fiers, M.A. - \ 2007
Wageningen University. Promotor(en): W.J. Stiekema, co-promotor(en): C.M. Liu. - [S.l.] : S.n. - ISBN 9789085045793 - 144
plantenontwikkeling - liganden - signaalpeptide - arabidopsis - brassica napus - signaaltransductie - genen - plant development - ligands - signal peptide - signal transduction - genes
Factors affecting the density of Brassica napus seeds
Young, L. ; Jalink, H. ; Denkert, R. ; Reaney, M. - \ 2006
Seed Science and Technology 34 (2006)3. - ISSN 0251-0952 - p. 633 - 645.
brassica napus - zaadproductie - oliezaden - kwaliteit - seed production - oilseeds - quality - protein-concentration - chlorophyll fluorescence - oilseed rape - degradation - selection
Brassica napus seed is composed of low density oil (0.92 and higher density solids (1.3-1.45 Seed buoyant density may potentially be used to determine seed oil content and to separate seeds with different oil contents, however, we have found that seeds with the lowest buoyant density had lower than expected oil contents. It is proposed that the low oil content observed in the lowest density seed is a function of air gaps or pockets within the seed coat with sufficient volume to disrupt the linear relationship between oil content and density. The air gaps make density-based selection of seeds for oil content more difficult. Seed solid and liquid components are essentially incompressible with increased hydrostatic pressure but air gaps in seeds may be compressed or filled if density sorting is conducted in a pressurised apparatus. Experimental evidence shows that compression of air pockets with hydrostatic pressure or filling of air pockets with solvent simplifies the relationship between seed density and oil Content. In spite of applied pressure it was observed that pressure was not sufficient to eliminate the buoyancy of all seeds with air pockets. Seeds containing air pockets may have been immature or improperly filled during maturation. Interrupted maturation, premature maturation or desiccation may potentially interrupt storage compound accumulation and lead to morphological changes. Chlorophyll fluorescence was used in this study as an indicator of seed maturity. An inverse relationship between chlorophyll content and oil content was observed, but was not observed for chlorophyll content and density. This relationship between oil content, chlorophyll content and density suggests that the lowest and highest density seed groups were enriched in immature seed.
Innovations in microspore embryogenesis in Indonesian hot pepper (Capsicum annuum L.) and Brassica napus L.
Supena, E.D.J. - \ 2004
Wageningen University. Promotor(en): Evert Jacobsen, co-promotor(en): J.B.M. Custers. - [S.I.] : S.n. - ISBN 9085041333 - 131
capsicum annuum - spaanse pepers - brassica napus - microvermeerdering - embryogenese - antherencultuur - haploïdie - kiemcellen - plantenveredeling - chillies - micropropagation - embryogenesis - anther culture - haploidy - germ cells - plant breeding
Teelthandleiding koolzaad - onkruidbestrijding
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - onkruidbestrijding - cultuurmethoden - teelt - akkerbouw - teelthandleidingen - rape - weed control - cultural methods - cultivation - arable farming - cultivation manuals
In dit hoofdstuk van de teelthandleiding koolzaad wordt ingegaan op de bestrijding van onkruid in koolzaad.
Teelthandleiding koolzaad : nieuwe ontwikkelingen
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - olieleverende planten - zaadoliën - brandstofgewassen - plantaardige oliën - akkerbouw - toepassingen - biobased economy - teelthandleidingen - rape - cultivation - cultural methods - oil plants - seed oils - fuel crops - plant oils - arable farming - applications - cultivation manuals
In deze teelthandleiding wordt ingegaan op plantaardige olie, biologisch afbreekbare smeermiddelen en brandstof uit koolzaad voor de toekomst.
Teelthandleiding koolzaad : ziekten en plagen
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - gewasbescherming - plagen - plantenziektebestrijding - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - plant protection - pests - plant disease control - arable farming - cultivation manuals
Er zijn veel organismen die koolzaad in meer of mindere mate kunnen beschadigen en daardoor de zaadopbrengst in ongunstige zin kunnen beïnvloeden. In het onderstaande wordt een opsomming met een korte beschrijving van de meest voorkomende beschadigers gegeven, terwijl van de belangrijkste tevens de bestrijdingswijze zal worden aangegeven
Teelthandleiding koolzaad
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
koolzaad - brassica napus - teelt - teelthandleidingen - cultuurmethoden - akkerbouw - olieleverende planten - biobased economy - rape - cultivation - cultivation manuals - cultural methods - arable farming - oil plants
In de teelthandleiding koolzaad vindt u informatie over de teelt, oogst en bewaring van koolzaad.
Teelthandleiding koolzaad - botanische gegevens
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - brassicaceae - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - arable farming - cultivation manuals
Koolzaad en raapzaad behoren met diverse kool soorten, knol of raap, koolraap, mosterdsoorten, radijs, rammenas, tuinkers en nog vele andere gewassen tot de familie van de Cruciferae (Kruisbloemigen). Deze familie ontleent haar naam aan het feit dat de vier bloem, kroonbladeren duidelijk in een kruis staan.
Teelthandleiding koolzaad - teeltvoorwaarden en grondbewerking
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - grondbewerking - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - tillage - arable farming - cultivation manuals
In deze teelthandleiding wordt ingegaan op teeltvoorwaarden, grondsoort, vruchtopvolging, grondbewerking en zaaibed bij koolzaad.
Teelthandleiding koolzaad - zaad en zaaien
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - zaaien - zaden - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - sowing - seeds - arable farming - cultivation manuals
In deze teelthandleiding wordt ingegaan op rassenkeuze, zaaizaadontsmetting en zaadbehandeling, zaaizaadhoeveelheid en standdichtheid, zaaitijd, zaaitechniek en zaaien zonder grondbewerking bij koolzaad.
Teelthandleiding koolzaad - bemesting
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - kunstmeststoffen - stikstofmeststoffen - bemesting - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - fertilizers - nitrogen fertilizers - fertilizer application - arable farming - cultivation manuals
Het doel van de bemesting van koolzaad is een aanvulling aan de bodemvoorraad te geven, zodat het gewas steeds over voldoende voedingsstoffen kan beschikken en zo een ongestoorde groei kan geven. Ingegaan wordt op bemesting met fosfaat, kali, stikstof en overige mineralen.
Teelthandleiding koolzaad - oogst en bewaring
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - oogsten - opslag (planten) - oogsttijdstip - zwadmaaien - maaidorsen - akkerbouw - teelthandleidingen - rape - cultivation - harvesting - volunteer plants - harvesting date - windrowing - combine harvesting - arable farming - cultivation manuals
In de praktijk wordt winterkoolzaad voor het grootste gedeelte eerst gezwadmaaid en na een velddroogperiode uit het zwad gedorst. Daarnaast wordt ook van stam gedorst. De oogsttijd, de oogstmethode en de droging en bewaring van het zaad hebben grote invloed op opbrengst en kwaliteit van koolzaad.
Teelthandleiding koolzaad - producten uit koolzaad
Bernelot Moens, H.L. ; Wolfert, J.E. ; PPO Akkerbouw, Groene Ruimte en Vollegrondsgroente, - \ 2003 2003 (2003)1 juli.
brassica napus - koolzaad - teelt - cultuurmethoden - landbouwproducten - akkerbouw - teelthandleidingen - rape - cultivation - cultural methods - agricultural products - arable farming - cultivation manuals
In deze teelthandleiding worden de producten genoemd die uit koolzaad worden verkregen. Koolzaad word geteeld vanwege oliewinning, het wordt gebruikt voor dierlijke en menselijke consumptie en er zijn technische en industriële toepassingen.
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