Linking leaf initiation to the aerial environment PDF
Preview Linking leaf initiation to the aerial environment
Linking leaf initiation to the aerial environment When air temperature is not the whole story Andreas Savvides i Thesis Committee Promotor Prof. Dr L.F.M. Marcelis Professor of Horticulture & Product Physiology Wageningen University Co‐promotors Dr W. van Ieperen Assistant professor, Horticulture & Product Physiology Wageningen University Dr J.A. Dieleman Researcher, Wageningen UR Greenhouse Horticulture Wageningen University & Research Centre Other members Prof. Dr N.P.R. Anten, Wageningen University, The Netherlands Prof. Dr K. Steppe, Ghent University, Belgium Dr D. Vreugdenhil, Wageningen University, The Netherlands Dr M. Chelle, INRA, France The research was conducted under the auspices of the C.T. de Wit Graduate School of Production Ecology & Resource Conservation ii Linking leaf initiation to the aerial environment When air temperature is not the whole story Andreas Savvides Thesis submitted in fulfillment of the requirements for the degree of doctor at Wageningen University by the authority of the Rector Magnificus Prof. Dr. M.J. Kropff, in the presence of the Thesis Committee appointed by the Academic Board to be defended in public on Monday 3 November 2014 at 4 p.m. in the Aula. iii Andreas Savvides Linking leaf initiation to the aerial environment: when air temperature is not the whole story, 162 pages. PhD thesis, Wageningen University, Wageningen, NL (2014) With references, summaries in Dutch and English ISBN 978‐94‐6257‐113‐6 iv To My Beloved Wife and Son v vi Contents List of abbreviations viii Chapter 1 General Introduction 1 Chapter 2 Meristem temperature substantially deviates from air 11 temperature even in moderate environments: is the magnitude of this deviation species‐specific? Chapter 3 Leaf initiation is solely dependent on the apical bud 35 temperature even under large bud‐plant temperature differences Chapter 4 Phenotypic plasticity to altered apical bud temperature: more 51 leaves‐smaller leaves and vice versa Chapter 5 Impact of light on leaf initiation: a matter of photosynthate 81 availability in the apical bud? Chapter 6 General Discussion 103 References 121 Summary 135 Samenvatting 139 Acknowledgments 143 Curriculum Vitae 147 List of publications 149 Education certificate 151 Funding 153 vii Abbreviations DMC Dry matter content DW Dry weight eair Air vapour pressure Ebud Apical bud transpiration rate ebud Saturation vapour pressure at bud surface Ebud area Apical bud transpiration rate per bud‐contained leaf area es Saturation air vapour pressure FLA Final leaf area FW Fresh weight LAR Leaves appeared per unit of time (leaf appearance rate) LED Leaf expansion duration LEDdd Leaf expansion duration in thermal time LER Leaf expansion rate (mean) LERdd Leaf expansion rate (mean) normalized for thermal time LIR Leaves initiated per unit of time (leaf initiation rate) LIRDD Leaves initiated per unit of thermal time LL Leaf length LUR Leaves unfolded per unit of time (leaf unfolding rate) LW Leaf width (maximum) PPFD Photosynthetic photon flux density RH Relative humidity (%) RLW Net absorbed longwave radiation (> 2800 nm) Rnet Net radiation absorbed by a body (0‐100 μm) RSW Shortwave radiation (< 2800 nm) SAM Shoot apical meristem SLA Specific leaf area Tair Air temperature Tbase Base temperature Tbud Shoot apical bud temperature Tceiling Temperature of the glass ceiling of the climate room Tleaf Leaf temperature Tmeristem Shoot apical meristem temperature Tplant Plant temperature U Wind speed VPD Vapour pressure deficit VPDbud‐air Vapour pressure difference between bud and air δbl Boundary layer thickness viii Chapter 1 General Introduction In higher plants, establishment, growth and reproduction are primarily dependent on the continuous activity of two different groups of undifferentiated cells, the root and shoot meristems. Shoot and root meristems are driving the above‐ and below‐ ground organ generation respectively (Barlow 1989). In indeterminate plant species the shoot apical meristem (SAM) is continuously producing shoot modules, i.e. the phytomeres (Barthélémy and Caraglio 2007). In Cucumis sativus L. (cucumber) plants, for example, a phytomer during the vegetative stage mainly consists of a leaf, an internode, an axillary meristem and a tendril while during the generative phase, phytomeres additionally consist of flower meristems. The SAM is, hence, the fountain and simultaneously the architect of the shoot. The SAM is a dome of cells (Fig. 1) usually surrounded by the already successively formed and folded primordial leaves. This creates a distinct structure that resides on the top of the shoot, the apical bud (Fig. 2). The formation of a new phytomer on the shoot is presignified when a new leaf primordium is initiated (projected) on this dome (Fig. 1). The fundamental importance of leaf initiation for plant growth and development led to the in‐depth, from cell‐to‐molecule, exploration of this process and the unravelling of its complex component‐ mechanisms (e.g. Lyndon 1994; Fleming et al. 1997; Ha et al. 2010; Besnard et al. 2011). Leaf initiation is taking place through the continuous proliferation of pluripotent cells in the SAM and the synchronous transition in the fate of a group of these pluripotent cells to determinate cells (Byrne 2012). The change in fate is associated with changes in gene expression and new patterns of cell division and expansion (Golz 2006). As these cells proliferate, new axes of growth are established lateral to the SAM resulting in an outgrowth (leaf primordium) from 1 Chapter 1 the flanks of the SAM (Golz 2006). The direction of this outgrowth and thus the positioning of the new leaf primordium on the SAM in relation to the earlier initiated primordia (i.e. phyllotaxis) are basically determined by auxin gradients (Reinhardt et al. 2003). Briefly, auxin once in the SAM, is absorbed by the existing developing primordia which are acting as auxin sinks depleting auxin from the surrounded tissue (Reinhardt et al. 2003). Therefore, auxin accumulates in the region of the SAM furthest from the previously formed primordia and, as a consequence, when auxin passes a critical threshold in this region a new primordium is initiated (Golz 2006). Consequently, leaf initiation and its spatial arrangement are determined by a complex signaling network between the SAM and the earlier initiated leaf primordia (Ha et al. 2010). While the spatial pattern of leaf initiation is mainly a matter of intrinsic plant decisions and less a matter of extrinsic (environmental) cues (Kuhlemeier 2007), the rate in which the process of leaf initiation is repeated is highly dependent on the environment (e.g. Hussey 1963a; Granier et al. 2002). Fig. 1. Stereo‐microscopic image of the shoot apical meristem (SAM), the latest (P1) and the earlier (P2) initiated leaf primordia after the dissection of the earlier initiated primordial leaves and tendrils in a cucumber plant. The scale bar represents 0.1mm. Leaf initiation rate (LIR; number of leaves initiated per day) is a widely‐ used measure of the number of leaves as well as the number of phytomeres 2
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