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Ontogenetic Studies on the Determination of the Apical Meristem In Ontogenetic studies on the determination of the apical meristem in racemose inflorescences D i s s e r t a t i o n Zur Erlangung des Grades Doktor der Naturwissenschaften Am Fachbereich Biologie Der Johannes Gutenberg-Universität Mainz Kester Bull-Hereñu geb. am 19.07.1979 in Santiago Mainz, 2010 CONTENTS SUMMARY OF THE THESIS............................................................................................ 1 ZUSAMMENFASSUNG.................................................................................................. 2 1 GENERAL INTRODUCTION......................................................................................... 3 1.1 Historical treatment of the terminal flower production in inflorescences....... 3 1.2 Structural understanding of the TF................................................................... 4 1.3 Parallel evolution of the character states referring the TF............................... 5 1.4 Matter of the thesis.......................................................................................... 6 2 DEVELOPMENTAL CONDITIONS FOR TERMINAL FLOWER PRODUCTION IN APIOID UMBELLETS...................................................................................................... 7 2.1 Introduction...................................................................................................... 7 2.2 Materials and Methods..................................................................................... 9 2.2.1 Plant material..................................................................................... 9 2.2.2 Measurements................................................................................... 9 2.2.3 Statistical analysis.............................................................................. 10 2.3 Results.............................................................................................................. 12 2.3.1 Ontogeny of the umbellets................................................................ 12 2.3.2 Measurements................................................................................... 12 2.4 Discussion ......................................................................................................... 17 2.5 Summary........................................................................................................... 19 2.6 Appendix........................................................................................................... 20 3 OPEN AND CLOSED INFLORESCENCES: MORE THAN SIMPLE OPPOSITES.................... 22 3.1 Introduction...................................................................................................... 22 3.2 A determinate inflorescence meristem does not imply a terminal flower....... 22 3.3 Histology of IMs support two kinds of open inflorescences............................. 26 3.4 What represses terminal flower formation in MC inflorescences?.................. 30 3.5 Concluding remarks.......................................................................................... 34 3.6 Summary........................................................................................................... 34 4 ONTOGENETIC COURSE AND SPATIAL CONSTRAINTS IN THE APPEARANCE AND DISAPPEARANCE OF THE TERMINAL FLOWER IN INFLORESCENCES....................... 36 4.1 Introduction...................................................................................................... 36 4.2 Material and Methods...................................................................................... 38 4.2.1 Plant material..................................................................................... 38 4.2.2 Measurements................................................................................... 41 4.2.3 Parameter calculation........................................................................ 43 4.2.4 Inflorescence bud age........................................................................ 45 4.2.5 Statistical analysis.............................................................................. 46 4.3 Results.............................................................................................................. 46 4.3.1 Inflorescence ontogeny by systematic groups .................................. 46 4.3.1.1 Berberidaceae..................................................................... 46 4.3.1.2 Papaveraceae-Fumarioideae .............................................. 49 4.3.1.3 Rosaceae............................................................................. 57 4.3.1.4 Campanulaceae....................................................................62 4.3.2 Comparison between closed and open inflorescences...................... 68 4.3.2.1 T test for the initial and final states of the meristems.........68 4.3.2.2 Correlation analysis............................................................. 68 4.4 Discussion..........................................................................................................72 4.4.1 Spatial and shape requisites of the inflorescence meristem (IM) for terminal flower (TF) production.................................................... 72 4.4.2 One common ontogeny for closed inflorescence and at least two ontogenies in open inflorescences............................................. 73 4.4.3 Histophysiological correlates............................................................. 77 4.4.4 Concluding remarks........................................................................... 79 4.5 Summary........................................................................................................... 80 4.6 Appendix........................................................................................................... 82 5 GENERAL CONCLUSIONS........................................................................................... 92 6 REFERENCES.............................................................................................................. 94 ACKNOWLEDGEMENTS...............................................................................................105 SUMMARY SUMMARY OF THE THESIS This thesis presents a comparative developmental study of inflorescences and focuses on the production of the terminal flower (TF). Morphometric attributes of inflorescence meristems (IM) were obtained throughout the ontogeny of inflorescence buds with the aim of describing possible spatial constraints that could explain the failure in developing the TF. The study exposes the inflorescence ontogeny of 20 species from five families of the Eudicots (Berberidaceae, Papaveraceae-Fumarioideae, Rosaceae, Campanulaceae and Apiaceae) in which 745 buds of open (i.e. without TF) and closed (i.e. with TF) inflorescences were observed under the scanning electron microscope. The study shows that TFs appear on IMs which are 2,75 (se = 0,38) times larger than the youngest lateral reproductive primordium. The shape of these IMs is characterized by a leaf arc (phyllotactic attribute) of 91,84° (se = 7,32) and a meristematic elevation of 27,93° (se = 5,42). IMs of open inflorescences show a significant lower relative surface, averaging 1,09 (se=0,26) times the youngest primordium size, which suggests their incapacity for producing TFs. The relative lower size of open IMs is either a condition throughout the complete ontogeny (‘open I’) or a result from the drastic reduction of the meristematic surface after flower segregation (‘open II’). It is concluded that a suitable bulge configuration of the IM is a prerequisite for TF formation. Observations in the TF-facultative species Daucus carota support this view, as the absence of the TF in certain umbellets is correlated with a reduction of their IM dimensions. A review of literature regarding histological development of IMs and genetic regulation of inflorescences suggests that in ‘open I’ inflorescences, the histological composition and molecular activity at the tip of the IM could impede the TF differentiation. On the other side, in ‘open II’ inflorescences, the small final IM bulge could represent a spatial constraint that hinders the differentiation of the TF. The existence of two distinct kinds of ontogenies of open inflorescences suggests two ways in which the loss of the TF could have occurred in the course of evolution. 1 ZUSAMMENFASSUNG ZUSAMMENFASSUNG Die Dissertation beinhaltet eine vergleichende Studie zur Entwicklung von Blütenständen, wobei der Schwerpunkt auf der Bildung bzw. dem Fehlen von Endblüten (EB) liegt. Morphometrische Veränderungen, die während der Ontogenie der Blütenstandsknospen am Infloreszenzmeristem (IM) auftreten, wurden erstmals ermittelt und quantifiziert, um eventuelle räumliche Zwänge, die das Ausfallen der EB erklären könnten, zu beschreiben. Die Studie umfasst die Infloreszenzontogenie von 20 Arten aus fünf Familien der Eudicotyledonae (Berberidaceae, Papaveraceae-Fumarioideae, Rosaceae, Campanulaceae and Apiaceae) mit offenen (ohne EB) und geschlossenen (mit EB) Blütenständen. Sie basiert auf der Analyse von 745 Infloreszenzknospen unter dem Rasterelektronmikroskop. Die Ergebnisse zeigen, dass sich Endblüten nur an Infloreszenzmeristemen entwickeln, die 2,75 (se = 0,38) Mal so groß sind wie ihr jüngstes Seitenprimordium. Ihre Form wird weiterhin von einem Primordienansatz von 91,84° (se = 7,32) und einer Wölbung von 27,93° (se = 5,42) bestimmt. Infloreszenzmeristeme offener Blütenständen zeigen dagegen mit nur 1,09-facher (se = 0,26) Größe des jüngsten Seitenprimordiums
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