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Comparative Analysis of Inflorescence Architecture In COMPARATIVE ANALYSIS OF INFLORESCENCE ARCHITECTURE IN AQUILEGIA SPECIES A Thesis Presented to the Faculty of California State Polytechnic University, Pomona In Partial Fulfilment Of the Requirements for the Degree Master of Science In Biological Sciences By Michael R. Speck 2021 SIGNATURE PAGE THESIS: COMPARATIVE ANALYSIS OF INFLORESCENCE ARCHITECTURE IN AQUILEGIA SPECIES AUTHOR: Michael R. Speck DATE SUBMITTED: Spring 2021 Department of Biological Sciences Dr. Bharti Sharma Thesis Committee Chair Professor of Biological Science __________________________________________ Dr. Valerie Mellano Professor of Plant Science __________________________________________ Dr. Paul Beardsley Professor of Biological Science __________________________________________ ii ABSTRACT The Aquilegia genus has undergone adaptive radiation over the last 1-5 million years. This has led to a wide array of diversity in flower size, plant height, and primary and secondary branching patterns of inflorescences. An inflorescence consists of flowers, branches, leaves, and bracts centered around the main stem. Some common inflorescence types in dicots and monocots include raceme, panicle, spike, thyrse, capitulum, and cymose. The Aquilegia genus is an example of a cymose inflorescence. Cymose patterns can be dichasial or monochasial and have primary, secondary, and tertiary branching. While a lot of recent effort has gone into the study of genes involved in the floral development (ABC model) in the Aquilegia, only a few studies have looked in detail into inflorescence development and intraspecific morphological differences in lateral organs such as bracts, and leaves. This study was aimed to track Aquilegia formosa and Aquilegia coerulea to understand heteroblasty in lateral organs and observe inflorescence development. I noticed conserved morphological patterns in simple bracts, trifoliate bracts, and leaves with/without a petiole at three to four main nodes on the stem in both species. I also observed some differences between the species; this included height, number of buds, the requirement of cold to flower, the ramification of inflorescence, and the number of days plants remain in the reproductive state. In the future, I suggest comparative morphological analysis including more species from this genus. iii TABLE OF CONTENTS SIGNATURE PAGE……………………………………………………………….……ii ABSTRACT……………………………………………………………………………..iii LIST OF TABLES………………………………………………………………...…….vi LIST OF FIGURES………………………………………………………………...…..vii CHAPTER 1: INTRODUCTION………………………………………………….……1 1.1 Inflorescence Types……………………...………………………………........1 1.2 (a) Review: Studies in Monocot (Grass) Model Systems………….………….7 1.2 (b) Review: Studies in Dicot Model Systems………………………….…….16 1.3 Aquilegia: A Dichasial, Basal Eudicot Model System…………..…………..23 CHAPTER 2: METHODS…………………………..…………………………………28 2.1 Growing Aquilegia Species……………………………………………….….29 2.2 Tracking Plants……………………………………………………………....30 2.3 Methods for Tracking 1-2-3 and 1-2-3-4 Patterns………………...…………32 2.4 Scanning Electron Microscopy Tissue Preparation for SEM…………….….42 CHAPTER 3: ANALYSIS………………………………………………………..……45 3.1 Analysis of A. formosa and A. coerulea……………………………..………45 3.2 Comparison of Primary Branches and Lateral Organs on Nodes in Stressed vs Non-Stressed Plant in A. coerulea in the 1-2-3 and 1-2-3-4 Patterns………..…..79 3.3 Analysis of Phyllotaxy of Primary Branches in A. formosa and A. coerulea…………………………………………...……………………..……91 iv 3.4 Scanning Electron Microscopy of Apical Meristems of A. coerulea and A. formosa………………………………………………………….……………….92 3.5 Analysis of Primary Branching in A. coerulea……………………..………..94 3.6 Tracking of A. formosa and A. coerulea………………………………...…...97 3.7 Differences between A formosa and A.coerulea……………….………….…98 CHAPTER 4: CONCLUSION AND FUTURE RESEARCH……………………...101 4.1 Conculsions from Research………………………………………………...101 4.2 Future Research……………………………………………………...…..…102 REFERENCES………………………………………………………………..……….106 v LIST OF TABLES Table 2.1. Steps involved in the dehydration of meristems for SEM……..…..…….…….43 Table 3.1. Comparison of leaf, bract, and petiole types in 1-2-3A and B pattern…….…46 Table 3.2. Comparison of leaf, bract, and petiole types in 1-2-3-4A and 1-2-3-4B….….53 Table 3.3. Comparison of most and least commonly found patterns….………….…...….60 Table 3.4. Comparison of leaf, bract, and petiole types in plants deviating from 1-2-3 and 1-2-3-4……………………………………………………………………………………68 Table 3.5. Comparison of frequencies of primary branching in A. coerulea…...…...……95 Table 3.6. Comparative analysis of A. coerulea and A. formosa…….…………….….....101 vi LIST OF FIGURES Figure 1.1. Monochasial cyme inflorescence diagram….……..……..……………………2 Figure 1.2. Dichasial cyme inflorescence diagram……….…….......……………………..3 Figure 1.3. Raceme inflorescence diagram………..……………..………...….…..………3 Figure 1.4. Thyrse inflorescence diagram……..……..……………..………..……………4 Figure 1.5. Panicle inflorescence diagram………...…………………..…..………………4 Figure 1.6. Spikelet inflorescence diagram……..…….………………...…..……………..5 Figure 1.7. Capitulum inflorescence diagram……..…………….………..…..…………...6 Figure 1.8. Meristem transitions in Zea mays……………..………………….…………..8 Figure 1.9. Meristem transitions in Brachypodium distachyon……..…….….………..…10 Figure 1.10. Meristem transitions in rice……..…...……………………..…..…………..12 Figure 1.11. Meristem transitions in wheat………………...…………..……………...…14 Figure 1.12. Meristem transitions in barley………..……..………………………...……15 Figure 1.13. Meristem transitions in Arabidopsis…….....…..…………………...………16 Figure 1.14. Meristem transitions in tomato…...…..….……...………………………….21 Figure 1.15. Stages of meristem transitions in Pisum sativum (Pea)……….……………23 Figure 1.16. Pictures of A. coerulea and A. formosa studied in this thesis …………….…24 Figure 1.17. Ramification of inflorescence in two Aquilegia species………….................................….......................................................................…25 Figure 1.18 Meristem transitions in A. formosa and A. coerulea ……..…..…….………26 Figure 2.1. Primary (1°) and secondary (2°) branching in A. coerulea and A. formosa ………………………………………...………………………….………………………29 vii Figure 2.2. Photographic tracking of Aquilegia species…….……..…………...……….31 Figure 2.3. A. coerulea ……...…………..…………..…..……...………………….……33 Figure 2.4. Observation 1 ………....……......…………………………………………....34 Figure 2.5. Observation 2 ……………………..…………………………………..……..35 Figure 2.6. Observation 3………….………………...………………….……………….36 Figure 2.7. Observation 4 ……………….……………….…………………...………….37 Figure 2.8. Observation 5 ……………….………………………….…………...…….…39 Figure 2.9. Observation 6……….……………...…………………...……………...……40 Figure 2.10. Observation 7…………….……...…...……….……………………………41 Figure 2.11. Observation 8………….……………………………...……………………42 Figure 2.12. Equipment used in the analysis of the SAM……………..……..…….…..…44 Figure 3.1. Pictures of node 1 in 1-2-3A pattern in A. coerulea..............,,,,,.....................47 Figure 3.2. Pictures of node 1 in 1-2-3B pattern in A. coerulea………..…..……………47 Figure 3.3. Pictures of node 1 in 1-2-3B pattern in A. formosa…………….…………….48 Figure 3.4. Pictures of trifoliate leaves and bracts at node 2 in 1-2-3A pattern in A. coerulea…..........................................................................................................................49 Figure 3.5. Pictures of trifoliate bracts leaves at node 2 in 1-2-3B pattern in A. coerulea…………………………………………….…………………………………….50 Figure 3.6. Pictures of trifoliate leaves and at node 2 in 1-2-3B pattern in A. formosa………….……………………………………………………………………….51 Figure 3.7. Pictures of trifoliate compound leaves at node 3 in 1-2-3A pattern in A. coerulea……………..……………………………………………………………………51 viii Figure 3.8. Pictures of trifoliate sessile leaves at node 3 in the 1-2-3B pattern in A. coerulea…………………………………………………………………………………..52 Figure 3.9. Pictures of trifoliate sessile leaves at node 3 in the 1-2-3B pattern in A. formosa…………………………………………………………………………………..53 Figure 3.10. Pictures of node 1 in 1-2-3-4A pattern in A. coerulea…….………………..54 Figure 3.11. Pictures of node 1 in 1-2-3-4B pattern in A. formosa…………..………….54 Figure 3.12. Pictures of node 2 sessile trifoliate bracts in the 1-2-3-4A pattern in A. coerulea……………..……………………………………………………………………55 Figure 3.13. Pictures of node 2 variations in the 1-2-3-4B pattern in A. formosa………..56 Figure 3.14. Picture of trifoliate leaves at node 3 in 1-2-3-4A pattern in A. coerulea….…57 Figure 3.15. Pictures of trifoliate leaves at node 3 in 1-2-3-4B pattern in A. formosa.…58 Figure 3.16. Picture of node 4 in 1-2-3-4A pattern in A. coerulea……..…………………59 Figure 3.17. Pictures of node 4 in 1-2-3-4B pattern in A. formosa…..……..…..………..59 Figure 3.18. Pictures of the most common pattern found in A. coerulea (1-2-3A)…….…62 Figure 3.19. Pictures of the second most common pattern found in A. coerulea (1-2- 3B)………………………………………………………………………………………..63 Figure 3.20. Pictures of the third most common pattern found in A. coerulea (1-2-3- 4A)……………………………………………………………………………………….64 Figure 3.21. Picture of the outlier plant showing dramatic heteroblasty in A. coerulea…………………………………………………………………………,……….65 Figure 3.22. Pictures of the most common pattern found in A. formosa (1-2-3-4B).……66 Figure 3.23. Pictures of the second most common pattern found in A. formosa (same as outliers) (1-2-3B)…………………………..…………………………………………….67 ix Figure 3.24. Pictures A. formosa showing the 1-2-3B pattern………..………………….70 Figure 3.25. A. formosa showing dramatic heteroblasty…………….…....……………...71 Figure 3.26. Picture of A. formosa……...…………………………………………….….74
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