Egyptian Wheat Salah Gowayed Department of Agrobiodiversity Institute of Crop Sciences University of Kassel Germany Egyptian Wheat Doctoral Dissertation Submitted for the degree of Doctor of Agricultural Sciences of the Institute of Crop Sciences of the University Kassel Presented by Salah Gowayed BSc, MSc, Egypt Witzenhausen, 3 December 2009 ii Die vorliegende Arbeit wurde vom Fachbereich Agrarwissenschaften der Universität Kassel als Dissertation zur Erlangung des akademischen Grades eines Doktors der Agrarwissenschaften (Dr. agr.) angenommen. Erster Gutachter: Prof. Dr. Karl Hammer Zweiter Gutachter: Prof. Dr. Christian Richter Tag der mündlichen Prüfung 3 December 2009 This work was approved by the Faculty of Agricultural Sciences, University of Kassel as a thesis to obtain the academic degree of Doctor of Agricultural Sciences (Dr. agr.). Referee: Prof. Dr. Karl Hammer Co-referee: Prof. Dr. Christian Richter Date of Examination: 3 December 2009 iii Table of Contents Content Pages List of Tables ««««««««««««««««««««««««««««««. vi List of Figures««««««««««««««««««««««««««««««. vii Dedication «««««««««««««««««««««««««««««««.. viii Acknowledgements «««««««««««««««««««««««««««« ix Summary x Zusammenfassung xi Chapter 1 Introduction …………………………………………………………………... 1 Problem Statement ««««««««««««««««««««««««......... 4 Aim and Scope ««««««««««««««««««««««««............... 5 Chapter 2 Egyptian Wheat: A Review………………………………………………….. 7 2.1 A Brief History of Wheat Classification «««««««««««««««««. 8 2.2.1 Domestication of Emmer Wheat «««««««««««««««««««..... 18 2.2.2 The Diversity Center of Domesticated Emmer Wheat ««««««««« 20 2.2.3 Origin and the Diversity Center of Ethiopian Wheat T. aethiopicum«««« 22 2.3 A Historical View of Agriculture and Egyptian Wheat «««««««««««... 24 2.3.1 Palaeoclimatic Conditions ««««««««..««««««««««««... 24 2.3.2 Egyptian History Phases««««««««««««««««««................. 25 2.3.2.1 Paleolithic or Old Stone Age««««««««««««««««««... 25 2.3.2.2 Neolithic Age 6000 BC (Neolithic Revolution)«««««««................ 26 2.3.2.3 Pharaonic Age and Following Phases 3200 BC ± 641A.D.««««««... 26 2.3.3 Emergence of Agriculture««««««««««««««««««............... 27 2.3.4 Egyptian Wheat in Antiquity«««««««««««««««««««. 27 2.3.4.1 Roman Empire and the Egyptian Wheat«««««««««««««« 28 2.3.5 Egyptian Agricultural Exports and External Relations in Antiquity««««....... 29 2.3.5.1 Trade with East Africa and the Arabian Peninsula«««««««««« 30 2.3.5.2 The Country of Nuba and Central and West Africa«««««««««.. 30 2.4 Morphological Studies««««««««««««««««««.......................... 32 2.4.1 Growth Analysis««««««««««««««««««............................. 32 2.4.2 Morphological Characteristics«««««««««««««««««««« 33 2.4.2.1 Inflorescence Architecture ««««««««««««««««««« 33 2.4.2.2 Structure of Inflorescence«««««««««««««««««««.. 36 2.4.2.3 Multiple Glumes««««««««««««««««««.................... 37 2.4.2.4 Grains««««««««««««««««««.................................... 38 2.5 Anatomical Studies««««««««««««««««««.............................. 40 2.5.1 Stem««««««««««««««««««................................................ 40 2.5.2 Leaf««««««««««««««««««................................................. 41 2.5.3 Grains««««««««««««««««««............................................. 43 2.5.4 Coleoptile««««««««««««««««««....................................... 44 2.5.5 First Leaf ««««««««««««««««««....................................... 45 Chapter 3 Materials and Methods ………………………………………………............ 47 3.1Plant Materials ««««««««««««««««««..................................... 48 3.2 Data Collections ««««««««««««««««««.................................. 50 iv 3.2.1 Morphological Studies ««««««««««««««««««............ 51 3.2.2 Anatomical Studies ««««««««««««««««««................. 51 3.3 Date Analysis ««««««««««««««««««...................................... 53 Chapter 4 Results and Discussion ………………………………………………............ 55 4.1 Morphological Studies««««««««««««««««««......................... 56 4.1.1 Quantitative Characters «««««««««««««««««««««.. 56 4.1.2 Qualitative Characteristics««««««««««««««««««« 57 4.1.3 Ramification and Multiple Glumes «««««««««««««««««« 62 4.1.3.1 Multiple Glumes««««««««««««««««.......................... 62 4.1.3.2 Ramifications««««««««««««««««.............................. 62 4.1.3.3 Influence of the Environmental Conditions on Ramification and Third Glume«««««««««««««««............................................. 63 4.1.3.4 Domestication««««««««««««««««................................ 71 4.2 Anatomical Studies««««««««««««««««...................................... 72 4.2.1 Cross-section of Terminal Internodes«««««............................................. 72 4.2.2 Cross-sections in Flag Leaf«««««............................................................ 77 4.2.2.1 Midrib Region«««««.............................«««««....................... 78 4.2.2.2 Mesophyll Region«««««.................................................................. 78 4.2.3 Cross-sections of Grains«««««............................................................... 85 4.2.4 Cross-sections of Coleoptile«««««........................................................... 90 4.2.5 Cross-sections of First Leaf«««««........................................................... 94 4.3 Taxonomic Studies«««««................................................................................... 99 4.4 Wheat in the Egyptian Flora«««............................................................................. 107 Chapter 5 Conclusions …………….............................…………….................................. 127 References«««««.............................«««««..................................................... 131 Curriculum Vitae Arabic Conclusion v List of Tables Table 1 Outline of the Dorofeev et al. (1979) classification system«««««««... 16 Table 2 Egyptian wheat excavations «««««««««««««««««««. 30 Table 3 Number of Egyptian wheat accessions collected ««««««««««« 48 Table4 Accession numbers and locations of Egyptian wheats used in the study (preliminary determination) ««««««««««««««««««.... 49 Table 5 List of Triticum species and their accessions (non-Egyptian) used in the study..50 Table 6 Quantitative characters of eight different Triticum taxa during the growing seasons of 2007 and 2008 «««««««««««««««««««« 59 Table 7 Qualitative characters of spikes, lemma and grains of eight different Triticum taxa. «««««««««««««««««««««««...... 60 Table 8 Morphological characters of glumes of eight different Triticum taxa«««... 61 Table 9 Anatomical structure of terminal internode of nine different Triticum taxa«.. 73 Table 10 Anatomical structure of midrib region of the flag leaf of nine different Triticum taxa««««««««««««««««««««««««« 79 Table 11 Anatomical structure of mesophyll region of the flag leaf of nine different Triticum taxa. ««««««««««««««««««««««««. 83 Table 12 Anatomical characters of various tissues of the mature grains of nine different Triticum taxa. ««««««««««««««««««««.. 86 Table 13 Anatomical structure of coleoptiles of ten different Triticum taxa««««« 91 Table 14 Anatomical structure of first leaf of ten different Triticum taxa«««««« 95 Table 15 Determination of Egyptian wheat«««««««««««««««««.. 99 Table 16 Determination of herbarium specimen of Egyptian wheat in the VIR Herbarium «««««««««««««««««««««««««« 99 Table 17 Scientific names of Triticum used in this study «««««««««««.. 106 Table 18 Cultivars released by Wheat Research Section of ARC with pedigree, year of release and grain yield potential««««««««««««««.......... 111 vi List of Figures Figure 1 Types of Triticum spike 35 Figure 2 Dendrogram resulting from cluster analysis of the morphological similarity among eight Egyptian wheat taxa««««««««««... 58 Figure 3 Varieties of Triticum turgidum convar. compositum with third glume... 64-67 Figure 4 Varieties of Triticum vavilovii with third glume«««««««««. 67 Figure 5 Varieties of Triticum dicoccon with third glume«««««««««. 67-68 Figure 6 Varieties of Triticum durum with third glume«««««««««« 68 Figure 7 Varieties of Triticum polonicum with third glume««««««««.. 69 Figure 8 Triticum aestivum with third glume«««««««««««««« 69 Figure 9 Presence of the third glume in different parts of spike ««««««« 69 Figure 10 True ramification ««««««««««««««««««««« 70 Figure 11 False ramification ««««««««««««««««««««... 70 Figure 12 False-true ramification««««««««««««««««««... 70 Figure 13 Influence the environmental conditions on ramification and third glume. 70 Figure 14 Transverse sections of terminal internode of nine different Triticum taxa taken at spike formation««««««««««««««««.. 74-77 Figure 15 Transvers sections of midrib region of the flag leaf of nine different Triticum taxa taken at spike formation ««««««««««««... 80-82 Figure 16 Transverse sections of mesophyll region of the flag leaf of nine different Triticum taxa taken at spike formation ««««««««« 84 Figure 17 Transvers sections of the mature grains of nine different Triticum taxa taken at dough ripe stage««««««««««««««««««. 87-89 Figure 18 Transverse sections of coleoptiles of ten different Triticum taxa taken at 5mm length ««««««««««««««««««««««. 92-93 Figure 19 Transverse sections of first leaf of ten different Triticum taxa taken at a full expansion «««««««...««««««««««««....... 96-97 Figure 20 Ways of distribution of tetraploidy wheat ««««««...««««... 106 vii Dedication Many thanks are to my God for giving me the opportunity to broaden my academic horizons. viii Acknowledgements This