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JULY 2008 Table of Contents ADDIS ABABA UNIVERSITY SCHOOL OF GRADUATE STUDIES DEPARTMENT OF BIOLOGY CHROMOSOME STUDY OF ELEVEN ENDEMIC ALOE SPECIES OF ETHIOPIA WITH EMPHASIS ON THE SATELLITES AND NUCLEOLI BY TEZERA TEMESGEN A THESIS SUBMITTED TO THE SCHOOL OF GRADUATE STUDIES OF ADDIS ABABA UNIVERSITY IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BIOLOGY (Applied Genetics) Addis Ababa JULY 2008 Table of Contents page Acknowledgment …………………………………………………………….. IV List of Tables ………………………………………………………………….. V List of figures ………………………………………………………………….....VI List of Appendices ……………………………………………………………. VIII Abstract ……………………………………………………………………….. IX 1. Introduction ……………………………………………………………… …. 1 1.1. Taxonomical position of Aloe species ……………………………………..1 1.2. Distribution, center of origin and endemism of Aloe species ……………...1 1.2.1. Distribution and endemism of Aloe species in Ethiopia …………. ….2 1.3. Habitat of Aloe species ………………………………………………... …..3 1.4. Botany and reproduction of the Aloe species ………………………… …..3 1.4.1. Botany of the Aloe species…………………………………………….3 1.4.2. Reproduction of the Aloe species ……………………………………..4 1.5. Uses of the Aloe species ……………………………………………… ….. 5 1.5.1. Therapeutic use of the Aloe species……………………………… …..5 1.5.2. Other uses of the Aloe species…………………………………… …...5 1.6. Biological description , ecology and distribution of Aloe species used in the present study ………………………………………………………….6 1.6.1. Aloe camperi Schweif ………………………………………………….6 1.6.2. Aloe otallensis Baker …………………………………………………..6 1.6.3. Aloe tewoldei Gilbert and Sebsebe ……………………………….........7 1.6.4. Aloe friisii Sebsebe and Gilbert ..............................................................7 1.6.5. Aloe kefaensis Gilbert and Sebsebe ........................................................7 1.6.6. Aloe harlana Retnolds ............................................................................8 1.6.7. Aloe debrana Christian ............................................................................8 1.6.8. Aloe schelpei Reynolds ............................................................................8 1.6.9. Aloe yavellana Reynolds ..........................................................................9 1.6.10. Aloe trigonantha Leach ........................................................................9 1.6.11. Aloe species from Bale .........................................................................9 2. Karyological studies of chromosomes ...........................................................9 2.1. karyotype ....................................................................................................10 2.1.1. Chromosome number ...........................................................................11 2.1.2. Chromosome size .................................................................................12 2.1.3. Chromosome morphology ....................................................................12 2.2. Factors that cause karyotype evolution.........................................................13 2.2.1. Polyploidy …………………………………………………………….13 2.2.2. Causes of karyotype evolution ………………………………………..14 3. Chromosomes of Aloe species ……………………………………………….15 3.1. Chromosome number and size ……………………………………………..15 3.2. Chromosome satellites ……………………………………………………..16 3.3. Polyploidy in Aloe species …………………………………………………17 3.4. Silver staining of nucleoli and nucleolus organizer regions ……………….18 3.4.1. Nucleolus ……………………………………………………………..18 3.4.2. Nucleolus organizer regions …………………………………………..19 3.4.3. Silver staining …………………………………………………………19 3.5 Chromosome C-banding …………………………………………………….20 4. Objectives of the study …………………………………………………………...22 4.1 General objectives ……………………………………………………………22 4.2 Specific objectives ……………………………………………………………22 4.3 Materials and methods ……………………………………………………….23 4.3.1. Plant materials …………………………………………………………...23 4.3.2. Pretreatment of roots for chromosome preparation ………………………23 4.3.2. Air-dry slide preparation …………………………………………………25 4.3.4. Silver staining …………………………………………………………….25 5. Results …………………………………………………………………………….27 5.1. A. friisii ……………………………………………………………………….27 5.2. A. otallensis …………………………………………………………………...30 5.3. A. tewoldei..........................................................................................................32 ii 5.4. A. camperi........................................................................................................34 5.5. A. yavellana.....................................................................................................36 5.6. A. schelpei........................................................................................................38 5.7. A. kefaensis ………………………………………………………………….39 5.8. A. trigonantha ……………………………………………………………….39 5.9. Aloe species from Bale ....................................................................................40 5.10. A. harlana ......................................................................................................41 5.11. A. debrana .....................................................................................................43 6. Discussion...............................................................................................................44 7. Conclusion and Recommendation..........................................................................48 7.1.1. Conclusion…………………………………………………………………48 7.1.2. ………………………………………………………………………..........48 8. References..............................................................................................................50 iii ACKNOWLEDGMENTS I would like to express my deepest gratitude to my research advisor Dr. Kifle Dagne for his unreserved follow up of this study throughout the study period and for photography of chromosomes pictures and critical reading of manuscript of my thesis. I would also like to extend my special thanks to my advisor Prof. Sebsebe Demissew for his help in the collecting the plant material and confirming the identification and for his valuable comments on the thesis. I am indebted to Yirgalem District Educational Office for giving me the chance to join postgraduate study and for paying me salary during the study time. I extend my heartfelt thanks to my family for providing the maximum care throughout this study, especially to my wife Agerenesh Bishaw and my children as they have been, beyond all, my success all the time. Finally, above all I thank the Almighty God who offered me everything and whose help made it possible for me to finish my study according to his will. iv List of Tables Page Table 1. Aloe species used in the present study and their sites of collection ……….. 22 Table 2 Chromosome number, karyotypic formula, and number of satellite chromosome. …………………………………………………………………. 28 Table 3. Number and frequency of nucleoli in each cell nucleus of the species and their frequency in percentage. ………………………………………….. 30 Table 4. Chromosome length of A. friisii, A. otallensis, A. tewoldei and A. camperi in µm. …………………………………………………………….36 v List of Figures Page Figure1. Pictures of Aloe species used in the present study: A. friisii; B. A. otallensis; C. A. tewoldei; D. A. camperi; E. A. yavellana;F. A.harlana; G. A. schelpei; H. A. trigonantha; I. Aloe species from Bale; J. A. debrana and K. A. kefaensis………………………………………………………………24 Figure 2. Chromosomes and nucleoli of Aloe friisii: A. Metaphase plate (2n = 14); B. Karyotype constructed from chromosomes in A; C. 13 moderately condensed metaphase chromosomes with four of the chromosomes showing satellites (arrows); D. Interphase nucleus with four nucleoli. Bar = 20 µm. ……29 Figure 3. Chromosomes and nucleoli of A. otallensis: A. metaphase spread (2n = 14); B. Early metaphase stage chromosomes, showing satellites on the long arms of four long chromosomes; C. Karyogram constructed from chromosomes shown in A; D. Interphase nucleus with four nucleoli. Bar = 20 µm. ………………..............................................................................31 Figure 4. Chromosomes and nucleoli of A. tewoldei: A. Metaphase chromosomes (2n = 14); B. Early metaphase chromosomes with satellites on the short arm of a pair of short and on the long arm of a pair of long chromosomes (arrows). C. Karyotype ………………………………………....33 Figure 5. Chromosomes and nucleoli of A. camperi: A. Metaphase chromosomes (2n = 14); B. Late prophase chromosomes with satellites on the long arm of two long chromosomes, and on short arm of a short chromosome (arrows); C. A karyotype constructed from chromosomes in A; D. Interphase nucleus with four nucleoli. Bar = 20 µm. …………………………………………………..35 vi Figure 6. Chromosomes and nucleoli of A. yavellana: A and B are cells with moderately condensed chromosomes showing satellites on the long and short chromosomes (arrows); C. Metaphase chromosome; D. karyotype constructed for chromosomes from chromosome C. E. Interphase nucleus with six nucleoli. Bar = 20 µm… ……………........................................................37 Figure 7. Chromosomes and nucleoli of A. schelpei: A. Metaphase chromosomes (2n = 14); B. Prophase chromosomes with six satellites, five of which are associated with two nucleoli (arrows); C. Two interphase nuclei with five and six nucleoli. D.Karyotypy constructed from A. Bar = 20 µm………………..38 Figure 8. Chromosomes and nucleoli of A. kefaensis: A. Metaphase Chromosomes (2n = 14) with satellites on the long arm of two long chromosomes (arrows); B. Prophase stage showing two satellite chromosomes associated with a nucleolus
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