ARISTOTLE UNIVERSITY OF THESSALONIKI FACULTY OF SCIENCES, SCHOOL OF BIOLOGY ...

284 Pages·2009·9.71 MB·English

ARISTOTLE UNIVERSITY OF THESSALONIKI
FACULTY OF SCIENCES, SCHOOL OF BIOLOGY
DEPARTMENT OF BOTANY

SHARAF MUSA AL-TARDEH
B. Sc. Biological Sciences
M.Sc. Food Quality Management

MORPHOLOGICAL AND ANATOMICAL ADAPTATIONS OF THE
PERENNIAL GEOPHYTE URGINEA MARITIMA (L.) BAKER
(LILIACEAE) TO THE MEDITERRANEAN CLIMATE

DOCTORAL DISSERTATION

Thessaloniki 2008
SHARAF MUSA AL-TARDEH
B. Sc. Biological Sciences
M.Sc. Food Quality Management

MORPHOLOGICAL AND ANATOMICAL ADAPTATIONS OF THE
PERENNIAL GEOPHYTE URGINEA MARITIMA (L.) BAKER
(LILIACEAE) TO THE MEDITERRANEAN CLIMATE

DOCTORAL DISSERTATION

Submitted to the School of Biology,
Faculty of Sciences, Aristotle University of Thessaloniki

Presented for public criticism on the 28th of November 2008
MORPHOLOGICAL AND ANATOMICAL ADAPTATIONS OF THE
PERENNIAL GEOPHYTE URGINEA MARITIMA (L.) BAKER
(LILIACEAE) TO THE MEDITERRANEAN CLIMATE

Figures on the cover page: (A) TEM micrograph of nectary of Urginea maritima
showing the nectary epithelium cell with nectar in crystallized form at the old stage;
(B) Photo of inflorescence stalk; (C) SEM micrograph of anther; (D) LM micrograph
of leaf with parallel veins and veinules; (E) TEM micrograph of leaf with sieve pore
between the sieve element; (F) LM micrograph of bulb scale revealing cells stained
red with Schiff’s reagent; (G) LM micrograph revealing cells stained brown-black
with Sudan Black B; (H) LM micrograph showing idioblastic cortical cells with
bundles of raphides and mucilage stained with TBO; (I) LM micrograph of root
showing the 10-arch vascular cylinder; (J) LM micrograph of root showing series of
idioblastic cells with bundles of raphides.
To my family

To my lovely wife “WOROD”

To those who defend their rights

SHARAF
ELABORATION OF THE DOCTORAL DISSERTATION
Place
Aristotle University of Thessaloniki, Faculty of Sciences, School of Biology
Department of Botany

Time
February 2004 – November 2008.

Examining committe
• Stylianos Delivopoulos, Professor, Department of Botany, School of Biology,
A.U.Th. Supervisor
• Thomas Sawidis, Associate Professor, Department of Botany, School of Biology,
A.U.Th. Cosupervisor
• Barbara-Evelin Diannelidis, Assistant Professor, Department of Botany, School
of Biology, A.U.Th. Cosupervisor

• Bosabalidis Artemios, Professor, Department of Botany, School of Biology,
A.U.Th. Examiner
• Eleftheriou Eleftherios, Professor, Department of Botany, School of Biology,
A.U.Th. Examiner
• Moustakas Michael, Associate Professor, Department of Botany, School of
Biology, A.U.Th. Examiner
• Pantis John, Associate Professor, Department of Ecology, School of Biology,
A.U.Th. Examiner

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© Sharaf Musa Al-Tardeh
© AUTh.

Morphological and anatomical adaptations of the perennial geophyte Urginea
maritima (L.) Baker (Liliaceae) to the mediterranean climate.
ISBN

Approval of the present Doctoral Dissertation by the Department of Botany, School
of Biology, Aristotle University of Thessaloniki does not imply the acceptance of
the opinions of the author (N. 5343/1932, law 202, paragraph 2).
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Sharaf Al-Tardeh Table of contents

TABLE OF CONTENTS
ELABORATION OF THE DOCTORAL DISSERTATION……………………....... i
TABLE OF CONTENTS…………………………………………………………...... iii
PROLEGOMENA ……...………………………………………………………......... vii
LIST OF ORIGINAL ARTICLES……….………....................................................... ix
LIST OF ABBREVIATIONS AND SYMBOLS………………………………..…… x
SUMMARY...………………………………………………………..…………......... xii
ΠΕΡΙΛHΨΗ…………………………………………………………………………... xvi
CHAPTER I: INTRODUCTION…………...……………………………………… 1
1.1 Mediterranean climate……………………………………………………………. 1
1.1.1 Introduction…………………...…………………………………………….… 1
1.1.2 Spatial patterns of temperature in the Mediterranean climate…………….…... 1
1.1.3 Spatial patterns of precipitation in the Mediterranean climate………………... 2
1.2 Geophytes……………………………………………………………………….... 3
1.2.1 Introduction…………………………………………………………………… 3
1.2.2 Taxonomy and origin of geophytes…………………………………………... 3
1.2.3 Diversity of geophytes………………………………………………………... 3
1. 2.3.1 Bulbous plants………………………………………………………......... 4
1.2.3.2 Tuberous plants…………………………………………………………… 4
1.2.4 The body of geophytes……………………………………………………...... 5
1. 2.4.1 Storage organs and tissues…………….…………………………………. 5
1.2.4.2 Aerial organs and tissues………..………………………………………… 6
1. 2.4.3 Roots……………………………………………………………………… 6
1.2.4.4 Tunics…………………………………………..…………………………. 7
1.2.5 Periodicity and dormancy of flower bulbs…………………………………….. 7
1.2.5.1 Definitions…………………………………………………………………. 7
1.2.5.2 Factors that affect bulb dormancy…………………………………………. 8
1.2.6 The flowering process……………………………………………………......... 9
1.2.7 Types of hysteranthous geophytes…………………………………………….. 9
1.2.8 Factors affecting bulb growth and development……………………………… 10
1.2.8.1 The size of the bulb……………………………………………………....... 10
1.2.8.2 Temperature………………………………………………………………... 10
1.2.8.3 Light……………………………………………………………………….. 11
1.2.8.4 Soil features……………………………………………………………....... 11
1.2.9 World production of geophytes……………………………………………..... 12
1.3 Chlorophyll contents and fluorescence…………………………………………... 13
1.4 Nectaries and nectar…………………………………………………………….... 17
1.4.1 Introduction…………………………………………………………………… 17
1.4.2 Definitions of nectary…………………………………………………………. 17
1.4.3 Basic types of floral nectaries.…..…………………………………………….. 18
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Sharaf Al-Tardeh Table of contents

1.4.4 Floral and extrafloral nectaries………………………………………………... 20
1.4.5 Gynopleural (septal) nectaries……………………………………………….... 20
1.4.5.1 Distribution of the gynopleural nectaries………………………………….. 20
1 4.5.2 Structure of the gynopleural nectaries…………………………………....... 21
1.4.5.3 Nectar secretion in the gynopleural nectaries…………………………….... 22
1.4.5.4 Fate of the gynopleural nectaries…………………………………………... 22
1.4.6 Nectar production and presentation…………………………………………… 23
1.4.6.1 Nectar secretion mechanism…………………………….............................. 23
1. 4.6.2 Nectar presentation………………………………………………………... 24
1.4.6.3 Fate of nectar and nectarines………………………………………………. 26
1.5 The summer flowering geophyte: Urginea maritima (L.) Baker (Liliaceae)…...... 27
1.5.1 Urginea maritima overview…………………………………………………... 27
1.5.2 Morphology…………………………………………………………………… 27
1.5.3 Growth, development and flowering……………………………………….… 28
1.5.4 Nectary of U. maritima……………………………………………………….. 29
1.5.5 Pollination in U. maritima as a model of hysteranthous geophytes………....... 29
1.5.6 Bulb production and/or propagation………………………………………….. 30
1.5.7 Horticultural usage……………………………………………………………. 32
1.5.8 Biochemical principles and their action………………………………………. 32
1.5.8.1 Toxic principle……………………………………………………………... 32
1.5.8.2 Defense mechanisms………………………………………………………. 33
1.5.8.3 Pharmaceutical advantages of U. maritima……………………………….. 34
1.6 Aim of the thesis…………………………………………………………………. 35
CHAPTER II: MATERIALS AND METHODS ……………………………….... 37
2.1 Materials………………………………………………………………………….. 37
2.1.1 Plant samples………………………………………………….……………..... 37
2.1.2 Apparatuses……….…………………………………………...……………… 38
2.1.3 Experimental design…………………………………………………………... 40
2.2 Methods…………………………………………………………………………… 42
2.2.1 Water content………………………………………………………………….. 42
2.2.2 Fixation………………………………………………………………………... 43
2.2.2.1 Solution preparations………………………………………………………. 43
2.2.2.2 Procedure…………………………………………………………………... 44
2.2.3 Sectioning……………………………………………………………………... 45
2.2.4 Staining techniques…………………………………………………………… 46
2.2.4.1 Staining with toluidine blue O (0.1%)…………………………………….. 46
2.2.4.2 Staining with Sudan Black B (1%)………………………………………... 46
2.2.4.3 Staining with Periodic acid-Schiff’s reaction (PAS reaction)…………….. 47
2. 2.4.4 Staining with uranyl acetate and lead citrate…………………………….. 48
2. 2.5 Scanning Electron Microscopy (SEM)……………………………………….. 48
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Sharaf Al-Tardeh Table of contents

2.2.6 Maceration…………………………………………………………………….. 49
2.2.6.1 Maceration of soft material………………………………………………... 49
2.6.2 Maceration of hard material…………………………………...…………….. 50
2.2.7 Leaf chlorophyll assay………………………………………………………... 50
2.2.8 Chlorophyll a fluorescence…………….……………………………………… 50
2.2.9 Morphometry…………………………………………………………………. 52
2. 2.10 Mathematical analysis of morphological and anatomical features of the
plant tissues………………………………………………………………….... 52
2.3 Statistical analysis………………………………………………………………... 52
CHAPTER III: RESULTS ………………..……………………………………….. 53
3.1 ANATOMICAL STUDIES ON THE ADVENTITIOUS ROOTS……………... 53
3.1.1 Root morphology ………………………………………………………..……. 53
3.1.2 Root anatomy………………………………………………………………….. 53
3.1.3 Root histochemistry…………………………………………………………… 56
3.1.4 Root morphometry……………………………………………………………. 56
3.2 Biomass and reserve allocation patterns within the bulb………………………… 58
3.2.1 Bulb morphology……………………………………………………………… 58
3.2.2 Bulb anatomy…………………………………………………………………. 58
3.2.3 Bulb histochemistry…………………………………………………………… 60
3.2.4 Bulb morphometry and reserves allocation patterns………………………….. 61
3.2.5 Biomass allocation patterns and water status…………………………………. 62
3.3 MORPHO-ANATOMICAL FEATURES OF THE LEAVES…………………. 65
3.3.1 Leaf anatomy………………………………………………………………….. 65
3.3.2 Leaf morphology and morphometry…………………………………………... 67
3.3.3 Leaf histochemistry…………………………………………………………..... 69
3.3.4 Leaf chlorophyll assay……………………………………………………….... 69
3.3.5 Chlorophyll a fluorescence……………………………………………………. 70
3.4 MORPHO-ANATOMY OF THE INFLORESCENCE…………………………. 73
3.4.1 Flower morphology……………………………………………………………. 73
3.4.2 Anatomy and development of the gynopleural (septal) nectary………………. 74
3.4.2.1 Young stage nectary……………………………………………………….. 74
3.4.2.2 Nectary at the intermediate stage…………………………………………... 75
3.4.2.3 Nectary at old stage……………………………………………………….... 77
3.4.3 The ovary and ovules………………………………………………………..…. 78
3.4.4 Obturator gland……………………………………………………………….... 79
CHAPTER IV: DISCUSSION…………………………............................................ 170
4.1 ADVENTITOUS ROOTS………………………………………………………... 170
4.1.1 Root system……………………………………………………………………. 170
4.1.2 Special adaptation of roots…………………………………………………….. 170
4.1.3 Velamen………………………………………… ……………………………. 171
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Sharaf Al-Tardeh Table of contents

4.1.4 Dimorphic hypodermis………………………………………………………... 172
4.1.5 Cortex…………………………………………………………………………. 173
4.1.6 The storage tissue……………………………………………………………... 173
4.1.7 Water economy………………………………………………………….......... 174
4.1.8 Raphides………………………………………………………………………. 175
4.1.9 Vascular cylinder…………………………………………………..………….. 177
4.2. BIOMASS AND RESERVE ALLOCATION PATTERNS WITHIN THE
BULB………………………………… …………………………………………… 177
4. 2.1 Morpho-anatomy of the bulb………………………………………………….. 177
4.2.2 Tunics…………………………………………………………………………. 178
4.2.3 Biomass, resources allocation and plant growth……………………………… 178
4.2.4 Histochemistry and the storage tissue………………………………………… 181
4.2.5 Defense mechanisms…………………………………………………………... 181
4.2.6 Water contents………………………………………………………………… 183
4.3 MORPHO-ANATOMICAL FEATURES OF THE LEAVES ………………….. 184
4.3.1 Leaf anatomy………………………………………………………………….. 184
4.3.2 Epidermis and cuticle…………………………………………………………. 184
4.3.3 Stomata as an adaptive strategy………………………………………………. 184
4.3.4 The mesophyll ground tissues……………………………………………........ 185
4.3.5 The vascular system…………………………………………………………… 186
4.3.6 Leaf morphology………………………………………………………………. 187
4.3.7 Leaf histochemistry……………………………………………………………. 189
4.3.8 Photosynthetic efficiency of the leaf of U. maritima…………………………. 190
4.3.8.1 Chlorophyll a content…………………………………………….………... 190
4.3.4.2 The basis of chlorophyll fluorescence measurements……………………... 191
4.4 MORPHO-ANATOMY OF THE INFLORESCENCE ………………………… 194
4.4.1 Gynopleural (septal) nectary………….……………………………………….. 194
4.4.1.1 Nectary epithelial cells……………………………………………………... 194
4.4.1.2 Subsidiary tissue…………………………………………………………… 195
4.4.1.3 Amyloplasts and nectar secretion………………………………………….. 196
4.4.1.4 Fate of the nectary…………………………………………………………. 199
4.4.1.5 Nectar presentation and pollinators………………………………………... 200
4.4.2 Contents of raphides………………………………………………………….. 201
4.4.3 The obturator gland…………………………………………………………… 202
CHAPTER V: CONCLUSIONS…………………………………………………… 203
ACKNOWLEDGEMENTS........................................................................................... 207
CHAPTER VI: LITERATURE CITED.…………………………………………... 208
ORIGINAL PUBLISHED ARTICLES………………………………………..…….. i

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