TAXONOMY AND BIOSYSTEMATICS OF THE CHENOPODIUM ALBUM AGGREGATE A thesis ...

293 Pages·2015·34.05 MB·English

TAXONOMY AND BIOSYSTEMATICS OF THE CHENOPODIUM ALBUM AGGREGATE
A thesis submitted to the University of Leicester for the
degree of Doctor of Philosophy
BY
Mohammad Reza Rahiminezhad Ranjbar
July 1995
Department of Botany
University of Leicester
UMI Number: U078397
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ACKNOWLEDGEMENT
First of all, I would like to thank my supervisors
Professor Clive Stace and Dr. Richard Gornall, whose expert
supervision, guidance and patience enabled me to perform this
research.
I would like to thank Dr. P. Hollingsworth, Dr. J. Bailey
and Dr. Y. Shi for their valuable technical advice. I am
grateful Dr. P. Uotila for expressing his important taxonomic
opinions and also I sincerely thank those people whose names
are mentioned in Appendix 1 and who provided me with
Chenopodium seed from all around the world.
I would like to thank Miss A. Conolly, Mr. P. Thompson
and Ms M. Hart for their friendly cooperation during my study
in the herbarium of the University of Leicester (LTR), and
similarly to all other members of the department of Botany.
Finally I would to say how much I owe my wife Lili, my
sons Hossein and Sina and my daughter Leyla, who were very
patient and devoted during my studies.
This study was funded by the Ministry of Higher Education
of Iran, and without its financial support this research would
not have been possible; I am indebted to them.
DEDICATION
To my mother and my wife Lili
STATEMENT
The contents of this thesis are based on work conducted by the
author in the department of botany, University of Leicester,
during the period April 1992 to March 1995.
The work recorded in this thesis is original, unless otherwise
acknowledged in the text or the references.
signed ,/
30 5 ^5
CONTENTS
CHAPTER 1 : INTRODUCTION 1
THE FAMILY CHENOPODIACEAE 1
HISTORICAL BACKGROUND 1
FEATURES 1
CLASSIFICATION 2
THE GENUS CHENOPODIUM L. 4
HISTORICAL BACKGROUND 4
CLASSIFICATION 4
FEATURES 12
THE SECTION CHENOPODIUM 13
HISTORICAL BACKGROUND 13
THE Ç. ALBUM AGGREGATE 16
HISTORICAL BACKGROUND 16
FEATURES 16
GENE FLOW AND HYBRIDIZATION 17
TAXONOMIC CONFUSION AND DIFFICULTIES 24
AIMS OF THIS STUDY 29
CHAPTER 2: CYTOLOGY 31
INTRODUCTION 31
CYTOLOGY IN CARYOPHYLLACEAE AND CHENOPODIACEAE 31
CYTOLOGY IN CHENOPODIUM 32
CYTOLOGY IN THE C. ALBUM AGGREGATE AND SOME
ALLIED SPECIES 33
AIMS OF THE PRESENT STUDY 36
MATERIALS AND METHODS 37
MITOTIC ROOT TIP CHROMOSOME COUNTS 37
RESULTS 38
DISCUSSION AND CONCLUSIONS 40
CHAPTER 3: MORPHOLOGY 42
INTRODUCTION 42
MORPHOLOGY IN THE GENUS CHENOPODIUM 43
MORPHOLOGY IN THE SECTION CHENOPODIUM 43
MORPHOLOGY IN THE C. ALBUM AGGREGATE 45
LEAF' CHARACTERS 46
INFLORESCENCE CHARACTERS 48
SEEC1 CHARACTERS 51
POLLEN CHARACTERS 53
AIMS: OF THE PRESENT STUDY 55
MATERIALS AND METHODS 55
RESULTS 57
GENERAL OBSERVATIONS 57
LEAF' MORPHOLOGY 58
C. Dolvsoermum 58
C. hvbridum 58
C. murale 58
C. alaucum 60
c. vu1 varia 60
c. fi ci fol i um 60
c. suecicum 63
c. novopokrovskvanum 65
c. striatum 65
c. bushianum 65
c. Quinoa 67
c. berlandieri 67
c. aiaanteum 67
c. ODulifolium 70
c. Drobstii 70
c. album 70
SEED MORPHOLOGY 77
DISCUSSION 82
LEAF-SHAPE 82
SEED MORPHOLOGY 85
CONCLUSIONS 85
CHAPTER 4: FLAVONOIDS 87
INTRODUCTION 87
BIOSYNTHESIS AND GENETIC CONTROL 89
FLAVONOIDS IN PLANT TAXONOMY 92
FLAVONOIDS AND PHYLOGENY OF PLANTS 94
BETALAINS AND FLAVONOIDS IN FAMILIES OF CARYOPHYLLALES 97
FLAVONOIDS IN THE CHENOPODIACEAE 99
FLAVONOIDS IN THE GENUS CHENOPODIUM 100
AIMS OF THIS STUDY 104
MATERIALS AND METHODS 1 04
EXTRACTION AND ISOLATION 110
IDENTIFICATION OF FLAVONOIDS 112
RESULTS 1 1 3
SUBSTITUTIONS 114
DISCUSSION 120
TAXONOMIC & EVOLUTIONARY CONCLUSIONS 124
CHAPTER 5: ENZYME ELECTROPHORESIS 126
INTRODUCTION 126
ISOZYME DATA AND PLANT SYSTEMATICS 126
ISOZYME DATA AND PLANT SPECIATION 130
ISOZYME DATA IN THE GENUS CHENOPODIUM 136
Taxonomy 138
Polyploidy 141
Crop-weed complexes 142
Infraspecific variation 145
Selection 147
AIMS OF THE PRESENT STUDY 1 48
MATERIALS 148
METHODS 152
EXTRACTION 153
GEL PREPARATION 153
GEL LOADING 154
GEL RUNNING 154
GEL STAINING 155
GENETIC INTERPRETATION 155
DATA ANALYSIS 158
RESULTS 158
AAT 158
AAT-1 1 59
AAT-2 160
AAT-3 1 61
GDH 170
IDH 175
MDH 184
MDH-1 184
MDH-2 184
MDH-3 185
PGD 195
PGD-1 195
PGD-2 196
PGM 206
PGM-1 206
PGM-2 207