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Fruit Breeding

About the Authors
Dr. Anil Kumar Shukla obtained his B.Sc. (Ag) and M.Sc
Ag (Hort.) from N.D. University of Agriculture and
Technology Kumarganj, Faizabad during 1993 and 1996
respectively. He completed his PhD from IARI, Pusa, New
Delhi in the year 2000. He has been awarded with Gold
Medal in B.Sc (Agri.) and he has second rank in ICAR in
JRF (1993). He was topper of ARS and SRF (1998). He
worked as Scientist and Sr. Scientist at CIAH, Bikaner. He
also served as Associate Professor at Maharana Pratap
University of Agriculture and Technology, Udaipur. Dr. Shukla also worked as
Principal Scientist (Horticulture) at Directorate of Research on Women in
Agriculture, Bhubaneswar, Odisha. Presently, he is head, ICAR-CAZRI, RRS,
Pali (Rajasthan). He is the recipient of N.E. Borlaug Fellowship for year 2008, He
also received Best worker Award by Vice-Chancellor, MPUAT, Udaipur. Fellowship
Award 2012, from Indian Society of Horticultural Research and Development,
Uttarakhand (Society). Fellowship Award 2012, from Hi-Tech Horticultural Society,
Meerut (Society), Fellowship Award 2012, from Confederation of Horticulture
Association of India, New Delhi (Society), Fellowship Award 2018, from Indian
Society of Arid Horticulture, Bikaner. Fellowhip Award 2019, International Society
for Noni Science Chennai, Fellowship Award 2019, Indian Academy of Horticultural
Science, Eminent Scientist Award 2016, from SVWS, Lucknow, Himadri Young
Scientist Award 2010 from GBPUAT, Pant Nagar, Uttarakhand and Indian Society
of Horticulture Research and Development, Uttarakhand.
Dr. Anil Kumar Shukla immensely contributed in developing two varieties of
Ber (Thar Sevika and Thar Bhubraj) and two varieties of guava MPUAT-S-1 and
MPUAT-S-2. He has developed several Agro techniques in Ber and Guava crops.
He has guided 02 Ph.D. and 01 M.Sc. (Ag) students as Major Advisor and 05
Ph.D. and 04 M Sc. (Ag.) students as Co-Advisor. He has published 70, research
papers, 38 book chapters, 76 research abstracts, 68 popular articles, 11 bulletins
and 03 books.
Dr. Arun Kumar Shukla graduated from T.D. College,
Jaunpur (Purvanchal University), M.Sc. (Ag.) Horticulture
and PhD from Department of Horticulture, Institute of
Agricultural Sciences, BHU, Varanasi. He has been
recipient of National Scholarship, ICAR-JRF, UGC-JRF/
SRF, Young Scientist-2015, Outstanding Scientist-2016,
Scientist of the Year Awards-2017 by Academic Research
Foundation, APJ Abdul Kalam Scientist, Dr BP Pal Scientist
Awards-2019 by Society of Tropical Agriculture. Awarded

with fellow-ISAH 2018 and Fellow-STA 2018. ICAR-NAIP Team award for
outstanding achievement of NAIP Project (Component-3) entitled “Integrated
farming system ….. Dhar districts of Madhya Pradesh”(As CoPI-IGFRI, Jhansi)
during Regional committee meeting, Goa, on 09.11.2012. Besides production
technologies, he is actively involved in genetic improvement of temperate fruits
He has published more than 180 publications including research paper, popular
articles, book, book chapter, training manual, leaflet/extension bulletins etc. At
present he is working as Principal Scientist (Fruit Science) at ICAR-IARI, Regional
Station, Amartara Cottage, Shimla.
Dr. M.B. Noor mohamed, is presently working as Scientist
(Agroforestry) at ICAR- Central Arid Zone Research
Institute, RRS, Pali Marwar (Rajasthan) with more than 5
years of experience in tree improvement, agroforestry,
carbon sequestration and breeding works in shrubs and
trees. He completed his UG, PG and Ph.D. in Forestry at
Forest College and Research Institute, TNAU,
Mettupalayam (Tamil Nadu). He has been awarded with
Gold medal in PhD (Forestry). He is actively involved in
germplasm collection, evaluation and improvement in trees and fruit crops. He
has published 20 research papers in international as well as national journals, 2
popular articles, 8 book chapters, 2 books, 1 training compendium and more than
20 abstracts. He has been honoured with Young Scientist Award, Innovative
Scientist Award and Best oral presentation award by various societies.
Dr. Akath Singh, Principal Scientist (Horticulture) at
ICAR- Central Arid Zone Research Institute with more
than 16 years of experience in production and improvement
of various fruit crops. He has remarkable research
achievements in strawberry, guava, peach and
pomegranate production and improvement. He contributed
in development of two guava cultivars, and one each in
lasora and karonda. He was actively participated in
finalizing national DUS guidelines for pomegranate. Selected
as Member of Indian scientist’s delegation visited to China.
He has been honoured with Young Scientist Associate Award, SARP Associate
Award, SDSH fellow by various societies and also granted ITS by DST for Spain
visit. He has around 85 publications in his credit including 35 research papers in
high impact national and international journals and author for three edited books,
18 book chapters and 7 technical bulletins. He has also guided two M.Sc. students.

Dr. Divya Tiwari completed her higher studies from Narendra
Deva University of Agriculture & Technology, Faizabad, U.P.
(India) and acquired B.Sc. (Ag.), M.Sc. (Ag.) and Ph.D. in
Horticulture (Fruit Science) Degrees in continuation during
2000-2010. She has also Awardee of The Best Teacher Award
(2016) and Young Scientist award (2017) and got recognition
as CHAI Fellow in 2019. She has published several research
papers, authored/edited books, book chapters, popular articles,
folders and manuals. She is also developed, introduced and popularized several
technologies. Among which fertigation and irrigation scheduling for fruit crops,
development of modified BBCH scale for mango and methodology for recording
the phenological observations was the most significant. Moreover, she has also
the experience of handling several projects which are either University funded or
externally funded.

NEW INDIA PUBLISHING AGENCY
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®
2
nd
Fully Revised and Enlarged Edition
Fruit Breeding
Anil Kumar Shukla
Arun Kumar Shukla
M.B. Noor mohamed
Akath Singh
Divya Tiwari

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ISBN : 978-93-89130-46-1 eISBN: 978-93-89992-41-0
Composed and Designed by NIPA
Distributed by NIPA GENX Electronic Resources and Solutions Pvt. Ltd. New Delhi

Dedicated to
Late Shree Yamuna Prashad Shukla
Agriculture Extension Teacher

Foreword to the First Edition
There are many challenges in fruit production to meet out the daily requirement
of fruits for growing population in the new millennium because of faster rate of
population growth, limited area under fruit crop, reduction in cultivable land,
poor yield potential, and susceptibility to insect pests and diseases. However,
for nutritional security of growing population, development of precocious, prolific,
biotic and abiotic stress resistant fruit varieties are essentially required.
There have been conscious efforts in fruit improvement through planned
hybridization and selection. Consequently, most of the outstanding present day
cultivars have resulted from chance seedling selection, some of these selections
are quite old. In response to new challenges need for better tree canopy resistant
to biotic and abiotic stresses, better shelf life and high productivity per unit area
still requires more attention. I am happy that concerted efforts made by the
authors to compile the work done in this direction are under one umbrella. I
congratulate the authors for bringing out such an useful publication. I hope the
book will be a valuable source of reference and text for students, teachers and
scientists associated with the discipline.
(G. Kalloo)

Preface of the Second Edition
The pressure of an ever-increasing population and periodic famine due to
unexpected flood and drought has forced and awakened the horticultural scientist,
to evolve new plant types for diversified use. Besides, some limitations in the
improvement of fruit crop such as long juvenile phase, high heterozygosity,
limited information on inheritance pattern, excessive fruit drop, parthenocarpy
and lesser number of seeds per fruit, hybridization, selection, mutation and other
tools of fruit breeding have resulted in the development of a number of varieties
in mango, grape, papaya, banana and guava for various purposes.
A vast literature is available on every aspects of fruit cultivation and many
excellent books and manuals have been written from time to time however,
compiled information on fruit breeding are not available under a single head.
The present book entitled “ Fruit Breeding Approaches and Achievements”
Revised and enlarged edition is ventured with the objective to provide latest
possible information on basic approaches in fruit breeding, breeding for biotic
stresses resistance, use of plant growth regulators in fruit improvement,
improvement of important fruit crops such as mango, banana, papaya, grape,
guava, citrus, ber, aonla, pomegranate, date palm, litchi, coconut, cashewnut,
pineapple, few under utilized fruits and temperate fruits in a broad spectrum.
It is with more than mere formality we thank Dr. A.K. Singh, Deputy Director
General (Horticultural Sciences), ICAR, New Delhi for inspiration given by
them in writing this book.
Authors express their deep sense of gratitude to Dr. O.P. Yadav, Director,
Central Arid Zone Research Institute, Jodhpur for giving more thrust to increase
quality publications. We are also thankful to Dr. Dipak Kumar Gupta and Dr.
R.S. Mehta for their critical comments and suggestions. Authors are thankful
to All Scientist and workers of Central Arid Zone Research Institute, Regional
Research Station, Pali (Rajasthan) for their help during writing this book.
Authors books, book chapters, research papers, popular articles and technical
papers used in writing of this book have duly been acknowledged in the text.The

xii Fruit Breeding
authors will appreciate receiving suggestions for further improvement of this
book. We hope that information presented here in the book would receive due
response from the teaches, students, researchers and other readers
.
Date: August, 2019
Anil Kumar Shukla
Place: Pali Marwar (Rajasthan) Arun Kumar Shukla
M.B. Noor mohamed
Akath Singh
Divya Tiwari

Preface of the First Edition
The pressure of an ever-increasing population and periodic famine due to
unexpected flood and drought has forced and awakened the horticultural scientist,
to evolve new plant types for diversified use. Besides, some limitations in the
improvement of fruit crop such as long juvenile phase, high heterozygosity,
limited information on inheritance pattern, excessive fruit drop, parthenocarpy
and lesser number of seeds per fruit, hybridization, selection, mutation and other
tools of fruit breeding have resulted in the development of a number of varieties
in mango, grape, papaya, banana and guava for various purposes.
A vast literature is available on every aspects of fruit cultivation and many
excellent books and manuals have been written from time to time however,
compiled information on fruit breeding are not available under a single head.
The present book entitled “ Fruit Breeding Approaches and Achievements”
Volume 1 is ventured with the objective to provide latest possible information
on basic approaches in fruit breeding, breeding for biotic stresses resistance,
use of plant growth regulators in fruit improvement, improvement of important
fruit crops such as mango, banana, papaya, grape, guava, citrus, ber, aonla,
pomegranate, date palm, litchi, coconut, cashewnut, pineapple and few
underutilized fruits in a broad spectrum.
It is with more than mere formality we thank Dr. G. Kalloo, Deputy Director
General (Horticultural and Crop Sciences), ICAR, New Delhi and Prof. R.K.
Pathak, Director, Central Institute for Subtropical Horticulture, Lucknow for
the inspiration given by them in writing this book. Authors are highly grateful to
Dr. S.N. Pandey, ADG (Horticulture and Plantation crops), Indian Council of
Agricultural Research, new Delhi for his encouragement and keen interest.
Authors express their deep sense of gratitude to Dr. D.G. Dhandar, Director,
Central Institute of Arid Horticulture, Bikaner for giving more thrust to increase
quality publications. We are also thankful to Dr. O.P. Awasthi, Dr. R.S. Singh,
Dr. D. Singh and Dr. D.K. Samdia for their critical suggestions. Authors are
thankful to Mr. D.C. Joshi, Mr. Sanjay Patil, Mr. M.K. Jain, B.R. Khatri and
Mr. Akhil Thukral of CIAH, Bikaner for their help during writing this book.
Thanks are due to Dr. J. Singh, Assistant Professor (Horticulture), Rajasthan
Agriculture University, Bikaner for thorough checking of the manuscript.

xiv Fruit Breeding
Authors books, book chapters, research papers, popular articles and technical
papers used in writing of this book have duly been acknowledged in the text.
The authors will appreciate receiving suggestions for further improvement of
this book. We hope that information presented here in the book would receive
due response from the teaches, students, researchers and other readers.
Date: April, 2004
Anil Kumar Shukla
Place: Bikaner Arun Kumar Shukla
B.B. Vashistha

Contents
Foreword to the First Edition .............................................................................. ix
Preface of the Second Edition ............................................................................. xi
Preface of the First Edition...............................................................................xiii
Abbreviations ............................................................................................ xxxi
1. Basic Approaches in Fruit Breeding .................................................. 1
1.1 Definition .............................................................................................. 1
1.2 Major problems in fruit breeding .......................................................... 1
1.3 Objectives of fruit breeding .................................................................. 2
1.3.1 For rootstock ........................................................................... 2
1.3.2 For scion cultivars ................................................................... 2
1.4 Steps in fruit breeding .......................................................................... 3
1.5 Modes of reproduction in relation to fruit breeding ............................. 3
1.6 Types of reproduction in fruits ............................................................ 3
1.6.1 Asexual reproduction ............................................................... 3
1.6.1.1 Reproduction by apomictic seed (apomixis) ............ 4
1.6.1.1.1 Classification of apomixis ........................ 4
1.6.1.1.1.1 Recurrent apomixis................ 4
1.6.1.1.1.2 Non recurrent apomixis ......... 4
1.6.1.1.1.3 Adventitive embryony............ 4
1.6.1.1.1.4 Vegetative apomixis................ 5
1.6.1.1.1.5 Apospory ............................... 5
1.6.1.1.1.6 Parthenogenesis ..................... 5
1.6.1.1.1.7 Apogamy................................ 5
1.6.1.1.1.8 Androgamy ............................ 5
1.6.1.1.1.9 Genetics of apomixis ............. 5
1.6.1.1.1.10 Advantages of apomixis....... 6
1.6.1.1.1.11 Disadvantages of apomixis .. 6
1.6.1.1.1.12 Reproduction by the
vegetative parts of the plants 6
1.6.1.2 Reproduction through specialized parts ................... 6
1.6.2 Sexual reproduction ................................................................. 7
1.6.2.1 Floral biology and development of gametophytes .... 7

xvi Fruit Breeding
1.6.2.2 Microsporogenesis and microgametogenesis ........... 8
1.6.2.3 Megasporogenesis and megagametogenesis ............ 8
1.6.2.4 Fertilization (syngamy) ............................................. 9
1.6.2.5 Post fertilization development ................................... 9
1.6.2.6 Significance of sexual reproduction ......................... 9
1.7 Modes of pollination ............................................................................. 9
1.7.1 Conditions and mechanism of autogamy (self-pollination) ... 10
1.7.1.1 Hermaphrodite or perfect flower............................ 10
1.7.1.2 Cleistogamy ............................................................ 10
1.7.1.3 Homogamy ............................................................. 10
1.7.2 Conditions and mechanism of allogamy (cross pollination) .. 10
1.7.2.1 Unisexuality ............................................................ 10
1.7.2.2 Dichogamy ............................................................. 10
1.7.2.3 Self incompatibility ................................................. 11
1.7.2.4 Male sterility ........................................................... 11
1.7.2.5 Heterostyly.............................................................. 11
1.7.2.6 Heteromorphism ..................................................... 11
1.7.2.7 Environmental factors ............................................ 11
1.7.3 Agencies of pollination ........................................................... 11
1.8 Self incompatibility ............................................................................. 12
1.8.1 Genetic control of self incompatibility................................... 12
1.8.1.1 Gametophytic self incompatibility .......................... 12
1.8.1.2 Sporophytic incompatibility .................................... 12
1.8.2 Mechanism of self incompatibility ......................................... 12
1.8.2.1 Pollen stigma interaction ......................................... 13
1.8.2.2 Pollen tube style interaction .................................... 13
1.8.2.3 Pollen tube ovule interaction ................................... 13
1.8.3 Methods of overcoming self-incompatibility ......................... 13
1.8.4 Advantages of self-incompatibility......................................... 14
1.8.5 Disadvantages ........................................................................ 14
1.9 Male sterility ....................................................................................... 15
1.9.1 Genetic male sterility.............................................................. 15
1.9.1.1 Sterility maintenance ............................................... 15
1.9.1.2 Fertility restoration ................................................. 16
1.9.2 Cytoplasmic male sterility ...................................................... 16
1.9.2.1 Sterility maintenance ............................................... 16
1.9.2.2 Fertility restoration ................................................. 16
1.9.2.3 Uses ........................................................................ 17
1.9.3 Cytoplasmic-genetic male sterility ......................................... 17
1.9.3.1 Sterility maintenance ............................................... 17
1.9.3.2 Fertility restoration ................................................. 18
1.9.3.3 Uses ........................................................................ 18

Contentsxvii
1.10 Centres of origin (primary and secondary) ........................................ 18
1.10.1 China centre of origin ............................................................ 18
1.10.2 Hindustan centre of origin ..................................................... 19
1.10.3 Central Asia centre of origin .................................................. 19
1.10.4 Asia minor centre of origin .................................................... 19
1.10.5 Mediterranean cetnre of origin............................................... 20
1.10.6 Central American centre of origin .......................................... 20
1.10.7 South American centre of origin ............................................ 20
1.11 Methods of fruit breeding ................................................................... 20
1.11.1 Introduction ........................................................................... 20
1.11.2 Procedure of introduction ..................................................... 22
1.11.3 Merits of introduction ............................................................ 22
1.11.4 Demerit .................................................................................. 22
1.12 Selection ............................................................................................. 22
1.12.1 Achievements ......................................................................... 23
1.13 Clone and clonal selection .................................................................. 23
1.13.1 Characteristics ....................................................................... 23
1.13.2 Genetic variation within clones .............................................. 24
1.13.3 Advantages ............................................................................. 25
1.13.4 Limitations.............................................................................. 25
1.13.5 Achievements ......................................................................... 25
1.14 Mutation .............................................................................................. 25
1.14.1 Procedure of mutation breeding ............................................ 26
1.14.2 General characteristics of mutation ....................................... 26
1.14.3 Mutagens................................................................................ 27
1.14.3.1 Physical mutagens .................................................. 27
1.14.3.2 Chemical mutagens................................................. 27
1.14.4 Achievements ......................................................................... 27
1.15 Polyploidy breeding ............................................................................ 27
1.16 Hybridization ....................................................................................... 28
1.16.1 Types of hybridization ........................................................... 28
1.16.1.1 Intervarietal hybridization ....................................... 28
1.16.1.2 Distant hybridization ............................................... 29
1.16.2 Procedure of hybridization programme ................................. 29
1.16.2.1 Choice of parents ................................................... 29
1.16.2.2 Evaluation of parents .............................................. 29
1.16.2.3 Emasculation .......................................................... 29
1.16.2.3.1 Types of emasculation ........................... 30
1.16.2.4 Bagging ................................................................... 30
1.16.2.5 Tagging ................................................................... 30
1.16.2.6 Pollination ............................................................... 30
1.16.2.7 Harvesting and storage of F1 seeds ....................... 30
1.16.3 Achievements.... .................................................................... 30

xviii Fruit Breeding
1.17 Biotechnological approach .................................................................. 31
1.17.1 Micropropagation ................................................................... 31
1.17.2 Conservation of germplasm ................................................... 31
1.17.3 Anther culture...... ................................................................. 32
1.17.4 Overcoming crossing barriers (embryo culture) ................... 33
1.17.5 Somaclonal variation .............................................................. 33
1.17.6 Somatic hybridization ............................................................ 33
1.17.7 Molecular biology techniques ................................................ 35
1.17.7.1 Uses ........................................................................ 35
1.17.7.2 Types of molecular marker technique .................... 36
1.18 Genetic engineering ....... ................................................................... 37
1.18.1 Gene transfer technology....................................................... 38
1.18.2 Important fruit crops in which transgenic plants
are reported ............................................................................ 38
1.18.3 Role of Genetic engineering ................................................... 39
1.19 References .......................................................................................... 40
2. Breeding for Biotic Stress Resistance ............................................. 49
2.1 Advantages of resistant breeding........................................................ 49
2.2 Pathogenicity ...................................................................................... 49
2.3 Breeding for insect resistance ............................................................ 50
2.3.1 Concept ............................................................. ..................50
2.3.2 Scales ................... .............................................................. 51
2.3.2.1 Immunity ................................................................ 51
2.3.2.2 High resistance ....................................................... 51
2.3.2.3 Low resistance ....................................................... 51
2.3.2.4 Susceptibility .......................................................... 51
2.3.2.5 High susceptibility................................................... 51
2.3.2.5.1 Host evasion ........................................... 51
2.3.2.5.2 Induced resistance .................................. 52
2.3.2.5.3 Escape..................................................... 52
2.4 Type of genetic resistance .................................................................. 52
2.4.1 Non preference ...................................................................... 52
2.4.2 Antibiosis................................................................................ 53
2.4.3 Tolerance................................................................................ 53
2.4.4 Genetics of resistance ............................................................ 54
2.5 Problems associated to breeding for resistance to insect .................. 54
2.6 Mechanism of insect resistance ......................................................... 55
2.7 Breeding for disease resistance .......................................................... 56
2.7.1 Mechanism of disease resistance ........................................... 56
2.7.2 Terminology ........................................................................... 56
2.7.2.1 Disease escape ........................................................ 56
2.7.2.2 Disease resistance................................................... 57
2.7.2.3 Susceptible reaction ................................................ 57

Contentsxix
2.7.2.4 Immune reaction..................................................... 57
2.7.2.5 Resistance ............................................................... 57
2.7.2.6 Tolerance ................................................................ 57
2.7.3 Vertical and horizontal resistance ........................................... 57
2.7.3.1 Vertical resistance ................................................... 57
2.7.3.2 Horizontal resistance............................................... 59
2.7.3.2.1 Hypothesis on the horizontal resistance ... 59
2.7.3.2.2 Evidences for horizontal resistance ......... 60
2.7.3.2.2.1 Constant ranking.................... 60
2.7.3.2.2.2 Historical ................................ 60
2.7.3.2.2.3 Geographical evidence ........... 60
2.7.3.2.2.4 Epidemiological evidence ...... 60
2.7.3.2.3 Components of horizontal resistance ........... 60
2.7.3.2.3.1 Tolerance ............................... 60
2.7.3.2.3.2 Slow development of disease 60
2.7.3.2.3.3 Morphological........................ 60
2.7.3.2.3.4 Functional .............................. 60
2.7.3.2.3.5 Hypersensitive reaction ......... 61
2.7.3.2.3.6 Escape mechanism ................ 61
2.8 Breeding methods for biotic stress resistance .................................... 61
2.8.1 Introduction ........................................................................... 61
2.8.2 Selection ................................................................................. 61
2.8.3 Hybridization .......................................................................... 61
2.8.4 Mutation ................................................................................. 62
2.8.5 Source of resistance for different diseases ........................... 62
2.8.6 Production of disease resistant plant by unconventional
breeding ................................................................................. 66
2.8.6.1 Basic technique in plant cell culture ....................... 66
2.8.6.2 Genetic engineering or Recombinant DNA
technology .............................................................. 66
2.9 Breeding for resistance to parasitc weeds .......................................... 66
2.10 References .......................................................................................... 66
3. Plant Growth Regulators in Fruit Breeding .................................... 69
3.1 Multiplication of hybrids/selections .................................................... 69
3.2 Root initiation ...................................................................................... 69
3.3 Seed germination/overcoming seed dormancy................................... 70
3.4 Substitution for chilling requirement .................................................. 70
3.5 Induction of male sterility ................................................................... 70
3.6 Shortening of breeding cycle in fruit crops ....................................... 70
3.7 Influence on reproductive growth ...................................................... 71
3.8 Control of fruit drop ........................................................................... 71
3.9 Making distant crosses ....................................................................... 72
3.10 References .......................................................................................... 72

xx Fruit Breeding
4. Breeding of Mango ............................................................................. 75
4.1 Centre of diversity .............................................................................. 76
4.2 Germplasm resources ......................................................................... 76
4.3 Problems in mango breeding .............................................................. 77
4.4 Objectives ........................................................................................... 77
4.5 Botany ................................................................................................. 77
4.6 Floral biology ...................................................................................... 78
4.7 Pollination ........................................................................................... 79
4.8 Inheritance pattern .............................................................................. 79
4.9 Pre selection criteria for dwarfness (Majumder et al., 1969) ............ 80
4.10 Breeding methods and achievements .................................................. 80
4.10.1 Introduction ........................................................................... 80
4.10.2 Selection ................................................................................. 80
4.10.3 Clonal selection ...................................................................... 80
4.10.4 Hybridization .......................................................................... 81
4.10.4.1 Interspecific hybridization ....................................... 81
4.10.4.2 Improved Hybridization technique............................ 81
4.10.4.3 Promising hybrid of mango ...................................... 82
4.10.5 Mutation ................................................................................. 82
4.10.6 Polyploidy .............................................................................. 82
4.10.7 Heterosis ................................................................................ 82
4.10.8 Biotechnological tool .............................................................. 83
4.11 Important wild species of mango and their uses ............................... 83
4.12 Important new cultivars of mango (Pandey et al., 2002) .................. 83
4.12.1 Important Commercial cultivars of mango for
different regions: .................................................................... 91
4.12.2 Evaluation of mango germplasm for various traits................ 91
4.13 Future thrust ....................................................................................... 92
4.14 References .......................................................................................... 92
5. Breeding of Banana............................................................................. 95
5.1 Centre of diversity .............................................................................. 96
5.2 Germplasm resources ......................................................................... 96
5.3 Objectives ........................................................................................... 97
5.4 Botany........................... .................................................................. 97
5.5 Floral biology .................................................................................... 101
5.6 Inheritance pattern ............................................................................ 101
5.7 Breeding methods and achievements ................................................ 102
5.7.1 Introduction ......................................................................... 102
5.7.2 Hybridization ........................................................................ 102
5.7.2.1 Interdiploid hybridization (2 x 2) .......................... 102
5.7.2.2 Triploid breeding................................................... 103
5.7.2.3 Tetraploid breeding ............................................... 104

Contentsxxi
5.7.3 Mutation ............................................................................... 104
5.7.4 Biotechnology ...................................................................... 105
5.8 Source of resistance ......................................................................... 105
5.9 Important hybrids of banana ............................................................ 105
5.9.1 Important varieties of Banana .............................................. 106
5.10 Important cultivars of banana with genome..................................... 109
5.11 References ........................................................................................ 109
6. Breeding of Papaya ........................................................................... 113
6.1 Center of diversity ............................................................................ 114
6.2 Germplasm resources ....................................................................... 114
6.3 Objectives ......................................................................................... 114
6.4 Botany ............................................................................................... 115
6.5 Floral biology .................................................................................... 115
6.6 Types of sex and sex inheritance ..................................................... 115
6.7 Pollination ......................................................................................... 117
6.8 Breeding methods and achievements ................................................ 117
6.8.1 Introduction ......................................................................... 117
6.8.2 Inbreeding and selection ...................................................... 118
6.8.3 Induction of polyploidy........................................................ 119
6.8.4 Hybridization ........................................................................ 119
6.8.5 Heterosis breeding................................................................ 120
6.8.6 Mutation ............................................................................... 120
6.8.7 Biotechnology ...................................................................... 120
6.8.8 Embryo culture .................................................................... 120
6.8.9 Somatic embryogenesis ....................................................... 121
6.8.10 Transgenic papaya ............................................................... 121
6.9 Important varieties and cultivars ...................................................... 121
6.9.1 Important species of papaya ................................................ 123
6.10 References ........................................................................................ 124
7. Breeding of Citrus ............................................................................127
7.1 Centre of diversity ............................................................................ 128
7.2 Germplasm resources ....................................................................... 128
7.3 Problems in citrus breeding .............................................................. 129
7.4 Objectives ......................................................................................... 130
7.5 Botany ............................................................................................... 130
7.5.1 Kumquats (Fortunella species) ............................................ 130
7.5.2 Poncirus ............................................................................... 131
7.5.3 Citrus ................................................................................... 131
7.6 Floral biology .................................................................................... 131
7.7 Inheritance pattern ............................................................................ 132
7.8 Breeding methods and achievements ................................................ 132
7.8.1 Introduction ......................................................................... 132

xxii Fruit Breeding
7.8.2 Selection ............................................................................... 133
7.8.3 Hybridization ........................................................................ 133
7.8.3.1 Techniques ............................................................ 133
7.8.4 Mutation ............................................................................... 134
7.8.5 Polyploid breeding........... ................................................... 135
7.8.6 Biotechnology in improvement of citrus ............................. 135
7.9 Important species and cultivars ......................................................... 136
7.9.1 Mandarin group.................................................................... 136
7.9.2 Lemon (
C. lemon)................................................................ 138
7.9.3 Acid lime (
C. aurantifolia) .................................................. 139
7.9.4 Sweet Orange ...................................................................... 141
7.9.5 Grape fruit (
C. paradisi) ...................................................... 143
7.10 Citrus Hybrids ................................................................................... 144
7.10.1 Hybrid of Poncirus .............................................................. 144
7.10.2 Hybrids of Fortunella ........................................................... 144
7.10.3 Interspecific hybrids ............................................................ 145
7.11 Future thrust ..................................................................................... 145
7.12 References ........................................................................................ 145
8. Breeding of Grape .............................................................................147
8.1 Centre of diversity ............................................................................ 148
8.2 Germplasm resources ....................................................................... 148
8.3 Objectives ......................................................................................... 148
8.3.1 Objectives of breeding for North Indian grapes .................. 148
8.3.2 Objectives of breeding for the Tropics ............................... 148
8.4 Botany ............................................................................................... 148
8.5 Floral biology and pollination ............................................................ 149
8.6 Inheritance pattern ............................................................................ 150
8.7 Breeding methods and achievements ................................................ 151
8.7.1 Introduction ......................................................................... 151
8.7.2 Selection ............................................................................... 152
8.8 Hybridization ..................................................................................... 153
8.8.1 Interspecific / Intergeneric hybridization ............................. 154
8.8.2 Intervarietal hybridization..................................................... 154
8.8.3 Hybridization technique........................................................ 155
8.9 Shortening the breeding cycle ............................................................ 156
8.10 Mutation ............................................................................................ 159
8.11 Polyploidy ......................................................................................... 159
8.12 Biotechnological tools ....................................................................... 160
8.12.1 Embryo rescue technique .................................................... 160
8.12.2 Genetic engineering/ plant transformation ........................... 160
8.12.3 Protoplast culture ................................................................. 160
8.12.4 Anther culture ...................................................................... 160
8.12.5 Somaclonal variation ............................................................ 160

8.13 Important species and cultivars of grape ......................................... 161
8.13.1 Other species of American grapes ....................................... 161
8.13.2 Asiatic species ...................................................................... 161
8.13.3 Carribean grape .................................................................... 162
8.13.4 Genus Muscadinia ................................................................ 162
8.13.5 Important native species of grape ....................................... 162
8.13.6 New cultivars of grape (
Source: Pandey et al., 2002) ......... 162
8.14 References ........................................................................................ 166
9. Breeding of Guava ............................................................................171
9.1 Centre of diversity ............................................................................ 171
9.2 Germplasm resources ....................................................................... 172
9.3 Objectives ......................................................................................... 172
9.4 Botany ............................................................................................... 172
9.5 Floral biology and pollination ............................................................ 172
9.6 Inheritance pattern ............................................................................ 173
9.7 Breeding methods and achievements ................................................ 173
9.7.1 Introduction ......................................................................... 173
9.7.2 Selection ............................................................................... 173
9.7.3 Hybridization ........................................................................ 174
9.7.4 Breeding for wilt resistance ................................................. 176
9.7.5 Mutation ............................................................................... 176
9.7.6 Biotechnological tools .......................................................... 176
9.8 Classification of guava cultivars ....................................................... 176
9.8.1 Characteristics of important species and cultivars .............. 177
9.8.2 Important varieties ............................................................... 179
9.9 References ........................................................................................ 183
10. Breeding of Ber .................................................................................185
10.1 Centre of diversity ............................................................................ 185
10.2 Germplasm resources ....................................................................... 186
10.3 Objectives ......................................................................................... 186
10.4 Botany ............................................................................................... 186
10.5 Floral biology .................................................................................... 188
10.6 Pollination ......................................................................................... 189
10.7 Breeding methods and achievements ................................................ 189
10.7.1 Selection ............................................................................... 189
10.7.2 Hybridization ........................................................................ 190
10.7.3 Biotechnological tools .......................................................... 190
10.8 Characteristics of important cultivars of ber ................................... 191
10.9 References ........................................................................................ 196
11. Breeding of Aonla..............................................................................199
11.1 Centre of diversity ............................................................................ 200
11.2 Distribution ....................................................................................... 200
Contentsxxiii

xxiv Fruit Breeding
11.3 Germplasm resources ....................................................................... 200
11.4 Breeding problems ............................................................................ 200
11.5 Objectives ......................................................................................... 201
11.6 Botany ............................................................................................... 201
11.7 Floral biology .................................................................................... 202
11.8 Breeding methods and achievements ................................................ 203
11.8.1 Introduction ......................................................................... 203
11.8.2 Selection ............................................................................... 203
11.8.3 Hybridization ........................................................................ 203
11.8.4 Mutation .............................................................................. 204
11.8.5 Polyploidy ............................................................................ 204
11.8.6 Biotechnological tools .......................................................... 204
11.9 Important cultivars/ varieties of Aonla ............................................. 204
11.10 Future line of work ........................................................................... 207
11.11 References ........................................................................................ 207
12. Breeding of Pomegranate ................................................................209
12.1 Centre of diversity ............................................................................ 209
12.2 Germplasm resources ....................................................................... 210
12.3 Objectives ......................................................................................... 210
12.4 Botany ............................................................................................... 210
12.5 Floral biology .................................................................................... 211
12.6 Inheritance pattern ............................................................................ 212
12.7 Breeding methods and achievements ................................................ 212
12.7.1 Introduction ......................................................................... 212
12.7.2 Selection ............................................................................... 212
12.7.3 Clonal Selection.................................................................... 213
12.7.4 Hybridization ........................................................................ 213
12.7.5 Mutation ............................................................................... 214
12.7.6 Biotechnological tools .......................................................... 214
12.8 Important varieties and cultivars of Pomegranate ............................ 215
12.9 References ........................................................................................ 218
13. Breeding of Date Palm .....................................................................221
13.1 Centre of diversity ............................................................................ 222
13.2 Distribution ....................................................................................... 222
13.3 Germplasm resources ....................................................................... 222
13.4 Objectives ......................................................................................... 222
13.5 Botany ............................................................................................... 222
13.6 Floral biology .................................................................................... 224
13.7 Pollination ......................................................................................... 224
13.8 Pollination Technique ........................................................................ 225
13.9 Metaxenia .......................................................................................... 226
13.10 Breeding methods and achievements ................................................ 226

13.10.1 Introduction ......................................................................... 226
13.10.2 Selection............................................................................... 227
13.10.3 Hybridization ........................................................................ 227
13.10.4 Biotechnological tool ............................................................ 228
13.11 Important cultivars of date palm grown in different
countries (Pareek, 1990) .................................................................. 228
13.12 References ........................................................................................ 229
14. Breeding of Litchi .............................................................................231
14.1 Centre of diversity ............................................................................ 232
14.2 Distribution ....................................................................................... 232
14.3 Germplasm resources ....................................................................... 233
14.4 Breeding objectives ........................................................................... 234
14.5 Botany ............................................................................................... 234
14.6 Floral biology and pollination ............................................................ 235
14.7 Inheritance pattern ............................................................................ 236
14.8 Breeding methods and achievements ................................................ 236
14.8.1 Introduction ......................................................................... 236
14.8.2 Selection............................................................................... 236
14.8.3 Hybrdization ......................................................................... 237
14.8.4 Biotechnological tools .......................................................... 238
14.9 Important cultivars and varieties of Litchi ....................................... 238
14.10 References ........................................................................................ 241
15. Breeding of Coconut .........................................................................243
15.1 Centre of diversity ............................................................................ 244
15.2 Germplasm resources ....................................................................... 244
15.3 Problems ........................................................................................... 244
15.4 Objectives ......................................................................................... 244
15.5 Botany ............................................................................................... 245
15.5.1 Tall palms ............................................................................. 245
15.5.2 Dwarf palms ........................................................................ 245
15.5.3 Intermediate types................................................................ 245
15.6 Floral biology and pollination ............................................................ 246
15.7 Breeding methods and achievements ................................................ 246
15.7.1 Introduction ......................................................................... 246
15.7.2 Selection............................................................................... 247
15.7.2.1 Selection of mother palm ..................................... 247
15.7.2.2 Collection of seed nuts ......................................... 248
15.7.2.3 Seedling selection ................................................. 248
15.7.2.4 Selection of prepotential palm .............................. 248
15.7.3 Hybridization ........................................................................ 249
15.7.3.1 Dwarf x Dwarf (DxD) ......................................... 249
15.7.3.2 Tall x Dwarf (T x D) ............................................ 249
Contentsxxv

xxvi Fruit Breeding
15.7.3.3 Dwarf x Tall (D x T) ............................................ 250
15.7.3.4 Tall x Tall (T x T) ................................................. 250
15.8 Important coconut cultivars (Anon., 1998) ..................................... 250
15.8.1 Hybrids of Coconut (Anon., 1998) ..................................... 251
15.8.2 Coconut varieties released by CPCRI and SAUs for
commercial cultivation ......................................................... 251
15.9 References ........................................................................................ 254
16. Breeding of Cashewnut ....................................................................255
16.1 Centre of diversity ............................................................................ 256
16.2 Distribution ....................................................................................... 256
16.3 Germplasm resources ....................................................................... 257
16.4 Objectives ......................................................................................... 257
16.5 Botany ............................................................................................... 258
16.6 Floral biology and pollination ............................................................ 258
16.7 Inheritance pattern ............................................................................ 258
16.8 Breeding methods and achievements ................................................ 259
16.8.1 Introduction ......................................................................... 259
16.8.2 Selection ............................................................................... 259
16.8.3 Hybridization ........................................................................ 259
16.8.3.1 Hybridization technique ........................................ 260
16.8.3.2 Achievements ....................................................... 260
16.8.4 Mutation ............................................................................... 260
16.9 Brief description of important varieties ............................................ 260
16.10 References ........................................................................................ 269
17. Breeding of Pineapple ......................................................................271
17.1 Centre of diversity ............................................................................ 271
17.2 Germplasm resources ....................................................................... 272
17.3 Objectives ......................................................................................... 272
17.4 Botany ............................................................................................... 272
17.5 Floral biology .................................................................................... 273
17.5.1 Classification of Pineapple ................................................... 273
17.6 Inheritance pattern ............................................................................ 274
17.7 Breeding methods and achievements ................................................ 274
17.7.1 Selection ............................................................................... 274
17.7.2 Hybridization ........................................................................ 275
17.7.3 Mutation ............................................................................... 275
17.7.4 Biotechnological tools .......................................................... 275
17.8 Major group and varieties of Pineapple ............................................ 276
17.9 References ........................................................................................ 276
18. Breeding of Underutilized Fruits ....................................................279
18.1 Bael ................................................................................................... 279
18.1.1 Centre of diversity ............................................................... 279

18.1.2 Distribution .......................................................................... 280
18.1.3 Germplasm resources .......................................................... 280
18.1.4 Objective .............................................................................. 280
18.1.5 Botany .................................................................................. 280
18.1.6 Breeding of underutilized fruits ............................................ 281
18.1.7 Important varieties and cultivars of Bael ............................. 281
18.1.8 References ........................................................................... 283
18.2 Jamun ............................................................................................... 284
18.2.1 Centre of diversity ............................................................... 284
18.2.2 Distribution .......................................................................... 285
18.2.3 Botany .................................................................................. 285
18.2.4 Floral biology ....................................................................... 285
18.2.5 Objectives ............................................................................ 286
18.2.6 Conservation and Breeding .................................................. 286
18.2.7 Breeding of underutilized fruits ............................................ 287
18.2.7.1 Selection ............................................................... 287
18.2.7.2 Morphological and Molecular Methods ................ 287
18.2.7.3 Hybridization ......................................................... 287
18.2.8 Important varieties and cultivars of Jamun ......................... 288
18.2.9 References ........................................................................... 290
18.3 Karonda............................................................................................. 291
18.3.1 Centre of diversity ............................................................... 291
18.3.2 Distribution .......................................................................... 292
18.3.3 Botany .................................................................................. 292
18.3.4 Floral biology ....................................................................... 292
18.3.5 Objectives ............................................................................ 293
18.3.6 Breeding of Karonda ............................................................ 293
18.3.6.1 Selection ............................................................... 293
18.3.6.2 Hybridization ......................................................... 293
18.3.7 Important varieties and cultivars of Karonda ...................... 294
18.3.8 References ........................................................................... 295
18.4 Phalsa ................................................................................................ 296
18.4.1 Centre of diversity ............................................................... 296
18.4.2 Distribution .......................................................................... 297
18.4.3 Botany .................................................................................. 297
18.4.4 Floral biology ....................................................................... 297
18.4.5 Objectives ............................................................................ 298
18.4.6 Breeding of Phalsa ............................................................... 298
18.4.7 Varieties and cultivars .......................................................... 298
18.4.8 References ........................................................................... 299
18.5 Khirni/Rayan ..................................................................................... 300
18.5.1 Centre of diversity ............................................................... 300
18.5.2 Distribution .......................................................................... 301
Contentsxxvii

xxviii Fruit Breeding
18.5.3 Botany and floral biology ..................................................... 301
18.5.4 Breeding of Khirni/Rayan ..................................................... 301
18.5.5 Varieties and cultivars .......................................................... 303
18.5.6 References ........................................................................... 303
18.6 Jackfruit ............................................................................................ 304
18.6.1 Center of diversity and distribution ..................................... 304
18.6.2 Botany .................................................................................. 305
18.6.3 Floral biology ....................................................................... 305
18.6.4 Objectives ............................................................................ 306
18.6.5 Breeding of Jackfruit ........................................................... 306
18.6.6 Important varieties and cultivars of Jackfruit...................... 306
18.6.7 References ........................................................................... 307
19. Breeding of Temperate Fruits.........................................................309
19.1 Apple ................................................................................................. 309
19.1.1 Center of origin and distribution .......................................... 309
19.1.2 Germplasm resources .......................................................... 310
19.1.3 Objective .............................................................................. 311
19.1.3.1 For Scion .............................................................. 311
19.1.3.2 For rootstocks ...................................................... 311
19.1.4 Botany .................................................................................. 311
19.1.5 Floral biology ....................................................................... 312
19.1.6 Inheritance pattern ............................................................... 312
19.1.7 Breeding Methods and achievements................................... 313
19.1.7.1 Introduction and selection .................................... 313
19.1.8 Important Varieties and cultivars ......................................... 313
19.1.8.1 Promising cultivars in major production
region of India ...................................................... 313
19.1.8.2 Varieties based on bearing habit (Ted, 2002) ........ 314
19.1.8.3 Pollinizing cultivars (Verma, 2013) ....................... 314
19.1.9 References ........................................................................... 315
19.2 Pear ................................................................................................... 316
19.2.1 Center of origin and distribution .......................................... 316
19.2.2 Varieties and Cultivars .......................................................... 317
19.2.3 References ........................................................................... 317
19.3 Peach ................................................................................................ 318
19.3.1 Centre of origin and distribution .......................................... 318
19.3.2 Germplasm resources .......................................................... 319
19.3.3 Objectives ............................................................................ 319
19.3.3.1 For Scion .............................................................. 319
19.3.3.2 For rootstocks ...................................................... 320
19.3.4 Botany .................................................................................. 320
19.3.5 Floral biology ....................................................................... 320
19.3.6 Breeding methods and achievements ................................... 321

19.3.6.1 Introduction .......................................................... 321
19.3.6.2 Selection ............................................................... 322
19.3.6.3 Hybridization ......................................................... 322
19.3.6.4 Mutation Breeding................................................. 322
19.3.7 References ........................................................................... 323
19.4. Plum .................................................................................................. 325
19.4.1 Center of origin and distribution .......................................... 325
19.4.2 Varieties and Cultivars .......................................................... 327
19.4.3 References ........................................................................... 327
19.5. Apricot .............................................................................................. 328
19.5.1 Center of diversity and distribution ..................................... 328
19.5.2 Cultivars and Varieties .......................................................... 329
19.5.3 References ........................................................................... 329
19.6. Cherry ............................................................................................... 331
19.6.1 Center of diversity and distribution ..................................... 331
19.6.2 Varieties and Cultivars .......................................................... 332
19.6.3 References ........................................................................... 332
19.7. Strawberry ........................................................................................ 333
19.7.1 Center of origin and distribution .......................................... 333
19.7.2 Important varieties and cultivars ......................................... 335
19.7.3 References ........................................................................... 336
Appendix ...............................................................................................337
Botanical name, family and origin of important fruit crops........................ 338
Contentsxxix

Abbreviations
2,4 – D2,4 Dichlorophenoxy Acetic Acid
ABM
Agrobacterium mediated
AES Agriculture Experiment Station
AFLP Amplified Fragment Length Polymorphism
AICRP All India Coordinated Research Project
APAU Andhra Pradesh Agricultural University
AP-PCR Arbitrarily Primered Polymerase Chain Reaction
BAC Bihar Agricultural College
BCKV Bidhan Chandra Krishi Vishwavidyalaya
CAP Cleaved Amplified Polymorphism
CHES Central Horticulture Experiment Station
CIAH Central Institute for Arid Horticulture
CISH Central Institute for Subtropical Horticulture
CPCRI Central Plantation Crop Research Institute
DBT Department of Biotechnology
DNA Deoxyribonucleic Acid
E M East Malling
EC Exotic Collection
EESEthyl Methane Sulphonate
EPElectroporation
FRS Fruit Research Station
FW-450 1,2 Dichloro- iso – butyrate
GA
3
Gibberellic Acid
GAU Gujarat Agricultural University
GBPUAT Govind Ballabh Pant University of Agriculture and Technology
HARP Horticulture and Agroforestry Research Programme
HAU Haryana Agricultural University
HETC Horticultural Experiment and Training Centre
HS Highly Susceptible
IARIIndian Agricultural Research Institute
IBAIndole Butyric Acid
ICIndigenous Collection
IIHRIndian Institute of Horticulture Research
IITAInternational Institute for Tropical Agriculture
IPM Integrated Pest management
IW Indigenous Wild Collection
KAU Kerala Agricultural University
LD Lethal Dose
MAS Marker Assisted Selection
M H Maleic Hydrazide
MMC Megaspore Mother Cell
MS Moderately Susceptible

xxxii Fruit Breeding
NAA Naphthalene Acetic Acid
NBPGR National Bureau of Plant Genetic Resources
NDUAT Narendra Dev University of Agriculture and Technology
NMU N-Nitrose-N-Methyl-Urea
NMUT N-Nitroso-N-Methyl Urethane
NRC National Research Centre
PB Particle Bombardment
PB+ AMB Particle Bombardment followed by
Agrobacterium Co-cultivation
PEG Polyethylene Glycol
PGM Polyethylene Glycol Mediated
PGR Plant Growth Regulator
PHB Polyhydroxy Butyrate
PMC Pollen Mother Cell
PRSV Papaya Ring Spot Virus
RResistant
RAPD Random Amplified Polymorphic DNA
RAU Rajendra Agricultural University
RFLP Restriction Fragment Length Polymorphism
RFRS Regional Fruit Research Station
SSusceptible
SADH Dimethylamino Succinamic Acid
SCAR Sequence Characterized Amplified Regions
SPAR Single Primer Amplification Reaction
SSCP Single-Strand Conformation Polymorphism
SSRP Single Sequence Repeat Polymorphism
STSSequenced Tagged Sites
TIBA 2,3,5, Tri iodobenzoic Acid
TNAU Tamil Nadu Agricultural University
TSSTotal Soluble Solids

Basic Approaches in Fruit Breeding
India is bestowed with a wide range of agro-climatic and soil conditions.
Therefore, almost all types of fruits can be grown in one or the other parts of
the country. India is the second largest producer of fruits next to China. In
India, fruit crops occupied 6.5 million hectares and it produces 96.75 million
metric tonnes of fruits. India has exported fruits worth Rs. 4817.35 crores
(692.01 USD Millions) (National Horticulture Board).
Improvement of fruit crops is difficult because of long gestation period, high
heterozygosity, scanty information on inheritance pattern, often cross pollination,
excessive fruit drop, parthenocarpy and less number of seeds restricting the
availability of hybrid seedlings for evaluation. Even though, planned hybridization
and clonal selections have been attempted in a number of fruit crops and these
efforts have resulted in the development of promising varieties (mango, grape, guava,
papaya, sapota, banana, etc). However, systematic and dedicated efforts are still
required for the development of ideal varieties through modern tools in fruit crops.
In future, search for desired gene, critical study of inheritance pattern and use
of biotechnological tools require due attention in combining ideal characteristics
in varietal improvement programme of fruit crops.
1.1. Definition
Fruit breeding is the manipulation of a biological system that requires many
generations to achieve results. It is also a dynamic, exciting and challenging
profession, operating under continually changing conditions.
1.2. Major problems in fruit breeding
It has long generation cycle of 2-10 years depending upon species and
cultivars.
It is not possible to have more recombination.
Fruit crop has long juvenile period due to that early assessment of strains
are not possible e.g. mango,
Madhuka latifolia, jack fruit etc.
1

2Fruit Breeding
Majority of the fruit species are highly heterozygous, therefore, large
population is required for selection.
Most of the fruit species are polyploid in nature e.g. ber, banana etc.
Polyembryony e.g. citrus, mango.
Parthenocarpy and seedlessness e.g. banana, pineapple etc.
Presence of sexual incompatibility e.g. mango, apple, pear, loquat etc.
More number of chromosomes, creating a big problem in genetical analysis
e.g. ber, mulberry.
Excessive fruit drop e.g. mango, citrus, grape etc.
Presence of single seed in single fruit, requiring more number of crossing
e.g. mango, litchi, mahua etc.
1.3. Objectives of fruit breeding
The objectives of fruit breeding vary with the fruit plants in relation to different
location and requirements. Even though, the main objective of fruit breeding
is to get maximum quality production per unit area with low cost and free from
biotic and abiotic stresses. The objectives may also vary with respect to breeding
for rootstock and scion.
1.3.1 For rootstock
Wide geographical adaptability.
Easily propagated, preferably through asexual means.
It should be compatible with most of the scion cultivars having strong
scion-stock union and more longevity.
It should be resistant to biotic and abiotic stresses.
For high density planting, it should have dwarfing effect without affecting
the productivity of scion cultivars.
Plants should not have brittle root system e.g. EM 9 rootstock of apple.
It should be free from suckering habit.
1.3.2 For scion cultivars
Dwarf growth stature.
Regular, precocious and prolific bearer per unit canopy area.
High productivity with good quality fruits.

Basic Approaches in Fruit Breeding3
Resistant to biotic and abiotic stresses.
Attractive fruit colour with pleasant aroma.
Suitable for processing and export.
Good keeping and transport quality.
1.4. Steps in fruit breeding
Introduction
Domestication
Germplasm
Collection
Natural variability
Mass of
variability
available
Mutation
Somaclonal variation
Polyploidy
Hybridization
Selection of ideal
ideotype (elite plants)
Performance
evaluation
Mass multiplication
and distribution
Created variability
1.5. Modes of reproduction in relation to fruit breeding
The reproduction system is principally responsible for perpetuation and
preservation of a particular genotype. Further, the mode of reproduction also
determines the genetic constitution of fruit crop whether it is homozygous or
heterozygous (Singh, 1990).
1.6. Types of reproduction in fruits
1.6.1 Asexual reproduction
By using apomictic seeds e.g. citrus, mango, apple.
By using vegetative parts of the plants e.g. mango, banana, guava, aonla,
bael, date palm, apple, peach, plum, etc.

4Fruit Breeding
1.6.1.1 Reproduction by apomictic seed (apomixis)
Apomixis refers to the development of embryo without the usual process of
meiosis and fertilization. The plants developed from apomictic seeds are true
to type (Raghaven,1986, Naumova, 1993, Nyrgen, 1954).
Obligate apomictic seeds develop with or without pollination but without
fertilization. e.g.
Malus spp. However, in case of facultative apomictic seeds
both types of embryo may develop i.e. zygotic as well as nucellar embryo
(Maheshwari and Sachar, 1963, Bhatnagar and Johri, 1972) e.g. citrus and
polyembryonic cultivars of mango.
1.6.1.1.1 Classification of apomixis
1.6.1.1.1.1 Recurrent apomixis
The embryo sac (female gametophyte) develops from the megaspore mother
cell where meiosis is disturbed (sporogenesis failed) or from adjoining cell
(megaspore mother cell disintegrates). The egg cell is diploid and embryo
develops directly from the diploid egg cell without fertilization. Generally,
somatic apospory, diploid parthenogenesis and diploid apogamy fall under
recurrent apomixis.
Example:
Rubus sp. (Raspberry), Malus hupehensis, Malus sikkimensis, Malus
sargenti
and Malus toringoides (Mitra, 1991, Vashishtha et al., 2004).
1.6.1.1.1.2 Non recurrent apomixis
The development of embryo takes place from haploid egg cell without
fertilization. Such type of apomixis rarely occurs. Generative apospory, haploid
parthenogenesis, haploid apogamy and androgamy fall under this category.
1.6.1.1.1.3 Adventitive embryony
This is also known as nucellar embryony, nucellar budding or polyembryony. In
this case more than one embryo develops in a single seed. In the seed both
types of embryo develops i.e. nucellar embryo from nucellar cell and zygotic
embryo from egg cell with the result of syngamy.
Example: Olour, Goa, Kurukkan, Bappakai, Vellaikolamban, Nileswar dwarf,
Salem, Bellary, Goakasargod, Mazagaon, Chandrakaran etc cultivars of mango
(Majumder and Sharma, 1991) and most of the species of citrus except
Citrus
medica
(citron), Citrus grandis (pummelo or shaddock) and Citrus latifolia
(Ghosh, 1991, Vashishtha et al., 2004).

Basic Approaches in Fruit Breeding5
1.6.1.1.1.4 Vegetative apomixis
This is not common in fruit crops. However, in some cases like Poa bulbosa
and some Allium, Agave and grass species produce vegetative buds or bulbils
instead of flower in the inflorescence.
1.6.1.1.1.5 Apospory
Some times when embryo sac develops from archisporial or from the nucellus
or from other cell. If it develops from haploid megaspore cell, it is known as
generative or haploid apospory. On the other hand, if it develops from diploid
cell i.e. nucellus of other cells it is termed as somatic or diploid apospory.
1.6.1.1.1.6 Parthenogenesis
It can be defined as development of embryo from egg cell with or without
pollination but without fertilization. Depending upon the ploidy levels of egg
cell, parthenogenesis can be halploid (non recurrent) and diploid (recurrent
type) e.g. mangosteen (
Garcinea mangostana).
1.6.1.1.1.7 Apogamy
Development of embryo from synergids or antipodal cells within the embryo
sac with or without pollination but without fertilization is termed as apogamy.
This type of apomixis is also grouped into haploid and diploid apogamy
depending upon the ploidy level of cell. Diploid apogamy is recurrent type
whereas, haploid aopgamy is non recurrent type.
1.6.1.1.1.8 Androgamy
Development of the embryo from male gametes inside or out side of embryo
sac is known as androgamy. Since the cells are haploid in nature therefore, it
comes under non recurrent type.
1.6.1.1.1.9 Genetics of apomixis
Stebbins (1958) stated that, as a rule, the apomictic condition is recessive to
sexuality, although polyploid apomicts show tendency towards dominance.
However, this recessiveness is not usually due to a monogenic difference. Since
there is frequent reversion of apomicts to normal sexuality or sterility or some
abnormal genetic behavior in crosses involving an apomict and an amphimictic
or involving two apomicts of diverse origins, it appears that a successful apomict
cycle is the result of an interaction of many gene which tend to break on
hybridization. It is only in the relatively simple type of apomixis like adventive
embryony and vegetative reproduction that simple genetic behaviour can be
expected. Recently, Vardy
et al. (1989) recorded three recessive genes with
additive effects which are responsible for parthenocarpy.

6Fruit Breeding
1.6.1.1.1.10 Advantages of apomixis
1. Faster multiplication of genetically uniform individual can be done with-
out any problem of segregation.
2. Heterosis or hybrid vigour can permanently be fixed in plants.
3. Efficient exploitation of maternal effect (if present) is possible from gen-
eration to generation.
1.6.1.1.1.11 Disadvantages of apomixis
Apomixis is nuisance when the breeder desires to obtain sexual progeny i.e.
self or hybrid.
1.6.1.1.1.12 Reproduction by the vegetative parts of the plants
In this method of reproduction, a new plant develops from a portion of the plant
body. Based on the different plant parts used it can be classified in to following
groups
1.6.1.2. Reproduction through specialized parts
Offshoot : date palm
Sucker : banana, pineapple
Runner : strawberry
Others : e.g. tuber, stolon, bulb, rhizome and corms etc. These are not common
in fruit crops.
1.6.1.2.1 Cutting
Stem cutting
Hard wood cutting : grape, fig, mulberry, guava, mango, quince,
currant, pomegranate etc.
Semi hard wood cutting : citrus, olive etc.
Soft wood cutting : crab apple, apple, peach, pear, plum, apricot,
cherry etc.
Herbacious cutting : Most common in ornamental plants.
Root cuttingapple, crab apple, blackberry, raspberry
Leaf cuttingNot common in fruit plants.
Layering Air layering: litchi
Stool layering: guava, apple rootstock, pear etc.
Trench layering: apple and pear rootstocks.

Basic Approaches in Fruit Breeding7
Compound layering: muscadine grapes.
Tip layering: blackberry, raspberry, gooseberry.
Grafting andmango, guava, cashewnut, aonla, bael, ber, citrus, grape, apple,
budding pear, plum, almond etc.
1.6.2 Sexual reproduction
It is also known as amphimixis. Multiplication is done through seed that develop
after normal meiosis followed by syngamy e.g. phalsa, karonda, papaya etc.
There are four important steps involved in reproductive process.
(i)Development of gametophytes and gametes
(ii)Pollination
(iii) Fertilization or syngamy
(iv)Post fertilization development, i.e. formation of embryo, seed and fruit.
1.6.2.1 Floral biology and development of gametophytes
A normal flower consists of calyx, corolla, androecium and gynoecium. The
last two parts of flowers have great significance in entire process of
reproduction. When androecium and gynoecium both are present in same flower
it is known as hermaphrodite or bisexual flower e.g. ber, citrus, guava, sapota,
etc.
When male (androecium) and female (gynoecium) parts of flower are present
in separate flower but on same plant it is known as monoecious condition e.g.
aonla, walnut, hazelnut, jackfruit etc.) but when they are present on different
plants separately it is known as dioecious condition e.g. papaya, date palm,
kiwi fruit etc. Andromonoecious condition is found in mango where male and
hermaphrodite flower are present in same panicle. Futher, the gynodioecious
condition is found in some cultivars of papaya (Pusa Delicious, Pusa Majestic
etc.)
On the basis of occurrence of flower, there are different types of inflorescence
found in fruit crops.
Racemose :
i)Catkin: mulberry, chestnut, walnut, pecannut.
ii)Corymb: pear.
iii)Raceme: wild cherry, currents, gooseberry, blackberry, blueberry, rasp-
berry.

8Fruit Breeding
Solitary: orange, quince, guava, peach, apricot, almond.
Cymose inflorescence: citrus, phalsa, sapota, strawberry, persimmon.
i)Fasicle: cherry, plum, ber.
ii)Panicles: grape, mango, litchi, loquat, pistachionut.
Anomolus inflorescence: apple.
Special types inflorescence:
i)Spadix: date palm, banana.
ii)Hypanthodium: fig.
1.6.2.2 Microsporogenesis and microgametogenesis
Production of microspore from pollen mother cell (PMC) is termed as
microsporogenesis. Generally, each anther has four pollen sacs which contain
numerous pollen mother cells (PMC’s). After meiosis, each PMC results into
four haploid cells or microspores. After thickening of the cell wall of microspore,
it is termed as the pollen grain.
On other hand, microgametogenesis refers to the production of male gametes.
During maturation of the pollen, the microspores nucleus divides mitotically
to produce a generative and a vegetative or tube nucleus. The pollen is generally
released in this binucleate stage. When the pollen reaches the stigma of flower
it is termed as pollination. Shortly after pollination, pollen grain starts
germination. The pollen tube enters in to stigma and grows through the style.
The generative nucleus now undergoes a mitotic division to produce two male
sperms. The pollen alongwith pollen tube is known as microgametophyte. The
process between microspore to microgametophyte is termed as
microgametogenesis. Pollen tube finally enters in to ovule through a small
pore, micropyle, and discharges the two sperms in to the embryo sac.
1.6.2.3 Megasporogenesis and megagametogenesis
Megasporogenesis can be defined as production of megaspore from megaspore
mother cells (MMC’s). Megasporogenesis occurs in ovule which are present
inside the ovary. A single cell in each ovule differentiates into a megaspore
mother cell. The megaspore mother cell undergoes meiosis to produce four
haploid megaspores. Three of them degenerate leaving one functional
megaspore per ovule.
Under the process of megagametogenesis, the nucleus of a functional megaspore
divides mitotically to produce four or more nuclei. The exact number of nuclei
and their arrangement varies considerably from one species to another species.

Basic Approaches in Fruit Breeding9
In most of the cases megaspore nucleus undergoes three mitotic divisions to
produce eight nuclei. Three of these nuclei move to one pole and produce
central egg cell and two synergid cell. One synergid is situated on either side of
the egg cell. Another three nuclei migrate to the opposite pole to give rise an
antipodal cell. Remaining two nuclei in the centre are known as polar nuclei
and after fusion forms the secondary nucleous. Megaspore develops into a
mature megagametophyte or embryo sac and the development of embryo sac
from a megaspore is known as megagametogenesis.
1.6.2.4 Fertilization (syngamy)
Fusion of the one of the two sperms with egg cell, producing a diploid zygote, is
known as fertilization or syngamy. The fusion of remaining sperm with the
secondary nucleus leading to the formation of a triploid primary endosperm
nucleus is termed as triplet fusion. When triplet fusion and syngamy takes
place simultaneously it is termed as the double fertilization.
1.6.2.5 Post fertilization development
A series of changes in the ovule follows as the fertilization is over. Synergids
and antipodal cells become disorganized, and the egg cell becomes covered
with a cell wall, known as oospore. This follows the development of seed. The
various parts of gynoecium develop into ovary-fruit, ovule-seed, integument
outer-testa (outer seed coat), integument inner tegmen (inner seed coat),
secondary nucleus after fertilization-endosperm (triploid), egg cell (after
fertilization) embryo, nucellus-perisperm (nutritive tissue like endosperm).
1.6.2.6 Significance of sexual reproduction
Sexual reproduction makes it possible to combine genes from two parents into
a single hybrid. Recombination of these genes produces a large number of
genotypes. This is an essential step in creating variation through hybridization.
Almost the entire fruit breeding is based on sexual reproduction except clonal
selection which is perpetuated by the vegetative means.
1.7 Modes of pollination
Transfer of pollen grains from anther to stigma is termed as pollination. When
the pollen grain from an anther falls on the stigma of same flower on same
plant (variety) it is termed as self-pollination or autogamy. But if pollen grain
from flower of a plant is transferred to the stigma of flower of another plant
(variety), it is known as cross pollination or allogamy. Third situation is known
as geitonogamy when pollen from flower of a plant falls on the stigma of other
flowers of the same plant (variety). The genetic consequences of geitonogamy
are the same as those of autogamy.

10Fruit Breeding
1.7.1 Conditions and mechanism of autogamy (self-pollination)
In self-pollinated species generally 5% cross pollination takes place. The degree
of cross pollination in self-pollinated species is affected by cultivars, location
and environmental conditions like temperature, humidity etc. Hermaphrodite
nature of flower, cleistogamy, and homogamy mechanism of flower favours to
self-pollination.
1.7.1.1 Hermaphrodite or perfect flower
In such type of flowers both male and female parts are present in a single
flower. It leads to self-pollination. Although in some cases perfect flower does
not allow the self-pollination due to self-incompatibility, sterility and
dichogamy. Self-pollinated fruits are guava, citrus etc.
1.7.1.2 Cleistogamy
Dehiscence of the pollen grains takes place before opening of the flower e.g.
grape, sapota.
1.7.1.3 Homogamy
The male and female parts of the flowers mature at same time which favours
to self pollination. e.g. citrus, peach and apricot.
1.7.2 Conditions and mechanism of allogamy (cross pollination)
In cross pollinated species 5-10% self pollination takes place.
The mechanisms which facilitate the cross pollination are as under.
1.7.2.1 Unisexuality
It is also known as dicliny.
i)Monoecious plants: jackfruit, aonla, walnut, hazelnut, pecannut
ii)Dioecious: papaya, kiwifruit, date palm etc.
iii)Andromonoecious: mango
1.7.2.2 Dichogamy
Stamens and pistils of hermaphrodite flowers may mature at different time
facilitating cross pollination. It is of two types (a) protandry in which
androecium matures first than gynoecium, e.g. walnut and coconut (b) protogyny
where gynoecium matures first than androecium e.g. annona, banana, plum,
pomegranate, avocado and fig etc.

Basic Approaches in Fruit Breeding11
1.7.2.3 Self incompatibility
It refers to the failure of pollen from a flower to fertilize the same flower or
other flowers on the same plant e.g. almond, apple, mango, pear, pineapple,
cherry, and apricot etc. The sporophytic incompatibility is found in mango
whereas gametophytic incompatibility is found in loquat.
1.7.2.4 Male sterility
Presence of nonfunctional pollen grain in the flower favours to cross pollination
e.g. triploid banana, J.H. Hale cutivar of peach, triploid apple varieties such as
Bramley’s Seedling, Blenheim Orange, King of Topkins etc and pear varieties
Beurre-De-Amanlis, Marguerite Marillat are mostly sterile (Crane and
Lawrence,1956).
1.7.2.5 Heterostyly
Presence of variable length of style refers to heterostyly condition. Which favours
to cross pollination. Flowers of pomegranate possess this condition.
1.7.2.6 Heteromorphism
Dimorphism and trimorphism also facilitate the cross pollination. e.g. pin type
of flowers are found in sapota and pomegranate whereas thrumb type of flowers
are found in almond.
1.7.2.7 Environmental factors
Strawberries grown in rich soils have been reported to produce sterile pollens
(Darrow,1957). Pear was found to be self-fertile in rich soil of California.
(Gourley and Howlett, 1955). Similarly, Bose pear is self fruitful in New York
and regularly parthenocarpic in South Africa (Reinicke, 1930).
1.7.3 Agencies of pollination
Based on the pollinating agent fruits can be grouped into (i) Anemophillus (by
wind), (ii) Hydrophillus (by water), (iii) Entomophillus (by insect). The wind
and insect are the major pollinating agents.
Fruits and their pollinating agents are as under
S. No.Pollinating agents Fruit crops
1. House fly mango
2. Hony bee, wasp and other insects apple, pear, plum, almond
3. Wind papaya, date palm, walnut, aonla
4. Bird banana and pineapple

12Fruit Breeding
1.8 Self incompatibility
It is the inability of a functional male and female gametes of the hermaphrodite
flowers to set seeds on self pollination.
1.8.1 Genetic control of self incompatibility
Incompatibility is generally controlled by a special gene at S-locus represented
by multiple allelic series in the population, each of these alleles control the
formation of a specific substance that determines the incompatibility reactions,
both in the pistil and pollen. Identical substances specified by identical genes
in pollen and pistil favours to prevent fertilization. Based on the timing and
mode of S-gene activity, the incompatibility reaction among homomorphic
angiosperm is categorized into two group.
(A)Gametophytic control of pollen reaction
(B)Sporophytic control of pollen reaction
1.8.1.1 Gametophytic self incompatibility
In this type of incompatibility, pollen is binucliate and pollen behaviour is
determined by the S allele present in each pollen and in this type of
incompatibility stigma is wet type. It means the incompatibility reaction of
pollen is determined by its own genotypes, and not by the genotype of the
plant on which it is produced. Generally, incompatibility reaction is determined
by a single gene having multiple alleles. Sometimes, polyploidy may lead to
the loss of incompatibility due to a competition between the two S alleles
present in diploid pollen. Important examples are pineapple, loquat, apple,
pear, plum, cherry, almond, apricot, some citrus and members of Solanaceae
family.
1.8.1.2 Sporophytic incompatibility
The incompatibility reaction of pollen is governed by the genotype of plant on
which the pollen is produced, not by the genotype of the pollen. It means the
incompatibility is imposed by the maternal genotype, due to that all the pollen
grains from a given plant behave similarly. Incompatibility occurs at the
stigmatic surface resulting in the inhibition of pollen germination. Pollens are
trinucleate and the stigmatic surface is dry e.g.
Mangifera indica.
1.8.2 Mechanism of self incompatibility
Based on the various phenomenon observed in self incompatible mating it can
be grouped in to three :

Basic Approaches in Fruit Breeding13
i)Pollen stigma interaction.
ii)Pollen tube style interaction.
iii)Pollen tube ovule interaction.
1.8.2.1 Pollen stigma interaction
This interaction occurs just after the pollen grains reach the stigma and generally
prevent pollen germination. In the gametophytic system, stigma surface is
plumose having elongated receptive cells and is commonly known as wet stigma.
Incompatibility reaction occurs at a later stage. There are clear cut serological
differences among the pollen grains with different S genotypes, such differences
have not been observed in sporophytic system.
In sporophytic system, stigma is papillate and dry covered with a hydrated
layer of proteins known as pellicle. There is evidence that the pellicle is involved
in incompatibility reaction. There are striking differences in the stigma antigens
related to the S allele composition. Within few minutes of reaching the stigmatic
surface, the pollen releases an exine exudates which is either protein or
glycoprotein in nature. This exudate induces immediate callose formation in
papilae (which are in direct contact with the pollen) of incompatible stigma.
Often callose is also deposited on the young protruding pollen tubes preventing
any further germination of the pollen. Thus, in the sporophytic system, stigma
is the site of incompatibility reaction.The incompatibility reaction of pollen is
probably due to the deposition of some compounds from anther tapetum on to
the pollen exine.
1.8.2.2 Pollen tube style interaction
In most of the gametophytic system, pollen grains germinate and pollen tubes
penetrate the stigmatic surface. But in the incompatible combinations, the
growth of pollen tube is retarded within the stigma.
1.8.2.3 Pollen tube ovule interaction
In some cases, pollen tube reaches the ovule and affect fertilization. However,
in incompatible combinations, embryo degenerate at early stage of development.
1.8.3 Methods of overcoming self-incompatibility
One of the following methods can be used for bringing partial fertility by
temporarily suppressing the incompatibility reaction.
Bud pollination - Application of mature pollens to immature non-re-
ceptive stigma i.e. 1-2 days prior to anthesis.

14Fruit Breeding
Surgical technique - Removal of stigmatic surface.
High temperatue - Exposure of pistils to temperature up to 60
0
C.
Irradiation - With x rays or g rays for single locus gametophytic incom-
patibility.
Double pollination - Incompatible pollen is applied as mixture with a
compatible pollen.
Pollination at the end of season.
Arora and Singh (1988) observed that in low chilling plum and peach
cultivars, methanol killed the mentor pallen and not helpful in overcom-
ing incompatibility barriers, however, frozen and thawed mentor pollen
(one which, if alive would be fully compatible with style receiving it)
improved fruit set in both intra and interspecific incompatibility.
In case of sporophytic incompatibility system the break down is comparatively
easy because the incompatibility reaction takes place between stigmatic surface
and pollen wall in comparison to gametophytic incompatibility in which reaction
starts when the pollen tubes has already travelled 1/3 to ½ length of styler
tissue (Arora, 1993).
1.8.4 Advantages of self-incompatibility
1. Where male sterility is nonexistent, self-incompatibility can alternatively
facilitate the production of F
1
hybrids.
2. Self fertility can be induced temporarily or permanently by mutation of
S allels to S
f
through artificial irradiation in clonally propagated orchard
species like cherry and apple (Lewis, 1956).
3. Seedless varieties, such as in pineapple, grape etc. can be evolved if
self-incompatibility is present.
1.8.5 Disadvantages
Variations in seed set due to poor fertility.
Poor preservation of genetic purity of improved varieties since cross
pollination is non-restricted.
Difficulties in development and maintenance of homozygous lines (inbreds)
which can be utilized for hybridization.
Uneven quality of fruits because of mixed planting of different varieties
based on their cross compatibility.

Basic Approaches in Fruit Breeding15
1.9 Male sterility
Male sterility is characterized by non-functional pollen grains, while female
gametes are functional. Male sterility can be classified into three groups genetic
male sterility, cytoplasmic male sterility, and cytoplasmic genetic male sterility.
1.9.1 Genetic male sterility
Like any other morphological traits, particularly mono and oligogenic, this type
of male sterility occurs in plant due to mutation of the fertility locus, situated on
a chromosomes within the nucleus. In this case cytoplasm is not involved in
bringing the sterility. There could be three possible genotypes for this locus and
only one of them is male sterile.
Fertile (R-line) = RR
Fertilie (B-line) = Rr
Sterile (A-line) = rr
1.9.1.1 Sterility maintenance
By crossing AxB lines sterile and fertile progenies are produced in equal
proportions.
rr x Rr
 Rr rr
(A-Line) (B-Line) 1 : 1
O O
Fertile Sterile
For the maintenance of sterile line the fertile plants need to be quickly removed
before the shedding of the pollen grains. The fertile plants can be removed in
early stage of plant growth by using marker gene.

16Fruit Breeding
1.9.1.2 Fertility restoration
Fertile line can be obtained by crossing A-line with R-line.
e.g.
(A-line) (R-line) (all O fertile progenies)
Uses: It can be used in hybrid seed production and genetical studies or for the
preservation of variability.
1.9.2 Cytoplasmic male sterility
It occurs due to the mutation of mitochondria or to some other cytoplasmic
factors outside the nucleus, resulting in the transformation of the fertile
cytoplasm in to a sterile one. Nuclear genes are not involved. Further, with two
types of cytoplasm i.e. sterile and fertile. At the most only two kinds of genotypes
are possible, one of them is sterile and another fertile. The fertile cytoplasm is
denoted by (F – B Line) and sterile cytoplasm is denoted by (f - A-Line)
1.9.2.1 Sterility maintenance
Due to two different types of genotype cytoplasmic sterility can be maintained
as under:
1.9.2.2 Fertility restoration
Since there is no third type of genotype which can act as R-line, as such
restoration of fertility is not feasible. However, this does not exhaust all the
possibilities of the use of cytoplasmic sterile lines.
Sterile
(A-line)
Fertile
(B-line)
(all O sterile progenies because they
inherited sterile cytoplasm from
sterile seed parents.)

Basic Approaches in Fruit Breeding17
1.9.2.3 Uses
As restoration is not possible, this type of sterility is useful only in crops where
the seed is not desired end product. This is important for horticultural crops
where vegetative parts are of economic value.
1.9.3 Cytoplasmic-genetic male sterility
Such sterility arises from the interaction of nuclear gene (s) and conditioning
sterility with sterile cytoplasm. The cytoplasmic genetic sterility is essentially a
cytoplasmic sterility with a provision for restoration of fertility. The fertility is
restored by (R) gene present in the nucleus. The combination of both nuclear
gene (s) and cytoplasmic factors determine the fertility or sterility in such
plants. Based on these combinations, there can be maximum of six types of
genotype and only one of them is sterile.
(i)RR genes with F cytoplasm RR F(Fertile R-line)
(ii) RR genes with f cytoplasm RR f(Fertile R-line)
(iii) Rr genes with F cytoplasm Rr F(Fertile)
(iv) Rr genes with f cytoplasm Rr f(Fertile)
(v) rr genes with F cytoplasm rr F(Fertile B-line)
(vi) rr genes with f cytoplasm rr f(Sterile A-line)
1.9.3.1 Sterility maintenance
As visualized by their genetic composition and cytoplasm, only [(rr) f] genotype
can maintain the sterility of A-line.

18Fruit Breeding
1.9.3.2 Fertility restoration
This is achieved by suitable restorer lines which can give rise to all fertile
progenies on crossing with A-line. Among the possible six genotypes, only
[(RR) F] and [(RR)f] are such restorer or R-line. They produced all fertile
progenies.
1.9.3.3 Uses
Cytoplasmic-genetic male sterile lines are of immense importance in exploitation
of hybrid vigour in crops where seed is desired end product.
1.10 Centres of origin (primary and secondary)
The concept of centres of origin was given by N. I. Vavilov based on his studies
of a vast collection of plants at the institute of plant industry, Laningrad. He
was the director of this institute from 1916 to 1936. According to Vavilov, crop
plants evolved from wild species in the areas showing great diversity and termed
them as primary centres of origin. But in some areas, certain crop species
show considerable diversity of forms although they did not originate there,
such areas are known as secondary centre of origin of these species. Eight
main centres of origin are recognized as proposed by Vavilov.
1.10.1 China centre of origin
This is one of the largest and oldest centres of origin. It includes mountaineous
parts of Central and Western China besides, neighbouring lowlands.

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a city of importance since the days of Hadrian, reached its greatest
splendour and glory under the Ottoman sultans. Here were planned, and
from here started, the expeditions westward and eastward, which increased
in strength, in efficiency, and in inspiring terror as the circle gradually
widened, until the star and crescent appeared under the walls of Vienna and
Cairo, on the shores of Italy and in the heart of Persia. No student of
Ottoman annals can fail to support the contention of the Sublime Porte after
the last Balkan war, that Adrianople is to the Osmanlis their sacred city.
From Lalashahin to Shukri pasha, the proudest and most precious memories
of the Osmanlis are in Adrianople, whose great mosque, still awe-inspiring
and altogether admirable in its decay, is typical both of what has been and
what is.
The decision of Murad was accepted by his successors. Even after the
capture of Constantinople, many an Ottoman sultan felt more at home in
Adrianople than in the imperial city. For more than a century the Osmanlis
directed their energies almost exclusively to European conquests. Whatever
they accomplished in Asia was the indirect result of their stupendous
successes in Europe. From first to last, the extension of Ottoman
sovereignty over the Moslems of Asia was by means of a soldiery gathered
and war-hardened in Europe, themselves Christian or of Christian ancestry,
in whose veins ran the blood of Greek and Roman, of Goth and Hun, of
Albanian and Slav.
V
In 1365, Murad received from the outside world the first
acknowledgement of his commanding position as heir apparent of the
Byzantine Empire. It was an overture from the flourishing republic of
Ragusa, on the Dalmatian coast, for a treaty guaranteeing freedom of trade
in the Ottoman dominions to the merchants of Ragusa. In return for
unrestricted commercial privileges, the republic offered to pay a large sum
annually, which the givers called a grant, but which was invariably accepted
by the recipients as tribute.
[275]
However it may have been at the beginning,

the grant soon became tribute, for after some years the existence of Ragusa
depended upon purchasing the benevolence of the Ottoman sultans. As the
helplessness of the Ragusans increased, the tribute became larger. If we
except the convention between the Genoese and Orkhan, of whose
provisions and character we know nothing, the Ragusan commercial treaty
is the first of the long series of treaties by which European cities and
nations purchased the right to trade in the Ottoman Empire and to sail the
high seas. Since in most cases the Osmanlis pledged themselves to nothing
except to refrain from robbing merchants or from preventing their trading,
the gifts exacted were nothing less than blackmail. After the sea-power of
the Osmanlis had been broken, the Barbary corsairs inherited the privileges
of this system which had been started in so small a way by the Ragusans.
Murad could not write. When the treaty with Ragusa was brought for his
signature, he put his hand in the ink and made the impression of his fingers
upon the paper. This is the origin of the tughra, which has ever since been
the official signature of the house of Osman.
[276]
VI
When Murad was settling himself in Adrianople, and laying plans for the
conquest of Macedonia and Bulgaria, he was menaced by a new crusade.
Despite its futile ending, or better, for that very reason, the expedition of
Amadeo of Savoy in 1366 commands our attention. For it furnishes, as does
the expedition of Admiral Boucicaut from Genoa in 1399, a striking
illustration of how easily the growing Ottoman power might have been
crushed by a resolute body of crusaders with a single aim, and of how
impossible it was to secure that oneness of purpose, owing to the ingrained
animosity of the East and West, of the Greek and Catholic Churches.
In 1361, when Lorenzo Celsi was elevated to the dogeship of Venice, the
Senate had made overtures to John Palaeologos for an alliance against
Murad.
[277]
This plan was frustrated by the successes of the Osmanlis in
Thrace. The Venetians held back, and allowed John to suffer the humiliation
of signing the treaty that made him a vassal of Murad. In the crusade that
ended in the disaster of the Maritza, the Venetian participation was half-
hearted, and it proved valueless. The Venetians were not even on hand to
prevent Murad from crossing the Dardanelles. In fact, there is every reason
to believe that they now began to look upon the Osmanlis as a valuable tool

in checkmating the ambition of Louis of Hungary to inherit the shortlived
empire of Stephen Dushan.
[278]
When he saw that Murad had come into Thrace to stay, and that there
was no hope from the Venetians, John Palaeologos turned to the
Hungarians. He made a secret visit to Buda to enlist the aid of Louis, and
made the usual promise that the Byzantines would return to the Roman fold.
[279]
On his return he passed through the principality of Sisman, who had
just inherited the lower portion of Bulgaria. Sisman, either at the suggestion
of Andronicus Palaeologos, who wanted to succeed his father, or in the
hope of winning favour with Murad, detained the emperor in the fortress of
Nicopolis on the Danube.
[280]
Amadeo VI of Savoy was one of the princes who had taken the cross
from Pope Urban V at Avignon on Holy Friday, 1363, for the crusade that
never materialized. The receipt of a letter from Louis of Hungary, informing
him of the imprisonment of his cousin (John’s mother was a princess of
Savoy), and pointing out the rapid spread of Ottoman power, caused
Amadeo to yield to the Pope’s continued and urgent solicitations.
[281]
With
some fifteen hundred soldiers, he embarked for the East on fifteen galleys.
After a stop at Negropont and Mitylene to get reinforcements, Amadeo
entered the Hellespont, and captured Gallipoli without difficulty. The
Osmanlis fled by night, abandoning the fortress.
[282]
But the Savoyards made no attempt to follow up this victory, or even to
keep Gallipoli. Instead of attacking the infidels, they sailed into the Black
Sea, and started a vigorous campaign against the Bulgarians. Sozopolis and
Burgas were captured, and several other important fortresses to the north.
The bravery of the crusaders was rivalled only by their cruelty. Their
bloodlust made such an impression upon the Bulgarians that they wanted
nothing to do with Franks bearing the cross. When the Savoyards laid siege
to Varna, Sisman gave up his prisoner to save the city.
John Palaeologos was borne back triumphantly to Constantinople. But
friction soon arose. When Amadeo urged upon his kinsman the necessity of
paying the price of his rescue and of the continued support of the crusaders
by fulfilling his promise to return to the Roman Church, he met with
stubborn refusal on the part of emperor and patriarch alike. In wild rage,
Amadeo withdrew to Pera, and began to fight the Greeks by sea and land.
The Constantinopolitans were so frightened that ‘they did not dare to show

their head out of doors’. Pressed on all sides by Osmanli and Bulgarian, as
well as by his deliverers, the wretched John saw no other way out than to
promise openly to abjure his errors and swear allegiance to the Pope.
Having wrung this promise from those whom he had come to defend,
Amadeo sailed away to Rome, where he reported to the Pope in full
consistory ‘how at his request the emperor of Constantinople and his people
desired to submit to the obedience and belief of the Holy Roman Church in
hope that the Church would aid them against the infidels who were too
strongly oppressing them’.
Urban and the cardinals listened without great interest to the Count of
Savoy’s recital of his success in preparing the ground for a reunion of the
churches. The story was getting to be an old one. John’s overture was
received with suspicion. Urban had got the same promise in the spring of
1366 in a letter from Louis, which reported the interview John had sought at
Buda.
[283]
To the envoy of Louis, who had arrived in Avignon just as Urban
was starting for Rome, the Pope gave a letter commanding the King of
Hungary to put off his crusade until the union of the churches was actually
accomplished.
[284]
VII
What lay behind the eagerness of Urban, at the beginning of his reign, to
revive the crusades? Was he burning with holy zeal to recover the sepulchre
of Christ from the hands of the Moslems? Was his heart set on protecting
Cyprus and Rhodes? Had he determined to leave no stone unturned to
protect the Byzantines and other eastern Christians from the encroachment
and persecution of Murad? His letters indicate that his chief interest was the
recovery of the lost power and glory of the papacy. There is the same
revelation in the letters of his immediate successor, Gregory XI. These two
popes had no catholic vision. They tried to keep their position as arbiters
between France and England and Spain at Avignon, and at the same time to
inherit the temporal power of the decaying Holy Roman Empire by
circumventing the Visconti of Milan. The great schism in the Western
Church, which so aided Murad and Bayezid in laying solidly the
foundations of an empire in Europe, was the outcome of the short-sighted
and purely selfish policy of these two popes. How far from the truth it is to
represent them as courageously, whole-heartedly, and persistently

endeavouring to awaken the interest and attention of Europe in the peril
from the East!
The fall of Adrianople and of Philippopolis should have been a warning
to Urban. He read in it, however, not a glorious opportunity to demonstrate
the solidarity of Christendom by driving the Moslems out of Europe and
rescuing fellow Christians from apostasy, slavery, and death, but an
occasion to force the schismatic Greeks to return to the Roman communion.
Of the popes of the fourteenth century, Urban had the greatest chance to
prove himself a worthy champion of Christ and civilization. For it was
during his reign that the Osmanlis began their conquests and their
proselytizing in Europe. At the beginning they could easily have been
checked. But it never occurred to Urban that there was a common interest
of Christendom higher than and outside of the Roman Church.
The fault lay not wholly with Urban and with Gregory. They reflected
the spirit of their age. But it does no credit to their personal character nor to
the high position which they held to say that they were the victims, rather
than the masters, of the prevailing bigotry and ignorance of their generation.
In the fourteenth century, the West had already begun to try to impose its
commerce, its customs, its laws, and its religion upon the East. There was
not, nor has there ever been since, a sympathetic ‘give and take’ between
Occident and Orient. In a mint, if the coin when stamped does not
correspond exactly to the mould, it is rejected. Similarly the West, when it
tries to put every eastern people through its mould and finds no exact
correspondence, rejects. Hence, on the one side, the scorn of the ‘I am
better than thou’: on the other side, a hatred born not only of fear and of
conviction of inferiority, but of a sense of injustice which is none the less
vital from a knowledge that the wrong is not, and will not be, righted.
Amadeo of Savoy, uncivilized, fanatical through ignorance, the fertile
breeding-ground of fanaticism, true and unchanged descendant of the
Fourth Crusaders, was a prophetic figure at Constantinople in 1366. He
represented the only possible type of deliverer for Byzantium. But
deliverance on his terms the Greeks would not accept. Death or Islam were
preferable. And who can blame them? Two years before Amadeo’s
expedition, the Greeks of Crete had risen in rebellion against their Venetian
overlords because an attempt had been made to impose upon them the Latin
faith and rites.
[285]
When they were hunted down and massacred for

refusing to worship after the western fashion, not only Pope Urban, but also
Petrarch, wrote to the Doge congratulating him upon his valiant and
successful efforts to save the Church of Christ in Crete!
[286]
In a letter to Pope Urban, Petrarch spoke with approval of the policy of
using the Ottoman menace to stamp out the Eastern heresy. ‘The Osmanlis
are merely enemies,’ he wrote, ‘but the schismatic Greeks are worse than
enemies. The Osmanlis hate us less, for they fear us less. The Greeks,
however, both fear and hate us with all their soul.’
[287]
These words of
Petrarch epitomize the feeling between the Eastern and Western Churches
during his own day, and, if what one can see with his own eyes in Jerusalem
and elsewhere is a fair example, up to the twentieth century.
[288]
If the European nations regarded the adherents of the Orthodox Church
(the term Greek in its religious sense must be taken to include all the
Balkan races) as ‘worse than enemies’, that is, than the Osmanlis, it is
equally true that the Osmanlis found from 1350 to 1500 that the hatred of
the Balkan races for the Latin Church was their most potent ally, not only in
the actual conquest, but in reconciling the conquered to their fate. One does
not want to detract from the genius of the early Ottoman sovereigns and
from the reputation for superb fighting ability so honestly won by the
Ottoman armies. But it must not be forgotten that each separate race in the
Balkans preferred the rule of the Osmanlis to that of their neighbours, and
that the one point in which the Balkan races were of the same mind was that
Ottoman domination was preferable to that of the Hungarians and the
Italians. For every crusade was a scheme for religious propaganda and
territorial aggrandizement, in just the same spirit as in modern times the
nations of Europe have exploited the misery of Ottoman Christians for the
purpose of securing concessions.
In spite of the fact that John Palaeologos was informed by the Patriarch
Philotheus that a mixed council of clergy and government officials,
presided over by the empress, had been held in June 1376, and had decided
against the reunion of the churches,
[289]
John persisted in his negotiations
with the Pope. Urban did all that he could to facilitate the visit of the
Byzantine emperor to Rome.
[290]
But at the same time he was writing to the
Venetians and to the Dalmatian cities to protect the Catholics of Cattaro
against the Serbian and Albanian heretics,
[291]
and was encouraging Louis
in his suicidal campaign against the Bulgarians.

In 1369, John Palaeologos left the government of Constantinople to his
elder son Andronicus, and set out for Rome, where, on October eighteenth,
he made his profession of faith in the presence of four cardinals, and
confirmed it by a golden bull. The next morning, at St. Peter’s, he formally
abjured the errors of the Orthodox Church before the high altar, with his
hands in those of the Pope.
[292]
The Pope accepted him as a ‘son of the
Church’, promised that he should be relieved of the Turk, and gave him
letters earnestly recommending his cause to the princes of Christendom.
[293]
Urban V was quick to use the prestige which he believed the adhesion of
John Palaeologos had given him. He announced broadcast the happy
consummation of his efforts, stating that the Byzantine emperor had done
homage to the Vicar of Christ in St. Peter’s.
[294]
But letters sent during the
same winter to the Greek clergy, urging them to accept the action of their
emperor,
[295]
and other letters from his secret correspondence of this year,
indicate how little faith he had in the Emperor’s sincerity or ability to fulfil
his promises. Was the abjuration in St. Peter’s a farce, in which Emperor
and Pope allowed themselves to trifle with holy things, each for the sake of
his immediate advantage?
John had hoped that his adhesion to the Roman Church would bring to
him grants of money, ships, and men from the Latin princes, and that an
army would rally around him to fight the Osmanlis. But not only did he
return from France ‘with empty hands’, but he was detained at Venice
because of debts owing to merchants. In vain he begged Andronicus to send
the money for his release. The son who had four years before been charged
with being party to his father’s imprisonment in Bulgaria was no more filial
at this humiliating crisis. He answered that there was no money in the
treasury, and that he could get nothing from the clergy. But his younger son,
Manuel, brought from Salonika the ransom.
[296]
John Palaeologos returned to his capital poorer than when he left. He
brought no help from Europe, and he had bound himself publicly by oath to
an obligation which he had known he could not fulfil. He had broken faith
with Murad, who during these years had been growing more and more
powerful. There was nothing for him to do but to make himself tributary to
Murad in order that he might enjoy ‘up to the end of his life’ his last
possessions in peace.
[297]
Three years later, in 1373, when his ambassador

John Lascaris failed in a second attempt to get aid from the Western princes,
[298]
the Byzantine emperor recognized Murad as his suzerain, promised to
do military service in person in Murad’s army, and gave to him his son
Manuel as hostage.
[299]
Urban died a few months after John’s visit to Rome. Gregory XI, who
succeeded him in December 1370, had little hope of carrying on further
negotiations with the Eastern Church; for the Greek ecclesiastics were
stubborn in their determination to maintain the absolute independence of
the patriarchate. The Genoese and Venetians were fighting bitterly in
Cyprus. In 1371, Gregory made a strong appeal to France, England, Venice,
and Flanders to co-operate with Genoa in saving the last Christians of the
Holy Land.
[300]
There was no response.
That Gregory realized clearly the peril to Christendom in the advance of
Murad’s armies is shown in two remarkable letters written to Louis of
Hungary in May and November 1372. His words were prophetic. He urged
Louis to resist the Osmanlis before they advanced farther into Europe. They
had already entered Serbia. He trembled to think what would happen if they
pushed through Albania and secured a port on the Adriatic. Unless Louis
entered without delay into an alliance with his Christian neighbours, how
could he protect his own kingdom and all Christendom from the
Mohammedan peril.
[301]
Seconding this warning to the King of Hungary,
the Pope commanded the Hungarian and Slavic archbishops to preach the
crusade in Hungary, Poland, and the Dalmatian cities. Everywhere special
boxes were placed in the churches for collecting funds. A tithe was levied
on the monasteries and abbeys of Hungary and Dalmatia. Louis, with five
of his most powerful nobles, took the cross, and swore to the Pope that he
would put an army in the field within a year.
[302]
Louis asked Venice for
triremes, but when the Venetians found that he intended them to be a
donation for ‘the common cause’, they found that they could not build
them.
[303]
Padua declined an invitation to guarantee the cost of
construction. The Hungarians did not fulfil their promises. In fact, there is
no evidence that they made any effort to acquit themselves of their oath.
When John Palaeologos made a last desperate appeal to the Pope, before
he entered into his third and final compact with Murad, Gregory, in
receiving the imperial envoy, burst into tears, and promised that he would
save Constantinople, if only the Byzantine emperor would cause his people

to renounce their heresies and return to the Roman Church. In 1375, he
wrote once more to Louis to inform him that Constantinople was in danger
of capture from Murad.
[304]
Letters in the same year to Edward of England
pictured the Ottoman advance and the peril of Christendom, urged a general
war against the Osmanlis, and asked for a subsidy to provide galleys ‘to
prevent the crossing into Europe of more Turks, because Constantinople is
in imminent danger’.
[305]
The letters of Gregory XI to the Christian princes
prove conclusively that the full import of Murad’s early successes was
understood by the Pope and was impressed upon both secular and
ecclesiastical authorities throughout Europe.
But both John and Gregory lost heart. Neither was able to fulfil the
compact made in Rome. Gregory could not unite Christendom to relieve the
Byzantines. John could not persuade the Byzantines to renounce, as he had
done, the ‘Greek heresies’. So, as we have seen, he became Murad’s vassal.
[306]
The Pope, involved in the quarrel of Emperor Charles IV and the Duke
of Bavaria with the Marquis of Brandenburg, and anxious over the
outcome, for the papacy, of the continual unrest in the Italian cities,
returned from Avignon to Rome in 1378. He died a few months later.
[307]
The struggle arising from the election of Gregory’s successor gave birth to
the ‘Great Schism’. This left Murad a free hand in subjugating the Balkan
peninsula.
VIII
The sources of information for the movements from the outside for the
relief of the Balkan Christians, and for the religious and political quarrels of
the Byzantines, are so numerous and so detailed that one is embarrassed by
too much material. Many interesting facts cannot even be mentioned. But
when we come to the beginning of the Ottoman conquest in Europe under
Murad and Bayezid, we find ourselves in the midst of what an eminent
Slavic historian has called ‘the most obscure and difficult period of South-
Slavic history’.
[308]
The chroniclers, whether they be Slavic, Rumanian, or
Ottoman, are so contradictory and so lacking in explicit statement that we
cannot speak with certainty of the sequence of events. The Byzantine
chroniclers, verbose to the point of weariness in detailing petty and trifling
quarrels and happenings, are almost silent concerning the momentous
events that marked the ruin of their empire. It is difficult to unravel the

twisted skeins, and find a thread to carry the story of the conquest from
1366 to 1389. When it is impossible to choose between contradictory
records, the geography of the field of action, with which one can gain a
first-hand knowledge, must be the final factor in determining the sequence
of conquest between the adoption of Adrianople by Murad as his capital and
the downfall of the Serbians at Kossova.
The occupation of Adrianople and Philippopolis was as severe a blow to
the Bulgarians as to the Byzantines. In spite of the fact, however, that Greek
and Bulgarian had a common interest in driving the Osmanli from Thrace,
or at the very least in checking his advance, there was no move made at this
time for an alliance. On the contrary, even when the Osmanlis were engaged
in the Thracian campaign, war arose between John V and Alexander. The
Byzantines captured Anchiale, and tried desperately to take Mesembria by
assault.
[309]
The Greek patriarch wrote to Czar Alexander, reminding him
of the sacredness of harmony and the necessity of accord at that critical
moment, but the letter was not backed by the good faith and good will of
the Byzantine emperor. Neither John nor Alexander attempted to give
assistance to the Serbian and Hungarian crusade that ended so disastrously
on the banks of the Maritza.
The conquest of Bulgaria up to the main Balkan range imposed itself
upon Murad as a corollary to the Ottoman dominion in Thrace, and the
undisturbed possession of Adrianople and Philippopolis. For the Bulgarians,
through centuries of varying fortunes, had grown accustomed to fighting for
the right to live in Thrace. Often had they been beaten back to the Balkans,
and as often pressed forward again to the Ergene. To win and lose
Adrianople and other Thracian cities was old history with them. They
always came back. Between 1362 and 1365, Murad had experience with
Bulgarian persistence and tenacity of purpose. They were masters again of
Kirk Kilisse, Midia, Bunar Hissar, and Viza when Murad made his change
of capital from Brusa to Adrianople. Yamboli had been strongly fortified by
Alexander. Bulgaria seemed as formidable and as forbidding to Murad’s
dream of empire as the emirates of Asia Minor.
Fortune again favoured the Osmanlis. Czar Alexander died in 1365,
[310]
leaving three heirs. To John Sisman fell middle and southern Bulgaria from
the Danube to the Rhodope Mountains and the Bulgarian pretensions in
Thrace. Old Tirnovo was his capital. Stracimir inherited western Bulgaria,

with Widin for capital, and the Bulgarian pretensions to the valley of the
Vardar and western Macedonia. (The Bulgarian remnant of eastern
Macedonia was in the hands of an independent Bulgarian prince,
Constantine, whose stronghold was Kustendil.) Dobrotich became master of
the Dobrudja and the upper Black Sea coast, where Bulgarian, Cuman and
Alan lived together with hardly any distinguishing characteristics.
The division of Bulgaria, at the moment when union was essential,
proved fatal. The sons of Alexander never joined to face the common
danger. So marked was the division of Alexander’s kingdom that thirty
years after the conquest the conquered territories were known as ‘the three
Bulgarias’.
[311]
Stracimir, jealous because Sisman seemed to have received the lion’s
share of Alexander’s inheritance, did not hesitate to make overtures to
Murad, offering to co-operate with the Osmanlis against his brother and to
share the portion of Sisman with them.
[312]
Before any agreement could be
made, however, Stracimir found himself face to face with a terrible danger
in the west, which soon caused him to forget both Sisman and Murad. Louis
of Hungary had interpreted his crusader’s commission as an authorization to
‘make war against the heretics’. It was a pretext to get possession of Widin,
which was essential to his ambitious project of adding Serbia to his
kingdom. He attacked the Bulgarians on the ground that they were enemies
of the Church and must be forced to acknowledge the supremacy of Rome.
Widin was captured and Stracimir imprisoned.
[313]
Stracimir’s dominions
were flooded with Franciscan missionaries, who were backed by a brutal
soldiery in their proselytizing efforts.
[314]
Two hundred thousand
Bulgarians abjured the orthodox heresy, and were re-baptized in the Latin
rite. This forcible conversion, which was purely a political matter, was as
objectionable to the Bulgarians as to the Cretans. They hated ‘with a perfect
hatred’ the Franciscans whom Pope Urban had sent, and the cause for which
they stood.
At the first opportunity, the Bulgarians of the west called in Sisman and
Vadislav of Wallachia. The Hungarians were driven out of Widin and the
Franciscans in the city massacred.
[315]
Louis was powerful enough to
wreak terrible vengeance. In 1370, Widin fell once more into his hands. The
Bulgarians of the western Balkans were subjected to such a relentless
persecution that they welcomed the Moslem conquest to secure freedom of

worship. Urban had incited Louis to this war, and had congratulated him
upon his laudable zeal in converting the heretics.
[316]
We have already spoken of the punishment that came to Sisman as a
result of the detention of John Palaeologos. The Italian crusaders on the
Black Sea coast were as powerful an aid to Murad’s empire-building as
were the Hungarian crusaders on the western frontier. The successors of
Louis reaped the bitter fruits of his insane policy. Louis and Amadeo of
Savoy contributed in no little measure to make possible the conquests of
Murad. When Amadeo withdrew from Bulgaria, he left the cities he had
captured to the Greeks. Sisman was compelled to expend his energy in
recapturing them. But Murad had already anticipated him in the important
fortress of Sozopolis, which commanded the entrance to the port of Burgas.
[317]
Shortly after the Ottoman occupation of Sozopolis, the Bulgarians were
everywhere dispossessed in Thrace, and the capture of Yamboli
[318]
forced
Sisman to follow the example of John Palaeologos. He became a vassal of
Murad. His sister Mara entered Murad’s harem, but with the stipulation that
she be allowed to retain her Christian faith.
Murad gladly gave his new ally and brother-in-law a strong Ottoman
army to co-operate in the attack upon the Hungarians. The Osmanlis helped
in driving Louis out of Widin. Sisman, like Cantacuzenos, first guided the
Osmanlis through the heart of his country. It was under the leadership of
Sisman that they saw the Danube, their river of destiny. When Sisman, even
with the help of the Osmanlis and Wallachians, could not gain possession of
Stracimir’s inheritance, he returned to Tirnovo. There he learned that
Lalashahin was planning an expedition westward, which seemed to be
intended against Sofia.
Sisman now realized that his position was critical and that the fate of
Bulgaria was at stake. In the early spring of 1371, he hurried into the Rilo
Mountains and sent out an appeal to the Serbian kral who was at that time
ruling in eastern Macedonia. Then he went to the relief of Ishtiman, which
was already menaced by the Osmanlis. Failing in this effort, Sisman fell
back to Samakov, where he was joined by the Serbians. Lalashahin led his
army from Ishtiman into the valley of the Isker. The two krals joined battle
with him in the plain of Samakov. The Ottoman victory was decisive.
[319]
The Serbians and Bulgarians fled into the recesses of Musalla, the highest

mountain in the Balkan peninsula, and of Popova Shapka. Sisman
disappeared after the battle.
[320]
The way to Sofia was open. All Bulgaria
lay at the feet of the conqueror. It is from the battle of Samakov that we
must date the destruction of an independent Bulgaria.
But Murad was not yet ready to follow up this decisive victory. The only
immediate result of the battle of Samakov was the submission of
Constantine, Bulgarian prince of Kustendil, in the upper valley of the
Struma. After the fall of Samakov, his position was untenable. Constantine
hurried to Murad’s camp, and did homage to the conqueror. Murad gave
back to him as vassal his principality.
[321]
With the wisdom that marked
every successive step of his progress in Europe, Murad refrained from
advancing beyond Samakov. He ordered Lalashahin to lead the army into
Macedonia, and to join Evrenos in the advance towards the Vardar.
IX
The dramatic death of Stephen Dushan, in 1355, just as he was starting
upon the expedition against Constantinople for which his whole life had
been a preparation, is recorded in the previous chapter. Stephen’s son was
so unfit to inherit the aspirations and carry on the work of his father that he
was called in derision by his people Nejaki, the weakling.
[322]
The nobles
and generals of Stephen Nejaki ignored him. Each man seized what
territory he could hold and defend against his neighbour. There was anarchy
in Macedonia and Serbia. The dissolution of Stephen Dushan’s conquests
resulted in a bloody and destructive civil war between cities and factions.
[323]
The dowager Czarina managed to preserve a semblance of prestige, if
not of authority, at Serres. But the ‘empire’ was no more. As local rulers,
Serbians stayed in the principal cities of Macedonia. There was
undoubtedly a Serbian element in the village population. Many villagers,
however, who acknowledged the overlordship of Stephen’s warriors and
other Serbian nobles, did not know then, any more than they know now, to
what race they themselves belonged. This has always been the Macedonian
problem.
The defeat of the crusaders on the banks of the Maritza in 1363 had been
a defensive battle on the part of the Osmanlis. There was no attempt to
invade Macedonia. While Murad was occupied in the subjugation of Thrace
and of southern Bulgaria, several efforts were made by the Byzantines to

come to an understanding with the Serbians. In 1364, the patriarch Callixtus
went to Serres to see Stephen’s widow, who had retired to a convent. His
purpose was to form an alliance. Soon after reaching Serres, Callixtus
succumbed to the hardships of the journey.
[324]
His effort came to nothing.
That Stephen’s son still held to the pretensions of his father and had no
intention of treating with the Byzantines, is demonstrated by a bull, dated
from Pristina in 1365, in which he calls himself ‘emperor of the Servians
and of the Greeks’.
[325]
Stephen Urosh, the ‘weakling’, died in 1367.
[326]
Uglesa, who usurped
the kralship of Serres and shared the ‘empire’ of Stephen Dushan with his
brothers and fellow adventurers, Vukasin and Goiko,
[327]
sent an embassy
to the patriarch Philotheos declaring that he would annul the bull of 1352,
by which Dushan had created an autocephalous Serbian Church,
[328]
and
would cause all the Serbians to return to the Orthodox allegiance.
[329]
After
three years of negotiation, precious time wasted with trifling formalities, the
reconciliation and union of the Serbian and Greek Churches was effected.
[330]
But, if we are to believe the authorities of Orbini, Uglesa, while he
was negotiating with the Greeks of Constantinople, had levied tribute upon
the Greeks of Salonika, and would have made himself master of Salonika,
had not his untimely death prevented the consummation of the great Serbian
dream.
[331]
At the time of the reconciliation with the Orthodox Church, Uglesa had
completed a plan of united action with his two brothers to oppose the
Ottoman invasion of Macedonia.
[332]
Uglesa had been informed that a great
army was gathered in Adrianople, which awaited the return of Murad from
Bulgaria to commence its march. Four weeks after the negotiations with the
Byzantines had been successfully concluded, in the early summer of 1371,
the Serbian army reached the Maritza at Cernomen,
[333]
between
Adrianople and Svilen.
[334]
This battle has been confused with the earlier
battle of 1363, and it is impossible to separate the accounts of the two
actions.
[335]
The Osmanlis were again victorious. Uglesa and Goiko were
drowned in the Maritza. Vukasin escaped from the field of battle only to be
killed by his servant for the gold chain he wore around his neck.
[336]
The battle of Cernomen lost Macedonia to the Serbians. The three
princes were killed. Most of the Serbian adventurers who had been the

companions of Stephen Dushan, and who had profited by his Macedonian
conquests, disappeared. The Osmanlis had no opposition in penetrating to
the valley of the Vardar.
The monk Isaias of Serres has left a graphic contemporary picture of the
Ottoman invasion of Macedonia. ‘Like the birds of Heaven, the Ishmaelites
spread themselves over the land, and never ceased murdering the
inhabitants or carrying them off into slavery. The country was empty of
men, of cattle, and of the fruits of the fields. There was no prince or leader:
there was no redeemer or saviour among the people. All faded away before
the fear of the Ishmaelites, and even the brave hearts of heroic men were
transformed into weak hearts of women. Rightly were the dead envied by
the living.’
[337]
The invasion of Macedonia in 1371-2 was as rapid and decisive a
campaign as the invasion of Thrace had been ten years before. Kavalla,
Drama and Serres were occupied by Khaïreddin and Evrenos.
[338]
Drama
and Serres were colonized, their churches converted into mosques, and they
soon became the residence of the owners of the timarets granted in eastern
Macedonia. These two cities have always been the strongholds of the
Mohammedan element in Macedonia, and the residence of the great
Moslem landowners. The cities and villages in the valleys of the Mesta and
the Struma acknowledged Murad as sovereign, and submitted without
resistance to Ottoman laws and Ottoman taxation.
[339]
Where-ever it was
safe to do so, Murad seized the lands, and appointed Ottoman governors. In
districts where pacification would have proved a difficult task, he allowed
Serbian chiefs to rule as his vassals.
With the same impetuosity that had carried them to the foothills of the
Rhodope Mountains after the capture of Adrianople, the Osmanlis crossed
the Vardar in 1372, and pushed their arms into Old Serbia, Albania, Bosnia,
and even to the mountains of Dalmatia, from which they could see the
Adriatic.
[340]
Other adventurous bands, eager to attract the attention, the
commendation, and the rewards of Murad, followed the footsteps of the
Catalans, traversed Thessaly, and appeared in the plains of Attica.
[341]
Murad destroyed the Macedonian empire of Stephen Dushan without
great effort. The Serbians remaining east of the Vardar, nobles and peasants,
became Ottoman subjects. In upper Serbia, they rallied round one of their
number, Lazar Gresljanovitch, whom they formally elected as successor of

the Serbian kings. But Lazar was so weak that he did not take the title of
emperor (tzar) or of king (kral), but called himself merely prince (knez).
[342]
To secure the existence of his kingdom or principality, he sought peace
with Murad, and, following the example of the Byzantine and Bulgarian
rulers, became vassal and tributary of the Ottoman emir.
[343]
X
Before the end of the year 1372, it was recognized that the Osmanlis had
come into the Balkans to stay. The conquest of Macedonia east of the
Vardar, following so closely upon the subjugation of southern Bulgaria and
the completion of the Thracian conquest, gave to Murad a preponderant
position in the Balkan peninsula. The Byzantine emperor and the Bulgarian
and Serbian princes were his tributaries. Wallachia, Bosnia, Albania, Epirus,
Thessaly, Attica and the Peloponnesus were now on the confines of the
Ottoman Empire, and menaced by Ottoman invasion.
In Europe, Murad was credited with having the intention of invading
Hungary. It was reported that he had made an alliance with the Tartars of
Russia to attack Hungary. The Tartars were to cross the Carpathians by way
of Moldavia into Transylvania, while Murad was to work his way up the
valley of the Danube.
[344]
Murad may have dreamed of such a project, just
as he had thought of making a supreme effort to enter Constantinople after
his first Thracian campaign. But, if he did, he was deterred by the same
well-grounded fear of moving too fast. Ten years before he had refrained
from committing a fatal error. He would continue to make haste slowly. The
early Osmanlis were not raiders. They were empire-builders. They
succeeded because they never forgot that their greatest problem was that of
assimilation. When they extended their conquests beyond the area of
possible assimilation, the period of decay automatically commenced.
The decade following the Macedonian campaign of 1371-2 was spent in
ottomanizing southern Bulgaria and eastern Macedonia, in completing the
assimilation of Thrace, in reorganizing the army, and in a rearrangement of
the system of distributing the timarets or military fiefs. Royal domains were
created, and lands were set aside for the support of the mosques and other
religious institutions in the form of inalienable endowments (vakufs).
The only move of Murad against the Hungarians was to send five
thousand archers, upon the request of the Senate, to help the Venetians in

their war against Louis.
[345]
After the Macedonian campaign, Murad turned his attention once more
to Byzantium. John, when he returned from his unsuccessful trip to Rome,
placated Murad by sending his third son, Theodore, to serve in the Ottoman
army. In 1373, John, passing over Andronicus, raised Manuel to the
imperial purple as co-emperor. The disloyalty of his eldest son in the
question of the emperor’s ransom from his Venetian creditors made it
natural that John should have selected Manuel to rule with him.
John was not wrong in his estimate of the character of Andronicus. The
disappointed prince entered into a conspiracy with Saoudji, son of Murad,
who had been entrusted with the command of the Thracian army while his
father was occupied in Anatolia. John and Manuel, according to some
accounts, were also in the field with Murad. So the moment was propitious.
The two sons raised the standard of revolt against their fathers.
[346]
Murad,
who hated his own son and feared him, crossed immediately into Thrace.
The army which was supporting the cause of the young princes abandoned
them, and the rebels fled to shut themselves up in Demotika.
[347]
Faced with starvation, the inhabitants of Demotika opened the gates of
their city to Murad. He exacted a most atrocious vengeance. The garrison
were bound hand and foot and thrown into the river. The young Osmanlis
and Greeks who had been led astray by the princes, were put to death.
Wherever possible Murad compelled fathers to act as executioners of their
sons. He set the example by tearing out Saoudji’s eyes, and then cutting off
his head.
[348]
It has been generally written that Murad intended that the same
punishment should be meted out to Andronicus. For the sake of
appearances, he did order John Palaeologos to have his son’s eyes put out.
But there was no order for execution. John Palaeologos consented to the
blinding of Andronicus and of his grandson and namesake, who was only
five years old.
[349]
The operation was not successfully performed. Both
Andronicus and his son, even if temporarily blinded, recovered their
eyesight. Some have explained this by stating that they were healed by a
Genoese physician.
[350]
There is recorded a beautiful story that Andronicus
owed the restoration of his sight to the empress, his mother, who visited
him daily in the tower of Anemas and was prodigal in her efforts to heal

him. He was in despair for some months, until one day he saw a lizard
climbing on a wall.
[351]
If Murad had really desired the death or total blindness of Andronicus,
he could easily have secured this result. While punishing his own son,
however, he saw to it that Andronicus escaped the consequences of the
same crime. Here we have a revelation of the far-sightedness and cold-
bloodedness of Murad. He killed his own son, because he feared his rivalry.
He spared the son of John Palaeologos in order to perpetuate the rivalry
between the emperor and his son. To have killed or incapacitated
Andronicus would have been from his view-point an act of folly rather than
of justice; for Andronicus, brilliant, adventurous, magnetic, was at the same
time a worthy exemplar of the name he bore, a name that stood for the acme
of unscrupulous conduct and contempt for ties of blood. Murad had only to
wait, and history would repeat itself. Internal dissensions in the family of
the Palaeologi had made the fortunes of Orkhan. Murad had no intention of
getting rid of Andronicus, in whom he saw the means of still further
enmeshing the Byzantine emperors.
[352]
The Byzantine historians record for the year 1374 another event, which
illustrates the power of Murad over John Palaeologos. Manuel, who had
resumed the government of Salonika, tried to induce the inhabitants of
Serres to recover their liberty by massacring the Ottoman garrison and the
Ottoman colonists. Serres, in spite of its prominent place in recent Serbian
history, was regarded by the Byzantines (as it still is by the Greeks of to-
day) as a city of their compatriots. We have no means of establishing the
grounds upon which Manuel believed it possible to restore the Byzantine
authority in the country between the Struma and the Vardar. The sequel
indicates that it was a wild and unfounded hope of a desperate man, and
shows how thoroughly in two years the Osmanlis had become masters of
the situation in Macedonia.
Murad, warned in time of the project, sent Khaïreddin pasha with a large
army to Serres. The Greeks implicated in the plot were promptly executed,
and Khaïreddin moved against Salonika. At the approach of the army,
Manuel fled by sea to Constantinople. John Palaeologos was so frightened
that he did not dare to receive in the imperial city the beloved son whom he
had raised to the dignity of co-emperorship. Manuel then went to Lesbos,
whose Genoese lord was his uncle by marriage. But the fear of Murad had

reached the Aegaean Sea. The fugitive was turned away. Staking all upon
the issue, Manuel went to Brusa and threw himself at Murad’s feet. The
time was not yet ripe to destroy the Palaeologi. Murad pardoned Manuel,
and sent him back to Constantinople. It was only after Manuel had
presented a letter from Murad, confirming the fact that forgiveness had been
granted, that the emperor of Byzantium dared to receive his son and heir
within the walls of Constantinople.
[353]
Pressed by the Venetians, John made in 1375 the mistake of giving them,
in exchange for three thousand ducats and the jewels which had been
pledged for his debts after the visit to Rome, the island of Tenedos.
[354]
The
strategic importance of Tenedos was so vital that the Genoese could not
allow this island to fall into the hands of their rivals. It is an axiom as old as
history that who holds Tenedos controls the entrance and exit to the
Dardanelles. Until the Black Sea dries up and the wheat-fields of Russia fail
to yield, there will be a ‘question of the Straits’.
The news of this grant to Venice meant but one thing to the Genoese.
There was feverish activity at Genoa. A fleet was manned, ostensibly for the
purpose of maintaining the Levant colonies against the Turks.
[355]
Pope
Gregory XI allowed the archbishop of Genoa to raise enormous sums by
questionable means for equipping and increasing the fleet.
[356]
Instead of
using this fleet to free the Aegaean and the Black Sea from the ever-
increasing Turkish pirates, or to attack the Osmanlis, the Genoese admiral
sailed to Constantinople. Aided by the Genoese of Galata and by Bayezid,
Andronicus had escaped from the tower of Anemas. When the fleet arrived
from Genoa, he gave to its admiral a golden bull, awarding Tenedos to
Genoa.
[357]
To Murad he offered his sister in exchange for help.
[358]
The
old story was repeated. After a month’s siege, Andronicus, by the aid of his
Ottoman and Genoese supporters, entered Constantinople. His father and
his two brothers, Manuel and Theodore, were imprisoned in the Tower of
Anemas, where he and his son had been shut up for two years.
[359]
The
foresight of Murad in regard to Andronicus was justified.
While Andronicus was besieging Constantinople, John V managed to
send word to the inhabitants of Tenedos to resist the Genoese and give
themselves to the Venetians. If this were not possible, they were to abandon
the island to the Turks rather than allow the Genoese to occupy it.
[360]

After a year’s imprisonment, the emperor, through the wife of his jailer,
succeeded in perfecting with Venetians residing in Constantinople a plan of
escape. But its execution was deferred when John discovered that his sons,
who were confined to separate rooms, could not be included in the rescue.
Later, the efforts of the Venetians were renewed upon the solemn promise
that Tenedos should revert to Venice. The plot was discovered. The
Venetians, availing themselves of the lucky chance that a Venetian fleet had
just arrived in the Golden Horn from the Black Sea, fled from
Constantinople, abandoning John Palaeologos to his fate.
[361]
Andronicus
IV was solemnly crowned in St. Sophia sole emperor of Byzantium.
After two more years of imprisonment,
[362]
John and his sons succeeded
in escaping in June 1379. They got across the Bosphorus, and took refuge
with Bayezid, who was again watching the course of events at Scutari.
Murad, still playing the game of pitting father against son, drove a hard
bargain. Andronicus must be pardoned once more, and given the
government of several cities, probably including Salonika.
[363]
John and
Manuel, as a price for freedom and restoration to the imperial throne,
agreed to pay an annual tribute of thirty thousand pieces of gold, furnish a
contingent of twelve thousand soldiers to the Ottoman army, and surrender
to the Osmanlis Philadelphia, the last Byzantine possession in Asia.
[364]
When the Philadelphians refused to assent to this shameful transaction,
John and Manuel joined the Ottoman army and fought against their last
Christian subjects in Asia to force upon them the Moslem yoke.
[365]
Thus did Murad hold to the lips of John Palaeologos the cup of
humiliation, nay, more, of degradation, until he drained the last bitter dregs.
We do not need to pass judgement upon John and Manuel. It is sufficient to
say that they drank and did not die!
The question of Tenedos brought Venice and Genoa into their most bitter
conflict of the century. The Visconti of Milan were allied to the Venetians,
while the Hungarians attacked them by land.
[366]
After initial successes, the
great Venetian admiral Pisani was beaten decisively in 1379. The Genoese
captured Chioggia, and held Venice at bay in her own lagoons. It was the
timely arrival of Charles Zeno and the fleet from the Levant that saved the
Adriatic republic.
[367]
In 1381, peace was made through the intermediary of
Count Amadeo of Savoy, on condition that the Senate surrendered Tenedos
to Amadeo, who guaranteed to demolish the fortress within two years. It

was also a stipulation of the treaty of Turin that Andronicus IV be
recognized as heir to John V.
[368]
Did the influence of Murad reach as far as
the peace negotiations in the capital of far-off Savoy? The Count of Savoy
fulfilled his promise. In 1383, the fortifications of Tenedos were rased, and
the inhabitants of the island removed to Crete and Negropont.
[369]
The war over Tenedos had kept open the Straits, but it helped Murad in
an inestimable degree to tighten the grip of the Osmanlis upon Thrace and
Macedonia. The Italian republics thought no more of driving the Osmanlis
out of Europe. From now on until they themselves see their possessions
wrested from them and their commerce in the Levant ruined by the
successors of Murad, the Venetians and Genoese are suitors for favours at
the door of the tent of the Moslem conqueror.
XI
While the struggle between the Palaeologi and the Venetian war with
Genoa and Hungary were strengthening Murad’s position in Europe, he
began to turn his attention, for the first time since the expedition against
Angora at the beginning of his reign, to the expansion of Ottoman authority
in Asia Minor. The antipathy of the South Slavs for the Hungarians, the
anarchy among the Serbians, the lack of leadership among the Bulgarians,
and the civil strife in the Byzantine imperial family made the period from
1376 to 1381 peculiarly appropriate for initiating a movement against the
emirates on the confines of his own state. Murad felt for the moment secure
in Macedonia and Thrace. The inhabitants of the conquered countries could
do nothing. There were no prospects of a crusade. Through the rapid
increase of the Ottoman race during the first fifteen years in Europe, and
through the vassalage of the Christian princes, which compelled them to
furnish contingents for war, Murad now had money and soldiers to confront
his nearer Anatolian rivals.
In 1360, after the capture of Angora and the defeat of the Galatian
village chiefs,
[370]
Murad did not lose his head. He was wise enough to fear
an attack on Kermian. Now he had only to threaten, thanks to the prestige
and actual power he had gained in Europe. The emir of Kermian was too
prudent to risk a war with the son of the rival whom he had despised. In
order to preserve his independence and at the same time his pride, he agreed
to give his daughter in marriage to Bayezid. The territories which Murad
coveted, and was ready to try to take by force, went with her as her

marriage portion. It was a munificent dot. The western and northern part of
Kermian became Ottoman. The most important city in the new territory was
Kutayia, the ancient Cotýaeum, a strategic point of great value. Its
remarkable citadel of countless towers is still standing.
The marriage of the emir of Kermian’s daughter to Bayezid was
celebrated at Brusa with much splendour. For the first time we hear of the
Osmanlis interested in matters of court and luxury. The simple warriors,
who had known nothing but the village council and the camp fire, were
becoming accustomed to the more formal and more complex life of the
Greek cities. With every victory and every extension of sovereignty, with
every addition to the army and to the body of civilian officials, the distance
between the sovereign and his people was widened. The ceremonial
evolved by the Ottoman court was that of Byzantium; the customs of the
higher classes, who were just beginning to realize their self-made rank,
were Byzantine, even to the veiling of women.
[371]
The Osmanlis had not
yet come into touch with the Arabs or Egyptians. If they received anything
from the Persians, it was by way of Constantinople.
The Ottoman occupation of Kutayia was a grave blow to the security of
the emirates of Tekke and Hamid. The emir of Hamid saw the hopelessness
of a struggle. He compounded with his pride by ‘selling’ to Murad, in 1377,
the territory between Tekke, Kermian, and Karamania. Several cities,
including Sparta and Kara-Agatch, became Ottoman, but most important of
all, Ak Sheïr, which brought the Osmanlis to the frontier of Karamania.
The purchase of this important territory extended the Ottoman state
south to the border of Tekke. In 1378, Murad made his only conquest by
arms from a rival emir in Asia. He invaded Tekke, and annexed the districts
at the south and south-west of the lake region. But he did not cross the
mountains to the Mediterranean, so the emir of Tekke still retained Adalia,
and Alaya was undisturbed.
For three years Murad devoted his energies to the pacification and
assimilation of these slices of Kermian, Hamid and Tekke. But none of the
three principalities had been extinguished. And Sarukhan, Aïdin and
Menteshe were untouched. There was still much to be accomplished in
western Asia Minor. But Murad preferred to return to Adrianople. He would
increase his power and prestige in Europe, recruit his armies in the Balkans,
and then come once more into Anatolia.

XII
To assure to the Osmanlis their preponderant position in the Balkan
peninsula, the possession of three cities was necessary. The capture of Sofia
meant the extension of Ottoman sovereignty over Bulgaria to the Danube.
Nish was the key to Serbia. Monastir was indispensable, if the Osmanlis
intended to be more than raiders west of the Vardar.
In 1380, Murad ordered the advance to the Vardar. Istip was captured,
and colonized in the same thorough way as had been done at Drama and
Serres. A large army under Timurtash crossed the Vardar, took Monastir by
assault through the marshes, and pushed north to Prilep.
[372]
Monastir and
Prilep became frontier fortresses of the empire. The conquest of Macedonia
was now complete. These cities were excellent bases of operation against
the Albanians to the west and the Epirotes to the south-west.
During the reign of Murad, the Osmanlis did not attempt a subjugation
of Albania and Epirus. They were, however, invited into these countries by
native princes.
Thomas, despot of Janina, used Ottoman mercenaries against the
Souliotes in 1382.
[373]
Two years later, after the assassination of Thomas,
the Albanians besieged Janina with Ottoman aid.
[374]
The civil war that
arose around the widow of Thomas prepared the way for the Osmanlis to
extend their rule to the Gulf of Arta.
In 1385, Khaïreddin pasha, who had occupied Okrida, the ancient
ecclesiastical seat of the Bulgarians, a day’s journey west of Monastir, was
invited by Charles Thopia, lord of Durazzo, to aid him in his war against
Balsa, the most powerful Frankish prince of Albania. Khaïreddin was glad
of the opportunity afforded by this overture. He crossed the mountains to
Elbasan, and then turned southward to meet Balsa. The first battle of the
Osmanlis in Albania was fought in the salt-wastes of Savra, on the left bank
of the river Devol. The Osmanlis faced fighting men who were fully their
equals in courage, in resourcefulness, in strength, and in willingness to
engage in a hand-to-hand struggle to death. The issue was long in doubt,
and the victory costly. Balsa and his ally and guest, Ivanitch, son of krai
Vukasin, were killed.
[375]
The Osmanlis gained one important result from
this battle. Albanian renegades joined their army in great numbers.
[376]
From that day to this the Albanian element in the Ottoman army, especially

among its officers, has been a source of strength which cannot be over-
estimated.
It is doubtful if the Osmanlis withdrew from Albania, even temporarily,
after the battle of Savra; for in 1388 the princess of Valona (Avlona) was so
hard pressed by the Osmanlis that she put her domains under the protection
of Venice.
[377]
In northern Albania, the invaders captured Croia and Scutari in 1386.
Scutari was given back by Murad in exchange for the addition of a member
of the ruling family of Zenta to his harem. From Croia, also, the Osmanlis
withdrew. Murad did not want to excite and alarm Venice at the moment
when Philippe de Mézières was preaching so vigorously and successfully a
new crusade.
[378]
The plain in which four tributaries join the Isker is the very heart of the
Balkan peninsula, almost equidistant from the Adriatic, the Aegaean, and
the Black Sea. Here the three great ranges of the West Balkan, the Central
Balkan, and the Rhodope Mountains converge, and three important rivers
find their source. The Struma flows south through Macedonia, the Isker
north-east through a canyon of the Balkans into the Danube, and the Nisava
north-west into the Morava. In the middle of the southern border of this
plain, under the shadow of a lofty mountain, lies Sofia.
The way to Sofia had been opened by the battle of Samakov. But its
occupation was not the next logical step to Murad until the valleys of the
Vardar and the Struma had been conquered. The occupation of Sofia was a
temptation splendidly resisted in 1371. In 1381 it was a necessity. For it
opened the path to trans-Balkan Bulgaria and to Serbia, and Murad was
now ready to extend his conquest to the Danube by way of the Isker and the
Morava.
The Slavic chronicles are silent concerning the fall of Sofia. From the
late Ottoman accounts, it would seem that the city was intermittently
besieged for several years. Then a young Osmanli, who had entered the city
as refugee, and had become the confidant and falconer of its commandant,
betrayed him. He urged his master in a chase some distance in front of his
followers, and fell upon him in a mountain gorge. The commandant was
bound to his horse, and taken a prisoner to Ishtiman. Indje Balaban, son of
the general of Osman who had besieged Brusa for ten years, brought his
army from Philippopolis, and paraded the commandant, garrotted, under the

walls of Sofia. The Bulgarians, discouraged and despairing of aid,
surrendered.
[379]
We can be certain neither of the name of the Bulgarian
commandant nor of the date of the surrender. But it was probably in 1385.
[380]
Bulgaria up to the main Balkan range was now Ottoman territory.
The fall of Nish, in the summer of 1386, marked the next extension of
Murad’s empire.
[381]
The Serbians did not yield without a struggle, as the
Bulgarians had done. Nish was taken by assault. Lazar secured peace only
by increasing the amount of his tribute and adding one thousand cavaliers to
his contingent in the Ottoman army.
[382]
Nish was sixteen days by carriage from Constantinople. Murad was now
master of four-fifths of the great Roman highway from Belgrade to the
Bosphorus; for Tchorlu, Demotika, Adrianople, Philippopolis, Ishtiman,
Sofia, and Nish were in his hands. Nish was also the point where the road
from Belgrade to Salonika turned southward. Practically all but the last
day’s journey of the road across the Balkan peninsula from Constantinople
to Durazzo on the Adriatic was Ottoman territory. In Asia Minor, Murad
held the ancient highway from Constantinople to Trebizond as far as
Angora, and the road which the pilgrims and Crusaders, Jerusalem-bent,
had travelled as far as Ak Sheïr. From Angora to Nish took twenty-five
days; from Constantinople to Durazzo seventeen days.
[383]
Twenty-five
years before, when Murad came to the chieftainship of the Osmanlis, the
Ottoman dominions could have been traversed in any direction in three
days.
XIII
The treaty concluded between the Byzantines and Genoese in 1386
affords a striking illustration of Murad’s power after the Nish campaign.
This treaty, whose text has been preserved, was signed by John and
Andronicus Palaeologos, the podesta of Pera, and the Genoese ambassador.
John Palaeologos bound himself to live in peace with his son Andronicus,
and to move his army against all the enemies of Genoa ‘except Morat bey
and his Turks’. The Genoese in turn promised to defend Constantinople
‘against all enemies of whatever nationality except the said Morat bey and
his Turks, who acted according to the will of the said Morat bey’!
Throughout the treaty, Murad is carefully excepted on both sides.
[384]

Genoa made a formal treaty with Murad in 1385. Favours were granted
to the Osmanlis who did business in Pera, in return for liberty to Genoese
merchants to reside and conduct business in the states of Murad. The treaty
recalls the friendship of the Genoese for Orkhan, and speaks of Murad as
‘the magnificent and powerful lord of lords, Moratibei, grand admiral
[385]
and lord of the admirals of Turchie’.
[386]
But in the very next year Genoa
secretly joined an offensive league with Cyprus, Scio (Chios) and Mytilene
‘against that Turk, son of unrighteousness and evil, and also of the Holy
Cross Morat bey, and his sect, who are attempting so grievously to attack
the Christian race’.
[387]
In the first year of Murad’s reign, the Venetian energy had become so
sapped by prosperity and luxury that the Senate passed a sumptuary law.
[388]
The recent triumph over Genoa had given them a belief in their
invincibility. Their self-sufficiency, and the growing disinclination to lay
aside the pen and ledger for the sword and shield, were alarming symptoms
of decay. The lesson of the Genoese at Chioggia was needed to teach the
Venetians that the struggle for existence never ceases.
In spite of their vital interest in the development of the Levant, and the
power that their wealth gave them in a generation when fighting strength
could be purchased so easily, Venice made no effort to oppose the progress
of Ottoman conquest. On the contrary, in 1368, long before an invasion of
Albania was imminent, the Senate negotiated with the Osmanlis for the
reddition of Scutari. This project was again taken up in 1384, in a tentative
way, during negotiations to fix the customs-duties of Venetian merchant-
vessels.
[389]
Following the example of Ragusa and Genoa, Venice
concluded, in 1388, a commercial treaty with Murad.
[390]
The traffic of the Italian republics with the Moslems had been
denounced by Gregory X in 1272, by Boniface VIII in 1299, by Urban V in
1366, and by Gregory XI in 1372.
[391]
In vain the popes exhorted; in vain
they threatened interdict and excommunication; in vain they held up to
execration the abominable slave traffic. Trade interests alone decided the
policies of the maritime cities. Their citizens never hesitated to cut each
other’s throats for the opportunity of selling goods. To them the crusades
were a purely commercial proposition. More than once the archives of
Venice reveal the approval of the Senate upon the action of merchants who
warned Moslem princes of the crusaders’ intentions. Guillaume d’Adam

declared with reason that the Saracens maintained their supremacy in the
Holy Land and Egypt through the support of the traders, who furnished
them with Christian slaves to keep up their armies.
[392]
Genoa passed laws
in 1315 and in 1340 against the slave traffic of the Black Sea,
[393]
but these
laws were never enforced.
[394]
Venice and Genoa turned a deaf ear to papal remonstrances and to papal
appeals for aid in crusades against the Osmanlis. For the sake of preserving
their commerce, they flattered Murad, and aided him, indirectly at least, to
subjugate the Christians of the Levant. Their children of the third and fourth
generation paid to the descendants of Murad the penalty of their greed.
They lost their commerce in trying to save it.
XIV
It was not until 1387 that Murad believed himself strong enough to
measure arms with Karamania. His son-in-law, Alaeddin, whose name is
reminiscent of the earlier glory of Konia, was emir of the most powerful
state in Anatolia. The Ottoman historians have represented Alaeddin’s
resistance of the encroachment of the Osmanlis, and his defiance of Murad,
as rebellion, and have been blindly followed in this by most of the
European historians. Such a conception of the conflict between the
Osmanlis and the Karamanlis is far from the truth. There is no record of
when and how Karamania had become subject to Murad. In fact, up to
1387, Murad had not yet extended his sovereignty over all of Tekke and
Hamid, the states which bordered Karamania on the west.
Neither Alaeddin himself nor his predecessors had ever acknowledged
the suzerainty of the house of Osman. From the standpoint of the
Karamanians, the Ottoman emir was not even primus inter pares of the
Turkish princes in Anatolia. Osman had probably not been known by name
to the founder of the house of Karaman. Orkhan never came into direct
contact with the Karamanlis. Murad, at the beginning of his reign, had
indirectly gained an advantage over the emir of Karaman in the successful
issue of his expedition against the Phrygian chiefs and the capture of
Angora. Fifteen years later his accessions of territory in Kermian, Hamid,
and Tekke brought him into rivalry with Alaeddin. But it was the prestige
and power gained by Murad in European conquests that made him a rival to
be reckoned with. The first acknowledgement of his growing strength was
the marriage alliance between the houses of these two emirs. Alaeddin,

however, did not by this marriage constitute himself a vassal of his father-
in-law. The letters of Murad to Alaeddin in the collection of Feridun are
couched in terms of equality.
Murad rallied his army at Kutayia for the first great Ottoman campaign
in Asia. He could not muster enough Osmanlis to undertake so formidable a
feat as the invasion of Karamania, and had to rely upon large contingents of
Greeks and Serbians, who were sent to him, in accordance with their
conventions, by his vassals, the emperor John and the kral Lazar.
[395]
The
Balkan soldiers, under the command of Bayezid, formed the left wing of the
Ottoman army.
Battle was joined in the great plain before Konia, which has so often
been the scene of Ottoman triumphs and reverses. The Ottoman historians
declare that Alaeddin was defeated, largely through the bravery of
Timurtash, and represent the battle of Konia as a decisive victory, which
‘put down the rebellion’. According to them, Alaeddin ‘sued for peace’.
Murad ‘forgave’ him, because he was moved by the tearful pleadings of his
daughter, Alaeddin’s wife.
[396]
But the net result of the costly expedition was the reconciliation of the
two emirs. The only result recorded by the Ottoman historians is that
Alaeddin kissed Murad’s hands! Murad withdrew to Kutayia without
annexing any portion of the Karamanian emirate, without booty, and
without promise either of tribute or military contingents for the European
wars. Had Murad actually accomplished more than merely holding his own
in the battle of Konia, the campaign would not have ended so profitlessly.
Granting the Ottoman victory, Murad’s conduct after the battle is
inconsistent with his whole life and character. We are compelled to discard
the story of a decisive victory. It must be that Murad, who had been able to
reduce to vassalage the Byzantines, the Bulgarians, and the Serbians, found
himself unable, even with the help of his European allies, to break the
power of this rival Anatolian emir.
XV
During the Karamanian campaign, Murad adopted the policy of treating
non-combatants in a friendly fashion. Strict orders were given to refrain
from violence and looting. Murad hoped to win the Karamanlis by
kindness, and to pave the way for a later assimilation. It was the first

campaign undertaken against fellow Moslems. The Serbian contingent, who
cared nothing for the success of this policy, and who claimed that they had
been promised booty in return for their services, did not obey the order. A
number of them were summarily executed.
[397]
When the survivors returned to their homes in the spring of 1388, they
complained bitterly of the way they had been treated, and declared that
service in the Ottoman army, for the Christian all risk and no gain, was
nothing less than a slavery leading to death. This discontent gave Lazar the
opportunity for which he had long been looking. He decided to profit by the
resentment of the Serbians against Murad,
[398]
and make a supreme effort
to free Serbia from the menace of the Ottoman yoke, which had grown very
real since the capture of Nish.
The Slavs of upper Serbia and of Bosnia realized the imminence of an
Ottoman invasion, and they were now ready—or at least they appeared to
be ready—to rally around Lazar. Up to this time the Serbians had never
recognized Lazar as the leader of the race.
The pan-Serbian alliance was made possible by the adhesion of Tvrtko,
kral of Bosnia. He had come into prominence after the battle of Cernomen
as a supporter of Lazar against the sons of Vukasin and other Serbian
chieftains who were dissatisfied with the election of Lazar. But in return for
his aid, he got under his control a large part of upper Serbia, including
Milesevo, which was the burial-place of St. Sava, apostle to the Serbians. In
1376, he crowned himself ‘king of Bosnia and Serbia’ on the tomb of St.
Sava, placing upon his head the two crowns, and changing his name to
Stephen. Neither Louis of Hungary nor Lazar was consulted by Tvrtko, and
he took no measures to secure their assent to his pretensions. After his
coronation, he conquered Cattaro, and fought successfully with Balza of
Albania.
[399]
In 1383 Tvrtko had become so powerful on the Dalmatian coast that the
Senate recognized him as ‘king of Serbia, Bosnia and the Riviera’, and
bestowed upon him the privilege of Venetian citizenship.
[400]
It was
evidently the intention of Venice to favour Tvrtko as an opponent to Louis
of Hungary, who had himself taken in 1382 the title of ‘king of Serbia,
Dalmatia and Bulgaria’.
[401]
Venice lost her grip upon or interest in the east
coast of the Adriatic for a few years immediately following the treaty of
Turin. We have already seen how in 1384 the Senate professed a

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Fruit Breeding Approaches And Achievements 2nd Anil Kumar Shukla

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