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Heat Stress Tolerance in Plants
Heat Stress Tolerance in Plants
Physiological, Molecular and Genetic

Perspectives
Edited by
Shabir Hussain Wani
Sher-e-Kashmir University of Agricultural Sciences and
Technology of Kashmir, India
Vinay Kumar
Modern College, Savitribai Phule Pune University, India
k
This edition first published 2020

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Library of Congress Cataloging-in-Publication Data
Names: Wani, Shabir Hussain, editor. | Kumar, Vinay (Biotechnologist),

editor.
Title: Heat stress tolerance in plants : physiological, molecular and
genetic perspectives / [edited by] Shabir Hussain Wani, Vinay Kumar.
Description: First edition. | Hoboken, NJ : John Wiley & Sons, Inc., 2020.
| Includes bibliographical references and index.
Identifiers: LCCN 2019032800 (print) | LCCN 2019032801 (ebook) | ISBN
9781119432364 (cloth) | ISBN 9781119432395 (adobe pdf ) | ISBN
9781119432388 (epub)
Subjects: LCSH: Plants--Effect of heat on. | Plants--Effect of stress on. |
Vegetation and climate--Research.
Classification: LCC QK755.5 .H43 2020 (print) | LCC QK755.5 (ebook) | DDC
581.4/2--dc23
LC record available at https://lccn.loc.gov/2019032800
LC ebook record available at https://lccn.loc.gov/2019032801
Cover Design: Wiley

Cover Images: Miaze plant damaged ©Aphakorn Fuengtee/Shutterstock, DNA structure isolated
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Set in 10/12pt WarnockPro by SPi Global, Chennai, India
Printed and bound by CPI Group (UK) Ltd, Croydon, CR0 4YY
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v
Contents
List of Contributors xiii

Foreword xix
About the Book xxi
About the Editor xxiii
1 Heat Tolerance in Cotton: Morphological,

Physiological, and Genetic Perspectives 1
Muhammad Tehseen Azhar, Shabir Hussain Wani,
Muhammad Tanees Chaudhary, Tariq Jameel,
Parwinder Kaur, and Xiongming Du
1.1 Introduction 1
1.1.1 Morphological and Physiological Traits 2
1.1.1.1 Seedling and Root Growth 3
1.1.1.2 Stomatal Conductance 3
1.1.1.3 Cell Membrane Thermostability 4
1.1.1.4 Canopy Temperature 5
1.1.1.5 Chlorophyll Content 6
1.1.2 Genetics and Molecular Basis of Heat Tolerance in
Cotton 8
1.1.3 Conventional Breeding Approaches 9
1.1.4 Modern Molecular Breeding Approaches 10
1.2 Conclusion and Future Prospects 12
References 12
vi Contents
2 Seed Priming as a Method to Generate

Heat-stress Tolerance in Plants: A
Minireview 23
Aditya Banerjee and Aryadeep Roychoudhury
2.1 Introduction 23
2.2 Mechanism of Heat Stress Injury in Plants 24
2.3 Seed Priming Generating Heat-stress Tolerance 26
2.4 Conclusion 27
2.5 Future Perspectives 27
Acknowledgments 28
References 28
3 How Effective Are Stress-associated Proteins

in Augmenting Thermotolerance? 33
Inès Karmous and Sandeep Kumar Verma
3.1 Introduction 33
3.1.1 Heat Shock Proteins (HSPs) 34
3.1.2 Proline 36
3.1.3 Dehydrins (DHNs) 37
3.1.4 Role of Metabolic Proteins in Thermotolerance 37
3.2 Conclusion 40
References 40
4 Biochemical and Molecular Markers:

Unraveling Their Potential Role in Screening
Germplasm for Thermotolerance 47
Ahmed Ismail, Kareem A. Mosa, Muna A. Ali, and
Mohamed Helmy
4.1 Introduction 47
4.2 Types of Markers 48
4.3 Morphological Markers 49
4.4 Molecular Markers 49
4.5 Biochemical Markers 53
4.6 Quantitative Trait Loci for Plant
Thermotolerance 54
4.7 Plant Metabolites Under Heat Stress 61
4.8 Antioxidant Enzymes and Heat Stress 63
4.9 Conclusion 68
References 70
Contents vii
5 Alteration in Carbohydrate Metabolism

Modulates Thermotolerance of Plant under
Heat Stress 77
Roseline Xalxo, Bhumika Yadu, Jipsi Chandra,
Vibhuti Chandrakar, and S. Keshavkant
5.1 Introduction 77
5.1.1 Heat Stress and Thermotolerance 79
5.1.1.1 Morphological Alterations 80
5.1.1.2 Anatomical Alterations 80
5.1.1.3 Physiological and Biochemical
Modifications 81
5.1.1.4 Cell Membrane Integrity 82
5.2 Carbohydrate as Protectives Molecules 83
5.2.1 Osmolyte 83
5.2.2 Thermoprotectant 84
5.3 Carbohydrates as Signaling Molecules 85
5.3.1 Reproductive Cell Development 85
5.3.2 Seed Development 86
5.3.3 Seed Germination and Yield Loss 87
5.4 Adverse Impacts of Heat Stress 87
5.4.1 Photosynthesis 87
5.4.1.1 Altered Carbon Assimilation 88
5.4.1.2 Chlorophyll Breakdown 88
5.4.2 Major and Minor Carbohydrate
Metabolism 89
5.4.3 Expression of Regulatory Genes 89
5.4.4 Enzyme Activity 90
5.5 Mechanisms Involved in Thermotolerance 93
5.5.1 Glucose and Heat-stress Tolerance 93
5.5.2 Sucrose and Heat-stress Tolerance 94
5.5.3 Fructan and Heat-stress Tolerance 95
5.5.4 Trehalose and Heat-stress Tolerance 96
5.5.5 Raffinose and Heat-stress Tolerance 97
5.6 Genetic Approaches/Strategies for Improving
Thermotolerance 97
5.6.1 Genetically Modified Crop Production 97
5.6.2 Transgenic Strategies 99
5.7 Conclusions and Future Perspectives 102
References 103
viii Contents
6 Transcriptomics to Dissect Plant Responses

to Heat Stress 117
Sagar Satish Datir
6.1 Introduction 117
6.1.1 Transcriptome Sequencing and Expression Profiling of
Genes Involved in Heat Stress Response 119
6.1.1.1 Rice (Oryza sativa L.) 119
6.1.1.2 Wheat (Triticum aestivum L.) 123
6.1.1.3 Maize (Zea mays L.) 125
6.1.1.4 Switchgrass (Panicum virgatum L.) and Ryegrass
(Lolium perenne L.) 126
6.1.1.5 Spinach (Spinacia oleracea L.) 128
6.1.1.6 Brassica rapa L. 129
6.1.1.7 Banana (Musa acuminate Colla) 130
6.2 Conclusions 131
References 132
7 Proteomics as a Tool for Characterizing the

Alteration in Pathways Associated with
Defense and Metabolite Synthesis 141
Reetika Mahajan and Sajad Majeed Zargar
7.2 What Is Proteomics? 142
7.3 Need of Proteomics in Post-genomic Era 143
7.4 Different Branches of Proteomics 143
7.5 Techniques Used in Quantitative Proteomics 145
7.5.1 Gel-based and Gel-free Methods 146
7.5.2 Label-based and Label-free Methods 149
7.6 Role of Proteomics in Studying Alteration in Pathways
Associated with Defense and Metabolite
Synthesis 150
7.7 Conclusion and Future Perspective 156
References 156
8 RNA World and Heat Stress Tolerance in

Plants 167
Usman Ijaz, Muhammad Amjad Ali, Habibullah Nadeem,
Lin Tan, and Farrukh Azeem
8.2 Plant microRNAs 168
Contents ix
8.3 Small Interfering RNA (siRNA) 177

8.4 Long Noncoding RNAs (lncRNAs) 178
8.5 Circular RNAs (circRNAs) 179
8.6 Conclusions and Future Perspectives 180
References 180
9 Heat Shock Proteins: Master Players for

Heat-stress Tolerance in Plants during Climate
Change 189
Annu Yadav, Jitender Singh, Koushlesh Ranjan, Pankaj
Kumar, Shivani Khanna, Madhuri Gupta, Vinay Kumar,
Shabir Hussain Wani, and Anil Sirohi
9.2 Classification of HSPs 192
9.2.1 HSP100 192
9.2.1.1 Structure of HSP100 192
9.2.1.2 Mode of Action: The HSP100 Chaperone Cycle 192
9.2.2 HSP90 195
9.2.2.2 Mode of Action: The HSP90 Chaperone Cycle 197
9.2.3 HSP70 198
9.2.3.2 Mode of Action: The Hsp70 Chaperone Cycle 198
9.2.4 HSP60 199
9.2.4.2 Mode of Action: The Hsp60 Chaperone Cycle 201
9.2.5 The Small Heat Shock Protein Family (sHSPs) 202
9.2.5.1 Structure of sHSP 202
9.2.5.2 Mode of Action: Small Heat Shock Proteins 203
9.3 HSPs Expression Under Heat Stress Condition 203
9.4 Conclusion and Future Prospects 205
References 205
10 The Contribution of Phytohormones in Plant

Thermotolerance 213
Sonal Mishra, Mansi Bhardwaj, Shakti Mehrotra, Aksar Ali
Chowdhary, and Vikas Srivastava
10.2 Protectants in Heat Stress Alleviation 215
10.2.1 Osmolytes 215
x Contents
10.2.2 Nutrients 215

10.2.3 Signaling Molecules 217
10.2.4 Polyamines 217
10.2.5 Phytohormones 218
10.3 Application of Hormones in HT Management 218
10.3.1 Auxin 219
10.3.2 Gibberellin 221
10.3.3 Cytokinin 222
10.3.4 Abscisic Acid 224
10.3.5 Ethylene 225
10.3.6 Salicylic Acid 225
10.3.7 Jasmonic Acid 227
10.3.8 Brassinosteroid 228
10.4 Conclusion and Prospects 229
Acknowledgements 229
References 230
11 Exploring In-built Defense Mechanisms in

Plants under Heat Stress 239
Giridara Kumar Surabhi and Jatindra Kumar Seth
11.2 Effect of Heat Stress on Crop Plants 240
11.2.1 Heat Stress Effects on Physiology and Cell
Structures 241
11.2.2 Heat Stress Effects on Vegetative Stages 242
11.2.3 Heat Stress Effects on Reproductive Stage 243
11.2.4 Heat Stress Effects on Yield 243
11.3 Threshold Temperature 244
11.4 In-built Defense System in Plants to Overcome High
Temperature Stress 245
11.4.1 Accumulation of Thermoprotectants 245
11.4.1.1 Heat Shock Proteins (HSPs) 245
11.4.1.2 Proline 246
11.4.1.3 Glycinebetaine (GB) 248
11.4.1.4 Abscisic Acid (ABA) 249
11.4.1.5 Salicylic Acid (SA) 250
11.4.1.6 Heat Stress Effects on Secondary Metabolism 255
11.4.2 Transcriptional Regulation 256
Contents xi
11.4.3 Role of Small RNAs (miRNAs) in Heat-stress

Tolerance 258
11.5 Conclusion 261
Acknowledgement 262
References 263
Index 283
xiii
List of Contributors
Muna A. Ali Haikou Experimental Station

Department of Applied Chinese Academy of Tropical
Biology, College of Sciences Agricultural Sciences
University of Sharjah (CATAS), Hainan Key
Sharjah, Laboratory of Banana Genetic
UAE Improvement
Haikou, Hainan Province
Muhammad Amjad Ali
China
Department of Plant
Pathology Aditya Banerjee
University of Agriculture Department of Biotechnology
Faisalabad, St. Xavier’s College
Pakistan (Autonomous)
Kolkata, West Bengal,
Farrukh Azeem
India
Department of Bioinformatics
and Biotechnology Mansi Bhardwaj
Government College ICAR-National Institute for
University Faisalabad (GCUF) Plant Biotechnology
Faisalabad, IARI Campus, New Delhi,
Pakistan India
and
xiv List of Contributors
Jipsi Chandra Xiongming Du

School of Studies in Institute of Cotton Research
Biotechnology of Chinese Academy of
Pt. Ravishankar Shukla Agricultural Sciences, State
University Key Laboratory of Cotton
Raipur, Biology
India Anyang, Henan,
China
Vibhuti Chandrakar
School of Studies in Madhuri Gupta
Biotechnology College of Biotechnology
Pt. Ravishankar Shukla Sardar Vallabhbhai Patel
University University of Agriculture &
Raipur, Technology
India Meerut, Uttar Pradesh,
India
Muhammad Tanees Chaudhary
Department of Plant Breeding Mohamed Helmy
and Genetics The Donnelley Centre
University of Agriculture University of Toronto
Faisalabad, Toronto, ON,
Pakistan Canada
Aksar Ali Chowdhary Usman Ijaz
Department of Botany Department of Bioinformatics
Central University of Jammu and Biotechnology
Samba, Government College
India University Faisalabad (GCUF)
Faisalabad,
Sagar Satish Datir
Pakistan
Biology Department
Queen’s University Ahmed Ismail
Kingston, ON, Department of Biotechnology,
Canada Faculty of Agriculture
Al-Azhar University
and
Cairo,
Department of Biotechnology Egypt
Savitribai Phule Pune
and
University
Pune, MS,
India
List of Contributors xv
Public Health Department, National Center for Natural

College of Applied Medical Resources
Science Pt. Ravishankar Shukla
Majmaah University University
Majmaah City, Raipur,
KSA India
Tariq Jameel Shivani Khanna
Department of Plant Breeding College of Biotechnology
and Genetics Sardar Vallabhbhai Patel
University of Agriculture University of Agriculture &
Faisalabad, Technology
Pakistan Meerut, Uttar Pradesh,
India
Inès Karmous
Biotechnology Laboratory, Pankaj Kumar
Department of Biology College of Biotechnology
Bolu Abant Izzet Baysal Sardar Vallabhbhai Patel
University University of Agriculture &
Bolu, Technology
Turkey Meerut, Uttar Pradesh,
India
Parwinder Kaur
UWA School of Agriculture Vinay Kumar
and Environment Department of Biotechnology
The University of Western Modern College of Arts,
Australia Science and Commerce,
Crawley, Western Australia, Savitribai Phule Pune
Australia University, Pune,
India
S. Keshavkant
School of Studies in Reetika Mahajan
Biotechnology Govt. Degree College Poonch
Pt. Ravishankar Shukla Jammu and Kashmir,
University India
Raipur,
India
and
xvi List of Contributors
Shakti Mehrotra Koushlesh Ranjan

Department of Biotechnology Department of Veterinary
Institute of Engineering and Physiology and Biochemistry
Technology College of Veterinary and
Lucknow, Animal Sciences, Sardar
India Vallabhbhai Patel University
of Agriculture and
Sonal Mishra
Technology
School of Biotechnology
Meerut, Uttar Pradesh,
University of Jammu
India
Jammu,
India Aryadeep Roychoudhury
Department of Biotechnology
Kareem A. Mosa
St. Xavier’s College
Department of Biotechnology,
(Autonomous)
Faculty of Agriculture
Kolkata, West Bengal,
Al-Azhar University
India
Cairo,
Egypt Jatindra Kumar Seth
Plant Molecular Biology and
and
‘OMICS’ Laboratory
Department of Applied Regional Plant Resource
Biology, College of Sciences Centre
University of Sharjah Bhubaneswar, Odisha,
Sharjah, India
UAE
Jitender Singh
Habibullah Nadeem College of Biotechnology
Department of Bioinformatics Sardar Vallabhbhai Patel
and Biotechnology University of Agriculture &
Government College Technology
University Faisalabad (GCUF) Meerut, Uttar Pradesh,
Faisalabad, India
Pakistan
List of Contributors xvii
Anil Sirohi Sandeep Kumar Verma

College of Biotechnology Biotechnology Laboratory,
Sardar Vallabhbhai Patel Department of Biology
University of Agriculture & Bolu Abant Izzet Baysal
Technology University
Meerut, Uttar Pradesh, Bolu,
India Turkey
Vikas Srivastava Shabir Hussain Wani
Department of Botany Mountain Research Centre
Central University of Jammu for Field Crops
Samba, Khudwani, Sher-e-Kashmir
India University of Agricultural
Sciences and Technology of
Giridara Kumar Surabhi
Kashmir
Plant Molecular Biology and
Jammu and Kashmir,
‘OMICS’ Laboratory
India
Regional Plant Resource
Centre Roseline Xalxo
Bhubaneswar, Odisha, School of Studies in
India Biotechnology
Pt. Ravishankar Shukla
Lin Tan
University
Haikou Experimental Station,
Raipur,
Chinese Academy of Tropical
India
Agricultural Sciences
(CATAS) Annu Yadav
Hainan Key Laboratory of College of Biotechnology
Banana Genetic Improvement Sardar Vallabhbhai Patel
Haikou, Hainan Province, University of Agriculture &
China Technology
Meerut, Uttar Pradesh,
Muhammad Tehseen Azhar
India
Department of Plant Breeding
and Genetics
University of Agriculture
Faisalabad,
Pakistan
xviii List of Contributors
Bhumika Yadu Sajad Majeed Zargar

School of Studies in Division of Plant
Biotechnology Biotechnology
Pt. Ravishankar Shukla Sher-e-Kashmir University of
University Agricultural Science and
Raipur, Technology of Kashmir
India Shalimar, Srinagar,
Jammu and Kashmir,
India
xix
Foreword
One of the prevalent environmental

stresses encountered by plants dur-
ing their important growth stages is
heat stress. Exposure to heat stress
for prolonged periods can even result
in plant death. Plants can be dam-
aged by either high day or high night
temperatures and by either high air
or high soil temperatures. Predictions
indicate that temperatures will inten-
sify by another 2–6 ∘ by the climax of
this century which will increase the likelihood of more fre-
quent and more severe heat stress in plants, begetting a serious
reduction in crop yields. The genetic basis of heat stress adapta-
tion is poorly understood. Conventional breeding methods have
met with limited success in improving the heat-stress tolerance
of important crop plants through interspecific or intergeneric
hybridization. Therefore, it is imperative to accelerate efforts for
unraveling the biochemical, physiological, and molecular mech-
anisms underlying heat-stress tolerance in plants. Heat stress
can induce diverse physiological and molecular responses in
plants that lead to the disturbance of various cellular metabolic
processes, impair membrane stability, cause protein denatura-
tion and thermal aggregation, and consequently affect plant
growth and development. The heat stress response is also illus-
trated by the inhibition of regular transcription and translation,
elevated expression of heat shock proteins (HPSs), and induc-
tion of heat tolerance. Recent advances in molecular genetics
xx Foreword
approaches have provided new insights into the plant heat stress
response. The QTL-based approaches allow loci to be identified
for markers that are linked to heat tolerance. Further, discovery
of novel genes and pathways, analyses of expression patterns,
and the determination of function of these genes during heat
stress adaptation will offer a basis for efficient engineering strate-
gies with the aim to enhance heat stress tolerance in plants.
I am delighted that the edited volume Heat-Stress Tolerance
in Plants: Physiological, Molecular and Genetic Perspectives is
being published. The chapters included in this book are nicely
written by scientists and researchers. This book describes recent
advances in heat-stress tolerance utilizing genetic and genomic
approaches and their application in important agricultural
crops. Recent advancements in plant omics and their impor-
tance and application in the development of heat-stress-tolerant
crops have also been included.
I congratulate the editors for unravelling this volume and hope
that this will be a useful reference material for researchers, stu-
dents, and policymakers.
9th May 2019 T. Mohapatra

New Delhi
xxi
About the Book
The global population is 7.3 billion and by 2050 it is expected to

reach 9.7 billion, while at the same time agricultural productiv-
ity is not increasing with the same speed, owing to the mounting
environmental constraints as a result of climate change. Heat
stress, which can be defined as a period in which temperatures
are hot enough for a sufficient period of time to cause irreversible
damage to a plant’s function or development, is one of the major
environmental stresses hampering plant growth and productiv-
ity severely. The projections about the increase in global ambient
temperatures over the next few decades are expected to increase
the intensity of the impact of heat stress on plants.
Owing to limited genetic basis for plant heat stress responses
and tolerance besides limited success of conventional breed-
ing approaches, a deeper understanding of the mechanisms
underlying plants’ responses to heat stress is needed. Recent
investigations confirmed that heat stress inhibits transcription
and translation, and increases the levels of heat shock pro-
teins (HSPs). During extreme heat stress conditions, signaling
pathways leading to apoptotic cell death are also activated. As
molecular chaperones, HSPs offer protection to cells against
the destructive effects of heat stress and augment survival. The
improved expression of HSPs is regulated by heat shock tran-
scription factors (HSFs). Several recent advances in identifying
gene(s) and/or pathways and their roles in heat stress responses
and tolerance have offered opportunities for researchers to
explore these gene(s) for engineering heat-stress tolerance in
major crops. Studies involving full genome profiling/sequencing,
and mutational and transgenic plant analyses have provided a
xxii About the Book
deep insight of the complex transcriptional mechanism that

operates under heat stress.
Through this book, we intend to integrate the contributions
from potential plant scientists targeting heat-stress tolerance
mechanisms using physiological, biochemical, molecular, and
genetic approaches.
xxiii
About the Editor
Dr. Shabir Hussain Wani is senior Assistant Professor at

Mountain Research Centre for Field Crops, Khudwani – 192101,
Sher-e-Kashmir University of Agricultural Sciences and Tech-
nology of Kashmir, Jammu and Kashmir, India. He received
a PhD in plant breeding and genetics on “transgenic rice for
abiotic stress tolerance” from the Punjab Agricultural Univer-
sity Ludhiana, India. After obtaining his PhD, he worked as a
research associate in the Biotechnology Laboratory, Central
Institute of Temperate Horticulture (ICAR), Srinagar, India. He
then joined the Krishi Vigyan Kendra (Farm Science Centre) as
program coordinator at Senapati, Manipur, India. He teaches
courses related to plant breeding, seed science and technol-
ogy, and stress breeding, and has published more than 100
papers/chapters in journals and books of international and
national repute. He served as guest editor and reviews editor for
the journal Frontier in Plant Science (2015–2018). He has also
edited several books on current topics in crop improvement
for abiotic stress tolerance published by Springer Nature and
CRC Press. His PhD research fetched first prize in the North
Zone Competition, at national level, in India. He was awarded
Young Scientist Award from the Society for Promotion of Plant
Sciences, Jaipur, India, in 2009. He is a fellow of the Society for
Plant Research, India. Recently, he also received Young Scien-
tist Award (Agriculture) 2015 from Society for Plant Research,
Meerut, India. He also served as visiting Scientist at Department
of Plant Soil and Microbial Sciences, Michigan State University,
USA under the UGC Raman Post Doctoral Fellowship program.
xxiv About the Editor
Currently, he is in charge of Wheat improvement program at

MRCFC Khudwani SKAUST Kashmir.
Dr. Vinay Kumar is an Associate Professor at the Post-

graduate Department of Biotechnology, Progressive Education
Society’s Modern College of Arts, Science and Commerce,
Ganeshkhind, Pune, India and a Visiting Faculty at the Depart-
ment of Environmental Sciences, Savitribai Phule University,
Pune, India. He obtained his PhD in Biotechnology from
Savitribai Phule Pune University (formerly University of Pune)
in 2009. For his PhD, he worked on metabolic engineering of
rice for improved salinity tolerance. He has published more
than 40 peer-reviewed research/review articles, contributed
18 book chapters and edited 4 books including this, published
by Springer and Wiley. He is a recipient of the Government of
India’s Science and Engineering Board, Department of Science
and Technology (SERB-DST) Young Scientist Award in 2011.
His current research interests include elucidating molecular
mechanisms underlying salinity stress responses and toler-
ance in plants. His research group is engaged in deciphering
the molecular mechanisms responsible plant abiotic stress
tolerance.
1
Heat Tolerance in Cotton: Morphological,

Physiological, and Genetic Perspectives
Muhammad Tehseen Azhar 1 , Shabir Hussain Wani 2 ,
Muhammad Tanees Chaudhary 1 , Tariq Jameel 1 ,
Parwinder Kaur 3 , and Xiongming Du 4
1
Department of Plant Breeding and Genetics, University of Agriculture, Faisalabad,
Pakistan
2
Mountain Research Centre for Field Crops, Khudwani, Sher-e-Kashmir University of
Agricultural Sciences and Technology of Kashmir, Jammu and Kashmir, India
3
UWA School of Agriculture and Environment, The University of Western Australia,
Crawley, Western Australia, Australia
4
Institute of Cotton Research of Chinese Academy of Agricultural Sciences, State Key
Laboratory of Cotton Biology, Anyang, Henan, China
1.1 Introduction
Cotton belongings to the genus Gossypium and four species,
namely G. hirsutum L., G. barbadense L., G. arboreum L., and
G. herbaceum L., that were grown independently for the purpose
of textile fiber (Fryxell 1992). Cotton is mainly grown in tropical
and sub-tropical areas having temperatures varying from 40 to
45 ∘ C (Ashraf et al. 1994). The phenological development and
accumulation of plant biomass depend upon temperature during
its growing season (Echer et al. 2014). Cotton sheds its flowers
and squares when the temperature reaches 36 ∘ C (Reddy et al.
1991). The plant reproduction, metabolism, and productivity
of cotton are adversely affected by high-temperature stress
(Demirel et al. 2014). The response of plants to heat stress
depends upon the duration and degree of heat stress (Hasanuz-
zaman et al. 2013). Furthermore, high-temperature stress is
closely related to water deficit and can be further exacerbated
by limited or unreliable availability of water in cotton-growing
Heat Stress Tolerance in Plants: Physiological, Molecular and Genetic Perspectives, First Edition.
Edited by Shabir Hussain Wani and Vinay Kumar.
© 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.
2 1 Heat Tolerance in Cotton
areas (Rizhsky et al. 2004). The presence of genetic variation

within a species is a prerequisite for a breeding program for the
development of genotypes responsive to heat stress (Azhar et al.
2009). All metabolic and biochemical activities of plants require
an optimum temperature range, which is termed the thermal
kinetic window (TKW). Plant temperature should be within
range of the TKW. The plant comes under heat stress if its
temperature goes below or above TKW. A TKW of 23.5–32 ∘ C
is required for proper plant growth (Sawan et al. 2009). Plant
breeders are continually screening cotton germplasm for heat
stress due to climate change. Like other physiological traits,
seed cotton yield (SCY) is considered a beneficial trait for the
development of cotton germplasm against high temperature
but it is complex, and affected by climatic conditions, and so
requires very specialized breeding programs, but even then one
cannot avoid environmental variations (Demİrel et al. 2016).
High temperature exerts negative impact due to inhibition
of photosynthesis process (DeRidder and Salvucci 2007). The
modifications induced by high temperature may be direct,
owing to changes in the physiological process, or can be indirect
by altering the developmental patterns. For example, developing
seeds may be affected by heat stress that may delay the germina-
tion or loss of vigor. Ultimately it will lead to reduced emergence
and seedling establishment (Wahid et al. 2007). The exposure of
cotton to high temperature causes shedding of 35% premature
flower and 50–75% of bolls in variable climatic conditions.
1.1.1 Morphological and Physiological Traits

One of the most economical and suitable ways to mitigate the
adverse effects of heat stress is to develop or identify heat-
tolerant cultivars. Under various heat stress regimes, plants
exhibit different kinds of survival mechanism, which include
various heat stress avoidance mechanisms, phonological alter-
ations like variable leaf angle, transpirational mechanism, or
changes in lipid membrane composition, and various mor-
phological changes (Niinemets 2010). Plants’ stomata closing
mechanism, reduced water loss, larger stomatal and trichoma-
tous densities and widening xylem vessels are the most common
1.1 Introduction 3
approaches followed by the plant under different conditions of

heat stress (Hasanuzzaman et al. 2013).
Research work about the use of various plant and physiological
traits is reviewed in the proceeding paragraphs.
1.1.1.1 Seedling and Root Growth

The germination and development of seedlings of crop plants
depend upon remaining with the optimum temperature range
of 28 to 30 ∘ C. The base temperature is about 12 ∘ C for seed
germination and 15.5 ∘ C for seedling growth. Cool temperature
between 2 and 4 ∘ C is a major problem in various places in
the United States, mainly across the Delta region of Missis-
sippi during germination and initial growth of the seedlings.
Genotypic differences have been observed for germination
and root development under cool soil temperatures (Mills
et al. 2005). While at the time of sowing in northern India, the
wind velocity and soil temperature are very high, resulting in
a rapid loss of soil moisture (Lather et al. 2001). McMichael
and Burke (1994) reveal that soil with a temperature range of
between 20 and 32 ∘ C is suitable for proper root growth and
development. Root temperature stress ranges between 35 and
40 ∘ C and badly affects the hydraulic conductivity and nutrient
uptake ability of the plant and causes low hormone synthesis,
and badly affects hormone transportation (Clark and Reinhard
1991; Burke and Upchurch 1995). Synthesis of cytokinins which
originate predominantly in roots is among the most sensitive
processes (Paulsen 1994). Many functions of roots – comprising
the uptake of nutrients and water, assimilation and synthe-
sis of metabolites, and translocation – are very sensitive to
temperature. Root temperature may be more critical than
shoot temperatures for plant growth because roots tolerate a
shorter range of temperature and are less adaptable to extreme
variations (Nielsen 1974).
1.1.1.2 Stomatal Conductance

Stomatal closures to reduce transpiration rates also reduce pho-
tosynthesis rates. Increased stomatal conductance helps with
the cooling of leaves by evaporation and thereby reduces ther-
mal stress. Optimum temperatures of < 30 ∘ C during daytime in
upland and Pima cotton are well below commonly occurring air
temperatures in most cotton-growing areas (Radin et al. 1994).

Stomatal conductance and net photosynthesis are inhibited by
moderate heat stress in many plant species, owing to decreases
in the activation state of rubisco (Crafts-Brander and Salvucci
2002; Morales et al. 2003). Stomata could be explored for the
development of heat tolerance in crop plants because they reg-
ulate the transpiration rate and determine the degree to which
water evaporation can cool down the leaves. If stomata open
wider in higher-yielding lines, transpiration may be increased
and photosynthetic rates improved because CO2 as well as
water vapors must diffuse through the stomata. A broader
examination of 16 species performed by Monteith (1995)
supported the theory that stomata respond not to humidity
levels but to respiration rates. Moneith (1995) also identified
that stomata from all investigated species had patchy stomata
closure in response to increased respiration rates to limit water
loss. Many studies were intended to assess the germplasm of
cotton and wheat by using stomatal conductance and pho-
tosynthesis as criteria for the identification of heat-tolerant
genotypes (Cornish et al. 1991; Lu et al. 1998; Ulloa et al. 2000;
Rahman 2005).
1.1.1.3 Cell Membrane Thermostability

The conventional screening methods of large germplasm are
time consuming and laborious but the success of any breeding
program depends on effective evaluation techniques (Asha
and Lal Ahamed 2013). Cell membrane thermostability (CMT)
protocol has been suggested by Sullivan (1972) as a depend-
able method for measuring heat tolerance by quantifying the
amount of electrolyte leakage from leaf disks after exposure
to heat stress, and this technique has been used efficiently
in potato (Coria et al. 1998), rice (Maavimani et al. 2014),
tomato (Golam et al. 2012), soybean (Martineau et al. 1979),
wheat (Blum et al. 2001), sorghum (Marcum 1998), and barley
(Wahid and Shabbir 2005). Membrane thermostability has been
reported to have a strong genetic correlation with grain yield in
wheat. Heritability of membrane thermostability in maize was
estimated to be 73%. CMT has been used in cotton as a suitable
screening and selection criterion for heat tolerance, owing to
its ability to distinguish heat-sensitive and -tolerant genotypes
1.1 Introduction 5
within the species (Singh et al. 2007). Marsh et al. (2000) found
a large portion of the variability for membrane stability to be
controlled by a small number of genes. Rahman et al. (2004)
indicate that exposure to high temperature preceding the CMT
test produces better distinction between heat-tolerant and
heat-susceptible cultivars; however, they caution regarding its
indirect selection on the basis of SCY under non-heat-stressed
environments, so Rahman et al. (2004) conclude that CMT
could be a useful physiological technique for screening large
sizes of germplasm of cotton for heat tolerance. In addition,
chlorophyll fluorescence, membrane leakage, and leaf extension
growth (Jamieson et al. 1995) were also the most practical,
dependable, but sensitive methods for quantifying the response
of cottons to high-temperature stress. This permeability acti-
vates a series of calcium-dependent protein kinases (CDPKs)
that spur cellular production of increased antioxidants (Wahid
et al. 2007). The effectiveness of CMT for field-grown leaf mate-
rial to determine difference in genotypes in heat tolerance may
be very useful in detecting the desirable genotypes for higher
temperatures (Singh et al. 2007). However, membrane stability
requires destructive sampling, and there is a large potential for
error inherent in the process of estimating membrane stability.
Similarly, measurements of chlorophyll fluorescence require the
use of expensive instrumentation and, in some cases, necessitate
dark adaptation of the leaf tissue, which limits the number of
plants that can be screened in a given day. The ability of cotton
to adjust its membrane structure under heat-stressed conditions
has been recognized as a significant physiological adaptation to
heat stress (Rahman et al. 2004).
1.1.1.4 Canopy Temperature

Owing to high temperature, nearly 65–70% of fruiting points
shed down because of heat-induced pollen sterility, which
causes a poor yield of seed cotton at harvest time (Taha et al.
1981; Hofmann et al. 1988; Baloch et al. 2000; Liu et al. 2006).
The relationship of canopy temperature (Tc) with transpiration
has been used for the detection of heat stress in crop plants
(Conaty et al. 2015). The development of heat tolerance in
cultivars through conventional breeding approach depends
upon the accessibility of an optimized protocol which could give
advantage to screen large number of varieties/lines (germplasm)

rapidly. A new physiological infrared technique that measures
the canopy temperature is becoming increasingly popular these
days as a way of screening large amount of germplasm for heat
tolerance (Singh et al. 2007). This technique had been used
by various workers in cotton and corn for the identification of
heat-tolerant lines which were used independently in breeding
programs (Wanjura et al. 2004; Karademir et al. 2012). The
same assay was used for drought tolerance in cotton and wheat
(Hatfield et al. 1987; Rashid et al. 1999; Blum et al. 2001). The use
of canopy temperature to assess plant growth and development
where there is less irrigation may be more reliable than air
temperature alone (Mahan et al. 2014). Previous studies have
evaluated the optimum thermal environment for growth and
metabolism, placing the thermal optimum for cotton at 28 ∘ C,
within a range of 25–31 ∘ C (Burke 1990; Hatfield and Burke 1991;
Mahan et al. 1995; Burke et al. 2004; Conaty et al. 2012). The PAR
model is used to detect the canopy temperature with optimal
rate of photosynthesis to produce different consequences of
plant architecture (Alarcon and Sassenrath 2015).
Crop management practices including suitable plant popu-
lation density (PPD) are an important technique for adjusting
canopy light distribution and increasing canopy photosyn-
thetic capacity of cotton grown in the field (Yao et al. 2016).
Enhancements in canopy photosynthetic capacity are essential
to increase yield (Zhang et al. 2004; Dong et al. 2012; Mao
et al. 2014; Dai et al. 2015). Canopy photosynthetic capacity is
also significantly affected by proper crop management prac-
tices (Dong and Liu 2010; Dong et al. 2012; Mao et al. 2014;
Dai et al. 2015).
1.1.1.5 Chlorophyll Content

Photosynthesis is extremely sensitive to the effect of increased
temperatures. Reduction in the photosynthetic rate has been
constantly shown in cotton, and is due to the decline in avail-
able carbohydrate supply to the developing bolls, leading to
increased rates of fruit shed (Guinn 1976; Hake and Silvertooth
1990; Sadras 1995; Brown and Oosterhuis 2004). Hall and Allen
(1993) hypothesize that cultivars displaying heat tolerance
during reproductive development should have the following
1.1 Introduction 7
characteristics: a high harvest index, high chlorophyll content

per unit leaf area, small leaves, and low leaf area per unit ground
area. Heat stress decreases photosynthesis, quantum efficiency,
and the chlorophyll content of plants, but increases stomatal
conductance. Photosynthesis is an important process that is
detrimentally affected by adverse environmental conditions
such as high temperature, drought, and salinity, etc. (Cothren
1999). Cotton grown under increased CO2 and higher temper-
atures has greater leaf area, increased photosynthetic capacity,
and a higher boll retention when compared to normal CO2 and
higher temperatures (Radin et al. 1987; Mauney et al. 1994). Heat
imposes negative impacts, such as reduced leaf water potential,
reduced leaf area, and premature leaf senescence, which has
negative impacts on total photosynthesis performance of plant
(Greer and Weedon 2013). Temperatures beyond the optimal
thermal limits of the species will hinder the photosynthetic
capability (Bibi et al. 2008). This is the result of photosynthesis
transitioning from noncyclic photophosphorylation pathways
to a cyclic phosphorylation pathway. During the vegetative
stage, high temperatures disturb the physiological mecha-
nisms in the leaves thereby decreasing CO2 assimilation rates,
damaging components of photosystem II (PSII) in thylakoid
membranes of the chloroplasts, and causing membrane damage
(Hall 2004). Cotton maintained significantly higher rates of
photosynthesis at 36 ∘ C with no reduction in Rubisco (ribulose-
1,5-bisphosphate carboxylase/oxygenase) activase activity at
CO2 concentrations of 450 parts per million (ppm), compared
to plants at the same temperature but with lower CO2 con-
centrations. Chlorophyll content has been used to assess high
temperature tolerance in Zea mays L. (Karim et al. 2000) and
Triticale (Grzesiak et al. 2003); however, use of chlorophyll
content as a screening technique for high temperature tolerance
in cotton is rare. But genotypic differences in cottons for
chlorophyll contents have been identified by various workers
(Pettigrew et al. 1993; Pettigrew and Meredith 1994; Pettigrew
and Meredith Jr 2012; Quisenberry et al. 1994; Pettigrew and
Turley 1998; Clement et al. 2013). These genotypic differences
and efficacy of the use of technique allow the cotton breeder
to initiate breeding work for the development of heat-tolerant
lines that enhance yield and fiber quality traits.
1.1.2 Genetics and Molecular Basis of Heat Tolerance

in Cotton
High-throughput molecular technologies assisted with
advanced phenotyping platforms provide new opportunities for
discovering marker-trait associations and can sustain genomic-
assisted breeding (Barabaschi et al. 2016). Understanding the
genetic basis of traits related to heat tolerance is essential for
the development of heat-tolerant genotypes. Basically, heat tol-
erance is not associated with a single trait; rather, it is a complex
and polygenic mechanism in field crops. Several morphological
and physiological characteristics – such as sterility of pollens,
electrolyte leakage, CMT, chlorophyll fluorescence, and heat
shock proteins (HSPs) in cotton, sorghum, wheat, and maize –
have been explored extensively (Joshi 1999). Heat-tolerance
responses have also been studied in Arabidopsis thaliana (Scharf
et al. 2001), Helianthus annuus L. (Coca et al. 1996), Nicotiana
tabacum L. (Rizhsky et al. 2002), and Solanum tuberosum L.
(Havaux 1993).
Plants respond to heat stress within 0.5–1 hour after expo-
sure to high temperature but it may vary in range of species,
by accumulating certain proteins like HSPs, moreover this
production of HSPs also depends on the temperature that the
plant experiences (Nakamoto and Hiyama 1999; Schoffl et al.
1999; Cottee et al. 2014). These HSPs and heat shock tran-
scription factors are synthesized by genes encoding heat stress.
While the rate of respiration increases, this may be due to
catabolism of assimilates and glycolysis, and it can also indicate
the up-regulation of stress-responsive genes. Besides, genes
associated with transport, signaling, and metabolism of sulfur
and nitrogen also enhance under stress (Cottee et al. 2014).
Burke and Chen (2015) report eight HSPs produced in the
dry land cotton leaves. The production of polypeptide was
quantified in cotton grown under control and heat-shocked
environment, and 11 different polypeptides were produced in
a leaf sample of heat shocked cotton, and eight of these HSPs
were similar to proteins accumulated in dry land cotton (Burke
et al. 1985). Demirel et al. (2014) studied 25 ESTs (express
sequence tags) out of which 16 were homologous to known
genes. The genes, namely TH1, FPGS, GhHS126, GhHS128,
1.1 Introduction 9
and IAR3 were up-regulated, while CTL2, RPS14, ABCC3,

LSm8, and CIPK expression was down-regulated but expres-
sions of PP2C and psaB-rps14 was not altered, owing to short-
term heat stress in cotton. The mechanism of HSPs under
high temperature is still unknown. Different studies show that
HSPs are molecular chaperones that have a role in protect-
ing cell proteins from denaturation under high temperature
(Bowen et al. 2002).
1.1.3 Conventional Breeding Approaches

Traditional breeding of heat-resistant crops basically based on
selection and common technique of selecting crops for heat
tolerance has been to grow breeding materials in a hot target
production environment and detect individuals/lines with a
higher yield (Ehlers and Hall 1998). Development of cotton lines
by conventional breeding methods reduces the loss of yield
reported due to heat stress. Selection of breeding lines for high
temperature always takes place in hot regions at critical stages
of crop plants (Mickelbart et al. 2015; Driedonks et al. 2016).
The researchers report that the optimum temperature of 23 ∘ C
is required for carbon fixation during photosynthesis and tem-
peratures above this may affect the photosynthetic processes of
cotton plant and adversely affect the yield of seed cotton and
fiber characteristics (Bibi et al. 2008; Song et al. 2014). Song et al.
(2014) reveal that high temperature affect the rate of pollen ger-
mination and boll setting. Numbers of potential heat-tolerant
lines of cotton were identified as having viable pollens after
the exposure of their pollens at 35 ∘ C for 15 minutes. The same
assay was used for the identification of heat-tolerant families
during segregating populations. In addition, Abro et al. (2015)
identified heat-tolerant genotypes based on relative cell injury
percentage (RCI %), heat susceptible index (HSI) value, and
boll retention percentage on individual genotypes. These are
simple, reliable, and inexpensive for screening methods to heat
tolerance. Development of heat resistance needs more attention
in comparison to other abiotic stresses. The information on
heat tolerance in cotton is scanty in the literature. Even in the
presence of all of complications in the mechanisms of heat resis-
tance, several heat-resistant inbred lines and hybrid varieties
of tomato have been improved and released for commercial

cultivation (Scott et al. 1995, 1996).
1.1.4 Modern Molecular Breeding Approaches

Conventional breeding is tedious, time-consuming, and depen-
dent on the environment as compared to molecular breeding,
particularly marker-assisted selection, which is highly efficient
and precise. Biotechnology has contributed significantly in
understanding and improving for high temperature tolerance in
crop plants. Various approaches – namely molecular markers,
correlation, co-segregation, and genetic variability – were being
used to understand genetic bases of thermotolerance (Maestri
et al. 2002). In order to cope with heat stress, plants use various
mechanisms, including maintenance of membrane stability,
scavenging of reactive oxygen species (ROS), production of
antioxidants, accumulation and adjustment of compatible
solutes, induction of mitogen-activated protein kinase (MAPK)
and CDPK cascades, and most importantly chaperone signaling
and transcriptional activation. All these mechanisms, which are
regulated at the molecular level, enable plants to thrive under
heat stress. Based on understanding such mechanisms, potential
genetic strategies to improve plant heat-stress tolerance include
traditional and contemporary molecular breeding protocols and
transgenic approaches (Wahid et al. 2007). With the advent of
DNA marker technique in the 1980s (Botstein et al. 1980) and
early 1990s, lot of restrictions linked to isozyme and morpho-
logical markers were conquered, and genetic mapping entered
a new and exciting era which promised to greatly enhance
the efficiency of crop breeding and genetics study. Hall (2001)
suggests that genetic engineering is a useful tool for ideotype
breeding, because a single gene can be inserted without chang-
ing the background of cultivars, and successful stories abound
in the literature concerning the development of isogenic lines
in a short period. In isogenic lines, differential expression of
an inserted gene can be evaluated in a range of environments
very precisely. Isogenic lines developed by traditional back-
crossing methods require several years to grow generations as
1.1 Introduction 11
compared to marker assisted selection (MAS). It is an efficient

approach for the improvement of a stress-tolerant crop. This
approach requires the availability of genetic markers linked with
stress-tolerant genes. Genetic markers associated in different
environmental stresses such as drought, heat, and salinity have
been identified in different crop species (Foolad 2005). For
example, four quantitative trait loci (QTLs) for heat tolerance
were identified in A. thaliana (Hong and Vierling 2000). Lin
et al. (1996) detected 14 random amplified polymorphic DNA
(RAPD) markers associated with heat-resistant characteristics
in tomatoes under heat stress with the use of the bulked seg-
regant analysis. In reaction to high temperatures, special HSPs
have been distinguished in various crop species. Experimental
data gained from transgenic, reverse-genetics, and mutation
attributes in noncereal species prove causal involvement of HSPs
in thermoresistance in crops (Queitsch et al. 2000). For instance,
HSP68, that is located in mitochondria and usually expressed
incorporately, was detected to have enhanced expression under
heat stress in barley, tomato, maize, potato, and soybean cells
(Newman et al. 1993). Likewise genotyping by sequencing (GBS)
is also a cost-effective method to detect and score thousands
of genome-wide single nucleotide polymorphism (SNP) from
diverse populations of crop plants (Elshire et al. 2011; Poland and
Rife 2012). In the same way, 11 QTLs were identified in maize by
using genetic marker RFLP (restriction fragment length poly-
morphism) responsible for the growth of pollen tube and pollen
germination in heat-stress conditions (Frova and Sari-Gorla
1994). Microarray technology has recently become a powerful
tool for the systematic analysis of expression (or transcriptome)
profiles of large numbers of genes that are induced or repressed
by heat treatment. An initial study in N. tabacum examined the
transcripts of 170 cDNAs from plants drought-stressed with or
without simultaneous heat stress. In current scenarios, several
techniques are available to plant scientists that could be utilized
for the development of new germplasm that could cope with
stressful environments, and ensure food security, because many
changes have already been made to the behavior of crops plants
which lead to reduced yields.
1.2 Conclusion and Future Prospects

Heat stress has become a major and wide problem for crop
production because it affects the development, growth, and
productivity of crop plants. However, the degree of occurrence
of heat stress varies greatly among climatic zones and depends
on the period and probability of high temperature and on the
timing of diurnal plants during high temperature. The global
ambient temperature is increasing day by day, owing to rapid
and increased emission of greenhouse gases including NO2
and CO2 from different industries and automobile sources
resulting in global warming. Therefore, plant adaptations and
responses against enhanced temperatures and the mechanisms
which are responsible for the identification or development of
heat-tolerant cultivars must be well understood for important
agricultural crops. Plant responses to high temperatures also
vary across and within species, as well as at different develop-
mental stages. Hence the development of more heat-tolerant
cotton has been a primary focus for breeders and geneticists,
and researchers examining heat-stress-related effects with
chemical ameliorants. Development of heat-tolerant cotton
is limited in that the domestication of wild type cotton into
its current lineages has reduced the genetic diversity for
thermotolerance.
References
Abro, S., Rajput, M.T., Khan, M.A. et al. (2015). Screening of cotton
(Gossypium hirsutum L.) genotypes for heat tolerance. Pakistan
Journal of Botany 47 (6): 2085–2091.
Alarcon, V.J. and Sassenrath, G.F. (2015). Optimizing canopy
photosynthetic rate through PAR modeling in cotton
(Gossypium spp.) crops. Computers and Electronics in
Agriculture 119: 142–152.
Asha, R. and Lal Ahamed, M. (2013). Screening of cotton
genotypes for heat tolerance. International Journal of
Bio-resource and Stress Management 4: 599–604.
Ashraf, M., Saeed, M.M., and Qureshi, M.J. (1994). Tolerance to
high temperature in cotton (Gossypium hirsutum L.) at initial
References 13
growth stages. Environmental and Experimental Botany 34 (3):

275–283.
Azhar, F.M., Ali, Z., Akhtar, M.M. et al. (2009). Genetic variability
of heat tolerance, and its effect on yield and fibre quality traits in
upland cotton (Gossypium hirsutum L.). Plant Breeding 128:
356–362.
Baloch, M.J., Lakho, A.R., Bhutto, H.U. et al. (2000). Estimates of
combining ability and genetic parameters for yield and fiber
traits in upland cotton. Pakistan Journal of Biological Sciences
3: 1183–1186.
Barabaschi, D., Tondelli, A., Desiderio, F. et al. (2016). Next
generation breeding. Plant Science 242: 3–13.
Bibi, A.C., Oosterhuis, D.M., and Gonias, E.D. (2008).
Photosynthesis, quantum yield of photosystem II and
membrane leakage as affected by high temperatures in cotton
genotypes. Journal of Cotton Science 12: 150–159.
Blum, A., Klueva, N., and Nguyen, H.T. (2001). Wheat cellular
thermo-tolerance is related to yield under heat stress. Euphytica
117: 117–123.
Botstein, D., R.L. White, M. Skolnick and R.W. Davis. 1980.
Construction of a genetic linkage map in man using restriction
fragment length polymorphisms. American journal of human
genetics. 32:314.
Bowen, J., Lay-Yee, M., Plummer, K.I.M., and Ferguson, I.A.N.
(2002). The heat shock response is involved in thermotolerance
in suspension-cultured apple fruit cells. Journal of Plant
Physiology 159: 599–606.
Brown, R.S. and Oosterhuis, D.M. (2004). High daytime
temperature stress effects on the physiology of modern versus
obsolete cotton cultivars. Summaries of Cotton Research: 63–67.
Burke, J.J. (1990). Variation among species in the temperature
dependence of the reappearance of variable fluorescence
following illumination. Plant Physiology 93: 652–656.
Burke, J.J. and Chen, J. (2015). Enhancement of reproductive heat
tolerance in plants. PLoS One 10: e0122933.
Burke, J.J. and Upchurch, D.R. (1995). Cotton rooting patterns in
relation to soil temperatures and the thermal kinetic window.
Agronomy Journal 87: 1210–1216.
Burke, J.J., Hatfield, J.L., Klein, R.R., and Mullet, J.E. (1985).
Accumulation of heat shock proteins in field-grown cotton.
Plant Physiology. 78: 394–398.
Burke, J.J., Velten, J., and Oliver, M.J. (2004). In vitro analysis of
cotton pollen germination. Agronomy Journal 96: 359–368.
Clark, R.B. and Reinhard, N. (1991). Effects of soil temperature on
root and shoot growth traits and iron deficiency chlorosis in
sorghum genotypes grown on a low iron calcareous soil. Plant
and Soil 130: 97–103.
Clement, J.D., Constable, G.A., and Conaty, W.C. (2013). CO2
exchange rate in cotton does not explain negative associations
between lint yield and fiber quality. Journal of Cotton Science
17: 270–278.
Coca, M.A., Almoguera, C., Thomas, T.L., and Jordano, J. (1996).
Differential regulation of small heat-shock genes in plants:
analysis of a water-stress-inducible and developmentally
activated sunflower promoter. Plant Molecular Biology 31:
863–876.
Conaty, W.C., Burke, J.J., Mahan, J.R. et al. (2012). Determining the
optimum plant temperature of cotton physiology and yield to
improve plant-based irrigation scheduling. Crop Science 52:
1828–1836.
Conaty, W.C., Mahan, J.R., Neilsen, J.E. et al. (2015). The
relationship between cotton canopy temperature and yield, fibre
quality and water-use efficiency. Field Crops Research 183:
329–341.
Coria, N.A., Sarquís, J.I., Peñalosa, I., and Urzúa, M. (1998).
Heat-induced damage in potato (Solanum tuberosum) tubers:
membrane stability, tissue viability, and accumulation of
glycoalkaloids. Journal of Agricultural and Food Chemistry
46: 4524–4528.
Cornish, K., Radin, J.W., Turcotte, E.L. et al. (1991). Enhanced
photosynthesis and stomatal conductance of Pima cotton
(Gossypium barbadense L.) bred for increased yield. Plant
Physiology 97: 484–489.
Cothren, J.T. (ed.) (1999). Physiology of the cotton plant. In:
Cotton: Origin, History, Technology, and Production, 207–268.
Wiley Blackwell.
Cottee, N.S., Wilson, I.W., Tan, D.K., and Bange, M.P. (2014).
Understanding the molecular events underpinning cultivar
differences in the physiological performance and heat tolerance
of cotton (Gossypium hirsutum). Functional Plant Biology
41: 56–67.
References 15
Crafts-Brander, S.J. and Salvucci, M.E. (2002). Sensitivity of

photosynthesis in a C4 plant, maize, to heat stress. Plant
Physiology 129: 1773–1780.
Dai, J., Li, W., Tang, W. et al. (2015). Manipulation of dry matter
accumulation and partitioning with plant density in relation to
yield stability of cotton under intensive management. Field
Crops Research 180: 207–215.
Demirel, U., Gür, A., Can, N., and Memon, A.R. (2014).
Identification of heat responsive genes in cotton. Biologia
Plantarum 58: 515–523.
Demİrel, U., Çopur, O., and Gür, A. (2016). Early-stage screening
for heat tolerance in cotton. Plant Breeding 135: 80–89.
DeRidder, B.P. and Salvucci, M.E. (2007). Modulation of Rubisco
activase gene expression during heat stress in cotton
(Gossypium hirsutum L.) involves post-transcriptional
mechanisms. Plant Science 172: 246–254.
Dong, K. and Liu, Y. (2010). Cross-cultural management in China.
Cross Cultural Management: An International Journal 17:
223–243.
Dong, H., Li, W., Eneji, A.E., and Zhang, D. (2012). Nitrogen rate
and plant density effects on yield and late-season leaf
senescence of cotton raised on a saline field. Field Crops
Research 126: 137–144.
Driedonks, N., Rieu, I., and Vriezen, W.H. (2016). Breeding for
plant heat tolerance at vegetative and reproductive stages.
Plant Reproduction 29: 67–79.
Echer, F.R., Oosterhuis, D.M., Loka, D.A., and Rosolem, C.A.
(2014). High night temperatures during the floral bud stage
increase the abscission of reproductive structures in cotton.
Journal of Agronomy and Crop Science 200: 191–198.
Ehlers, J.D. and Hall, A.E. (1998). Heat tolerance of contrasting
cowpea lines in short and long days. Field Crops Research
55: 11–21.
Elshire, R.J., Glaubitz, J.C., Sun, Q. et al. (2011). A robust, simple
genotyping-by-sequencing (GBS) approach for high diversity
species. PLoS One 6: 19–79.
Foolad, M.R. (2005). Breeding for abiotic stress tolerances in
tomato. In: Abiotic Stresses: Plant Resistance Through Breeding
and Molecular Approaches (ed. P. Harris), 613–684. New York:
The Haworth press Inc.
Frova, C. and Sari-Gorla, M. (1994). Quantitative trait loci (QTLs)

for pollen thermotolerance detected in maize. Molecular and
General Genetics 245: 424–430.
Fryxell, P.A. (1992). A revised taxonomic interpretation of
Gossypium L. (Malvaceae). Rheedea 2: 108–116.
Golam, F., Prodhan, Z.H., Nezhadahmadi, A., and Rahman, M.
(2012). Heat tolerance in tomato. Life Science Journal
9: 1936–1950.
Greer, D.H. and Weedon, M.M. (2013). The impact of high
temperatures on Vitis vinifera cv. Semillon grapevine
performance and berry ripening. Frontiers in Plant
Science: 4.
Grzesiak, S., Grzesiak, M.T., Filek, W., and Stabryła, J. (2003).
Evaluation of physiological screening tests for breeding drought
resistant triticale (x Triticosecale wittmack). Acta Physiologiae
Plantarum 25: 29–37.
Guinn, G. (1976). Water deficit and ethylene evolution by young
cotton bolls. Plant Physiology 57: 403–405.
Hake, K. and Silvertooth, J. (1990). High temperature effects on
cotton. Cotton Physiology Today Newsletter 1: 1–4.
Hall, A.E. 2001. Heat stress and its impact. http://www.plantstress
.com/Articles/heat_i/heat_i.htm, accessed 8th July 2019.
Hall, A.E. (2004). Breeding for adaptation to drought and heat in
cowpea. European Journal of Agronomy. 21: 447–454.
Hall, A.E. and Allen, L.H. (1993). Designing cultivars for the
climatic conditions of the next century. International Journal
of Crop Science: 291–297.
Hasanuzzaman, M., Nahar, K., Alam, M.M. et al. (2013).
Physiological, biochemical, and molecular mechanisms of heat
stress tolerance in plants. International Journal of Molecular
Sciences 14: 9643–9684.
Hatfield, J.L. and Burke, J.J. (1991). Energy exchange and leaf
temperature behavior of three plant species. Environmental and
Experimental Botany 31: 295–302.
Hatfield, J.L., Quisenberry, J.E., and Dilbeck, R.E. (1987). Use of
canopy temperatures to identify water conservation in cotton
germplasm. Crop Science 27: 269–273.
Havaux, I. (1993). Rapid photosynthetic adaptation to heat stress
triggered in potato leaves by moderately elevated temperatures.
Plant, Cell & Environment 16: 461–467.
References 17
Hofmann, W.C., Kittock, D.L., and Alemayehu, M. (1988). Planting

seed density in relation to cotton emergence and yield.
Agronomy Journal 80: 834–836.
Hong, S.W. and Vierling, E. (2000). Mutants of Arabidopsis
thaliana defective in the acquisition of tolerance to high
temperature stress. Proceedings of the National Academy of
Sciences 97: 4392–4397.
Jamieson, P.D., Brooking, I.R., Porter, J.R., and Wilson, D.R. (1995).
Prediction of leaf appearance in wheat: a question of
temperature. Field Crops Research 41: 35–44.
Joshi, A.K. (1999). Genetic factors affecting abiotic stress
tolerance in crop plants. In: Handbook of Plant and Crop Stress
(ed. M. Pessarakli), 795–826. CRC Press.
Karademir, E., Karademir, Ç., Ekinci, R. et al. (2012). Screening
cotton varieties (Gossypium hirsutum L.) for heat tolerance
under field conditions. African Journal of Agricultural Research
7: 6335–6342.
Karim, M.A., Fracheboud, Y., and Stamp, P. (2000). Effect of high
temperature on seedling growth and photosynthesis of tropical
maize genotypes. Journal of Agronomy and Crop Science
184: 217–223.
Lather, B.P.S., Saini, M.L., and Punia, M.S. (2001). Hybrid cotton
retrospect and prospects in Indian context. National Journal of
Plant Improvement 3: 61–68.
Lin, J.J., Kuo, J., Ma, J. et al. (1996). Identification of molecular
markers in soybean comparing RFLP, RAPD and AFLP DNA
mapping techniques. Plant Molecular Biology Reporter
14: 156–169.
Liu, Z., Yuan, Y.L., Liu, S.Q. et al. (2006). Screening for
high-temperature tolerant cotton cultivars by testing in vitro
pollen germination, pollen tube growth and boll retention.
Journal of Integrative Plant Biology 48: 706–714.
Lu, Z., Percy, R.G., Qualset, C.O., and Zeiger, E. (1998). Stomatal
conductance predicts yields in irrigated Pima cotton and bread
wheat grown at high temperatures. Journal of Experimental
Botany: 453–460.
Maavimani, M., Jebaraj, S., Raveendran, M. et al. (2014). Cellular
membrane thermostability is related to rice (Oryza sativa L.)
yield under heat stress. International Journal of Tropical
Agriculture 32: 201–208.
Maestri, E., Klueva, N., Perrotta, C. et al. (2002). Molecular

genetics of heat tolerance and heat shock proteins in cereals.
Plant Molecular Biology 48: 667–681.
Mahan, J.R., McMichael, B.L., and Wanjura, D.F. (1995). Methods
for reducing the adverse effects of temperature stress on plants:
a review. Environmental and Experimental Botany 35:
251–258.
Mahan, J.R., Young, A., Payton, P. et al. (2014). Effect of differential
irrigation on accumulation of canopy temperature-based heat
units in cotton. Journal of Cotton Science 18: 129–136.
Mao, L., Zhang, L., Zhao, X. et al. (2014). Crop growth, light
utilization and yield of relay intercropped cotton as affected by
plant density and a plant growth regulator. Field Crops Research
155: 67–76.
Marcum, K.B. (1998). Cell membrane thermostability and
whole-plant heat tolerance of Kentucky bluegrass. Crop Science
38: 1214–1218.
Marsh, J.L., Walker, H., Theisen, H. et al. (2000). Expanded
polyglutamine peptides alone are intrinsically cytotoxic and
cause neurodegeneration in Drosophila. Human Molecular
Genetics 9: 13–25.
Martineau, J.R., Specht, J.E., Williams, J.H., and Sullivan, C.Y.
(1979). Temperature tolerance in soybeans: evaluation of a
technique for assessing cellular membrane thermostability.
Crop Science 19: 75–78.
Mauney, J.R., Kimball, B.A., Pinter, P.J. et al. (1994). Growth and
yield of cotton in response to a free-air carbon dioxide
enrichment (FACE) environment. Agricultural and Forest
Meteorology 70: 49–67.
McMichael, B.L. and Burke, J.J. (1994). Metabolic activity of
cotton roots in response to temperature. Environmental and
Experimental Botany 34: 201–206.
Mickelbart, M.V., Hasegawa, P.M., and Bailey-Serres, J. (2015).
Genetic mechanisms of abiotic stress tolerance that translate
to crop yield stability. Nature Reviews Genetics 16: 237.
Mills, A.J., Cowling, R.M., Fey, M.V. et al. (2005). Effects of goat
pastoralism on ecosystem carbon storage in semiarid thicket,
Eastern Cape, South Africa. Austral Ecology 30: 797–804.
Monteith, J.L. (1995). A reinterpretation of stomatal responses
to humidity. Plant, Cell & Environment 18: 357–364.
References 19
Morales, D., Rodriguez, P., Dell’Amico, J. et al. (2003).

High-temperature preconditioning and thermal shock
imposition affects water relations, gas exchange and root
hydraulic conductivity in tomato. Biologia Plantarum 47: 203.
Nakamoto, H. and Hiyama, T. (1999). Heat-shock proteins and
temperature stress. In: Handbook of Plant and Crop Stress
(ed. M. Pessarakli), 399–416. New York: Marcel Dekker.
Newman, T.C., Ohme-Takagi, M., Taylor, C.B., and Green, P.J.
(1993). DST sequences, highly conserved among plant SAUR
genes, target reporter transcripts for rapid decay in tobacco.
Plant Cell 5: 701–714.
Nielsen, K.F. (1974). Roots and root temperatures. Proceedings of
Plant Root Environment: 293–333.
Niinemets, Ü. (2010). A review of light interception in plant stands
from leaf to canopy in different plant functional types and in
species with varying shade tolerance. Ecological Research 25:
693–714.
Paulsen, G.M. (1994). High temperature responses of crop plants.
In: Physiology and Determination of Crop Yield,
(physiologyandde) (eds. K.J. Boote, J.M. Bennett, T.R. Sinclair
and G.M. Paulsen), 365–389.
Pettigrew, W.T. and Meredith, W.R. (1994). Leaf gas exchange
parameters vary among cotton genotypes. Crop Science
34: 700–705.
Pettigrew, W.T. and Meredith, W.R. Jr. (2012). Genotypic variation
in physiological strategies for attaining cotton lint yield
production. Journal of Cotton Science 16: 179–189.
Pettigrew, W.T. and Turley, R.B. (1998). Variation in
photosynthetic components among photosynthetically diverse
cotton genotypes. Photosynthesis Research 56: 15–25.
Pettigrew, W.T., Heitholt, J.J., and Vaughn, K.C. (1993).
Gas exchange differences and comparative anatomy among
cotton leaf-type isolines. Crop Science 33: 1295–1299.
Poland, J.A. and Rife, T.W. (2012). Genotyping-by-sequencing for
plant breeding and genetics. The Plant Genome 5: 92–102.
Queitsch, C., Hong, S.W., Vierling, E., and Lindquist, S. (2000).
Heat shock protein 101 plays a crucial role in thermotolerance
in Arabidopsis. Plant Cell 12: 479–492.
Quisenberry, J.E., McDonald, L.D., and McMichael, B.L. (1994).
Responses of photosynthetic rates to genotypic differences in
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CHAPTER XXV.
THE DISTURBING ELEMENT.
"We must have a Baby Show before the summer is over," said Mrs. Greville,
as she sat in the garden by Rachel's side darning stockings, "and next year a
Flower Show. The village has been dreadfully neglected, evidently, by the last
Rector."
"I don't think so," said Rachel. "He was a splendid visitor and was looked upon
as a father by the people."
"Perhaps; but there is nothing social by which to draw the people together.
Now a Women's Institute ought to be started."
"But you must remember there is not a single helper in the parish with the
exception of a very nice farmer's daughter, who conducts the whole Sunday
School. It would be impossible to start things here that need helpers."
But Mrs. Greville was not to be put off.
"I expect if helpers were really looked for they would be found. I'll look round
the parish and see what I can do."
Rachel sighed.
"You won't make the people want things that they can't have, will you?" she
said.
"Certainly not. But there is no need to say they can't have them. Anyhow I am
here to help; and anything in the way of writing I am sure you could do. There
are two workers at once."
"Oh, you must not think of me as a worker," said Rachel. "I can only just live."
She had been more conscious of her weakness since her mother-in-law's
advent than ever before.
"My dear, of course you cannot be a worker in the ordinary sense. But believe
me you would feel all the better with something to interest you a little. It never
does anyone any good to lie and think about themselves all day. I found that
out when I was ill; and no sooner did I begin to think of my work than I felt
better at once."
Rachel was silent. She wondered how much longer she could endure the
intense activity of her mother-in-law. She thought she would have to take to
her bed; but she knew the air was the only thing for her.
"Let me see," continued Mrs. Greville. "There's a Mrs. Simpkins at the bottom
of the hill who has a fine boy, and next door is that poor thriftless woman
whose baby is nothing but skin and bones. It will do the mothers a world of
good having to weigh their babies; they'd be ashamed then of their little puny
creatures. We must certainly have the Show before the summer is over. When
shall it be?"
"Whenever you like," said Rachel wearily. Then she added, "Ask Luke."
"Oh it won't make any difference to Luke. It is a question for his mother and
wife to decide. Shall we say at the end of August?"
"Luke has a clerical meeting here one day in August. You had better ask him."
"Will the clergy be here to tea?"
"Yes, and they come to lunch too."
"Come to lunch! How ridiculous! In these days to give lunch is quite

unnecessary I am sure. Why it can't be done under—"
"Oh don't say that to Luke," interrupted Rachel. "It's such a delight to him to be
able at last to offer hospitality to his brother clergy. He is quite looking forward
to it."
"Well I call it very unnecessary. What are they going to have? I hope you are
not preparing a spread. Because you have a rather better living than your last
it does not mean that you must forget the necessity of being economical."
Rachel laughed hysterically. Here it was again! Eggs at 4d and margarine

instead of butter!
"Don't let us talk of economy yet," she said. "Let us enjoy, anyhow for a time,
being able to give something to one's neighbours."
"And pray who is going to do the cooking?"

"Emma. In her last place she had to cook for clerical lunches and knows just
what is necessary. And," she added laughing, "I mean to give them a good
lunch."
"What do you mean by a good lunch?"
Mrs. Greville sarcastically. "Do you intend to give them chickens? They are
nine shillings each at Trowsby."
"No I don't mean to give chickens. They are probably tired of them in the
country. But they shall have a good lunch, I am determined about that, for
Luke's sake."
"My dear you are talking like a very silly woman;" said Mrs. Greville gravely.
"Am I? I am sorry, but for once in my life I mean to be silly. So you must
forgive me," said Rachel.
Mrs. Greville said no more, but came to the conclusion that Rachel was a little
light-headed, particularly as the conversation had set her laughing weakly.
She supposed her presence was too exciting for her, so in a few minutes she
picked up her work and made for the house, and before long Rachel caught
sight of her making her way towards the gate with a note book and pencil in
hand.
"She is going to look after the babies," sighed Rachel. "I hope she won't make
those poor mothers too miserable by her comments." Then her mind reverted
to one of Mrs. Greville's remarks.
"I wonder," she thought, "if she really imagines that I lie and think of myself all
day! A woman with a husband and child, not to mention a mother-in-law and
the housekeeping, has scarcely time to lead an idle life even in thought. Little
Pat's clothes would be enough to keep me busy even if I had nothing else to
do." And Rachel, who had laid down her work while talking to Mrs. Greville,
took it up again. It was not long before she heard the gate open and her
mother-in-law's somewhat heavy tread on the gravel.
"You are back sooner than I expected," said Rachel. Mrs. Greville looked tired
and dispirited.
"Yes. I found my suggestion of a Baby Show rather upset the women. They
seemed to take offence and were up in arms at once. That thriftless creature
with the puny baby was almost rude to me, so I felt it would be as well to put
the show off for a time."
Rachel sighed a sigh of relief.
"I'm sorry that you have had your walk for nothing," she said, "but on the other
hand it would be such a pity to create an ill feeling. Besides I would so much
rather that our first gathering should be a tea in the garden. They would get to
know us then and Luke could talk to them. I am bent, when I feel a little
stronger, on inviting the whole parish."
"Another expense," said Mrs. Greville.
"I know. But it need not be an annual affair; just a kind of house warming."
"I wonder how much it would cost."
Rachel, who had been anticipating managing the tea herself directly she felt
strong enough, had noticed the disappointed expression on Mrs. Greville's
face when her plan of the baby show had failed; she evidently felt a little
depressed. Should she put the whole management of the tea into her mother-
in-law's hands? She knew Mrs. Greville would not be happy unless she had
plenty to do, In fact she was so restless that she would make work if it were
not provided for her. It would be a disappointment, but anything was better
than to have an unhappy, restless, person in the house.
When Rachel asked Mrs. Greville if she would undertake the tea for her she
cheered up at once. Taking out her pocket book she put down the probable
number that they might expect, and began to discuss at once how much
bread would be wanted, and what the whole affair would cost.
"I shall give them margarine," she said. "It would be quite absurd to go to the
expense of butter." Rachel would have given them butter but she knew it
would be useless to suggest this.
"And the tea might be sugared also altogether. It is much easier to have it all
ready mixed with the milk."
"But some may not like sugar in their tea," objected Rachel.
"Well they mustn't mind it just for once," answered Mrs. Greville. "It is by far
the best way when you have a large party. Then I almost think the cake and
bread and butter had better be divided up before they come, and put on their
plates. They will all fare alike then, and they'll know exactly how much they
can have. If they don't want to eat it all they can take it away with them."
"But they are not by any means all children. I want them to be treated just as
we should treat our own friends," Rachel expostulated. She began to wish she
had never asked her mother-in-law to undertake the tea.
"But," said Mrs. Greville, "you see you have not had much experience in these
matters and have no idea how much easier it is if you arrange everything
beforehand. Method is everything on these occasions."
"But I want to make it really nice for the people," said Rachel eagerly. "It's
better not to have the tea at all than in any way to hurt their feelings. I don't
think for instance the men would at all like the plan of the food being piled on
to a plate. Besides I want them to have sandwiches, and they would not be
nice all mixed up with cake. I do hope you don't mind," she added, seeing that
Mrs. Greville, who was considered such a first-rate caterer in Trowsby, was a
little flushed at the thought that her suggestions were not liked. "I fancy that
people in a country village would not be used to that kind of thing; besides
there will not be very many of them. I do want them to have a dainty tea."
"Dainty? How do you mean?" said Mrs. Greville. She was evidently put out.
"I mean the same kind of tea that we would give to our own personal friends.
Of course, rather thicker sandwiches as they would not appreciate the thin
ones, but I am thinking more of the way it is all arranged."
"Well," said Mrs. Greville shortly, "if you will tell me exactly how you wish
things to be done I will follow your directions."
Rachel wished that the subject of the tea had never been mentioned. The
slight discussion had tired and rather worried her.
"Shall we talk about it another day?" she said, "there is plenty of time, and,"
she added, "of course you may be right. I have had no experience in such
things and you have had so much."
Mrs. Greville was slightly mollified. She was not used to having her plans
criticised. She preferred criticising the plans of others.
"Well my dear," she said, "I don't think you need worry about it. I promise you
that your guests will have enough to eat; and after all that is the main thing."
Rachel did not think so. She thought that the manner in which the food was
presented to them was as important as the food itself; but she did not say so.
"You can leave it all in my hands perfectly happily," added Mrs. Greville. "I will
take the whole responsibility of it so don't worry yourself. Why, you look quite
flushed even at the thought of it. There is really no need to be anxious," and
Mrs. Greville moved towards the house notebook and pencil in hand.
Rachel sighed.
CHAPTER XXVI.
LUKE TELLS RACHEL ABOUT HIS DREAM.
"What is Luke doing all day in his study? That sedentary life is very bad for
him and he ought to be visiting his people."
Rachel, who was trying on a little frock for Pat which she had been making,
looked up amused.
"He has visited every one of his people a dozen times since he came here. In
fact he was afraid of over visiting them."
"But then what is he doing all day? He is young and strong and ought to be at
work," said Mrs. Greville, standing in front of Rachel as if she was in some
way to blame. Her face was anxious and perplexed. She could scarcely
understand work without movement, and rush.
"You may be quite sure he is not wasting his time," said Rachel; then hoping
to distract her mother-in-law's thought she held up the baby to be admired.
"Doesn't he look sweet," she said. "He shall wear this little frock when we
have the villagers to tea. He must look his best."
But once Mrs. Greville was on the war path nothing would move her.
"It isn't as if he had an intellectual congregation to preach to as he had at

Trowsby," she said. "Of course then he had to study and fit himself for the
work. But here they are only uneducated people. I am sure he cannot want all
this time to prepare for them."
"But don't you know that a man like Luke must find it much more difficult to
speak to such people? He has to make everything so plain and to choose the
simplest subjects. I am quite sure he finds it much harder to prepare his
sermons here than he did at Trowsby."
"I don't understand it at all," said Mrs. Greville, moving away to the window.
"And why he should object to me going into his study, I can't imagine. He does
not say in so many words that he prefers my absence to my presence, but I
see it worries him. Of course I used to leave him quite alone at Trowsby, but
then I knew every moment was precious. But here there ought to be plenty of
time for a chat with his mother now and then."
"I must make the sleeves a little shorter," said Rachel. "Don't you think so?"
She hoped that Mrs. Greville would feel interest in the baby's frock and would
not continue the conversation which was getting embarrassing. But no. It was
of no use trying to turn her thoughts.
"I think it's very sad to see a young man getting so idle and losing all interest
in his people," she said. "It's quite different to what it was at Trowsby. Of
course, he worked far too hard before his marriage and, indeed, after it, too,"
she added rather grudgingly. "But I would far rather he worked too much, than
too little."
Rachel's colour rose. She felt indignant.
"But he is working very hard," she said. "He is studying; and you know the
complaint is that the clergy don't study and so can't help their people in these
difficult times. Visiting their parishioners is only part of the work after all."
"Well, all I can say is that things are very different to what they were. And I
never remember a time in which he did not welcome his mother." Mrs. Greville
did not recollect how carefully she kept watch over the dining-room, which
then stood for his study, so that he should never be interrupted. "Of course,"
she added, "I don't suppose I can quite expect things to be exactly the same
as they were before his marriage. I don't imagine he objects to you going into
his study at any time."
Rachel, glancing at her mother-in-law, was aware of a certain pathetic

eagerness displayed in the expression of her face as she waited for the
answer. The slight indignation that Rachel was conscious of feeling passed
away, and she was glad that she could assure Mrs. Greville that she never
interrupted her husband unless it was absolutely necessary as she knew he
would not like it.
"But," she said, "don't you think that he may have been sent here so as to give
him time to study before some great and important post is offered him? That is
what I love to think. I could not bear to believe that he would be here all his
life. When I am stronger, I hope we shall go to some town where he can have
a larger sphere of work. That is one of the things that reconciles me to this
country living."
"Well," said Mrs. Greville, "I feel that he ought to be doing something, and not
spending all those hours in his study."
"He is doing something."
But Mrs. Greville was only half convinced.
Rachel was feeling stronger so that the conversation with her mother-in-law
did not trouble her as it might have done. It almost amused her.
That Luke could possibly be wasting his time in his study was so absurd an
idea that it made her smile. She had sometimes wondered herself at the
amount of writing he was engaged in doing; but she had not asked him about
it, as one of his peculiarities was the fact that he did not care to be questioned
about his work. That she would know in time she had not a doubt. That he
was doing anything that she did not know about did not make her either
curious or restless. Apparently it had that effect on her mother-in-law.
Rachel had noticed that since coming to live with them Mrs. Greville had
shown a propensity for wishing to know exactly what everyone in the house
was doing. She wandered from room to room just to see what was going on.
The servants were not the only people who found it trying. Rachel felt it
getting on her own nerves. She put it down to a restlessness arising from Mrs.
Greville's illness, but this did not make it less trying. She could seldom enjoy
little Pat for long by himself. Mrs. Greville, under the impression that she was
helping, would come into the nursery while Rachel bathed the child, giving her
various hints as to the best way of doing it. Or she would take up a great deal
of her time giving out her theories on the bringing up of children, while Rachel
was busy putting him to bed. It was just as much as Rachel could do in her
present state of health to dress and undress the child, and at times she had to
pass over what was to her such a happy duty to Polly, so that to be talked to,
and to have to listen to various theories on the training of children, was tiring
in the extreme. But she felt she must bear it without complaining as Mrs.
Greville had so little in which to occupy her time, and if she was shown that
her advice was not welcome she would grow melancholy.
But till to-day Rachel had had no idea that Luke had found it trying. His study
door was open that evening when after putting Pat to sleep she went down
stairs. At the sound of her footstep, he called her in, closing the door after her.
He was looking a little worried and distressed.
"I'm afraid I have hurt my mother," he said, "but I find it so difficult to get on
with my work while she is in the room. Three times this afternoon she has
interrupted me at the most unfortunate moment. I try not to let her know it, but
I can't make out what she wants. I fancy it's only a chat, and I can't chat about
nothing in the middle of my work."
"She has not enough to do," said Rachel, "and feels dull I expect."
"That's unfortunate," said Luke, running his fingers through his hair. "I'm sorry
for her. But I had to tell her I was doing some important work to-day and that I
was sorry to say I could not have a chat just then. I'm afraid I've hurt her. I
think," he added, "that my mother has altered a little since her illness. She
used to have such tact."
"It would help matters if you could tell her what the important business is," said
Rachel, laughing.
"I don't much want to tell her," he said. "I'm afraid she would be constantly
referring to it and I could not stand that. You see dear," he said looking down
at his wife with shining eyes, "I believe I have found my work."
"Your work?" said Rachel mystified.

"I believe my prayers are being answered and that I am being shown why I
have been sent here. Sit down. I want to tell you something." He drew a chair
up to the window for her and sat down by her side.
"I never told you that just before we left Trowsby I had a dream. It was the
most painful dream I have ever had and I can't think of it even now without a
shudder. I'd rather not tell you what it was; but it was nothing more or less
than a looking glass in which I saw myself for the first time, and I want you to
know that I believe we left Trowsby for my sake as well as for yours."
Any great emotion was bad for Rachel, and when Luke turned and looked at
her he was distressed to see how white she had become.
"Am I tiring you dearest?" he said, with concern.
"No. I'm only so very glad to hear what you say," she answered. "The thought
that you had to leave your interesting work for me, has been such a burden on
my mind."
"Well it need be a burden no longer. I believe it was for the sake of your bodily
health that we came here but also for my soul. You see, I doubt if your soul
wants the discipline that mine does. Anyhow, my spiritual life was not in a
healthy condition; I needed to take a back seat, if you understand what I
mean. I am not sure that my work was not becoming my god, and I was too
much in the lime-light. Anyhow, now I can work for God and for Him alone. I
think He has given the work that is best for me."
Rachel, in her weak state could not prevent the tears of sympathy from falling,
but she brushed them away with her hand, and Luke did not see them.
"What is the work?" she asked.
"I'm writing a book for young men. When I was in Trowsby I often felt the need
of one which faced the difficult problems of the day honestly and wisely. I am
trying to contend earnestly for the faith. Of course it means the need of much
prayer and study."
Rachel held out her hand and Luke grasped it in his own.
"You see how much better it is for me than preaching," he explained. "I don't
mean to put my name to the book, and shall never know what it has done for
people. God only will know that, so there will be no chance of conceit and
pride getting the better of me in my work. I do not see how I can forget God in
the work as I need Him every moment. I feel I am contending with the devil
and he does not leave me alone. But no-one knows but God—and now," he
added, "my wife."
Rachel had never loved her husband or admired him more than she did at that
moment. She sat and looked at him with her heart in her eyes; but she did not
speak.
"I meant to keep it a secret," he said, "till I could put the finished book into
your hands. But now you know. I have sent the first few chapters to Sharnis
and he is pleased with them and quite hopes to undertake its publication. But I
don't want anyone to know about it. Now do you think you can help me about
my mother? I don't feel as if I could be constantly asked questions about it. It's
such a sacred thing to me that I cannot talk about it. Besides, as I'm not
putting my name to the book, it is important that no-one should know that I'm
writing it."
"She would be so pleased that she would find it difficult to keep the fact to
herself."
"That's just it."
"What a pity though that she will never know," said Rachel.
"Perhaps sometime after its publication I may tell her, but not now. But can
you manage to keep her from disturbing me without hurting her? A woman
can generally do this kind of thing better than a man. I would not hurt my dear
mother for the world if I could help it."
They neither of them heard the door open and then softly close again, and
Luke never knew that his mother had turned away from his study murmuring:
"Luke was talking to Rachel about me! His mother! I never thought he would
talk to anyone about his mother." The pain was almost too much to bear.
CHAPTER XXVII.
RACHEL'S GARDEN PARTY.
Rachel noticed a marked difference in her mother-in-law's behaviour to her

after finding her talking with Luke in his study. She was perplexed as to its
cause and it worried her. Moreover, she found that her efforts to keep her out
of the study during Luke's busy time there were unavailing, and she came to
the conclusion that it would hurt her feelings less if Luke mentioned the
subject to her himself. Her pride would not be wounded. She told Luke so.
But, manlike, he felt that to burn Pussy's paws was by far the better plan. But
Rachel, unwilling to give him a hint as to his mother's strained feelings in
respect to herself, determined to do nothing in the matter unless she found a
particularly favourable moment in which to speak. Having suffered herself in
the past from jealousy she began to suspect that the same wretched enemy to
all happiness was attacking her mother-in-law; and instead of despising her
for giving way, she felt the situation to be particularly pathetic.
Since the day on which Luke heard of his wife's delicacy, he had been filled
with anxiety about her and was continually thinking of her comfort and good.
In fact, Rachel's health was now his first concern, and plans which would in
any way give her too much to do or think about were put on one side at once,
even though they might be suggested by his mother. His wife and his book
were at present the absorbing interests in his life and everything else fell into
the background.
Rachel had long ago come to the conclusion that Luke found it difficult to
concentrate his mind on many things at the same time. When he was at
Trowsby, his parish and Church took up all his thoughts to the exclusion of
almost everything else. In fact, he suffered from a want of seeing things in
their right proportion. And now that his mind was full of anxiety for his wife, his
mother was somewhat put on one side. It distressed Rachel, for she knew the
signs of the terrible pain caused by jealousy and felt sure she saw them
written on her mother-in-law's face. They had in fact changed places since
leaving Trowsby. Luke now consulted his wife about matters of interest instead
of his mother; and though Rachel loved that it should be so, she felt Mrs.
Greville's position to be very difficult.
Some women would have been triumphant over the reversed positions but
Rachel was above all such unworthy feelings; she could only see the pathetic
side and it distressed her. Remembering too her own fight with the sin that
brings such misery on those who give way to it, she longed to give Luke a hint
as to how matters stood, and to ask him to be very careful; but she knew Luke
would no more understand his mother's feelings than he would have
understood those of his wife in the old days; and she felt it would not be fair to
his mother to enlighten him. It would inevitably make her fall in his eyes; and
up till now, he thought her perfection.
So Rachel came to the conclusion that the only course to take was to behave
to Mrs. Greville as if nothing was the matter, and to ignore the severity of her
expression of face and voice when she spoke to her. But the anxiety and
constant effort not to say anything that might offend or hurt was not good for
her and was tiring.
As the time drew near for the parish garden party, Mrs. Greville's thoughts
became engaged with preparations for the event, and at times she seemed to
forget that she was not pleased with her daughter-in-law.
She had taken the whole affair off Rachel's hands and was thoroughly
enjoying the responsibility of it. Luke was thankful that it meant no effort for his
wife.
"If it hadn't been for my mother being here," he said, "we could not have had
it. The doctor said you were to have perfect rest."
But Rachel was disappointed, though at the same time she was thankful that
her mother-in-law had something to interest her and to fill up her time.
That it was Mrs. Greville's party was quite evident when the day dawned.
Rachel was amused when her mother-in-law told her that she had better stand
at the gate and welcome the people, as she herself would be engaged seating
them at the tables. Rachel had quite intended to be at the gate.
"But," she said, "I think it would be better not to have tea the moment they
arrive. They might walk about the garden and have games first."
But Mrs. Greville had arranged in her own mind that tea was to be the first
item of the programme and she did not wish it altered.
"It's far better to get the business of tea well over first," she said, "then we
shall be free to think of games," and Rachel gave way. She felt it was not
worth while discussing the matter.
About sixty people with their children arrived and were shown at once to their
seats by Mrs. Greville. Rachel saw them looking at the plates before them with
a smile, and heard them making jokes with one another. But after Luke had
appeared and had said a word of welcome and tea began in earnest, they
were silent. The business of eating did not seem in their estimation to need
the effort of talking. Suddenly Rachel heard Mrs. Greville say:
"Mind you all make a good tea, and what you don't want you can put in your
pockets and take home with you. I won't look!" There was a laugh and the
sound of it sent the colour into Rachel's face. The talking too began in real
earnest and did not cease till tea was over; but the words that Rachel
occasionally heard convinced her that Mrs. Greville's tactless remark was not
liked by the men; and she noticed that no-one save the woman with the puny
baby took advantage of the invitation to take the food home.
Rachel did what she could to take away the impression made by her mother-
in-law's mistake, by noticing the many babies that the mothers had brought
with them. And the subject of babies was so absorbing that when Luke took
the men off for a game of bowls, the women were quite contented to sit and
tell Rachel of the various complaints their children had suffered from.
They were not allowed however to sit still for long. Mrs. Greville bustled them
about from one part of the garden to the other dividing them into groups and
setting them to games. The mother with the puny baby alone refused to
budge. She was too tired, she explained, and Rachel, looking at her as she
talked, came to the conclusion, that probably the reason why the baby was so
thin and weird looking was because the money went in drink. She tried to gain
the mother's confidence and to help her, but the woman simply sat and cried
and complained of her hard life. One of the women who had been playing
games and was a little jealous at seeing Mrs. Grot having such a long talk with
the Rector's wife, informed her spitefully, that she was a "bad lot," and left the
baby alone in the house while she went to spend her evenings at the public
house. Rachel came to the conclusion that the village was not such a little
heaven below as she had imagined; but that the great enemy of souls was as
busy there as elsewhere.
CHAPTER XXVIII.
"THINK OF RACHEL."
Gwen came to stay at the Rectory. She was much impressed with two things.
First she was terribly upset at the look of Rachel. She had had no idea that
she had grown so thin and weak. It gave her quite a shock, and she felt
disinclined to admire or appreciate anything of her surroundings.
But she was equally struck with the alteration in Luke. She could find nothing
with which to find fault in his behaviour to Rachel. He was as thoughtful for her
as before he had been negligent.
"What have you done to Luke to change him so?" she asked her sister one
evening, as they were sitting in the garden after supper.
"I have done nothing, but you were blind."
"Then if you are not the cause of his alteration I must be," said Gwen. "I
believe it was the talk I had with him when I told him the truth about himself.
But I must confess I never expected it to have such an effect."
Rachel laughed.
"You had better not remind me of your delinquencies," she said. "I don't think I
ever felt more angry with anyone in my life than I did with you that day.
However I have forgiven you so we will say no more about it."
"One thing I can't understand," said Gwen. "What could have induced you to
have that tiresome old lady to live with you? It isn't as if she were fond of you
for I'm sure she rather dislikes you; that is to say if her manner to you means
anything. Why did you let her come?"
"I don't think she dislikes me. She has a very kind heart, but she has been ill
and finds it a little difficult to be always nice. You must not blame her."
"But I do blame her. Why did you have her?"
"I knew it would make Luke happier. You see she was all alone in rooms in
Trowsby."
"I can't see why Luke should be made happy at your expense."
"But you must remember that before I arrived on the scene she had Luke to
herself. She is devoted to him, and must have missed him dreadfully when we
left Trowsby."
"Well it seems to me that my next move will have to be to tackle her, as my

words take such effect. But look there's Luke. Why is he tearing down the
drive on his bicycle at such a rate?"
"There's a fire in the village," he shouted, and was gone before there was time
to ask questions.
Both girls rose.
"Yes, I smell the smoke," said Gwen. "It comes from the bottom of the hill. I
shall run down and see it."
Rachel stood looking in the direction of the fire and had just made up her mind
to follow Gwen slowly, when Mrs. Greville appeared.
"I saw Luke rushing off on his bicycle," she said. "What is the matter?"
"It's a fire in the village and I'm afraid it must be near those tumble down
cottages in which Mrs. Simpson and Mrs. Grot live. If the fire reaches them
they will burn like tinder."
"Well then I shall go and see if I can't help," said Mrs. Greville. "If anyone is
hurt I may be of use. I shall put a few things in my bag in readiness. I've some
old linen and some carron oil," and she hurried into the house followed by
Rachel, who made her way into the nursery and stood for a few minutes
looking down at her sleeping boy.
It was a hot evening and he had kicked off the coverlet in his sleep. Rachel
kissed the tiny feet gently, holding them in her hand tenderly. Then she
covered him up again.
"You'll take care of him I know," she said to Polly as she moved away. "I'm
going down to see if there is anything to be done in the village. I'm afraid
several cottages must be in danger."
Then as she left the room she looked back once more at her sleeping boy,
saying, "I'm so glad, Polly, that I can trust you implicitly. I know you would
never leave the house under any circumstances, whatever Emma may do."
Polly looked at her with the brightest of smiles. She knew there was no need
for words. Pat was her one thought.
Arrived at the bottom of the hill Rachel found herself in the midst of an excited
crowd, and was dismayed to see that the flames had not only reached Mrs.
Simpson's cottage but were approaching the one next to it, belonging to Mrs.
Grot.
"The Rector has sent a boy off on his bicycle for the fire engines," explained a
woman to Rachel in answer to her questions. "But I doubt if they'll come in
time."
"Is Mrs. Grot here?"
"No Ma'am, she's half a mile away at the 'Three Swans' and has the key with
her. Her baby is still in the cottage."
Rachel was horror struck.
"But somebody must save the baby," she cried. "There's time yet. Where's the
Rector?"
Before the woman could answer a cry rang through the crowd, and Mrs. Grot
rushed in among them shrieking, "My baby! my baby!"
"You just follow me," said the woman to Rachel, "and I'll find the Rector for
you. It's no good that woman 'holloring' like that. It won't mend matters; the
Rector is the one to help." As she pushed her way through the crowd Rachel
caught sight of Luke placing a ladder against the cottage wall.
"There he is Ma'am," said the woman. "He knows what to do, for the poor mite
is in that front upstairs room with the small window."
"Here's Mrs. Grot with the key," shouted a man near by. "Hold on, sir, for a
moment, and I'll unlock the door, we may be able to get in this way."
But alas it was too late. A thick cloud of smoke issued from the door and drove
the man back. A groan came from the crowd.
Rachel caught sight of Gwen rushing towards Luke and catching hold of his
arm.
"You mustn't go Luke," she cried. "You'll be killed. Think of Rachel."
At that moment Rachel saw her husband was on the first step of the ladder.
He was going to save the baby! Her heart gave an exultant leap, and then
almost stood still.
At Gwen's words Luke took a hurried glance at the crowd beneath and his
eyes rested for one moment on his wife's face. Though it was deadly white
there was something in its expression that nerved his steps. She was helping
him, not for the first time by any means, to do what was his duty, and this time
he knew it was at the risk of his life.
He looked down at Gwen as he mounted, with a smile, saying:
"I am thinking of Rachel. That is why I am going to save the baby."
In another moment he was forcing the window out, which gave way easily, and
disappeared within the room.
The silent crowd stood spellbound, but Gwen in a panic ran to the side of her
sister. Her voice alone was heard, crying:
"Why didn't you call him back, why did you let him go?"
Call him back! Rachel would not have called him back for the world. It was just
the brave thing she expected him to do, and knew he would do.
She stood calm and still while the crowd surged around her murmuring.
Suddenly they saw Luke at the window. He dropped a small bundle into the
arms of someone below. A cheer went up but was suddenly checked by the
sound of timber falling and a lurid flame soaring up into the air. There was only
one explanation to give—the floor of the room in which the baby had been
sleeping had given way, and the terror-stricken crowd knew that their Rector
must be lying buried among the ruins.
At that moment the engines arrived, but Rachel stood dazed and unconscious
of what was going on around her. She did not move or speak even when the
women gathered about her, neither did she hear their words of sympathy and
commiseration.
"You'd better go, my dear," said an old woman with tears streaming down her
face, "you'd better go. It don't do any good to him or to you to stay. If your
husband, God bless him, has gone to glory why he's with the Lord, and if not
they'll bring him safe home to you."
"To think," cried a hard-faced woman, "that he's done it to save a drunkard's
baby, that's already half starved to death. He'd have done better to leave it
alone."
The old woman turned sharply at the words.
"That baby, I take it, is as dear as the rest of us to the Lord. But come, my
dear," she said again, addressing Rachel, "go home, I beg of you. You'd be
best at home. And the Lord be with you."
But Rachel did not stir till a touch on her arm suddenly aroused her. She
looked round and met the eyes full of anguish of Luke's mother. No words
were spoken, but the silent cry found its way at once to Rachel's heart and
awoke her from her dazed condition.
"Mother!" she said softly, then taking her arm in hers she slowly moved away.
No words were spoken, as they mounted the hill. The only sounds were
Gwen's sobs as she followed behind them.
On arriving at the house they made their way automatically into Luke's study
and sat down silently together on the sofa—the two women who loved Luke—
clinging to one another, and listening for that sound of all sounds, which
strikes a chill into the bravest heart.
CHAPTER XXIX.
IN THE LIGHT OF THE MOON.
Gwen, in the drawing-room, had hidden her face in the sofa cushion. She had
not wanted to hear the dreadful steps of the men carrying in the stretcher.
She was feeling frightened and unstrung and as if she had lost Rachel. Since
that fearful moment when the floor of the cottage gave way and Luke was
buried in its ruins, her sister had not spoken a word to her. Her dazed
condition frightened Gwen; and she almost dreaded seeing her.
She knew where she probably was at this moment; for no sooner had the men
who had carried in Luke's body left, than Rachel had gone up into the room,
where they had laid him. It was now 10 o'clock and she was still there, and
there was a deathly silence in the house. Gwen opened the door and listened.
Not a sound could be heard. All was still. At last feeling it unbearable she
crept upstairs hoping to find Polly in the nursery.
Polly was sitting by Pat's little crib, crying. It was a relief to Gwen to hear her
sobs. It broke the terrible silence.
"Is Mrs. Luke still there?" she asked, glancing in the direction of the room in
which she knew they had laid him.
Polly could not speak for her tears; but nodded.
"It's ten o'clock," said Gwen. "She ought to be coming to bed. She'll be quite ill
to-morrow. Do you think you could go in and persuade her to come? Tell her I
want to speak to her. Or where is Mrs. Greville? Perhaps she would go."
"She ain't fit Miss Gwen," said Polly, "and I daren't go to her. She's in her room
and maybe in bed for all I know. I guess I'd better go myself."
Gwen stood outside the room while Polly went softly in. The very fact of the
door being opened frightened her; all her nerve had left her and she hoped
that Polly would not be long.
She was not gone a minute; she came out of the room on tiptoe, closing the
door softly behind her. She was trembling so that at first she could not speak
and had to lean against the wall for support.
"Oh what is it? What is it?" cried Gwen in a panic, turning white.

Heat Stress Tolerance in Plants Physiological Molecular and Genetic Perspectives Kumar Full Chapter

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Heat stress tolerance in plants:

physiological, molecular and genetic

Physiological, Molecular and Genetic

This edition ﬁrst published 2020

Limit of Liability/Disclaimer of Warranty

Library of Congress Cataloging-in-Publication Data

Names: Wani, Shabir Hussain, editor. | Kumar, Vinay (Biotechnologist),

Cover Design: Wiley

Set in 10/12pt WarnockPro by SPi Global, Chennai, India

List of Contributors xiii

1 Heat Tolerance in Cotton: Morphological,

2 Seed Priming as a Method to Generate

3 How Eﬀective Are Stress-associated Proteins

4 Biochemical and Molecular Markers:

5 Alteration in Carbohydrate Metabolism

6 Transcriptomics to Dissect Plant Responses

7 Proteomics as a Tool for Characterizing the

8 RNA World and Heat Stress Tolerance in

8.3 Small Interfering RNA (siRNA) 177

9 Heat Shock Proteins: Master Players for

10 The Contribution of Phytohormones in Plant

10.2.2 Nutrients 215

11 Exploring In-built Defense Mechanisms in

11.4.3 Role of Small RNAs (miRNAs) in Heat-stress

Muna A. Ali Haikou Experimental Station

Jipsi Chandra Xiongming Du

Public Health Department, National Center for Natural

Shakti Mehrotra Koushlesh Ranjan

Anil Sirohi Sandeep Kumar Verma

Bhumika Yadu Sajad Majeed Zargar

One of the prevalent environmental

9th May 2019 T. Mohapatra

About the Book

The global population is 7.3 billion and by 2050 it is expected to

deep insight of the complex transcriptional mechanism that

About the Editor

Dr. Shabir Hussain Wani is senior Assistant Professor at

Currently, he is in charge of Wheat improvement program at

Dr. Vinay Kumar is an Associate Professor at the Post-

Heat Tolerance in Cotton: Morphological,

areas (Rizhsky et al. 2004). The presence of genetic variation

1.1.1 Morphological and Physiological Traits

approaches followed by the plant under diﬀerent conditions of

1.1.1.1 Seedling and Root Growth

1.1.1.2 Stomatal Conductance

temperatures in most cotton-growing areas (Radin et al. 1994).

1.1.1.3 Cell Membrane Thermostability

1.1.1.4 Canopy Temperature

advantage to screen large number of varieties/lines (germplasm)

1.1.1.5 Chlorophyll Content

characteristics: a high harvest index, high chlorophyll content

1.1.2 Genetics and Molecular Basis of Heat Tolerance

and IAR3 were up-regulated, while CTL2, RPS14, ABCC3,

1.1.3 Conventional Breeding Approaches

of tomato have been improved and released for commercial

1.1.4 Modern Molecular Breeding Approaches

compared to marker assisted selection (MAS). It is an eﬃcient

1.2 Conclusion and Future Prospects

growth stages. Environmental and Experimental Botany 34 (3):

Crafts-Brander, S.J. and Salvucci, M.E. (2002). Sensitivity of

Frova, C. and Sari-Gorla, M. (1994). Quantitative trait loci (QTLs)

Hofmann, W.C., Kittock, D.L., and Alemayehu, M. (1988). Planting

Maestri, E., Klueva, N., Perrotta, C. et al. (2002). Molecular