Breeding Plants For Future Climates

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Chapter Title Breeding Plants for Future Climates

Copyright Year 2020
Copyright Holder Springer Nature Singapore Pte Ltd.
Corresponding Author Family Name Ali
Particle
Given Name Qasim
Suffix
Division Department of Botany
Organization/University Government College University
Address Faisalabad, Pakistan
Author Family Name Shahid
Particle
Given Name Sumreena
Suffix
Author Family Name Hussain
Particle
Given Name Abdullah Ijaz
Suffix
Division Department of Chemistry
Author Family Name Shehzad
Particle
Given Name Faisal
Suffix
Author Family Name Perveen
Particle
Given Name Rashida
Suffix
Division Department of Physics
Organization/University University of Agriculture
Author Family Name Habib
Particle
Given Name Noman
Suffix
Author Family Name Ali
Particle
Given Name Shafaqat
Suffix
Division Department of Environmental
Sciences and Engineering
Author Family Name Iqbal
Particle
Given Name Naeem
Suffix
Author Family Name Waseem
Particle
Given Name Muhammad
Suffix
Organization/University Allama Iqbal Open University
Address Islamabad, Pakistan
Author Family Name Hussain
Particle
Given Name Makhdoom
Suffix
Division Department of Zoology
Organization/University Government CollegeUniversity
Abstract At present, the erratic environmental conditions along with ever-increasing
human population have created a problem for the agricultural researchers to
fulfill the world food demand, with present rate of increase in crop production.
Increase in world mean temperature along with shortage of freshwater has
further worsened this situation. Due to increasing greenhouse effect in last few
decades, it has increased the average world temperature more than 1.5 oC that
increased the evapotranspiration and created a problem of aridity in some areas
of the world. It is estimated that till 2050, there is a need to double the world
food production with a rate of 2–5% per year but present rate of only 0.9%. It
is only possible by increasing the crop production area or by crop production
per capita. The former one is not possible in present environmental conditions
due to limited sources of freshwater. However, it is possible to achieve the
latter one through different ways, when there is a limited supply of water. It is
possible through the selection from the available germplasm that can perform
better with better production under the changing environmental conditions,
and it seems the important and foremost way to deal with the problems of
world food demand. Crop breeding for the selection of varieties with better
production under changing and stressful environmental conditions is gaining
interest. In this regard, the selection of crop varieties against different abiotic
stresses for better production seems to be the most important one. Normally, the
breeding for the selection of stress-tolerant crop varieties is based on different
agronomic traits (traits of interest) such as plant biomass production, yield
attributes, and different stress tolerance indices that are considered necessary
ones. The stress tolerance mechanism in plants is a phenomenon of different
cellular physio-biochemical attributes. However, it is a complex mechanism
because the stress tolerance in crops is multigenic and complex mechanism
and is purely under the control of genetics. In this regard, present interest and
focus of researchers in the study of quantitative trait loci (QTLs) has played
an important role for the selection and development of stress-tolerant crop
varieties in a short time. Though the fruitful success regarding the QTL-based
selection has been achieved, but due to the complexity in multigenic nature of
stress tolerance, the behavior of the crop varieties changes under the changing
environmental conditions. The present chapter is a comprehensive update
regarding selection of stress-tolerant crop varieties for better production under
the changing environmental conditions. So, in the future in view of changing
environmental scenario, it is necessary to find out or develop the crop varieties
that can perform better with better production under such conditions that will
be fruitful to fulfill the world food demand for ever-increasing world population
at present and in the near future.
Keywords (separated Environmental stresses - Crop plants - Breeding - Physiological traits -
by “ - ”) Agronomic traits - QTLs
Chapter 27 1
Breeding Plants for Future Climates 2
Qasim Ali, Sumreena Shahid, Abdullah Ijaz Hussain, Faisal Shehzad, 3

Rashida Perveen, Noman Habib, Shafaqat Ali, Naeem Iqbal, 4
Muhammad Waseem, and Makhdoom Hussain 5
Contents 6
27.1 Introduction 000 7

27.2 Breeding Against Water Stress 000 8
27.3 Breeding Against Salt Stress 000 9
27.4 Breeding Against Temperature Stress 000 10
27.5 Breeding Against Heavy Metal Stress 000 11
27.6 Quantitative Trait Locus (QTL)-Based Selection of Crop Genotypes Under 12
Different Environmental Stresses 000 13
27.7 Development of Tolerant Crop Genotypes by Manipulating QTLs 000 14
References 000 15
Abstract At present, the erratic environmental conditions along with ever- 16

increasing human population have created a problem for the agricultural researchers 17
to fulfill the world food demand, with present rate of increase in crop production. 18
Increase in world mean temperature along with shortage of freshwater has further 19
worsened this situation. Due to increasing greenhouse effect in last few decades, it 20
has increased the average world temperature more than 1.5 oC that increased the 21
evapotranspiration and created a problem of aridity in some areas of the world. It is 22
Q. Ali (*) · S. Shahid · F. Shehzad · N. Habib · N. Iqbal

Department of Botany, Government College University, Faisalabad, Pakistan
A. I. Hussain
Department of Chemistry, Government College University, Faisalabad, Pakistan
R. Perveen
Department of Physics, University of Agriculture, Faisalabad, Pakistan
S. Ali
Department of Environmental Sciences and Engineering, Government College University,
Faisalabad, Pakistan
M. Waseem
Department of Botany, Allama Iqbal Open University, Islamabad, Pakistan
M. Hussain
Department of Zoology, Government CollegeUniversity, Faisalabad, Pakistan
© Springer Nature Singapore Pte Ltd. 2020

M. Hasanuzzaman (ed.), Plant Ecophysiology and Adaptation under Climate
Change: Mechanisms and Perspectives I,
https://doi.org/10.1007/978-981-15-2156-0_27
Q. Ali et al.
23 estimated that till 2050, there is a need to double the world food production with a
24 rate of 2–5% per year but present rate of only 0.9%. It is only possible by increasing
25 the crop production area or by crop production per capita. The former one is not
26 possible in present environmental conditions due to limited sources of freshwater.
27 However, it is possible to achieve the latter one through different ways, when there
28 is a limited supply of water. It is possible through the selection from the available
29 germplasm that can perform better with better production under the changing envi-
30 ronmental conditions, and it seems the important and foremost way to deal with the
31 problems of world food demand. Crop breeding for the selection of varieties with
32 better production under changing and stressful environmental conditions is gaining
33 interest. In this regard, the selection of crop varieties against different abiotic
34 stresses for better production seems to be the most important one. Normally, the
35 breeding for the selection of stress-tolerant crop varieties is based on different agro-
36 nomic traits (traits of interest) such as plant biomass production, yield attributes,
37 and different stress tolerance indices that are considered necessary ones. The stress
38 tolerance mechanism in plants is a phenomenon of different cellular physio-
39 biochemical attributes. However, it is a complex mechanism because the stress tol-
40 erance in crops is multigenic and complex mechanism and is purely under the
41 control of genetics. In this regard, present interest and focus of researchers in the
42 study of quantitative trait loci (QTLs) has played an important role for the selection
43 and development of stress-tolerant crop varieties in a short time. Though the fruitful
44 success regarding the QTL-based selection has been achieved, but due to the com-
45 plexity in multigenic nature of stress tolerance, the behavior of the crop varieties
46 changes under the changing environmental conditions. The present chapter is a
47 comprehensive update regarding selection of stress-tolerant crop varieties for better
48 production under the changing environmental conditions. So, in the future in view
49 of changing environmental scenario, it is necessary to find out or develop the crop
50 varieties that can perform better with better production under such conditions that
51 will be fruitful to fulfill the world food demand for ever-increasing world population
52 at present and in the near future.
53 Keywords Environmental stresses · Crop plants · Breeding · Physiological traits ·

54 Agronomic traits · QTLs
55 27.1 Introduction
56 Climate can be defined as the change in weather conditions in a given region for a
57 long period of time that range from decades to millions of years. The climate change
58 may be concise to a specific region or across the entire earth (Sahney et al. 2010).
59 Farmers have developed ways to the changing climatic conditions but the changing
60 weather conditions in a way are major problems for the proper crop growth and AU1
27 Breeding Plants for Future Climates
production (Adger et al. 2007). Changing environmental conditions are major threat 61
to annual food production that causes the problem of malnutrition and food security. 62
Furthermore, these changing environmental conditions could lead to different abi- 63
otic stresses like drought (Hendrix and Glaser 2007; Lobell et al. 2008). Among 64
AU2 environmental issues are the changes in rainfall patterns, which could cause drought 65
in one area and flooding in others with a significant decline in crop production and 66
final yield (Hendrix and Glaser 2007) that causes the problem of food security for 67
ever-increasing world population (Gleditsch et al. 2006; Brown and Funk 2008). 68
AU3 Due to increased urbanization a huge amount of greenhouse gasses have been 69
AU4
added in environments that have resulted in annual increase in temperature. It is 70
estimated that the average increase in temperature is about 0.8 °C globally, while in 71
Pakistan the average temperature increase is 0.5 °C that results in an increase in 72
number of heat shocks per year (El-Sharkawy 2014). This increase in temperature 73
has been resulted in decreased plant growth due to increased transpiration rate and 74
low availability of irrigation water (El-Sharkawy 2014). 75
AU5 The gradual increase in earth’s temperature can cause the melting of the ice caps, 76
rise on the sea level, and many other environmental challenges. The main reason 77
behind this is the accumulation of the greenhouse gases and in the future it could be 78
more dangerous for the health, safety, and welfare of the ecosystem (Mahmood 79
et al. 2010). High temperature can result in increased growth for the crops growing 80
in cooler but in most of cases the negative results in crop yield are expected under 81
high temperature. High temperature results in water shortage in crops growing in 82
the areas with low rainfall. High temperature directly affects the different physio- 83
logical and biochemical processes in the plants (Karl et al. 2009). There are four 84
major compounds that are responsible for the greenhouse effect. These compounds 85
include CO2 that took play their 50–55% role in greenhouse effect. The other com- 86
pounds include chlorofluorocarbons (CFCs), methane, and nitric oxide (NO2) 87
(Nepstad et al. 2009; Smith and Gregory 2013). Furthermore, increased deforesta- 88
tion along with high atmospheric CO2 in the environment may result in more 89
increase in mean atmospheric temperature and problems could be more severe. 90
High temperature significantly affects the plant growth especially at early growth 91
stages. Normally, the temperature ranging from 35 to 45 °C results in heat stress 92
(Mahmood et al. 2010; Ulukan 2011). Heat stress directly affects the plant metabo- 93
lism that leads to the damage to the cellular membranes due to overproduction of 94
reactive oxygen species (Yousfi et al. 2010). 95
Human activities are affecting earth’s natural environment since after the indus- 96
trial revolution around 1750. This significantly affected the ratio between incoming 97
solar radiations and outgoing infrared radiations that are the major part of earth 98
energy balance. The increased concentration of all greenhouse gases in the atmo- 99
sphere has further increased the atmospheric temperature. CO2 is increased due to 100
industrialization, urbanization, and increased consumption and burning of fossil 101
fuels. Agriculture, landfills, and natural gas distributions have increased the concen- 102
tration of methane in the environment. NO2 have been increased due to increased 103
use of fertilizers and fossil fuels. All these factors collectively increased the effect 104
of global warming that result in annual increase in temperature of the atmosphere 105
Q. Ali et al.
106 (IPCC 2007). Global temperature has been significantly increased during the twen-
107 tieth century (Alexander et al. 2006). Average global temperature has been risen
108 1.1°F just in past thirty years (Hansen et al. 2006). There is a direct correlation
109 between annual increase in temperature and increased concentration of greenhouse
110 gasses. It has been estimated by the IPCC that the human activities have increased
111 the concentration of greenhouse gasses up to 70% from 1970 to 2004. The IPCC
112 states that the rise of global temperatures is due to human activity and predicts the
113 average global temperature can rise 2 to 11.5°F by 2100 (IPCC 2013).
114 At present due to rapid increase in population, it increased the demand for food
115 with better nutritional quality. Along with the world population increase, the chang-
116 ing environmental condition has further created a problem to fulfill this demand.
117 There is a need of 2.9% in crop production till 2050 to fulfill the food demand of
118 ever-increasing population. Further food production, storage, and marketing are
119 also directly linked to weather extremes and climatic changing conditions. Global
120 food security can be defined as when all the people have access to safe and nutri-
121 tious food to meet their dietary needs (FAO 1996). World food supply is dependent
122 on few crops especially cereals, oil seeds, sugars, and soybeans. The demand for
123 these crops is increasing with increasing population, but due to change in environ-
124 mental condition and adverse effects of abiotic stress, the growth and production of
125 these crops is affected adversely (Curtis and Halford 2014). Yields of these crops are
126 directly depending on environmental conditions. Sudden change in the environment
127 and harsh climatic conditions reduce the crop production.
128 Climatic changes have significant effects on water resources, human health, and
129 food security especially in developing countries (Magadza 2006). Huge literature
130 has been cited to study the effect of the environment on crop yield especially in
131 cereals like wheat, maize, and rice. Environmental changes are one of the most sig-
132 nificant factors that directly affect the crop growth and production (Magadza 2006).
133 During past few decades climatic conditions have changed so rapidly that signifi-
134 cantly affected the annual food production. Changes in rainfall patterns and annual
135 increase in atmospheric temperature due to global warming have significantly
136 decreased the crop production and the problem is getting more severe in the coming
137 future (Reddy and Pachepsky 2000). Climatic changes are a global phenomenon but
138 developing countries are affected more due to its dependence on agriculture. Food
139 security and availability of water are highly susceptible to changing environmental
140 conditions. Due to these changing environmental conditions and annual increase in
141 temperature, it is estimated that the Himalayan glaciers will disappear by 2035. This
142 rapid melting of glacier will result in floods in some areas and water shortage in
143 others (Misra 2014). Many studies have revealed that the climatic change has direct
144 effect on food production (Janjua et al. 2010; Kirby et al. 2016). It is estimated that
145 the wheat production of South Asian countries will decrease up to 50% by 2050
146 (Mahmood et al. 2010). It is stated by Peterson Institute that global warming will
147 decrease the agricultural productivity by 10–25 % in almost all developing coun-
148 tries and if it remains unchanged it will also reduce the agricultural capacity.
149 Therefore climate changes have serious threat to food production if the problem is
150 not resolved (Cline 2007). Due to global warming, the areas suitable for crop growth
are becoming too hot and too dry and the problem is becoming more severe in the 151
coming future (Hatfield et al. 2008; Karl et al. 2009; Gornall et al. 2010; Muller 152
et al. 2011). 153
Any external factor that negatively or positively affects the plant growth, metab- 154
olism, reproductive mechanism, as well as yield and productivity is termed as 155
“stress.” Stresses can be categorized into two main classes such as abiotic (environ- 156
mental) and biotic (biological) stresses. Abiotic stresses include nonliving factors 157
such as water, temperature, salinity, light, and heavy metal stress that may be 158
imposed by the environment on plants while biotic stresses are caused by living 159
organisms such as insect, diseases, or any other plants that may be exposed to the 160
plant during its lifetime (Rhodes and Nadolska-Orczyk 2001; Kranner et al. 2010) 161
(Tables 27.1 and 27.2). 162
Abiotic stresses are result of nonbiological factors that alter the growth, repro- 163
duction, and life cycle of plants. Some sources of abiotic stresses are changes in 164
water status, temperature, salinity, soil nutrient levels, etc. These stresses are classi- 165
fied into external environmental or nutritional stresses that can either positively 166
(eustress) or negatively (distress) affect plant growth and metabolism. These stresses 167
can affect the plants through disturbance in its life cycle by disturbing its cellular 168
mechanism at physiological and molecular levels (Peterson et al. 2001; Shao et al. 169
2009; Kranner et al. 2010). 170
Among different abiotic stresses, drought or water deficiency is one of the major 171
hindering factors affecting all the growth and metabolic processes by dehydration 172
that ultimately reduced the crop productivity (Zhu 2001; Sazzad 2007). The fore- 173
most effect of water stress is an imbalance in osmotic and metabolic activities 174
resulting in closure of stomata and losses of turgidity. So, water stress restricts the 175
uptake of CO2 that represses the cell growth and, therefore, reduces the process of 176
photosynthesis (Shinozaki and Yamaguchi-Shinozaki 2000). The prolonged expo- 177
sure of plants to these water deficit conditions results in permanent wilting and 178
ultimately the death of plants. 179
All plant species require optimal temperature for their normal growth and metab- 180
olism. Therefore, this optimal temperature is an important factor for distribution of 181
plant species on the planet. The optimal of daytime temperature for plants ranges 182
between −70 and 60 °C that is significantly affected by local topology (Sazzad 183
2007; Kranner et al. 2010). Another major factor that disturbs the life cycle of plants 184
from germination (beginning) to reproduction is temperature stress. The changes in 185
temperature can cause uncontrollable damages or even plant death. Temperature 186
stress also causes the initiation of other stresses such as drought and salinity. 187
Temperature stress is divided into two types, i.e., cold (low temperature) and heat 188
(high temperature) stress. The former can cause unmanageable physical and 189
mechanical damages to plants that result in severe cell disruption, while latter one 190
reduces the growth and nutrient uptake. Freezing stress also results in freezing inju- 191
ries. It is the result of cell desiccation that occurs when water moves from cells to 192
intercellular spaces due to high water potential outside the cell. Cold stress leads to 193
molecular precipitation, membrane damage, cell lysis, and production of reactive 194
oxygen species, especially at molecular level. Another most important temperature 195
t1.1 Table 27.1 Studies showing the selection of drought-tolerant crop genotypes when grown under different agroclimatic conditions
t1.2 Experimental No. of lines/ Studied
t1.3 Crop type conditions Studied characteristics genotypes used Selected tolerant lines area References
t1.4 Beta vulgaris Greenhouse Growth and yield parameters, sugar, proline, Ten SC C2×S7, SC C2×S10, Iran Taghizadegan
t1.5 experiment and LRWC populations and SC C2×S11 et al. (2019)
t1.6 and two
t1.7 control
t1.8 varieties
t1.9 Oryza sativa Glass house Relative water content, osmotic potential, Two varieties PR-115 India Khan et al.
t1.10 L. membrane damage, chlorophyll content, (2017)
t1.11 phenolic acids, MDA, and total phenolic
t1.12 content
t1.13 Zea mays L. Pot Growth and yield, lipid peroxidation rate and Three hybrids Dong-Dan 80 China Anjum et al.
t1.14 experiment lipoxygenase activity, antioxidant activity, (2017a, b)
t1.15 osmolyte accumulation profile
t1.16 Triticum Greenhouse Number of days to heading (DTH), days to 96 genotypes LM29, LM22, LM04, South Mwadzingeni
t1.17 aestivum L. experiment maturity (DTM), productive tiller number (TN), LM77, LM71, LM23, Africa et al. (2016)
t1.18 plant height (PH), spike length (SL), spikelet LM100, LM27, LM85,
t1.19 per spike (SPS), kernels per spike (KPS), LM96, LM03, LM31,
t1.20 thousand kernel weight (TKW), and grain yield LM35, LM44
t1.21 (GY) and proline content (PC)
t1.22 Beta vulgaris Field Root yield, brix % age, brown sugar yield, 17 genotypes Cauvery and Srenada Bangladesh Ganapati et al.
t1.23 experiment chlorophyll content, and root yield (2016)
t1.24 Triticum TOL, STI, SSI, HM, GMP, MP, modified STI 13 hybrids Bio-9681 and HQPM-7 Rajasthan Kumar et al.
t1.25 aestivum L. under optimum conditions (K1STI), and STI (2015)
t1.26 under stress condition (K2STI)
t1.27 Brassica Field Stress sensitivity index (SSI), tolerance index 28 cultivars Nk fair and Oase Iran Majidi et al.
t1.28 napus L. experiment (TOL), MP, GMP, and STI (2015)
t1.29 Solanum Lab SSI, STI, TOL, GMP, MP, YI, and YSI 15 cultivars C11, C15, and C10 Jeddah Metwali et al.
t1.30 lycopersicum experiment (2015)
Q. Ali et al.
Experimental No. of lines/ Studied
Crop type conditions Studied characteristics genotypes used Selected tolerant lines area References
t1.31 Beta vulgaris Greenhouse Seven drought indices including SSI, STI, TOL, Four cultivars Hayola 408 and Hayola Iran Naderi and
t1.32 experiment MP, HARM, GMP, and yield loss ratio (S) 308 Emam (2014)
t1.33 Solanum Pot SSI, relative drought index (RDI), MP, STI, SI, 41 tomato G125, G105, and G104 Serbia Brdar-Jokanović
t1.34 lycopersicum experiment dry weight yield index, superiority index, GMP, population et al. (2014)
t1.35 harmonic mean (HM), drought resistance index
t1.36 (DI), modified stress tolerance index (MSTI),
t1.37 abiotic tolerance index (ATI), stress
t1.38 susceptibility percentage index (SSPI),
t1.39 sensitivity drought index (SDI), relative
t1.40 decrease (RD)
t1.41 Brassica Pot Plant height, SFW, SDW, RL, RFW, RDW, LA, 11 canola KS-75 and Rainbow Pakistan Haq et al.
t1.42 napus L. experiment leaf Na+, leaf K+, K+/Na+ cultivars highly tolerant, Shiralee, (2014a, b)
t1.43 DGL, Westar, KH-65,
t1.44 and Legend as
t1.45 moderately tolerant
t1.46 Helianthus Field Mean productivity (MP), STI, GMP, HARM, 56 landraces Anghane 4, Marand- Iran Gholinezhad
t1.47 annuus experiment SNPI, YI, DI, modified stress tolerance index in Dizaj-Ghalami, et al. (2014)
t1.48 optimum irrigation (MPSTI), and modified Salmas-Sadaghian,
t1.49 stress tolerance index in moderate and severe Gharagoz1, Saribaglou 5
t1.50 stress (MsSTI)
t1.51 Zea mays L. Protein NIR, starch, oil, protein Kjeldahl, 13 landraces L3, IP2, and IP8 Serbia Ignjatovic-Micic
t1.52 tryptophan, QI, and kernel type et al. (2014)
t1.53 Solanum Laboratory Germination rate and percentage, shoot and 120 lines 19905, 19906, LA0716, Pakistan Shamim et al.
t1.54 lycopersicum experiment root length and their ratio, shoot and root fresh and LA0722 (2014)
t1.55 and dry weight, shoot and root moisture
t1.56 content, shoot to root ratio, and relative water
t1.57 content
(continued)
Table 27.1 (continued)
t1.58 Beta vulgaris Field Mean productivity, STI, GMP, STI, yield 15 cultivars Karun, NK Aviator, and Iran Aliakbari et al.
t1.59 experiment stability index, and modified stress tolerance NK Octans (2014)
t1.60 index
t1.61 Zea mays L. Field Germination %, overall survival % 62 maize 2, 13, 18, 28, 31, 38, 39, USA Meeks et al.
t1.62 experiment inbred lines 42, 47, 50, 54, 57, 58, 61 (2013)
t1.63 and their
t1.64 hybrid test
t1.65 cross
t1.66 Triticum Field STI, SSI, TOL, HAM, GMP, MP, YI, YSI, x DI, 15 genotypes Saji and Rizhav Iran Dehbalaei et al.
t1.67 aestivum L. experiment ATI, stress non-stress product index (SNPI), (2013)
t1.68 MSTI, and SSPI were calculated based on grain
t1.69 yield under drought (Ys) and irrigated
t1.70 conditions (Yp)
t1.71 Zea mays L. Field WUE, GMP, SSI, YSI, Ypi, LR, and SEN 135 F2 and F3 KDV2/CML444-14 and Kenya Ngugi et al.
t1.72 experiment progenies KDV2/CML440-224 (2013)
t1.73 Helianthus Field 1000-kernel weight, biological weight, grain Five cultivars Favorit Iran Rafiei et al.
t1.74 annuus L. experiment yield, harvest index (HI), and oil percentage (2013)
t1.75 Beta vulgaris Field MP, GMP, and STI Two 111-HSF.42 Iran Rajabi et al.
t1.76 experiment populations (2013)
t1.77 Triticum Field Yield, biomass and HI, yield, biomass and 50 lines hs65 India Ramya et al.
t1.78 aestivum L. experiment harvest index (HI) (2016) AU6
t1.79 Helianthus Field Yield stability index (YSI), stress susceptibility 8 cultivars Azargol Iran Taherabadi et al.
t1.80 annuus L. experiment index (SSI), tolerance (TOL), mean (2013)
t1.81 productivity (MP), geometric mean productivity
t1.82 (GMP), stress tolerance index (STI), and
t1.83 harmonic mean (HAM)
Q. Ali et al.
t1.84 Zea mays L. Field Average daily precipitation, daily high Five inbred Tx205, C2A554-4, and USA Chen et al.
t1.85 experiment temperature, relative water content, plant lines B76 (2012)
t1.86 height, plant growth
t1.87 Triticum Field STI, GMP, MP, SSI, TOL, YI, YSI, drought 13 landraces WC- 4953S (20), Iran Farshadfar et al.
t1.88 aestivum L. experiment response index (DRI), DI, MSTI, relative WC-47572 (19), and (2012)
t1.89 drought index (RDI), ATI, and SSPI WC-47574 (4)
t1.90 Zea mays L. Field STI, SSI, TOI, MP, GMP, HM, and golden 8 hybrids KSC704 hybrid and H4 Iran Moradi et al.
t1.91 experiment mean (GM) (2012)
t1.92 Triticum Field MP, GMP, tolerance, SSI, and STI 18 genotypes G18, G14, G11, and G4 Iran Mohammadi
t1.93 aestivum L. experiment et al. (2012)
t1.94 Helianthus Laboratory GSI (germination stress index), PHSI (plant 7 genotypes S473, S471, and S475 Thailand Saensee et al.
t1.95 annuus L. experiment height stress index), RLSI (root length stress and hybrid Pacific 77 (2012)
t1.96 index), DMSI (dry matter stress index), RWCSI
t1.97 (relative water content stress index)
t1.98 Triticum Field Photosynthetic pigments (chl. a, b, 5 varieties KW, UP 2752, and PBW India Chakraborty and
t1.99 aestivum L. experiment carotenoids), antioxidative defense mechanism 343 Pradhan (2012)
t1.100 (enzymatic and nonenzymatic), membrane
t1.101 stability, lipid per oxidation
t1.102 Helianthus Pot Growth parameters, MDA, H2O2, and proline Two cultivars Aydın Turkey Baloğlu et al.
t1.103 annuus L. experiment contents, antioxidant activity (2012)
t1.104 Solanum Greenhouse Growth and yield parameters as well as water Four genotypes Pakmore, VF, and Saudi Wahb-Allah
t1.105 lycopersicum experiment use efficiency drought-tolerant Arabia et al. (2011)
t1.106 breeding line (L 03306)
t1.107 Fragaria × Pot Chlorophyll, carbohydrates, proline content, Two cultivars Kurdistan Iran Ghaderi and
t1.108 ananassa experiment relative water content, membrane stability Siosemardeh
t1.109 (2011)
(continued)
t1.110 Brassica Field Chlorophyll content, leaf relative water content, Four cultivars Elite Isfahan Sepehri and
t1.111 napus L. experiment leaf area Golparvar
t1.112 (2011)
t1.113 Beta vulgaris Field Stress susceptibility index (SSI), tolerance 23 genotypes Modena, Geronimo, Iran Rad and
t1.114 experiment index (TOL), stress mean productivity (MP), Elite, Syn-4, and Abbasian
t1.115 geometric mean productivity (GMP), stress SLM046 (2011a, b)
t1.116 tolerance index (STI), yield index (YI), and
t1.117 yield stability index (YSI)
t1.118 Triticum Field Relative water content, growth, and yield Two cultivars Inqlab-9l Faisalabad Akram (2011)
t1.119 aestivum L. experiment components
t1.120 Triticum Field Yield potential and stability under stressful 14 wheat Gidara-II, Sarıçanak-98, Turkey Kilic and
t1.121 drum L. experiment conditions cultivars Balcalı-2000, Yagbasanlar
t1.122 Altıntoprak-98, (2010)
t1.123 Aydın-93, and Harran-95
t1.124 Triticum Field SSI, STI, YC, YSI, MP, and GMP 24 cultivars 1, 6, 7, 19, 20, and 21 Iran Talebi et al.
t1.125 drum L. experiment (2009)
t1.126 Triticum Field Yield and yield components 25 varieties Sarsabz and Kiran-95 Pakistan Mirbahar et al.
t1.127 aestivum L. experiment (2009)
t1.128 Triticum Laboratory Ys, Yp, SSI, STI, TOL, GMP, MP, and MSTI 42 genotypes Sardar, Sabalan, and Iran Shahryari et al.
t1.129 aestivum L. experiment 4057 (2008)
t1.130 Beta vulgaris Glasshouse Growth and yield components as well as Four cultivars Hayola 401, Hayola 308 Iran Naderikharaji
t1.131 photosynthetic attributes et al. (2008)
t1.132 Beta vulgaris Field Growth attributes Nine genotypes MSTC2 and 7233.P3 Iran Mohammadian
t1.133 experiment et al. (2005)
Q. Ali et al.
t2.1 Table 27.2 Studies showing the selection of salt-tolerant crop genotypes when grown under different agroclimatic conditions
t2.2 No. of lines/
t2.3 Experimental genotypes Studied
t2.4 Crop type conditions Studied characteristics used Selected tolerant lines area References
t2.5 Avena sativa Laboratory Seed germination, shoot and root Seven NDO-2 and UPO-212, salt tolerant India Chauhan et al.
t2.6 experiment length, dry weight of shoot and cultivars and moderately salt tolerant, (2016)
t2.7 root, total dry weight of seedling respectively
t2.8 Zea mays L. Greenhouse Growth parameters, Two S2000 Pakistan Gul et al.
t2.9 experiment photosynthetic pigments, relative cultivars (2016)
t2.10 water contents, and Na+ and K+
t2.11 concentrations
t2.12 Triticum Hydroponic Membrane stability (MS) and its 20 cultivars Moderately tolerant cultivars Turkey Kaya and
t2.13 aestivum L. experiment injury (M1) (Dagdas-94, Eraybey, Altay-2000, Arısoy (2016)
t2.14 Mesut, Somez-01, Bayraktar 2000,

t2.15 Gun-91, and Ekiz), highly tolerant
t2.16 cultivars (Karahan-99, Gerek-79,
t2.17 Konya-2002, and Kinaci-97)
t2.18 Oryza sativa L. Pot Physiological, parameters, and 12 cultivars Niewdam Gs.no.00621 Thailand Chunthaburee
t2.19 experiment antioxidant enzyme et al. (2016)
t2.20 Zea mays L. Hydroponic Root length, shoot and root dry 13 maize SC2 andAD3 lines Argentina Collado et al.
t2.21 mass, relative water content, leaf inbred lines (2016)
t2.22 water loss, stability of membrane,
t2.23 tolerance index (TOL)
t2.24 Brassica napus Field Number of pods per plant, pod 14 cultivars Modena and Opera Iran Ahmadzadeh
t2.25 experiment length, pod weight, kernel et al. (2015)
t2.26 number per pod, 100 kernel
t2.27 weight, and kernel yield
t2.28 Lactuca sativa Hydroponic Growth; stomatal conductance; Five Parris Island lettuce cultivar Romania Bartha et al.
t2.29 Na+, Ca2+, and K+ content in cultivars (2015)
t2.30 shoot; proline
(continued)
No. of lines/
Experimental genotypes Studied
Crop type conditions Studied characteristics used Selected tolerant lines area References
t2.31 Piper nigrum Greenhouse Growth parameters, chlorophyll 26 cultivars CV. paramo Iran Bavani et al.
t2.32 experiment content, leaf gas exchange, (2015)
t2.33 RLWC, and STI
t2.34 Oryza sativa L. Pot Shoot, root parameters, Na+ /K+ 22 cultivars Pachaperumal, Periavellai, At 303, Sri Lanka Pradheeban
t2.35 experiment Adakari, Bg 406, and CO 10 et al. (2015)
t2.36 Brassica napus Hydroponic Growth , Na+, K+, and their ratio 11 cultivars Con-II, Con-III, Dunkeld, and Oscar Pakistan Haq et al.
t2.37 system (2014a, b)
t2.38 Triticum Greenhouse Protein assay, peroxidase, Na+, Four Barley cultivar “Kavir” and wheat Iran Izadi et al.
t2.39 aestivum L. and experiment K+, and the K+/Na+ ratio, proline cultivars cultivar “Bam” (2014)
t2.40 Hordeum each
t2.41 vulgare
t2.42 Oryza sativa L. Pot TOL, SSI, MP, GMP, STI, HM, Eight Dasht Northern Kamyab-Talesh
t2.43 experiment YI, YSI cultivars Iran et al. (2014)
t2.44 Helianthus Pot Germination Three Sandaomei China Luan et al.
t2.45 annuus experiment cultivars (2014)
t2.46 Piper nigrum Lab Germination and growth Three Anaheim Chili Tunisia Hassen et al.
t2.47 experiment parameters cultivars (2014)
t2.48 Triticum drum Field and pot Germination and growth attributes 119 Malta2 Japan Turki et al.
t2.49 experiment landraces (2014)
t2.50 Solanum Hydroponic Growth, proline content, K+/Na+ Six cultivars Mozart and Desiree Netherlands Jaarsma et al.
t2.51 tuberosum accumulation, H2O2, and relative (2013)
t2.52 water content
Q. Ali et al.
No. of lines/
t2.53 Triticum drum Greenhouse Height of plant, length of Ten cultivars Boomer and PGS Iran Kahrizi et al.
t2.54 experiment peduncle, area of flag leaf, area of (2013)
t2.55 penalty leaf, number of leaf,
t2.56 length of spike, no. of grains, and
t2.57 weight of grains
t2.58 Triticum Pot Chlorophyll content (Chl.), leaf Ten cultivars Sehar-06, Lu-26, NARC-09, BARC- Pakistan. Rao et al.
t2.59 aestivum L. experiment relative water content (LRWC), 09, and Pirsbak-09 (2013)
t2.60 Na+/K+, activities of APX, GPX,
t2.61 SOD, CAT, GPX, H2O2, and
t2.62 membrane stability index

t2.63 Solanum Hydroponic Na+ accumulation Six cultivars Desiree and Russett Burbank Netherlands Jaarsma et al.
t2.64 tuberosum (2013)
t2.65 Brassica napus Field Yield, component characters, and Eight DSM12 and Hyola401 Iran Valiollah
t2.66 experiment nutrient compositions genotypes (2013)
t2.67 Triticum Lab Germination, seedlings 20 lines/ “Chamran,” “Yavarous,” “83-3,” Iran Shekoofa et al.
t2.68 aestivum L. experiment establishment, and respiration cultivars “Taro3,” “Cross Boullani,” “Cross (2013)
t2.69 Aadl,” and “Dabira”
t2.70 Hordeum Lab Germination, growth, leaf Three Nosrat Iran Somayeh et al.
t2.71 vulgare experiment chlorophyll, proline rate cultivars (2012)
t2.72 Abelmoschus Pot 1000-grain weight, grain yield, 39 Acc. No. 019232, Acc. No. 000010- Pakistan Haq et al.
t2.73 esculentus experiment biological yield, number of tillers, genotypes 10237, Chinese Red, Ikra III, and Acc. (2012)
t2.74 number of fertile tillers, spike No. 015371
t2.75 length, salinity susceptibility
t2.76 index (SSI), and salt tolerance
t2.77 index (STI)
(continued)
No. of lines/
t2.78 Capsicum Greenhouse Growth parameters, chlorophyll, Five Awlad Haffouzz and Korba Tunisia Kaouther et al.
t2.79 annuum experiment proline cultivars (2012)
t2.80 Brassica napus Pot Proline metabolism Two Dunkled Pakistan Saadia et al.
t2.81 experiment cultivars (2012)
t2.82 Brassica napus Hydroponic Tolerance, mean productivity 12 cultivars Craker and Amica Iran Toorchi et al.
t2.83 culture stress, susceptibility index, (2012)
t2.84 geometric mean, productivity
t2.85 stress, tolerance index
t2.86 Brassica napus Greenhouse Root and shoot fresh and dry Five SLM046 Iran Bybordi and
t2.87 experiment weight, relative water content, cultivars Tabatabaei
t2.88 Na+/K+, lipid per oxidation, CAT, (2010)
t2.89 SOD, POD, APX, GSG, GSSG
t2.90 Triticum Lab Germination, growth, sugar, Five HD2689 India Datta et al.
t2.91 aestivum L. experiment protein, chlorophyll, proline varieties (2009)
t2.92 Oryza sativa L. Greenhouse Growth and physiological Seven Co39 and Moroberekan Pakistan Haq et al.
t2.93 experiment parameters varieties (2009)
t2.94 Brassica napus Greenhouse Chlorophyll fluorescence, Ten Dunkeld and Hayola 401 Iran Saeedipour
t2.95 experiment photosynthetic parameters cultivars/ (2009)
t2.96 accessions
t2.97 Triticum Greenhouse Shoot Na+, K+, K+/Na+ ratio, 15 cultivars Kavir,Niknejad, Chamran, and Falat Iran Goudarzi and
t2.98 aestivum L. 1000-grain weight, grain yield, Pakniyat (2008)
t2.99 biological yield, number of tillers,
t2.100 number of fertile tillers, spike
t2.101 length, SSI, STI
t2.102 Triticum Laboratory No. of seeds, percent of callus Four LU-26 Islamabad Akhtar (2006)
t2.103 aestivum L. experiment formation, and regeneration cultivars
Q. Ali et al.
No. of lines/
t2.104 Helianthus Field Growth and yield, K+, Na+, Ca+, Ten CRN-1435, HU-777, Super-25, and Pakistan Ahmed et al.
t2.105 annuus experiment and Mg+ concentration genotypes cultivar 6451 (2005)
Q. Ali et al.
196 stressor is extremely high temperature that can cause heat shocks. It can cause wilt-
197 ing by increasing the evaporation rate from leaves, and its prolonged exposure leads
198 to permanent wilting or even death of plants (Thomashow 1998, 1999; Pearce 1999;
199 Sazzad 2007) (Tables 27.3 and 27.4).
200 Like other abiotic stresses, salt stress also substantially decreases the growth as
201 well as yield of crop plants and in many cases by 50% or more reduction all around
202 the world. The foremost effect of salinity is that it disturbs the seed germination and
203 viability. The presence of excess salt or ions around the roots decreases the water
204 uptake potential of plants that ultimately reduces the growth through disturbed
205 metabolism (Sazzad 2007; Kranner et al. 2010; Machado and Serralheiro 2017).
206 Salinity is the cause of excessive quantity of water-soluble salts such as sodium
207 carbonates, sodium sulfate, potassium sulfate, sodium nitrate, calcium sulfate, mag-
208 nesium sulfate, sodium chloride, and magnesium chloride that not only negatively
209 alters the different processes of plant metabolism and as a result both positively and
210 negatively triggers its growth and metabolism (Sazzad 2007; Machado and
211 Serralheiro 2017). Plants need all these salts in an optimal amount for regulating
212 their normal metabolic activities. The presence of these salts above and below their
213 optimal level may cause cell disruption. A number of these salts can help the plants
214 in regulation of growth and metabolic activities while their excess amount may be
215 toxic for plants (Sazzad 2007). For example, if sodium chloride is present in an
216 optimal amount in the soil, it will improve the plant growth and metabolism up to
217 some extent while at higher concentration it may inhibit the seed germination and
218 seedling establishment as well as plant growth and development (Tester and
219 Davenport 2003; Shabala et al. 2015). Plants can be divided into two classes on the
220 basis of their salt tolerance mechanism, i.e., halophytes (salt tolerant) or glyco-
221 phytes (salt sensitive). Halophytes can tolerate the negative effects of salinity by
222 decreasing the uptake of salt from soil and by reducing the concentration of salt in
223 cytoplasm as well as in cell wall through exclusion or vacuole storage. Glycophytes
224 (salt sensitive) are less able to tolerate high salt level and can accumulate high salt
225 concentration in cytosol that is toxic for plant metabolism (Munns et al. 2006;
226 Sazzad 2007).
227 The elements with density greater than 5 g cm−3 are categorized as heavy metals.
228 These mainly include Co, Cu, Cd, Pb, Fe, Ni, Mn, Zn, and Mo. These elements in
229 some cases are essential for plants as they directly or indirectly affect the growth,
230 metabolism, senescence, as well as many energy-generating processes in studied
231 plant species. Many of these elements can stimulate various growth and metabolic
232 activities in plants if they are present in low concentration in soil (Emamverdian
233 et al. 2015). On the other hand, their concentration above threshold level in the soil
234 is toxic to plants either causing the stress that is known as heavy metal stress. Heavy
235 metal stress disturbs the plant growth by disturbing the activities of various enzymes
236 through higher levels of free radicals or by replacing the uptake and mobility of
237 many nutrients and essential elements. The toxic effects of these metals are directly
238 depending on the nature of these metals and type of plant species as well (Jozefczak
239 et al. 2012; Emamverdian et al. 2015).
t3.1 Table 27.3 Studies showing the selection of temperature stress-tolerant crop genotypes when grown under different agroclimatic conditions
t3.2 No of lines/
t3.3 Type of Experimental genotypes
t3.4 Crop type stress conditions Studied characteristics used Selected tolerant lines Studied area References
t3.5 Glycine max Low and Lab Growth and physiological 64 cultivars CZ 5225 LL and USA Alsajri et al.
t3.6 L. high experiment parameters as well as GS47R216 sensitive and (2019)
t3.7 temperature cumulative stress tolerant to LT while
t3.8 stress response indices cultivars 45A-46 and
t3.9 5115LL tolerant and
t3.10 sensitive to HT,
t3.11 respectively
t3.12 Oryza sativa Cold stress Greenhouse Germination %age, 40 rice CT6749-36-7-2-M-M, Egypt Elgamal et al.
t3.13 L. germination index, accessions Milyang 80, PSRM1- (2018)
t3.14 seedling height, root 17-4B-13, Ryong Sung 25

t3.15 length, no. of leaves or
t3.16 seedlings
t3.17 Vigna radiata High Field Growth, morphology, and 41 elite mung EC693357, EC693358, India Sharma et al.
t3.18 temperature experiment yield traits bean lines EC693369, Harsha, and (2016)
t3.19 stress ML1299
t3.20 Oryza sativa Heat stress Field Days to 50% flowering, 1217 IR 87606-109-2-2 and IR Bangladesh Masuduzzaman
t3.21 L. experiment days to maturity, plant 86991-146-2-1-1 et al. (2016)
t3.22 height, no. of spikelets/
t3.23 spike, fertility%, plant
t3.24 yield
t3.25 Zea mays L. Cold stress Growth Germination rate, index, 22 inbred CO439, CO438, CO450, Korea Farooqi and Lee
t3.26 chamber root length, and seed lines CO435, and CO445 (2016)
t3.27 vigor index values
(continued)
No of lines/
Type of Experimental genotypes
Crop type stress conditions Studied characteristics used Selected tolerant lines Studied area References
t3.28 Glycine max Chilling Growth room Germination attributes, Six domestic Jutro and Nawiko Lublin Borowski and
t3.29 L. stress and pot leaf electrolyte leakage, soybean Michalek (2014)
t3.30 experiment free proline content and cultivars and
t3.31 catalase activity Canadian-bred
t3.32 photosynthetic rate, leaf cultivars
t3.33 area
t3.34 Daucus carota High Field Quantitative, qualitative, 38 inbreds IPC-4, IPC-8, IPC-11, New Delhi Saha et al.
t3.35 subsp. sativus temperature experiment and morphological traits IPC-13, IPC-124 (2016)
t3.36 stress
t3.37 Triticum Heat stress Field Phenology Five cultivars Faisalabad-2008 and Pakistan Sattar et al.
t3.38 aestivum L. experiment Iqlab-91 (2015)
t3.39 Zea mays L. Cold stress Growth parameters such 33 hybrids DKC6697, DKC6804, MississippiState Wijewardana
t3.40 as plant height, leaf and M2V707 et al. (2015)
t3.41 length, leaf area, root
t3.42 development and root
t3.43 growth parameters,
t3.44 nutrient uptake
t3.45 Zea mays L. High Tunnel and Days to 50% tassel, days 21 inbred YH-1898 and YH-1921 Pakistan Rahman et al.
t3.46 temperature field to 50% silk, plant height, lines (2017)
t3.47 stress conditions cob height, no. of viable
t3.48 seeds
t3.49 Strawberry Heat stress Greenhouse LRWC, chlorophyll 15 cultivars “Elsanta,” “R. Hope,” and Turkey Kesici et al.
t3.50 experiment content, heat stress “Camarosa” (2013)
t3.51 tolerance
Q. Ali et al.
No of lines/
t3.52 Triticum Cold stress Field Day to maturity (DM), 15 cultivars Sayson, Kavir, pishtaz Iran Habibi et al.
t3.53 aestivum L. experiment grain number per head, (2012)
t3.54 plant height, 1000-kernel
t3.55 weight (1000-KW), and
t3.56 grain yield
t3.57 Oryza sativa Cold stress Field Germination and seedling Six cultivars Tarom, Domsiya, Hybrid, Iran Soleymani and
t3.58 L. experiment stage Nemat, Gerde, Unda Shahrajabian
t3.59 (2012)
t3.60 Brassica High Field Yield and yield attributes Two cultivars Hyola401 Iran Faraji et al.
t3.61 napus L. temperature experiment (2008)

t3.62 stress
t3.63 Solanum High Pot experiment Photosynthetic pigments, Two Nagcarlang Cuba Camejo et al.
t3.64 lycopersicum temperature gas exchange, genotypes (2005)
t3.65 stress fluorescence, and
t3.66 electrolyte leakage
t3.67 Oryza sativa Cold stress Lab Germination parameters 24 genotypes BR-IRGA 410 and IRGA Brazil Cruz and Milach
t3.68 L. experiment 416 (2004)
t3.69 Oryza sativa High Pot experiment Percentage fertility, Nine cultivars Nipponbare and Japan Matsui and
t3.70 L. temperature anther dehiscence, and Akitakomachi Omasa (2002)
t3.71 stress size
t3.72 Brassica Heat stress Lab Fatty acid content, yield Three Oscar Australia Aksouh et al.
t3.73 napus experiment components cultivars (2001)
(continued)
No of lines/
t3.74 Solanum Heat stress Greenhouse Gas exchange, leaf 46 clones LT-5, S. acaule Canada Midmore and
t3.75 tuberosum experiment chlorophyll fluorescence (P1498066), and Prange (1991)
t3.76 net assimilation rate S. circaefolium (PI
t3.77 (NAR) and relative 498116)
t3.78 growth rate (RGR)
t3.79 parameters, and total dry
t3.80 weight (TDWT)
Q. Ali et al.
t4.1 Table 27.4 Studies showing the selection of heavy metal-tolerant crop genotypes when grown under different agroclimatic conditions
t4.2 No. of lines/
t4.3 Experimental genotypes Studied
t4.4 Crop type conditions Studied characteristics used Selected tolerant lines area References
t4.5 Zea mays L. Pot experiment Growth, leaf gas exchange, and Two cultivars Dong-Dan 80 China Anjum et al.
t4.6 photosynthetic pigments (2017a, b)
t4.7 Zea mays L. Greenhouse Growth parameters, Two cultivars S2000 Pakistan Gul et al.
t4.8 experiment photosynthetic pigments, relative (2016)
t4.9 water contents, and Na+ and K+
t4.10 concentrations
t4.11 Zea mays L. Hydroponic Growth and photosynthetic Seven hybrids 32-B-33 and 23-T-16 Pakistan
Akhtar et al.
t4.12 experiment attributes (2017)
t4.13 Triticum Field Growth and physiological Ten Ulbinka, Zaulbinka, and Omskaya-18 Kazakhstan Alybayeva
t4.14 aestivum L. experiment attributes genotypes et al. (2016)
t4.15 Solanum Glass house Yield attributes and Cd 100 9086, Roma, Sitara TS-01, Pakistan Hussain et al.
t4.16 lycopersicum accumulation in root, shoot, and genotypes pak0010990, CLN-2123A, Picdeneato, (2015)
t4.17 leaf 0.006231, and 7035
t4.18 Vigna radiata Pot experiment Germination, growth and Three M-6 and M-8 Pakistan Hasnain et al.
t4.19 photosynthetic attributes cultivars (2011)
t4.20 Vigna radiata Laboratory Growth parameters and stress Eight Tolerance response of genotypes: India Samantaray
t4.21 experiment tolerance indices cultivars Dhauli > PDM-116 > LGG-407 > et al. (1998)
t4.22 K-851 > TARM-22 > TARM-1 >
t4.23 TARM-21 > TARM-26.
Q. Ali et al.
240 Plants have the ability to survive in the stressful environmental conditions that is
241 plant species and growth stage specific. It also depends on the type and the duration
242 of the stress. To cope with these adverse environmental conditions, plants have the
243 ability to develop specific mechanisms that ultimately help the plants to complete
244 their life cycles. These include some structural changes that lead from morphologi-
245 cal to physiological adaptations (Jaleel et al. 2008; Hund et al. 2009; Gupta and
246 Huang 2014). Morphological adaptations include the reduced leaf area, leaf rolling,
247 early life cycle completion (escape), as well as the reduction in growth rate.
248 Anatomical adaptations include the cuticle formation and pubescence that also play
249 a significant role in plant stress tolerance by reducing the transpiration and evapora-
250 tion rate. Similarly, plants also have developed some physiological mechanisms that
251 help plants in ameliorating the adverse effects of different abiotic stresses by con-
252 trolling the metabolic activities. These include the stomatal regulation, osmotic
253 adjustment, oxidative defense mechanism (activity of enzymatic and nonenzymatic
254 antioxidants), accumulation of various osmolytes (carbohydrates, betaines, proline,
255 and other amino acids), as well as osmoprotectors through the production of chemi-
256 cal messengers in response to environmental stimuli that help in scavenging the
257 ROS and regulate the osmotic potential of cell under stressful environmental condi-
258 tions (Apel and Hirt 2004; Krishnan et al. 2008). Knowledge about responses of
259 plants to these abiotic stresses and their stress tolerance mechanisms could be help-
260 ful in understanding the different adaptations of plants to extreme environmental
261 conditions that are also relevant to the management and breeding programs
262 (Mohammadi 2018).
263 Different strategies are being used to counteract these problems that include the AU7
264 manipulation of important agronomic characteristics, better use of irrigation water
265 through different ways, use of chemicals/biochemical through different modes
266 (such as amino acids, plant-based extract, plant growth regulators, biostimulants),
267 fertilizers, pesticides, as well as the use of biophysical methods. Among these, the
268 breeding for selection of stress-tolerant genotypes with better production under dif-
269 ferent environmental stresses is an important one and gaining interest. To fulfill the
270 future world food demand for ever-increasing world population, the agriculturalists
271 are mainly relying on crop breeding.
272 Breeding for stress tolerance and better crop production is in practice from
273 ancient times that is based on different agronomic traits and yield attributes. In old
274 times the breeding was mainly to obtain the better production and to develop high-
275 yielding crop cultivars. At present due to environmental changes, and the lesser
276 availability of freshwater for irrigation, there is a dire need to develop stress toler-
277 ance crop cultivars with better production under stressful environmental conditions
278 to fulfill the demand of ever-increasing world population. Primarily for the develop-
279 ment of higher yielding crop varieties, the major focus was on the agronomic traits,
280 but with the development of modern tools and invents of molecular biology tech-
281 niques, the breeding has become a multidisciplinary subject. Agronomic traits are
282 the basic parameters for the selection of stress-tolerant crop cultivars. The parame-
283 ters are also being employed for horticulture crops as well. The major focus for the
selection of stress-tolerant genotypes from available germplasm through breeding is 284

the selection against water stress tolerance (Mohammadi 2018). 285
27.2 Breeding Against Water Stress 286
AU8 The selection for water stress tolerance is mainly based on stress susceptibility 287
index (SSI), stress tolerance index (STI), relative drought index (RDI), stability 288
index (SI), drought resistance index (DI), modified stress tolerance index (MDTI), 289
stress susceptibility percentage index (SSPI), sensitivity drought index (SDI), abi- 290
otic tolerance index (ATI), drought response index (DRI), relative drought index 291
(RDI), modified stress tolerance index in optimum irrigation (MPSTI), modified 292
stress tolerance index in moderate and severe stress (MsSTI), modified STI under 293
optimum conditions (K1STI) and STI under stress conditions (K2STI), and stress 294
sensitivity index (SSI) which is based on plant growth and biomass production. 295
Along with these indices, the mean productivity (MP), geometric mean productivity 296
(GMP), yield index (YI), yield stability index (YSI), dry weight yield index (DWYI), 297
harvest index (HI), superiority index, grain yield under drought (Ys), and irrigated 298
conditions (Yp) are also necessary traits. For example, in wheat various drought- 299
tolerant cultivars have been selected on the basis of different agronomic traits such 300
as STI, SSI, TOL, DRI, DI MSTI, ATI, and SSPI that are further used for breeding 301
(Shahryari et al. 2008; Talebi et al. 2009; Rad and Abbasian 2011a, b; Farshadfar 302
et al. 2012; Mohammadi et al. 2012; Dehbalaei et al. 2013). In tomato selection, the 303
agronomic traits that have been focused are SST, RDI, STI, SI, DWYI, STI, ATI, 304
SSPI, SSI, and TOL, and different drought-tolerant cultivars were selected on the 305
basis of these indices (Brdar-Jokanović et al. 2014; Metwali et al. 2015). In another 306
experiment on corn that was performed under field conditions, the two corn hybrids 307
such as KSC704 and H4 were selected out of eight hybrids as drought tolerant on 308
the basis of various agronomic attributes such as STI, SSI, and ROL (Moradi et al. 309
2012). Majidi et al. (2015) reported that out of 28 canola cultivars that were grown 310
under field water deficit conditions, only two cultivars were selected as drought- 311
tolerant ones, cultivars that were based on different stress tolerance indices such as 312
AU9 SSI, TOL, and STI. Furthermore, out of eight sunflower cultivars, grown under field 313
conditions to find out the tolerant were more tolerant to water stress was the Azargol, 314
selected on the basis of various stress tolerance indices such as TOL, SSI, and STI 315
(Taherabadi et al. 2013). In a laboratory experiment, seven sunflower genotypes 316
were grown in Petri dishes to find drought-tolerant potential of these genotypes on 317
the basis of germination stress index (GSI), plant height stress index (PHSI), root 318
length stress index (RLSI), dry matter stress index (DRSI), as well as relative water 319
content stress index (RWCSI) as reported by Saensee et al. (2012). Four genotypes 320
(S473, S471, and S475 and a hybrid Pacific 77) were reported as drought tolerant. 321
In parallel with agronomic characteristics, yield attributes were also considered as 322
important to select drought-tolerant varieties/genotypes such as MP, GMP, YI, YSI, 323
DWY, superiority index, Ys, and Yp. For example, in wheat many drought-tolerant 324
Q. Ali et al.
325 cultivars/varieties have been identified on the basis of different yield indices such as
326 YI, YSI, GMP, MP, and HI (Talebi et al. 2009; Dehbalaei et al. 2013; Ramya et al.
327 2016; Aliakbari et al. 2014). Mirbahar et al. (2009) selected two wheat varieties
328 (Sarsabz and kiran-95) out of the 25 genotypes that were grown under field water
329 deficit conditions. These varieties were selected based on plant yield and yield com-
330 ponents that showed more tolerance against water stress. The agronomic and yield
331 attributes are the basis of different physiological alteration mechanisms. So, the
332 selection based on the agronomic and physiological attributes, taken together, is
333 found better than that of only considering the agronomic traits. Many studies have
334 been reported about the selection of drought-tolerant varieties/genotypes on the
335 basis of some physiological parameters such as germination attributes, biomass pro-
336 duction, and plant photosynthetic activity, amino acid profile, and enzymatic and
337 nonenzymatic antioxidative activities (Meena et al. 2016). For example, three maize
338 hybrids were grown in pots to find out the drought tolerance potential of three maize
339 hybrids on the basis of extent of reduction in growth, yield, extent of lipid peroxida-
340 tion, and osmolyte accumulation (Anjum et al. 2017a, b). They reported that these
341 biochemical and physiological attributes helped them to develop drought-tolerant
342 hybrid Dong-Dan 80 that can survive in water deficiency with minimum yield
343 reduction. In another study, conducted by Khan et al. (2017), they reported that out
344 of two rice varieties one variety (PR-115) was selected as drought tolerant when
345 grown in pots under water stress. The selection of drought-tolerant variety was also
346 based on the different physiological parameters such as relative water content,
347 osmotic potential, membrane damage, chlorophyll content, phenolic acids, MDA,
348 and total phenolic content. For example, in rapeseed, selection of two wheat culti- AU10
349 vars as tolerant ones out of four cultivars when grown in glass house water deficit
350 conditions (Naderikharaji et al. 2008). They made the selection based on growth
351 and yield components as well as photosynthetic activities of these varieties.
352 Chakraborty and Pradhan (2012) considered antioxidative defense mechanism,
353 membrane stability, lipid peroxidation, as well as photosynthetic activities for the
354 selection of stress-tolerant genotypes out of five wheat varieties that were grown in
355 field under water deficiency. They selected three varieties (KW, UP 2752, and PBW
356 343) as tolerant ones depending on abovementioned physiological attributes. A
357 greenhouse experiment was conducted to find out the drought tolerance potential of
358 four tomato genotypes based on growth and seed yield as well as water use effi-
359 ciency when grown under water shortage. Pakmore VF and the breeding line L
360 03306 showed good growth and yield performance and are selected to use for breed-
361 ing programs as drought-tolerant genotypes (Wahb-Allah et al. 2011). Two straw-
362 berry cultivars (cv. Kurdistan and Selva) were grown in pots to find out their drought
363 tolerance potential based on the various physiological attributes including chloro-
364 phyll, proline, and carbohydrate content, membrane stability index, as well as leaf
365 relative water content. The cultivar Kurdistan was selected as tolerant one to use in
366 breeding programs (Ghaderi and Siosemardeh 2011). Antioxidative and physiologi-
367 cal responses of two sunflower cultivars (Musala and Aydın) were studied and their AU11
368 efficiency to grow under PEG-induced water stress was checked as reported by
Baloğlu et al. (2012). The cultivar Aydin was nominated as drought-tolerant cultivar 369
in terms of growth, MDA, H2O2, and proline contents as well as antioxidant activity. 370
27.3 Breeding Against Salt Stress 371
Salinity stress is a type of osmotic stress that includes the specific ion toxicity, espe- 372
cially of Na+ in sodicity that not only disturbs the membrane potential but also cel- 373
lular ion homeostasis. Along with ion toxicity, it also disturbs plant water relations 374
especially the cause of physiological drought. Plants have developed specific mech- 375
anisms to avoid the toxicity of these toxic ions but this mechanism is species and 376
cultivar specific (Munns 2005). This mechanism includes the ion exclusion at dif- 377
ferent organization levels (Munns 2005), accumulation in vacuole, and discrimina- 378
tion in the uptake of these toxic ions. The excessive toxic levels of these salts in the 379
rooting medium compete with essential macro- and micronutrients such as Na+ that 380
competes with K+, Ca2+, Mg2+, etc. So, breeding for toxic ion discrimination is an 381
effective one to develop the salt-tolerant crop genotypes. Others include the exces- 382
AU12 sive accumulation of osmotica. Water relation through osmotic adjustment, better 383
antioxidative defense mechanism, and better performance in different physio- 384
biochemical attributes. Presence of one of the above mechanisms results in better 385
crop production. Haq et al. (2012) reported that in okra the salt-tolerant genotypes 386
were selected based on different yield attributes along with SSI and STI. They 387
reported that among 39 genotypes, seven were selected as tolerant ones. A hydro- 388
ponic experiment was conducted by Jaarsma et al. (2013) to identify the salt-tolerant 389
genotypes in potatoes. They found that among seven different genotypes grown 390
under salt stress cultivars “Mozart and Desiree” were selected as tolerant ones and 391
the selection was based on different growth attributes, K+/Na+ ratio, proline content, 392
H2O2, as well as relative water content. In durum wheat, different salt-tolerant culti- 393
vars were selected on the basis of different growth and yield attributes. Two salt- 394
tolerant cultivars (Boomer and PGS) were selected, out of ten cultivars (Kahrizi 395
et al. 2013). A hydroponic experiment was conducted by Kaya and Arısoy (2016) 396
who reported that 20 wheat cultivars were grown under salt stress to select the salt- 397
tolerant genotypes. They reported that eight genotypes were moderately tolerant 398
and four were highly salt tolerant, selected against salt stress based on membrane 399
stability (MS) and membrane injury (MI) indices. Antioxidant activity, photosyn- 400
thetic attributes, and lipid peroxidation were recommended. Similarly, based on 401
abovementioned attributes, among ten wheat genotypes, five were identified as tol- 402
erant ones that further for breeding program (Rao et al. 2013). A field study was 403
conducted by Ahmadzadeh et al. (2015) to evaluate the effect of salinity stress on 404
yield and yield components of fourteen rapeseed cultivars. Among all studied culti- 405
vars, they reported two high-yielding cultivars as tolerant ones on the basis of dif- 406
ferent growth, yield, and membrane stability indices. Growth and ionic relations 407
(K+, Na+, Ca+, and Mg2+) of ten sunflower genotypes grown in field under saline 408
Q. Ali et al.
409 environment were studied to select the salt-tolerant genotypes. Four genotypes were
410 selected as tolerant ones based on abovementioned attributes (Ahmed et al. 2005).
411 A laboratory experiment was conducted under saline conditions using various oat
412 cultivars to find out the changes in germination and growth parameters. They found
413 that among seven cultivars, NDO-2 and UPO-212 were selected as highly salt toler-
414 ant and moderately tolerant, respectively, based on their germination and growth
415 behavior (Chauhan et al. 2016). Various physio-biochemical attributes were studied
416 to evaluate salt-tolerant cultivars of rice grown in pots under saline conditions.
417 Based on studied attributes cultivar Niewdam Gs.no.00621 was nominated as toler-
418 ant one as reported by Chunthaburee et al. (2016)). Various salt-tolerant wheat cul-
419 tivars were also selected based on germination attributes, growth, and chlorophyll
420 and proline contents in wheat (Datta et al. 2009), barely (Somayeh et al. 2012), and
421 pepper (Hassen et al. 2014). Based on Na+ accumulation various salt-tolerant potato
422 cultivars were selected when grown in a hydroponic experiment as depicted by
423 Jaarsma et al. (2013). Similarly, a pot study was conducted on rice to evaluate the
424 salt-tolerant genotypes based on various stress tolerance and yield indices such as
425 TOL, SSI, MP, GMO, STI, HM, YI, and YSI (Kamyab-Talesh et al. 2014). In
426 another study, conducted by Toorchi et al. (2012) reported that various canola culti- AU13
427 vars were identified as salt-tolerant ones based on TOL, MP and SI, GM through the
428 agronomic and physiological mechanisms are considered important ones but these
429 are purely controlled by genetics. In this regard selection based on QTLs is consid-
430 ered short term and easy way from the available germplasm (Table 27.5).
431 27.4 Breeding Against Temperature Stress
432 In present era due to erratic environmental conditions, especially the atmospheric
433 temperature, it is necessary to develop crop genotypes that can better perform under
434 such conditions. The main focus of breeders is on the development of genotypes
435 that can perform under increasing temperature because the global temperature is
436 rising rapidly every year. Along with the high temperature, the chilling stress is also
437 disturbing the crop production in specific areas. In case of temperature stress, heat
438 shock proteins (HSPs)/chilling stress protein is considered important one. In rice
439 various stress-tolerant genotypes were selected against high temperature stress AU14
440 when grown under field conditions and these genotypes were selected on the basis
441 of different growth and yield attributes such as days to 50% flowering, days to matu-
442 rity, plant height, no. of spikelets/spikes, fertility, as well as plant yield
443 (Masuduzzaman et al. 2016). In a pot experiment conducted by Matsui and Omasa
444 (2002), it was reported that various rice cultivars were selected as tolerant ones
445 under high temperature stress, based on percentage fertility, anther dehiscence, and
446 size. Changes in gas exchange attributes, leaf chlorophyll florescence, relative
447 growth rate (RGR), and total dry weight (TDW) of various tomato clones grown in
448 a greenhouse under high temperature stress were reported by Midmore and Prange
449 (1991). Based on these attributes, they selected three clones and reported that they
Table 27.5 Studies showing the QTL-based selection of stress-tolerant crop genotypes when t5.1
grown under different agroclimatic conditions t5.2
Stress t5.3
Crop type type Studied QTLs Studied attributes References t5.4
Zea mays L. Drought 69 studied; 21 were Agronomic traits Zhao et al. (2018) t5.5
stress stable t5.6
Oryza sativa Cold 35 QTLs were Spikelet fertility, Wainaina et al. t5.7
L. stress identified; two were panicle length, spikelet (2018) t5.8
stable number per panicle, t5.9
panicle number, panicle t5.10
weight, culm length, t5.11
heading date t5.12
Triticum Heat Two marker-trait Grain yield and its Tadesse et al. t5.13
aestivum L. stress associations were components (2018) t5.14
detected on chromosome t5.15
no 4A and 5A t5.16
Triticum Salt 19 QTLs were detected; Maximum root length Ren et al. (2018) t5.17
aestivum L. stress 11 were detected under (MRL), root dry weight t5.18
normal condition and six (RDW), shoot dry t5.19
under salt stress weight (SDW), total dry t5.20
condition. The other two weight (TDW), and the t5.21
QTLs for ratio of TDW ratio of TDW of wheat t5.22
of wheat plants between plants between salt t5.23
salt stress and control stress and control t5.24
(TDWR) (TDWR) t5.25
Oryza sativa Cold 17 QTLs were Spikelet fertility Liang et al. t5.26
L. stress identified; three were (2018) t5.27
tolerant (qCT3.12, t5.28
qCT6.7, and qCT9.6) t5.29
Hordeum Salinity CM72 QTL on Germination, seedling Angessa et al. t5.30
vulgare L. stress chromosome 3H emergence, and first leaf (2017) t5.31
full expansion growth t5.32
stages t5.33
Solanum Heat 13 QTLs were identified Pollen viability, pollen Xu et al. (2017) t5.34
lycopersicum stress number, style length, t5.35
L. anther length, style t5.36
protrusion, female t5.37
fertility, and flowering t5.38
characteristics, i.e., t5.39
inflorescence number t5.40
and flowers per t5.41
inflorescence t5.42
Triticum Heat 24 QTLs were identified Grain yield (GY) and its Bhusal et al. t5.43
aestivum L. stress components, (2017) t5.44
physiological t5.45
parameters t5.46
Triticum Salt 49 QTLs were identified Salt tolerance and Hussain et al. t5.47
aestivum L. stress and 2 were salt tolerant micronutrient (2017) t5.48
concentrations at t5.49
seedling stage t5.50
(continued)
Q. Ali et al.
Stress
Crop type type Studied QTLs Studied attributes References
t5.51 Hordeum Salt 11 tentative and 11 Gas exchange and Liu et al. (2017)
t5.52 vulgare L. stress significant QTLs were stomatal regulation
t5.53 identified and 1 for traits
t5.54 salinity tolerant
t5.55 Oryza sativa Salt 34 QTLs were identified Shoot length, root Rahman et al.
t5.56 L. stress and eight were stable length, shoot fresh and (2017)
t5.57 under salt stress root dry and fresh
t5.58 biomasses
t5.59 Zea mays L. Salt Luo et al. (2017)
t5.60 stress
t5.61 Brassica Salt 45 QTLs were aSTR, salt tolerance Lang et al. (2017)
t5.62 napus L. stress identified, seven for rank; SPAD,
t5.63 aSTR, seven for SPAD, chlorophyll; EC,
t5.64 nine for EC, six for electric conductivity;
t5.65 RDW, two for SOD, RL, root length; SH,
t5.66 three for SP, three for seed height; SP, solution
t5.67 SH, two for RL, three protein; SOD,
t5.68 for LFW, and three for superoxide dismutase;
t5.69 LDW LDW, leaf dry weight;
t5.70 RDW, root dry weight;
t5.71 LFW, leaf fresh weight
t5.72 Triticum Drought 98 QTLs were identified Different agronomic Gahlaut et al.
t5.73 aestivum L. stress that were distributed on traits along with (2017)
t5.74 20 different drought sensitivity
t5.75 chromosomes, 66 QTL index (DSI)
t5.76 agronomic traits, and 32
t5.77 QTLs, which affected
t5.78 drought sensitivity index
t5.79 Indica rice Fe, Zn, 21, 30, and 21 QTLs Growth parameters, Meng et al.
t5.80 and Al were identified for Fe, biomass production, and (2017)
t5.81 Zn, and Al toxicity metal tolerance score
t5.82 tolerance, respectively (MTS)
t5.83 Triticum Heat Three stable QTLs were Fv/Fm (maximum Sharma et al.
t5.84 aestivum L. stress identified quantum efficiency of (2017)
t5.85 photosystem II)
t5.86 Oryza sativa Heat Five QTLs were STI and SSI for % Shanmugavadivel
t5.87 L. stress identified; two were spikelet sterility, STI et al. (2017)
t5.88 stable and SSI for yield per
t5.89 plant
t5.90 Oryza sativa Salt 20 QTLs were detected, Growth and biomass Bizimana et al.
t5.91 L. stress two (qSESI12.1 and production (2017)
t5.92 qSESF12.1) on
t5.93 chromosome; 12 were
t5.94 salt tolerant
t5.95 Lens Drought QTL(QSRLIV-61.63), Different root and shoot Idrissi et al.
t5.96 culinaris stress QTL(QLRNIII-98.64) traits (2016)
t5.97 Medik.
(continued)
Stress
t5.98 Solanum Drought 19 QTLs were identified specific leaf area (SLA), Lounsbery et al.
t5.99 lycopersicum stress on chromosome no. 9 SDW, yield, and (2016)
t5.100 L. maturity traits
t5.101 Zea mays L. As stress 14 QTLs were identified Yield, production, and Fu et al. (2016)
t5.102 phytoremediation
t5.103 Zea mays L. Salt Eight QTLs were Shoot fresh weight, Cui et al. (2015)
t5.104 stress detected. Five were shoot dry weight, tissue
t5.105 associated with salt water content,
t5.106 tolerance concentration of Na+
t5.107 and K+, and their ratio
t5.108 in shoot
t5.109 Triticum Salt 12 QTLs were identified Growth attributes and Turki et al.
t5.110 durum stress biomass production and (2015)
t5.111 salt tolerance indices
t5.112 Pisum Drought Ten QTLs were detected Relative water content Iglesias-García
t5.113 sativum L. stress in soil (RWCS) and et al. (2015)
t5.114 relative water content in
t5.115 leaves (LWCL)
t5.116 Oryza sativa Drought qDTY1.1, qDTY2.2, Yield and yield Kumar et al.
t5.117 L. stress qDTY3.1, qDTY3.2, components (2014)
t5.118 qDTY6.1, and
t5.119 qDTY12.1
t5.120 Oryza sativa Cold Five QTLs were Cold tolerance at Ranawake et al.
t5.121 L. stress detected germination (CTG) and (2014)
t5.122 early seedling (CTS)
t5.123 stages
t5.124 Solanum Drought 18 QTLs were detected Promptness index, Qie et al. (2014)
t5.125 lycopersicum stress radical root length,
t5.126 L. coleoptile length, and
t5.127 lateral root number at
t5.128 germinating stage and
t5.129 seedling survival rate
t5.130 Zea mays L. Drought 45 QTLs were identified Yield and yield Nikolic et al.
t5.131 stress on all chromosomes components (2013)
t5.132 except chromosome 9,
t5.133 five for grain yield per
t5.134 plant and 40 for eight
t5.135 yield components
t5.136 Oryza sativa Heat Two QTLs were Ye et al. (2012)
t5.137 L. tolerance identified on
t5.138 chromosome 1 and
t5.139 chromosome 4; 1 QTL
t5.140 was found tolerant one
(continued)
Q. Ali et al.
Stress
t5.141 Oryza sativa Salt 17 QTLs were RNC, root Na+ Wang et al.
t5.142 L. stress identified, five QTLs concentration; RKC, (2012)
t5.143 might represent novel root K+ concentration;
t5.144 salt tolerance loci, and a SNC, shoot Na+
t5.145 major salt-tolerant QTL concentration; SKC,
t5.146 was also detected shoot K+ concentration;
t5.147 STR, salt tolerance
t5.148 rating
t5.149 Triticum Heat Three QTLs were Heat susceptibility Paliwal et al.
t5.150 aestivum L. stress identified that were index (HSI) of thousand (2012)
t5.151 associated with heat grain weight
t5.152 tolerance (HSITGW), (2) HSI of
t5.153 grain fill duration
t5.154 (HSIGFD), (3) HSI of
t5.155 grain yield (HSIYLD),
t5.156 and (4) canopy
t5.157 temperature depression
t5.158 (CTD)
t5.159 Hordeum Salt Five significant QTLs Concentration of Na+ Zhou et al.
t5.160 vulgare L. stress were identified for and K+ in root and shoot (2012)
t5.161 salinity tolerance
t5.162 Oryza sativa Salt Three salt-tolerant QTLs Salt tolerance Islam et al.
t5.163 L. stress were identified parameters (2011)
t5.164 Zea mays L. As stress Three QTLs were As concentration in the Ding et al. (2011)
t5.165 detected four tissues
t5.166 Glycine soja Salt A significant QTL was Salt tolerance rating Tuyen et al.
t5.167 Sieb. & stress identified (STR) and leaf (2010)
t5.168 Zucc. chlorophyll content
t5.169 (SPAD)
t5.170 Oryza sativa Cd stress Only a single QTL was Grain Cd concentrations Ishikawa et al.
t5.171 L. detected (2009)
t5.172 Helianthus Water Several QTLs were Yield-related traits Kiani et al.
t5.173 annuus L. stress identified (2009)
t5.174 Zea mays L. Drought Umc 11 on chromosome Morphophysiological Tuberosa et al.
t5.175 stress 1 and csu133 on traits and yield (2004)
t5.176 chromosome 2
t5.177 Solanum Salt Five QTLs were Salt tolerance Foolad et al.
t5.178 lycopersicum stress identified parameters (2001)
t5.179 L.
t5.180 Helianthus Water 19 QTLs were detected Photosynthesis and Hervé et al.
t5.181 annuus L. stress water status traits (2001)
450 were tolerant to high temperature stress. Alsajri et al. (2019) reported that different
451 soybean cultivars were selected as high temperature tolerant on the basis of various
452 growth and physiological parameters. Three cultivars of canola were grown in lab
453 under heat stress to find out their tolerance. The cultivar “Oscar” was selected as
tolerant one that was based on oil quality and different yield attributes (Aksouh 454
et al. 2001). Changes in growth, morphology, and yield traits in different tempera- 455
ture stressed mung bean cultivars were observed to select the different tolerant cul- 456
tivars and five cultivars were selected as the tolerant ones based on these changes 457
AU15 (Sharma et al. 2016). In a greenhouse experiment conducted by Kesici et al. (2013), 458
they reported that out of 15 cultivars that were subjected to heat stress to sort out 459
various tolerant cultivars, two cultivars were selected as tolerant ones against heat 460
stress. In another study conducted by Wijewardana et al. (2015), it was reported that 461
various rice hybrids were subjected to cold stress to find out their chilling tolerance 462
potential. Among 33 hybrids, three were selected as stress tolerant depending on 463
different growth parameters and nutrient uptake. Cruz and Milach (2004) reported 464
that out of 24 rice genotypes grown in field under cold stress, three hybrids were 465
selected as stress-tolerant hybrids to check their stress tolerance potential based on 466
different germination parameters. 467
27.5 Breeding Against Heavy Metal Stress 468
AU16 Due to industrialization, the agricultural lands are being contaminated due to toxic 469
effluents and a rich source of heavy metals, acid, and dyes that are severely affecting 470
the crop production as well as health risk for the consumers depending on their soil 471
concentration. High concentration of heavy metals in soil disturbs the nutrient 472
uptake especially the essential ones. At cellular level after the uptake of heavy met- 473
AU17 als the disturbance in ion homeostasis takes place. So, in future due to rapidly 474
increasing in heavy metal, contaminated areas, it is necessary to select the geno- 475
types that have heavy metal stress tolerance mechanism with the ability to reduce 476
heavy metal stress including losses in production. Gul et al. (2016) conducted a 477
greenhouse experiment to assess the interactive effects of heavy metal stress (Cd 478
and Cu) on ecophysiological responses of two maize cultivars. They reported that 479
cultivar S2000 was nominated as tolerant one while the cultivar Okmass was identi- 480
AU18 fied as sensitive one. In another study that was conducted on two maize cultivars 481
grown in pots under Cd and As stress to develop tolerant cultivars, they found that 482
the cultivar Dong Dan and Run Nong 35 were identified as tolerant and sensitive 483
cultivars, respectively, and the selection was based on alteration in different growth, 484
gas exchange, as well as photosynthetic attributes (Anjum et al. 2017a, b). Changes 485
in growth and photosynthetic responses of different maize hybrids were observed to 486
evaluate tolerant hybrids when grown in hydroponic culture under Cd stress. They 487
reported that among seven hybrids, two were selected as tolerant ones. The differ- 488
ential responses of maize hybrids due to Cd stress will be useful in selection and 489
breeding for Cd-tolerant genotypes (Akhtar et al. 2017). A field experiment was 490
conducted on various wheat genotypes grown under Zn and Cd stress to find out the 491
stress-tolerant genotypes based on different growth and physiological attributes as 492
reported by Alybayeva et al. (2016). In a glass house experiment conducted by 493
Hussain et al. (2015), it was found that out of 100 tomato genotypes, eight were 494
Q. Ali et al.
495 selected as tolerant ones, when grown under Cd stress, and selection of these stress-
496 tolerant genotypes was made on the basis of different yield parameters as well as
497 accumulation of Cd in root, shoot, as well as leaf. In mung bean different stress-
498 tolerant genotypes were selected to use in breeding program when grown under Ni
499 stress, and these tolerant genotypes were selected on the basis of various growth
500 parameters and different stress tolerance indices (Samantaray et al. 1998). Hussain
501 et al. (2015) reported that three mung bean cultivars were grown in greenhouse
502 under boron stress to find out their tolerance against stress based on germination,
503 growth, and photosynthetic responses against boron stress. Based on the results, two
504 cultivars, i.e., M-6 and M-8, were selected as tolerant ones.
505 27.6 Q
uantitative Trait Locus (QTL)-Based Selection
506 of Crop Genotypes Under Different Environmental
507 Stresses
508 In genetics, quantitative trait loci (QTLs) have gained great interest by the breeders
509 for over a century as they play the most important role in natural variation in design-
510 ing the phenotype in almost all eukaryotes, including crop plants. A number of
511 traits, those that are important agriculturally, i.e., growth, stress tolerance, as well as AU19
512 plant yield, are controlled by a variety of genes in a given population and these
513 genes are segregated according to Mendelian fashion. Mathar in 1949 defined these
514 traits as “polygenes.” Afterward in 1975, Gelderman called these polygenes as
515 QTLs. It can be defined as a specific region of DNA that is responsible for pheno-
516 typic expression of organisms and also associated with quantitative trait (Dhingani
517 et al. 2015). Several techniques/methods have been employed for the detection of
518 these genomic regions that are responsible for the phenotypes in quantitative term.
519 These methods vary from a simple statistical test or models that include interaction
520 between multiple markers (Martínez et al. 2016). A QTL may be single that shows
521 major effect or present in the form of clusters of genes on genome that have signifi-
522 cant minor effects on a specific trait. The number of QTLs present on a particular
523 chromosome affects the changes in the phenotypic trait that explain the genetic
524 makeup of that trait. For example, it explains how plant height can be controlled by
525 a small number of genes having large effect or by a large group of genes having
526 small effect collectively in a total expression. A variety of QTLs have been found
527 involved with abiotic stress tolerance in crop plants that are associated with com-
528 plex stress-related traits, such as growth, yield, and biomass production under
529 drought, salinity, and heat stress. So, there is a need of developing QTLs that are
530 responsible for improvement of agronomic as well as physiological traits in a num-
531 ber of crop plants such as maize, wheat, and rice subjected to different abiotic
532 stresses (Yao et al. 2016; Gahlaut et al. 2017). As it is well known that developing
533 stress-tolerant crop cultivars is the main focus of breeders especially for better pro-
534 duction under changing environmental conditions that cause the large economic
AU20 losses almost 70% of over the world. Under well-known abiotic stresses at any stage 535
of plant growth disturbs the cellular biochemical reactions that as a result alter the 536
morphology and physiology of plants (Sahoo et al. 2014) and this mechanism is 537
purely genetically controlled that regulate up and down regulations and under 538
adverse environmental conditions (Mizoi and Yamaguchi-Shinozaki 2013). 539
However, the mechanism is stress type, plant species, and growth stage specific 540
AU21 (Gall et al. 2015). In this regard, QTLs play significant learning plant genetic behav- 541
ior and tells how to improve them under stressful conditions (Bo et al. 2015) and to 542
understand the physiological and the genetic elements play significant role in source 543
sink relationship when grown under stressful environment (Welcker et al. 2007). In 544
evolutionary biology, QTL study plays a significant role in finding the location of 545
genes of interest on a specific chromosome. Study of QTLs is an important tool for 546
understanding the phenotypic and genotypic behavior of plants under stressful envi- 547
ronment (Soda et al. 2015). 548
27.7 D
evelopment of Tolerant Crop Genotypes by 549
Manipulating QTLs 550
Quantitative trait locus (QTL)-based selection for stress tolerance is gaining interest 551
due to its role in solving the complex genetic mechanism controlling the stress- 552
tolerant crop plants. QTL-based selection in crop plants is focusing especially on all 553
of the morphological, physiological, and yield potential taken together. These 554
mechanisms are considered important ones but these are purely genetically con- 555
trolled. In this regard, QTL-based selection is considered a short term and easy way, 556
from the available germplasm. For example, in rice 69 QTLs were identified for 557
some agronomic traits such as plant height, for the induction of drought tolerance, 558
ear weight, cob weight, 100-kernel weight, and ear length in two F2:3 populations 559
both under stressed and non-stressed conditions. Out of these 69 QTLs, 52 were 560
identified under water deficit conditions, while 21 were found stable and showed 561
tolerance against water stress (Zhao et al. 2018). In a study conducted by Gahlaut 562
et al. (2017) it was reported that in wheat 98 QTLs were identified on chromosome 563
no. 20 and 66 QTLs for agronomic traits were selected and out of them 32 QTLs 564
affected the drought sensitivity index. Lounsbery et al. (2016) reported that in 565
Solanum lycopersicum L. plants 19 QTLs were identified on chromosome no. 9 that 566
were associated with drought tolerance potential and the identification of tolerant 567
QTLs was based on few traits such as SDW, yield, and maturity traits. Ten different 568
QTLs were identified using composite interval mapping (CIM) and multiple inter- 569
val mapping (MIP) to check the drought tolerance potential of pea plants focusing 570
based on relative water content in soil (RWCS) and leaves (LWCL) as reported by 571
Iglesias-García et al. (2015). Kumar et al. (2014) developed some high-yielding 572
drought-tolerant rice varieties based on QTL analysis. Fourteen QTLs were detected 573
using grain yield as selection criteria. Out of these fourteen QTLs, six were identified 574
Q. Ali et al.
575 as high-yielding ones. In maize, 45 drought-tolerant QTLs (five for grain yield per
576 plant and 40 for eight other yield components) were detected associated with yield
577 and different yield components (Nikolic et al. 2013). A hydroponic experiment was
578 conducted to identify some QTLs associated with salt-tolerant potential of some
579 inbred wheat lines. A total of 19 QTLs were detected, 11 under normal condition
580 and six under salt stress condition, and other two QTLs were identified for
581 TDWR. The selection was based on MRL, RDW, SDW, TDW, and TDWR (Ren
582 et al. 2018). In another study conducted by Angessa et al. (2017), they reported that
583 two barely varieties were grown under salt stress to check their salt tolerance poten-
584 tial based on QTL analysis. A total of 12 QTLs were detected on chromosomes AU22
585 2H–6H based on seed germination, seedling emergence, and first leaf full expansion
586 growth stages, but only one QTL showed stability both under controlled and stress-
587 ful conditions, i.e., CM72 QTL, on chromosome 3H. Different recombinant inbred
588 rice lines were genotyped using QTL analysis to check their stability when grown
589 under salt stress. A total of 39 QTLs were detected, and eight were stable QTLs
590 based on some growth parameters such as shoot length, root length, shoot fresh, and
591 root dry and fresh biomasses (Rehman et al. 2017). Salt-tolerant potential of two
592 rice varieties was checked based on QTL analysis. Five QTLs represented novel salt
593 tolerance loci and a major salt-tolerant QTL was also detected based on RNC, RKC,
594 SNC, SKC, and STR (Wang et al. 2012). Zhou et al. (2012) reported that five sig-
595 nificant QTLs were identified for salinity tolerance in two rice varieties based on
596 concentration of Na+ and K+ in root and shoot. Furthermore, Bizimana et al. (2017)
597 identified 20 QTLs for salinity tolerance in two rice varieties. Among 20 QTLs, two
598 were associated with salt tolerance based on growth and biomass production. In
599 parallel with drought and salinity, QTL-based selection was also found helpful in
600 developing new crop genotypes tolerant to other stresses such as temperature (heat
601 and cold) and heavy metal stresses. For example, in wheat 24 QTLs were identified
602 associated with heat tolerance based on grain yield (GY) and yield components and
603 also on some physiological parameters as well (Bhusal et al. 2017). In another study
604 conducted by Shanmugavadivel et al. (2017), they reported that two rice varieties
605 (Aus cultivar and IR64) were crossed to check their heat tolerance potential based
606 on QTL analysis. Five QTLs were identified associated with STI and SSI for %
607 spikelet sterility, such as STI and SSI for yield per plant. Among these QTLs, two
608 were tolerant ones. Liang et al. (2018) identified different QTLs in rice grown under
609 cold stress. They reported that out of 17 QTLs, three were stable based on spikelet
610 fertility. Wainaina et al. (2018) crossed two rice varieties for the identification and
611 validation of QTLs for cold tolerance at the booting stage and other agronomic
612 traits. They identified 35 QTLs for abovementioned traits. Among these, two were
613 associated with cold tolerance potential. QTL analysis of rice was done to check
614 their heavy metal tolerance potential at the seedling stage. A total of 21, 30, and 21
615 QTLs were identified for Fe, Zn, and Al toxicity tolerance, respectively (Meng et al.
616 2017). Ishikawa et al. (2009) reported that in rice only a single QTL was identified
617 associated with Cd stress toxicity. In maize, 14 QTLs were identified that were
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AU1 Refs. “Gleditsch et al. (2006); Alexander et al. (2006); Hansen
et al. (2006); Peterson et al. (2001); Thomashow (1999);
Metwali et al. (2015); Rehman et al. (2017); Metwali et al.
(2015); Bartha et al. (2015); Luo et al. (2017)” are cited in the
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AU23 References “Aliu et al. (2013), Bano et al. (2015), Basnayake
et al. (2012), Hazra et al. (2008), Hu et al. (2016), Irfan et al.
(2005), Jiang et al. (2018), Khan et al. (2007), Khedher &
Ewing (1985), Latake et al. (2015), Li et al. (2011), Menezes
et al. (2014), Oladzad-Abbasabadi et al. (2018), Rykaczewska
(2017), Sipos & Prange (1986), Tobeh & Jamaati-e-Somarin
(2012), Vahidi et al. (2013), Yang et al. (2018), Zhang et al.
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