Scientia Horticulturae 304 (2022) 111302

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Plant hormone-mediated stress regulation responses in fruit crops- a review

Vinaykumar Rachappanavar a, b, *, Arushi Padiyal b, Jitender Kumar Sharma c,
Satish Kumar Gupta a
a
MS Swaminathan School of Agriculture, Shoolini University of Biotechnology and Management Science, Himachal Pradesh 173212, India
b
Department of Seed Science and Technology, Dr YSP UHF, Nauni, Solan, Himachal Pradesh 173230, India
c
School of Agricultural Science, College of Agriculture, Baddi University, Himachal Pradesh 173205, India

A R T I C L E I N F O A B S T R A C T

Keywords: One of the challenges being faced by the fruit crop production globally is that regional climate regimes are
Signalling becoming more unpredictable every year. The exact impact of these abiotic stresses on fruit crops is difficult to
Cascade predict and it depends on numerous other climacteric factors that are all always counted in predictive models.
Transduction
Plant hormones can improves tolerance against abiotic stresses by inhibiting or reducing the active oxygen
Endogenous and exogenous
species (AOS) accumulation, electrolyte leakage, induce the expression of stress-specific genes, etc. Among the
Transcription factors
major hormones produced by the plants, ABA, SA, JA and ET are known to play major roles in mediating plant
defence response against various abiotic stresses. Typically ABA has major role in contrast to abiotic stresses
because environmental condition such as drought, heat stress, cold, chilling, salinity and heavy metals are known
to trigger increase in ABA levels. Crosstalk networks among phytohormones mediated by diverse key regulators
have been extensively investigated and documented in various research investigations. Many canvasser’s rec­
ommended the use of phytohormones to counteract abiotic stress because of increasing stress tolerance through
conventional breeding methods requires time and a diverse cultivation environment for validation of perfor­
mance and heritability. In this review, we summarize the roles of the major phyto-hormones present in fruit
crops in regulating abiotic stresses responses with special focus on the significance of crosstalk between various
hormones at biochemical and molecular level in generating a sophisticated and efficient stress response.

1. Introduction was observed by abiotic stresses on several occasions in different fruit

Fruit crops are highly vulnerable to different environmental stresses.
These stresses are grouped into two main categories- biotic and abiotic-
based on the nature of stimulants. Biotic stress caused by various fungi,
bacteria, viruses, nematodes, insects, and weeds. Whereas the second
group generally linked with the physiographic, climatic, and edaphic
components of the environment affects plant development and pro­
ductivity (Verma, 2020). In “ The State of Food and Agriculture 2007”,
FAO (Food and Agriculture Organization) reported that less than 3.5%
of the total global land area remains free from abiotic stress for culti­
vation. The concern about these stresses in horticultural crops is
important due to reducing annual production year after year by climate
change (Bray, 2002). Plants have to face naturally occurring stresses
throughout life without a decline in growth and development (Suzuki
et al., 2014; Ali et al. 2020). Up-to 50-70% overall productivity decline
crops (Mittler, 2006). These stresses may cause morphological, physio­
logical, biochemical, and molecular changes in plants (Dos Reis et al.,
2012; Ahmad and Prasad, 2012). Morphological changes like retarded
shoot and root growth, flower drop, low fruit setting, deformed fruit
shape observed during abiotic stress conditions. Nutrient content and
their proportion may change with the intensity of stress. At the physi­
ological level, Photosynthetic rate, respiration, transpiration, and
assimilation rate are affected by these environmental pressures (Har­
fouche et al., 2014; Vinocur and Altman, 2005). To survive under un­
favorable environmental condition plants activate defence mechanism
by producing antioxidant enzymes, hormones and other biochemical
leading to neutralize the negative impact of abiotic stress (Wolters and
Jurgens, 2009; Herbette et al., 2007; Jogawat et al., 2021).
The exact impact of abiotic stresses on fruit crops is difficult to
predict and it depends on numerous other climacteric factors that are all

* Corresponding author at: MS Swaminathan School of Agriculture, Shoolini University of Biotechnology and Management Science, Himachal Pradesh 173212,
India.
E-mail address: vinayrachappanavar@shooliniuniversity.com (V. Rachappanavar).

https://doi.org/10.1016/j.scienta.2022.111302
Received 19 March 2022; Accepted 23 June 2022
Available online 6 July 2022
0304-4238/© 2022 Elsevier B.V. All rights reserved.
V. Rachappanavar et al.
always counted in predictive models. In some environmental circum­
stances, different abiotic stresses appear together and cause greater crop
loss. In this situation, studying individual stress separately without
considering other minor abiotic stress factors is not satisfactory because
plant response is unique and can be predicted the same result by each
stress factor applied individually (Mittler, 2006 and Choudhury, 2017).
Plants have root system architecture (RSA) to respond to changing
environment and it facilitates lateral root branching enables plants to
Fig. 1. Phytohormones and related compounds.
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Scientia Horticulturae 304 (2022) 111302
withstand unfavorable environmental conditions such as salinity,
moisture deficit, and heavy metal toxicity (Lavenus et al., 2013).
Moreover, abiotic and biotic components typically interact in a given
ecosystem (Pandey et al., 2017). Focusing on fruit crops concerning
abiotic stress is important because fruit crops are considered as cash
crops and also provide essential dietary elements in human diet (Shan­
non and Grieve, 1998).
FAO reports that about 90% of dietary vitamins supplied by fruit
V. Rachappanavar et al.
Fig. 1. (continued).
crops. According to the global burden of disease study, nearly 3.4
million deaths can be attributed to a shortage of fruit and vegetable
production and consumption yearly (FAO, 2017). Therefore need to
grow a vast amount of fruit crops under diverse stress conditions be­
comes the most important goal for the current agriculture scenario to
provide a properly balanced diet. Since 2000, extensive studies were
conducted to know the exact impact of climate change encompasses all
abiotic stress on agricultural production and they find some solution to
resolve these problems but only few research articles pay attention to
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Scientia Horticulturae 304 (2022) 111302
fruit crops in terms of quality fruit production under abiotic stress
condition (Parajuli et al., 2019). In that suggestion, increasing phyto­
hormones biosynthesis and external application is the main concern to
increase the tolerance against abiotic stress in fruit crops with a less
economic burden. To maintain the normal plant physiological activities
in plants, it is essential to understand clearly how these stress factors
respond to different hormones at different stages of plant growth (Wien,
1997).
Traditional agronomical approaches do not have the potential to
V. Rachappanavar et al.
Fig. 1. (continued).
completely overcome the impact of environmental stresses. Therefore,
several molecular approaches like marker-assisted selection (MAS), as­
sociation mapping, QTL mapping, and whole-genome sequencing used
to identify the stress tolerant genes. Which helps to develop abiotic
stress tolerant new cultivars by incorporating these genes in existing
high yielding varieties. Till now, limited work was accomplished at
molecular level to know stress tolerance in fruit crops (Gong et al.,
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Scientia Horticulturae 304 (2022) 111302
2010). Still, now only a few QTLs and genes are identified in fruit crops
which involved in stress level regulating activities. The introduction and
improvement of stress tolerance through a conventional breeding
method does not provide the required result because of the linkage drag
of unknown characters from donor parents (Asins, 2002). Whereas
phytohormones plays same role in abiotic stress regulation at a partic­
ular concentration directly or indirectly without any negative impact on
V. Rachappanavar et al.
Fig. 1. (continued).
producing plants. The current review presents an overview on role of
phytohormones in mitigating abiotic stresses in fruit crops.
2. Phytohormones
Plant hormones are bimolecular compounds required in small
quantities (Fahed et al., 2015) to regulate normal physiological activ­
ities (Fraire-Vela´zquez et al., 2011; Bhatt et al., 2020). These organic
molecules manage the abiotic stress by forming complex signaling cas­
cades from stimuli perception to abiotic specific gene expression (Aze­
vedo et al., 2012). They play key roles in plants by modifying growth
and development, source and sink relationship, and allocation of nu­
trients to different organs based on the need to acclimatize the plant to
the environment (Wani et al., 2016). They include gibberellin acid (GA),
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Scientia Horticulturae 304 (2022) 111302
auxin (AUX), cytokinin (CK), abscisic acid (ABA), ethylene (ET), sali­
cylic acid (SA) and jasmonates (JAs) involved in different stress regu­
lation activities directly or indirectly. The phytohormones and related
compounds which identified in different crops were presented in Fig. 1.
Different phytoPhytohormones synthesis and site of action may be the
same or different (Peleg et al., 2011; Rhaman et al. 2021). They act as a
chemical messenger with highly complex regulation mechanism, these
biomolecules allow plants to maintain normal growth plasticity even
under stress conditions (Xu et al., 2016; Khan et al., 2020a). Various
studies (Table 1) have showed that different plant hormones can
improve plant tolerance against chilling, high temperature, salt,
drought, heavy metal and UV-light like abiotic stresses (Ogawa et al.,
2005). Phytohormones regulates active oxygen species (AOS) accumu­
lation and electrolyte leakage in plant cells by inducing stress-specific
V. Rachappanavar et al.
Fig. 1. (continued).
genes expression (Kang et al., 2004). Furthermore, DELLA proteins work
collaterally with GAs, which leads to an increase in tolerance against
environmental stresses (Achard et al., 2009). However, the identifica­
tion of novel genes and its stress responses may prove to be a vibrant
target for engineering abiotic stress tolerance in fruit crops. More in­
formation is needed to make clear regarding the regulation of abiotic
stress at hormonal level in fruit crops along with how and which hor­
mones regulate different abiotic stresses.
2.1. Role of GA in plant defense response
GAs are primarily growth-regulating phytohormone involved in
various developmental processes, including germination, leaf expansion
(Claeys et al., 2012), stem elongation, flowering (Hyun et al., 2016) and
trichome and reproductive organs development (Claeys et al., 2012).
Recently researcher identified the role of GA under stress condition. GA
signaling and metabolism is a vital factor for environmental
stress-tolerant in resilient phenotypes. It modifies the physiological
activiteis in plants with regulating sink and source relationships (Iqbal
et al., 2014). The methylerythritol phosphate pathway synthesizes the
gibberellic acids in plastids from trans-geranylgeranyl diphosphate, thus
both GA and ABA share common precursors during biosynthesis
(Banerjee et al., 2016). The pleiotropic action of GAs is modified by the
degradation of DELLA proteins which belongs to the GRAS protein
family (Colebrook et al., 2014). DELLAs may regulate the gene expres­
sion by acting as transcriptional activators in multiplexes with other
transcription factors (TFs) (Hirano et al., 2012). In the same way, they
also act as inhibitors by sequestering gene activating TFs (Feng et al.,
2008). This proves that, GA signaling crosstalk with other pathways.
DELLAs have been observed to interact with the other phytohormone
cascades at the molecular level (Bai et al., 2012) during stress conditions
(Bai et al., 2010). In recent years, several tissue-specific GA dioxyge­
nases have been identified as CsGA2ox1, CsGA2ox2, CsGA20ox2,
CsGA3ox2, CsGA3ox3, and CsGA2ox4 in Camellia sinensis, these genes
are utilized as a candidate marker gene for abiotic stress tolerance
breeding (Pan et al., 2017). DREB2, an AP2/ERF family protein was
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Scientia Horticulturae 304 (2022) 111302
found to be induced by ABA, but suppressed by overexpression of the
DREB2 gene may cause dwarfness in rice with GA deficient symptoms.
Normal growth restored by exogenous application of GA. Expression of
OsGA20ox1, OsGA20ox2, OsGA3ox2, and OsGA20ox4 was significantly
declined in the plants under stress condition especially to salt stress
(Tang et al., 2019).
2.2. Role of auxin in plant defense response
Auxin is an essential phytohormone that regulates cell elongation,
cell division, differentiation, apical dominance, tropical response, and
other biological activities in plants. Broadly IAA (indole-3-acetic acid)
biosynthesis occurs through tryptophan dependent and tryptophan-
independent pathways (Wright et al., 1991). The maintenance of
endogenous auxin at the proper level is important to maintain cellular
activities (Seidel et al., 2006). Based on the major intermediates
involved in IAA biosynthesis, Tryptophan dependent pathways can be
categorized into indole-3-acetamide (IAM), indole-3-acetaldoxime
(IAOx), tryptamine (TAM), and indol-3-pyruvic acid. (IPA) pathway. A
uxin play major rule during embryogenesis and seed development by
critically regulating organ polarity and root initiation (Chapman and
Estelle, 2009; Zhao et al., 2010). Several researchers found that auxin
also has a key role against biotic and abiotic stress conditions in plants.
Genes involved in various auxin-related pathways show variation in
expression during stress conditions. Auxin stimulates the initiation of
lateral roots from primary roots at seedling stage in plants by promoting
lateral root primordia (LRP) initiation through activation of root peri­
cycle cells present opposite to xylem (Lavenus et al., 2013) with the help
of multiple IAA and ARFs (Auxin Response Factors; Okushima et al.,
2007). ARFs directly and to the auxin response elements to regulate the
transcription of auxin-responsive genes. These ARFs contains (1) a
variable middle transcriptional regulatory region (MR), (2) N-terminal
DNA-binding domain (DBD) and (3) a C-terminal dimerization domain
(CTD) (Zhang et al., 1998). AuxRE region of auxin- promoter directly
target by DBD domain and regulate the expression of these regions. The
members of Aux/IAA and ARF class showed variation in expression
V. Rachappanavar et al. Scientia Horticulturae 304 (2022) 111302

Table 1
Role of different phytohormones in combating different abiotic stresses in fruit-crops.
Phytohormone Fruit crops Effects against abiotic stresses Refs.

1 ABA Grape wine Improves drought tolerance by upregulating certain key genes like chlorophyll synthesis, HSPs and other Nassar et al. (2004)
defence-related genes
2 ABA Mango Increase water stress tolerance Zaharah and Razi (2009)
3 ABA Citrus improves water stress tolarance by increasing ABA concentration in young leaves Arbona and
Gómez-Cadenas (2008)
4 ABA Crabapple improves water stress tolarance by increasing ABA concentration in young leaves Bai et al. (2013)
5 ABA peas icreased accumulation of ABA in fresh leaves improves water tolarance Jackson et al., (1988)
6 ABA Citrus increased anoxia tolerance Gómez-Cadenas et al.
(1996)
7 ABA Almond regulate xylem sap movement under water stress condition Wartinger et al. (1990)
8 ABA Citrus Negutiviely regulate water stress Alferez et al. (2005)
9 ABA Apple Promote drought resistance by overexpressing pRD29A::PYL9 gene Hou et al. (2020)
10 ABA Strawberry Maintain fruit quality even under salt and drought stress condition Perin et al. (2019)
11 ABA Grape Exogenous ABA improves resistance againest various abiotic stresses Wang et al. (2016)
12 ABA sweet cherry Increase the expression of PacSnRK2 and PacPP2C gene families againest dehydration stress Shen et al. (2017)
13 ABA Apple Improve salt and droght stress resistance by upregulating MdSAT1 gene expression Yang et al. (2021)
14 ABA Apple enhance drought tolerance and WUE by overexpression of RNA binding protein MhYTP1 Guo et al. (2019)
15 ABA Chinese crab Lowers the inflex of Cd heavy metal into plant cell by inhibiting 9-cis-epoxycarotenoid dioxygenase Zhang et al. (2019)
apple (MhNCED3) activity
16 ABA Apple Regulat both osmotic and salt stress by overexprssing MdCAIP1 gene Liu et al. (2019)
17 ABA Sweet orange ncrease Late Embryogenesis Abundant (LEA) proteins expression under salt and water stress condition Pedrosa et al. (2015)
18 Auxin Banana Regulate salt stree Hu et al. (2015)
19 Auxin Banana Heat stress regulation Hu et al. (2010)
20 Auxin Apple minimize drought stress by maintaining root development rate and architecture Shi et al. (2014)
21 Auxin Apple lowers salt stress Pauwels and Goossens
(2011)
22 Auxin Apple regulate photooxidative and heat stress by crosstalking with ethylene Torres et al. (2013)
23 Brassinosteroids Fruit crops Regulate chilling injury in varous fruit crops Fariduddin et al. (2014)
24 Brassinosteroids Fruit crops Application of 24-EBR augment osmo-regulation material and minimize damage caused by lipid Xia et al. (2009)
peroxidation and reactive oxygen species
25 Brassinosteroids sweet cherry lowers oxidative damage caused by cold stress when stored at 1◦ C Roghabadi and Pakkish
(2014)
26 Brassinosteroids Fruit crops Regulate vsrious abiotic stresses by cross-talking with ethylene Jiroutova et al. (2018)
27 Brassinosteroids Banana Minimize thermo-stress by regulating photosynthesis process Gonzalez-Olmedo et al.
(2004)
28 Brassinosteroids Orange Enhances cold stress by regulating antioxidant enzymes Ghorbani and Pakkish
(2014)
29 Brassinosteroids Orange play important role in response to cold stress by increased accumulation of secondary metabolites Jiang et al. (2021)
30 Brassinosteroids Strawberry Exogeneous application of 24-EBL minimize cold damage by increasing malondialdehyde in leaves Balcı et al. (2021)
31 Brassinosteroids Grapewine Enhance chilling stress tolerance by regulating sugar, protein and free proline accumulation Xi et al. (2013)
32 Brassinosteroids Balete tree Alleviate high temperature stress by increasing glyoxalase (Gly I and GlyII) and antoxidants (CAT, SOD, Jin et al. (2015)
GPX, etc.) activites.
33 Brassinosteroids Peach Avoid cold stress by decreasing proline dehydrogenase and enhanced Δ1-pyrroline-5-carboxylate Gao et al. (2016)
synthetase activities
34 Brassinosteroids loquat improve the tolerance againest salinity stress by increasing soluble sugars, pigments and leaf and root K+ Sadeghi and Shekafandeh
content (2014)
35 Brassinosteroids Apple minimize water stress by regulating photosynthesis mechanism Kumari and Thakur (2019)
36 Brassinosteroids tea crabapple exogenous BL application alleviate salt stress tolerance by eliminating reactive oxygen species (ROS) Su et al. (2020)
production by strengthening superoxide dismuatase and catalase activities
37 Brassinosteroids Grape improves the fruit self life under cold storage by elicitation of antioxidant enzymes including ascorbate Pakkish et al. (2019)
peroxide, superoxide dsmutase, catalase and peroxidase
38 Ethylene Apple Regulate water stress Wang and Steffens (1985)
39 Ethylene Apple Improves cold tolerance by upregulating MdERF1B–MdCIbHLH1 genes Wang et al. (2021)
40 Ethylene Apple salt stress tolerance by upregulating MdNAC047 gene An et al. (2018)
41 Ethylene Apple uoregulate aluminium stress tolerance by upregulating MdMYC2 gene An et al. (2017)
42 Ethylene Papaya Overexpression of CpERFs observed under exogenous application of ethylene Adkins et al. (1997)
43 GA Apple Improves germination under cold stress Wani et al. (2014)
44 GA Toon Induce tolerance to chilling stress Hu et al. (2018)
45 GA datepalm The exogenous application significantly promote plant growth under strss condition Khan et al. (2020b)
46 GA Apple Improves salt tolerance by upregulating MdBZR1 and MdBZR1-2like Transcription Factors Wang et al. (2019)
47 Brassinosteroids Banana Improves heat tolerance @0.2µM Nassar et al. (2004)
48 JA Apple Regulate sunburn by regulating ABA biosynthesis Torres et al. (2013)
49 JA Fruit crops Exogenous methyl jasmonate improves antioxidant activites and increases secondary metabolites Reyes-Díaz et al. (2016)
production
50 Jasmonic acid Apple Increase water stress tolerance by regulating xylem sap movement Yanping et al. (2004)
51 Jasmonic acid Peach Alleviate chilling injury by promoting phospholipid remodeling Chen et al. (2019)
52 Jasmonic acid Banana alleviate polyethylene glycol-mediate water stress and reduces oxidative stress Mahmood et al. (2012)
53 Jasmonic acid Siberian Exogeous application of MeJA increase ascorbate peroxidase (APX),glutathione reductase (GR) and Li et al. (2017)
crabapple monodehydroascorbate reductase (MDHAR) activities and decreaseoxidized gluthione content in cells
54 SA Apple Burst of SA content was noticed againest heat and photooxidative stress Torres et al. (2013)
55 SA Banana Minimize heat and photooxidative stress in leaves Mahouachi et al. (2014)
56 SA Grape Improves cold stress tolerance Mutlu et al. (2013)
57 SA Kiwifruit external application of methyl salicylate avoid cold stress by regulating ABA and ethylene production Zhang et al. (2003)
(continued on next page)

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Table 1 (continued )
58 Salicyclic acid Orange Mitigate the heat stress Abd El-Naby et al. (2020)
59 Salicyclic acid Apple improves the thermotolerance by overexpressing MdVQ37 gene Dong et al. (2021)
60 Salicyclic acid Grapewine Alleviates decrease in photosynthesis under heat stress condition Wang et al. (2010)
61 Salicyclic acid Red bayberry minimize the water stress by increasing photosynthesis ratem superoxide dismutase activity, catalase Ying et al. (2013)
activity, stomatal conductivity and methane dicarboxylic aldehyde content
62 Salicyclic acid Strawberry Amelorate salt stress by increasing leaf water relative content (LWRC) in plants Karlidag et al. (2009)
63 Salicyclic acid Banana Exogenous application of SA significantly improves the anthocyanin and proline content under water Bidabadi et al. (2012)
stress condition
64 Salicyclic acid Strawberry SA incombitaion with Fe nano-particles minimize the drought stress under in-vitro condition Akbar Mozafari et al.
(2018)
65 Salicyclic acid Peach Alleviates chilling iinjury by activating heat shock proteins Wang et al. (2006)

during cold stress conditions in Arabidopsis (Hannanh et al., 2005).
Cheong et al. (2002) showed fluctuation in auxin-related genes expres­
sion, including IAA2, IAA3, GH3-like protein, and SAUR-AC1, a+fter
damage caused by the environment (Kovtun et al., 1998). During
oxidative stress conditions, expression of PIN1, and PIN3 downregulated
(Pasternak et al., 2005). Genome-wide study shows that the gluta­
redoxin (GRX) gene family exhibits differential expression patterns
under different phytohormones treatment. Among them, OsGRX8, a
member of GRX, has expression during different abiotic stress conditions
(Garg et al., 2010). WES1, encoding auxin conjugating enzyme, is
induced by drought, heat, cold, and oxidative stress conditions (Park
et al., 2007). Under ABA, drought, and salt treatments, numerous auxin
transport genes showed differential regulation (Shen et al., 2010).
2.3. Role of abscisic acid in plant defense response
Abscisic acid (ABA) is a small organic molecule and classified as
sesquiterpene (Roychoudhury et al., 2013). The C-15 ABA skeleton is
commonly found in the biosynthetic precursors for abscisic aldehyde,
xanthoxin, abscisic alcohol, phasic acid, 8′ -hydroxy-ABA, and dihy­
drophaseic acid. The level of ABA increased in the plant system as stress
signals appeared (Roychoudhury and Basu, 2012). The formation of
carboxylic-acid from abscisic acid oxidation is the last step of ABA
biosynthesis. The process of ABA synthesis is carried out by an abscisic
aldehyde oxidase (AAO3) enzyme. AAO3 protein consists of molybde­
num (Mo) as a cofactor and activated through MoCosulfurase reaction
(Tan et al., 1997). It plays an important role in mediating stress re­
sponses (Yang et al., 2011). When environmental condition unfavorable
for plants growth, the ABA biosynthesis (Kim et al., 2010) and accu­
mulation in various plant tissues increases drastically and bind to its
receptors to initiate signal transduction leads to stress response at the
cellular level (Ng et al., 2014). Diverse ZEP genes regulate the ABA
biosynthesis in different species and plant growth stages but NCED genes
have a major role in drought stress conditions in tomato (Burbidge et al.,
1990), bean (Qin and Zeevaart, 1999) and avocado (Chernys and Zee­
vaart, 2000). A remarkable rise in the NCED transcript level observed
under induced dehydration conditions (Xiong and Zhu, 2003). But the
NCED expression not varied by exogenous ABA application (Iuchi et al.,
2010). Exogenous ABA regulates the ZEP, AAO3, and MCSU genes. Both
ABA degradation and biosynthesis have a significant role in ABA
expression and adjusting plant stress responses and developmental
strategies. ABRE and DRE like cis-elements are promoters of
stress-inducible ABA genes (Xiong et al., 2001; Bray, 2002).
2.4. Role of ethylene in plant defense response
Ethylene has the same activities as ABA for senescence but it has a
different site of action and regulates diverse biotic and abiotic stresses
(Brodersen et al., 2005). It is biosynthesized from methionine amino
acid with the help of S-adenosyl methionine (SAM) and 1-aminocyclo­
propane-1-carboxylic acid (Kende et al., 1993). Various environmental
stresses promote ethylene biosynthesis (Khan et al., 2013). Abiotic stress
promotes the ethylene production by modulating the activities of ACC
synthase and ACC oxidase. Underwater deficit condition, increase in
ethylene level observed in avocado and orange (Upreti et al., 1998).
Abeles et al. (1992) revealed that ethylene regulates the PR gene tran­
scription in response to pathogen attack, wounds, and drought stress.
(Sessa et al., 1995). One of the subfamilies of AP2/ERF transcription
factors, Ethylene Responsive Factor VII group (ERF-VII) participates in
growth and development processes and stress response mechanisms in
plants (Yao et al., 2016). These proteins contain a conserved APETALA2
(AP2) domain necessary for protein-DNA interaction (Papdi et al.,
2015). At the N-terminal, ERF-VII has highly conserved motif
MCGGAI/V (N-degron) to regulate protein turnover and initiate a pro­
tein degradation pathway through an oxygen-sensing mechanism
(Gibbs et al., 2015). The ERF-VII proteins not only participate in regu­
lation under hypoxia stress but are also involved in biotic and abiotic
stress and hormone response. At the vegetative stage, they enhance the
drought-responsive genes expression to recovery from dehydration
stress (Fukao et al., 2011).
2.5. Role of salicylic acid in plant defense response
Salicylic acid is a phenolic compound and the most common
pathway in plants for SA synthesis is the phenylalanine pathway and
isochorismic acid pathway (An et al., 2011). SA artificial derivatives
used for preparation of aspirin, codeine, morphine, digitalis and taxol
like drugs. SA concentration in various crops is about 1 mg per 1 kg of
plant biomass (Ahmad and Prasad, 2012b). It involved in different
plants physiological and molecular activities, such as seed germination,
seedling growth respiration, stomatal aperture, and senescence (Boat­
wright and Pajerowska-Mukhtar, 2013; Arif et al., 2020). Most of the
research evidence shows that SA can enhance tolerance in plants against
various external stresses including salt stress, osmotic stress, chilling,
heavy metal stress (Chen et al., 2016), and high light by activating the
antioxidant system to protect photosynthesis (Gururani et al., 2015).
Salicyclic acid has a key role in plants to gather hypersensitive response
(HR) or systemic acquired resistance and it also modulates various
physiological responses such as thermogenesis, ion absorption, and
programmed cell death during stress conditions.
2.6. Role of jasmonic acids in plant defense response
Jasmonates (JA) are derivatives of fatty acids, including jasmonic
acids (JA), jasmonate isoleucine conjugate (JA-IIe), and methyl jasmo­
nate (MeJA) as key compounds (Ruan et al., 2019). It is an endogenous
signaling molecule involved in the various developmental process and
also in stress mitigating activities (Llanes et al., 2016). JA is closely
associated with plant resistance to abiotic stress by the physiological and
molecular response. Under stress condition, JA alleviates the antioxi­
dant system, accumulation of isoleucine, methionine, soluble sugars,
regulate the stomatal activities (Acharya and Assmann, 2009), expres­
sion of JA-associated genes (JAZ, AOS1, AOC, LOX2 and, CO11) (Hu
et al., 2017) and interaction with other phytohormones (Yang et al.,
2019).

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2.7. Role of brassinosteroids in plant defense response
Brassinosteroids (BRs) are a group of steroidal hormones that play
pivotal roles in wide ranges of developmental phenomena including cell
division and elongation, photomorphogenesis, reproductive develop­
ment, and leaf senescence and also external stress response (Choudhary
et al., 2012; Ahammed et al., 2020; Hussain et al., 2020). BRs increase
the activities of catalyzing and decrease the activities of peroxidase and
ascorbic acid oxidase under stress conditions (Ahammed et al., 2013)
and also regulate secondary metabolites (Vardhini et al., 2006). Bras­
sinosteroids regulate physiological activities in plants by 28-homobras­
sinolide (HBL) and 24-epibrassinolide (EBL) isomers (Siddiquiet al.,
2018). These role of these isomers in abiotic stress regulation is reported
in several crops (Kolomeichuk et al., 2020). Under salt stress condition,
BRs maintain the PSII photochemistry efficiency and leaf osmotic po­
tential. It induces methylation and epigenetic regulation under salt
stress condition (Alam et al., 2019). Exogenous application of 24-epi­
brassinolide (EBL) upregulate antioxidation and detoxification system
and decrease the accumulation of H2O2, NO and MDA in plant cells
(Sharma et al., 2017). 24-EBL can restore the stress induced by
im-idacloprid (IMI) challenged environment by avoiding Cr accumula­
tion in plants (Jan et al., 2020).
2.8. Hormonal crosstalk in plant defense mechanism
The signaling pathways of different phytohormones are known to
interact among themselves in various situations, such as biosynthesis of
hormone-responsive transcription factors to regulate plant defense re­
action. To regulate the balance between seed dormancy and germina­
tion, the cross talk of ABA with GA, mediated by DELLAs is a key
mechanism for evading early abiotic stress conditions (Khan et al.,
2020a). The signaling pathways of SA and JA are known to interact at
various points to regulate the numerous biotic stress antagonistically
(Bari and Jones, 2009). Different studies evidenced that SA-facilitated
suppression of JA-responsive genes like PDF1.2, LIPOXYGENASE 2
(LOX2), and VEGETATIVE STORAGE PROTEIN (VSP) was abolished in
npr1 mutant plants (Spoel et al., 2003). Both JA and ET pathways sta­
bilize EIN3 and thus exhibit resistance to necrotrophs (Zhu et al., 2011).
Besides, ET also suppresses the ABA action in seed and thereby improves
the germination by releasing dormancy (Arc et al., 2013). Auxin asso­
ciate with ethylene to regulate root development under drought and
salinity stress conditions. Furthermore, benzothiadiazole S-methyl ester
(BTH), SA analog, treated Arabidopsis suppress the several
auxin-responsive genes. SA signaling minimizes the TRANSPORT IN­
HIBITOR RESISTANT 1 (TIR1)/ AUXIN SIGNALING F-BOX (AFB) gene
expression (Wang et al., 2007a, 2007b).
The role of cytokinins in stress response has been reported by several
studies (Reusche et al., 2013). They are also known to crosstalk with SA
and others signaling cascade to regulate plant defenses. For instance,
ARABIDOPSIS RESPONSE REGULATOR 2 (ARR2) (cytokinin–activated
transcription factor), a type B ARR, interacts with TGA3 (bZIP-type
transcription factor) and promotes SA defense responses in an
NPR1-dependent manner (Choi et al., 2010). A majority of the time, ABA
suppresses the CYTOKINE OXIDASE. ARABIDOPSIS HISTIDINE KINASE
(AHKs) functions as cytokinin receptors and regulates drought and
salinity response and also ABA signaling, while AHK2 and AHK3 decent
osmotic stress response and ABA signaling (Tran et al., 2007). ARABI­
DOPSIS HISTIDINE PHOSPHOTRANSFER PROTEINs (AHPs) negatively
control responses to drought stress by losing activity of AHP genes
resulted in the up-regulation of ABA-responsive genes and thus a strong
water stress tolerance phenotype (Nishiyama et al., 2013).
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3. Hormone functions and cross-talk toward stress tolerance in
fruit crops
3.1. Hormonal regulation of growth responses under water stress
condition
Farooq et al. (2009a, 2009b) reported that depend on the intensity
and duration of the water stress, 13 to 94% reduction in crop yield is
recorded in several crops. Water is a fundamental component which
requires to maintain normal physiological activities and membrane
transport processes in the plant (Jones and Tardieu, 1998). Since fruit
products are usually sold on a fresh weight basis in the market and yield
is predominantly determined by water content (Marcelis et al., 1998). In
arid and semiarid tropics, plant experiences drought stress either when
the insufficient water supply to roots or increased rate of transpiration
with temperature (Mubarik et al., 2021). Due to low precipitation, high
temperature, high wind velocity, and low soil moisture storage capacity
and unequal distribution of water availability during the life cycle of a
crop plant that restricts the complete expression of the genetic potential
of the plant (Sinha and Watson, 2007). Plants adopted morphological
(reduced leaf area, leaf molding, increased waxy content, reduced stem
length, high root volume and density, less number of branches, erect
shoot, efficient rooting system, small plant size, early maturity), physi­
ological (water use efficiency, leaf water potential, relative water con­
tent, osmotic adjustment, controlled transpiration, efficient stomatal
activity, and osmotic adjustment) and biochemical responses (antioxi­
dants, accumulation of proline, ABA, polyamine, trehalose, glycine,
betaine storage of carbohydrate at cellular and organism levels and
increasing of nitrate reductase activity) to mitigate or minimize the ef­
fect of drought stress (Conesa et al., 2016; Ozturk et al., 2021). Drought
is responsible for the drastic reduction in productivity (Medici et al.,
2014). Flooding also produces the same impact as drought on fruit crops
by creating an anaerobic respiration zone around the root (Issarakraisila
et al., 2007) and upsurges the soil-borne diseases (Rao and Li, 2003).
Thereby, relevant reports about several hormones influence the water
stress regulation in fruit crops that have been produced and some are
presented next.
Some studies show that ABA-induced auxin regulates the root growth
and root cell differentiation (Xu et al., 2016; Gupta et al. 2020) and
modulate proton secretion at root tips under drought stress condition.
Also, ABA-mediated auxin transporter inhibits lateral root growth under
moderate water stress conditions (Shkolnik-Inbar and Bar-Zvi, 2010). At
the genetic level, ABI3 (ABA insensitive 3) interact with Aux/IAA or ARF
proteins and LR growth inhibited in mutants of ABI3 mutants by
attenuation of auxin responses. In contrast, ABI4 overexpression harms
LR development. Some of the research work provided evidence that
drough t can affect the expression of core abscisic acid signaling con­
stituents, equivalent to ABA, PYR/PYL/RCAR ABA receptors, subclass III
SnRK2 protein kinases and protein phosphatases 2C (PP2C) (Weiner
et al., 2010). ABA non-functional mutants are extra vulnerable to
drought and salinity as compared to transgenic plants which can induce
much hormonal response (Qin and Zeevaart, 2002). PYR/PYL/RCARs,
PP2C, SnRK2s, and bZIP transcription are incorporated by the
ABA-facilitated signaling pathway during water stress conditions (Fujita
et al., 2014). Zhang and Huang (2010) proved that OsPIN3t, auxin influx
carrier involved in drought stress conditions and overexpression of
OsPIn3t upregulate drought-responsive genes, OsDREB2A and OsAP37
and improve the tolerance against drought. ABA is known to regulate the
balance between intrinsic growth and external responses. AtABCG25
acts as ABA transporter in cell membrane and export ABA from the
cytoplasm to outside of the cell. Reduced transpiration observed when
AtABCG25 overexpressed. Kuromori et al. (2016) observed that
AtABCG25 over-expression stimulated a local ABA response in guard
cells and increase the drought tolerance, probably resulting from the
maintenance of water content.
Jasmonic acid also regulates the water loss under drought conditions
V. Rachappanavar et al.
by regulating stomatal opening and closing (Savchenko et al., 2014).
The JA concentrate increases in different tissues of plants rapidly
following drought stress and return to a normal level if stress periods are
prolonged. Jasmonate ZIM-domain proteins (JAZ) are regulators (typi­
cally repressors) in the JA signaling pathways. OsJAZ1 play a negative
role for stress tolerance in rice under drought condition with the ABA
and signaling pathways (Fu et al., 2017). Seo et al. (2011) demonstrated
that OsbHLH148 acts as a transcriptional regulator and up-regulates
OsDREB1 and OsJAZ under dehydration condition. Furthermore, Ge
et al. (2010) found that JA accumulation could promote leaf senescence,
improve plant survival by preventing excessive water loss. Foliar
application of MEJA on soybean plants can improve water stress toler­
ance with an increase in sugars, phenolic compounds, and flavonoids in
leaves (Mohamed et al., 2017).
Mango orchids present in the rainfed area can survive for a long
period by adopting drought-tolerant mechanisms (Chacko, 1986) and
they set fruit when daily average temperature more than 20◦ C. So they
need water stress along with high temperatures for flowering (Schole­
field et al., 1986). Severe flooding for more than 2 weeks can signifi­
cantly reduce root dry weight (Larson et al., 1991), net CO2 assimilation,
stomatal conductance, and transpiration rate(Schaffer, Azuma and
Madura, 1995). Grapes plants develop large-sized vessels in the xylem to
transport water under drought conditions (Serra et al., 2014). Grafting
high yielding commercial varieties scions on drought-tolerant rootstocks
helps to overcome the water stress (Kodur et al., 2010). ABA deposit
with prolonged water stress in different plant organs (Torrey, 1976),
Initially ABA concentration is an increase in stressed roots, and later it is
transported to the leaves to reduce the photosynthesis and partially
transpiration. Pomegranate is considered to be both winter hardy and
drought-tolerant plants (Aseri et al., 2008) but they need regular opti­
mum irrigation during the dry summer season (Sulochanamma et al.,
2005). Mild to severe drought stress during the reproductive stage may
reduce the fruit size by 10–50% in cluster apple. Whereas 50 ppm GA3
application along with wheat straw mulch gave the highest net income
(Patel et al., 2010) with good fruit setting during water stress conditions.
Custard apples are very sensitive to salt in irrigation water and should
not exceed an electrical conductivity of 800 μS/cm (George et al., 2004).
Ber plants are comparatively more resilient for water stress by producing
pubescent leaves with a large number of stomata arranged in the
interveinal regions, whereas the adaxial surface is glabrous, with few
comparatively sunken stomata. (Clifford et al., 2002). The date palm
tree has naturally evolved as drought and salt-tolerant plants through
selection (Zaid and deWet, 2002) with an adaptation capacity more than
barley for saltwater (Furr and Armstrong, 1975). In litchi, ABA content
in leaves increased with the drought (Stern et al., 2003) by inhibited
CO2 assimilation in litchi leaves (Damour et al., 2010).
Application of paclobutrazol (gibberellin biosynthesis inhibitor) can
induce stress protection in apple (Swietlik and Miller, 1983), strawberry
(Navarro et al., 2007), and peach (George and Nissen, 1992) under
drought conditions. In flood soil, PM ATPase stimulates the accumula­
tion in putrescine to support cell homeostasis and nutrient acquisition
(Bertini et al., 1997) with a reduction in superoxide radicals and H2O2
accumulation (Yiu et al., 2009). GA interferes with the ethylene and ABA
biosynthesis by upregulating ethylene response factor (ERF) domain
proteins SNORKEL1 and SNORKEL2 to minimize the negative effect of
excess water. During drought, IAR3 (IAA-Ala Resistance3) involved in
lateral root production, free auxin production (Kinoshita et al., 2012). It
stimulates the gibberellin production and inhibits the ABA synthesis for
plant accumulation under submerged conditions (Colebrook et al.,
2014). ABA level shows a decrease with an increase in hydration, but PA
or DPA levels either increase or remain unaltered (Zhou et al., 2005).
ABA restrict their water loss through controlled transpiration by closing
stomata at high temperature and increased root hydraulic conductivity
(Thompson et al., 2007). It regulates the influx or efflux of K+ balanced
by the flux of anions regulates guard cell volume. ABA and root growth
relationship different from the ABA and shoot relationship, as increase
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the level of ABA promote root growth and maintain low water potential
((Biddington and Dearman, 1985; Watts et al., 1981). The decline in
cytokinin concentration under soil moisture deficit conditions observed
in grapevines (Satisha et al., 2005) and partial root drying harmed the
xylem cytokinin concentration with a distinct increase in xylem sap pH
(Stoll et al., 2000). During water stress conditions enhances ethylene
production in orange (Ben-Yehoshua and Aloni, 1974), avocado (Adato
and Gazit, 1974), and in many other plant species leads to increased
senescence of fruits/ leaves and reduced growth. The ethylene produc­
tion indirectly depends on the magnitude of stress, stress intensity, and
stress duration (Upreti et al., 1998, 2000).
Under flood, condition roots trigger the ACC synthesis, which later
oxidized to ethylene when it reaches shoots and cause nastic movements
of the leaves and promote aerenchyma tissue formation to mitigate
oxygen deficiency (Colmer et al., 2004). External application of poly­
amines ameliorates the drought stress in pomegranate by increasing
proline concentration (Amri and Mohammadi, 2012) Polyamines sta­
bilize membranes and forming potential of higher buffering and anti­
oxidant capacity (Chaitali and Sengupta, 2014). The balance of auxin
levels (IAA) required for the adaptive recovery growth of litchi plants
after release from the stressed environment. Although the levels of cy­
tokinins decreased and levels of ABA increased (Krishna, 2012).
Some signaling molecules include inositol (Sengupta et al., 2008);
polyamines (Rangan et al., 2014), and hormones like ethylene (Xiong
et al., 2014). Abscisic acid (Saradhi et al., 2000), and methyl jasmonate
(Wu et al., 2008) involved in minimizing the water stress with minimum
biomass loss. They may induce the production like protective proteins
such as dehydrins; late embryogenesis abundant (LEA) proteins; heat
shock proteins (HSPs) (Juszczak and Bartels, 2017). LEA proteins,
dehydrin, lipid transfer proteins saturate enzymes, RAB (responsive to
ABA) are associated with the protection of cellular structures (Cushman
and Bohnert, 2000). AREB1/ABF2, AREB2/ABF4, and ABF3 like bZIP
transcription factors are induced by dehydration, salinity, or ABA
treatment under drought condition (Yoshida et al., 2010).
Genomics and crop physiology studies of waters stress provide ac­
curate information to breeders for plant improvement (Tuberosa and
Salvi, 2006). Plants have sets of drought regulatory genes, which acti­
vated during the water-deficient condition to minimize the stress con­
dition (Wohlbach et al., 2008). In plants at the initial phase of drought,
water stress stimulate the production of ABA, which induce the
expression of genes (Ding et al., 2016) like, RD29A, RD29B, KIN2 and
RAB18, rd22 (Abe et al., 1997) and PYL8 (Lim et al., 2013). Mutants of
Arabidopsis erecta were studied under controlled conditions to know the
response of ERECTA gene complementation (Masle et al., 2005). SUBA-1
allele induces the negative regulation of ethylene in plants to survive
under prolonged submerged condition (Xu et al., 2016). Overexpression
of the ESKIMO1 gene in Arabidopsis (Bouchabke-Coussa et al., 2008)
and pro biosynthesis gene in chickpea (Cicer arietinum; Bhatnagar-Ma­
thur et al., 2009) were observed during artificial drought condition.
Several transcription studies show, how plants recover after mild
drought condition as compared to complete irretrievable inhibition of
photosynthesis under severe drought (Watkinson et al., 2003). Most of
the time, MYC, MYB, DRE-binding protein (DREB)/ C-repeat-binding
factor (CBF), ABA-binding factor (ABF) and mainly cis-elements
ABA-responsive element (ABRE) transcription factors present in ABA
biosynthesis cycle are involved in drought stress management (Bartels
and Sunkar, 2005; Yadav et al., 2020). Most of the genes involved in
drought stress management are identified by dehydration and/or os­
motic treatments (Shinozaki and Yamaguchi-Shinozaki, 2007). Path­
ways involved in this process were discovered with the help of
Arabidopsis and rice plants (Nakashima et al., 2014) withholding water
to create artificial stress at a certain level to know the expression of
different genes (Bouchabke et al., 2008). One of the most effective
methods is progressive drought (pDr), in which water is withheld until
plants show drying or wilting symptoms (Ning et al., 2010) but it does
not create the same condition as observed under field condition.
V. Rachappanavar et al.
Automated controlled drought screens like PHENOSIS are developed to
create the same drought condition as occurred in the field and used in
screening different genotypes of species to now the genes involved in
drought conditions(Granier et al., 2007). These results suggested that
water stress in fruit crops could modify the phytohormonal activities and
modify the concentration of different hormones to maintain the normal
physiological activities in plants.
3.2. Hormonal regulation of growth responses under high-temperature
stress condition
The temperature has a major influence next to moisture for plant
growth and development. In optimum temperature, the plants may grow
faster and comes to flowering even earlier than normal. Those plants
grown under cool conditions, they produce more upright leaves, but
under warm humid conditions, they are more horizontal to downward
(Ravi et al., 2013). Under subtropics, banana plant development is
slower than in the tropics irrespective of yield and bunch size and they
need optimum temperature around 21–22◦ C and stops above 38 or 39◦ C
and below 9 or 10◦ C for proper. When temperature crosses the sunburn
critical temperature (>38◦ C) causes chlorophyll degradation and ba­
nana choking. Pomegranate can tolerate drought by morphological
and/or physiological modification which enables them to avoid or
postpone desiccation in the semiarid zone (Save et al., 1995). Custard
apple normally requires a hot and dry climate during flowering and high
humidity for a good fruit setting. Even they can tolerate extreme tem­
peratures (below freezing and up to 40◦ C). Soil temperatures below
10◦ C may cause severe chilling injury leading to loss of cell membrane
stability (Levitt, 1980).
Most of the time-temperature in the plant is regulated by the tran­
scription regulators like HSFs by activation of many heat shock
responsive genes and heat shock proteins. Genes that are responsible for
minimizing heat shock are conserved near the N-terminus called the
DNA-binding domain (DBD). A large number of HSFs and Hsps genes are
identified in different crops from available genome sequence informa­
tion and they involved in heat shock regulatory network (Hu et al.,
2017; Jha et al. 2021). there are 25 HSFs, 29 sHsps, 26 Hsp70, 9 Hsp90,
and 10 Hsp100 family genes that were identified in rice crop, 35 DcHCFs
in carrot, 21HSFs in Arabidopsis thaliana, 28 in Populus trichocarpa, 16 in
Medicago truncatula, 52 in Brassica rapa, 25 in Zea mays, and 26 in
Glycine max, respectively (Huang et al., 2015). When temperature
crossed 45◦ C, the HsfA1 gene activated in the tomato plant to minimize
the impact of high temperature (Mishra and Singh, 2011). Whereas,
Hsp101 (mitochondrial transcription termination factor-related protein)
in Arabidopsis enhanced the thermo-tolerance via mitochondrial
oxidative damage control (Kim et al., 2012). Nearly 5% of tran­
scriptomic activity observed and chaperones conquer only a minor part
during heat stress conditions (Larkindale and Vierling, 2008). They
mainly involved in metabolism, calcium, and phytohormone signaling,
protein phosphorylation(Al-Saidi and Elagib, 2017). For maintaining
photosynthesis and transpiration under stress conditions, Chloroplast
protein synthesis transcription factor accumulation is mandatory (Ristic
et al., 2008). Under high-temperature stress condition, polycationic
nature of polyamines stabilize the membrane structural integrity and
facilitates their strong binding to nucleic acids, proteins, and mem­
branes (Childs et al., 2003).
3.3. Hormonal regulation of growth responses under chilling stress
condition
Low-temperature stress in plants is an inevitable environmental
factor that can reduce growth, development (Thakur et al., 2010) and
photosynthesis efficiency (Farooq et al., 2009b). Chilling may causes
dehydration in plants by limiting water uptake and stomatal activities
(Mahajan and Tuteja, 2005). It is a type of external damage caused by
low temperature as a result of oxidative burst. The rate of chilling injury
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depends on the compound of the level of low temperature and duration
of exposure. Plants differ in their response to chilling (0-15◦ C) and
freezing (< 0◦ C) (Xin and Browse, 2000). In general, initially chilling
symptoms appear in the cell membrane by forming flexible liquid
crystalline cell membranes into a solid gel phase (Farooq et al., 2009b).
Whereas in fruit crops, chilling may cause a delay in fruit ripening and
fruit deformation. Low temperature during storage may cause browning,
reddening, woolliness, and mummification (Lurie and Crisosto, 2005).
Incidence of frost during fruit development forms ice crystals in sub­
cellular space and damages the plant cell activities, in severs condition,
complete plant death may also occur (Levitt, 1980). The intensity of
frost damage also depends on the physiological condition of plants
(Proebsting and Mills, 1978). It affects cell membrane integrity, ion
leakage, biomolecule activities, and gene expression (Dhanapal and
Crisosto, 2013). Strong relation observed between the sensitivity of cold
stress in plants and the proportion of desaturated phosphatidylglycerol
across different plant species (Sun et al., 2008). In fruit crops, chilling
can suppress the photosynthesis by reducing photosystem II efficiency
(Allen et al., 2000).
Different phytohormones have a role in cold stress is yet to be dis­
cussed in detail. ABA acts as a messenger in the regulation of water level
in plants (Mahajan and Tuteja, 2005). ABA-responsive genes are regu­
lated by several ABA-dependent or ABA-independent transcription fac­
tors (Chinnusamy et al., 2004). It was reported that freezing tolerance
accumulation depends on the genotype of plants (Galiba et al., 1993).
ABA could mediate some sugar signals in plant cells, which helps to
regulate the further ABA biosynthesis (Leon and Sheen, 2003). Same as
ABA, auxins involved in defense response against abiotic stresses in
plants (Wolters and Jurgens, 2009). It more intricately involved in the
various developmental process like fruit development, root initiation in
grafting, root growth, and development (Depuydt and Hardtke, 2011).
Members of the auxin-responsive GH3 gene family also has cold
stress-responsive activities. OsGH3-2 found to modulate both endoge­
nous free IAA and ABA homeostasis and differentially affect cold toler­
ance in rice (Du et al., 2012). However, the role of auxin remains unclear
under stress conditions in fruit crops. Brassinosteroids avoid ions
leakage in chilled stressed rape seeds (Janeczko, 2011), whereas it im­
proves the antioxidant defense in young grapevines to avoid the nega­
tive impact of cold stress (Xi et al., 2013) by biosynthesis of de novo or
activation of enzymes (Bajguz, 2000). When chilling sensitive banana
plants were treated with SA (0.5mM) caused an increase in SOD, CAT,
and APX activities when plants are induced to very low temperature
(Kang et al., 2003). Wang and Li (2006) also showed that SA treated
grapevines decreased lipid peroxidation and electrolyte leakage and
induced cold tolerance.
GA also plays an important role in the low-temperature condition. It
enhance the bolting and flowering in Limonium gmelinii plants when GA
sprayed at 500 mg L− 1 (Guda et al., 2000). GA inhibits the sucrose
formation from starch and mannitol (Ogasawara et al., 2001). This
outcome suggests that the GA application enhances the rapid tissue
enlargement and differentiated to resist the cold stress in plants
(Jacobsen and Olszewski, 1993). Chilling causes discoloration to the
peel of the ripened fruit by coagulating latex in latex ducts. The exog­
enous application of methyl jasmonate enhanced soluble protein and
antioxidant content under cold stress in banana plants (Zhao et al.,
2013a). Cold stress also enhances the ethylene biosynthesis (Zhao et al.,
2014). Zhao et al. (2009) reported that ethylene levels in plants and cold
tolerance are positively correlated up to a certain level.
Chilling can induce the expression of jasmonic acid biosynthesis
genes, including allene oxide synthase (AOS1), oxide cyclase (AOC), and
lipoxygenase (LOX2). Recent studies show that, in banana, two MYC2
TFs are activated following the exogenous application of MeJA in the
cold storage condition and enhances the expression of an inducer of CBF
expression (ICE-CBF) cold-sensitive pathways genes (Zhao et al., 2013b,
2013c). In MeJA induce chilling tolerance, the MaMYC2 transcription
factor in coordination with MaICE1 reduces lipoxygenase activity and
V. Rachappanavar et al.
increases antioxidant synthesis. Cao et al. (2011) found that ascorbate
peroxidase (APX), catalase (CAT) and superoxide dismutase (SOD) ac­
tivities in MeJA treated loquat fruits.
To manage the chilling injury, SA and MeSA treatments are given,
which are easy to set up, less expansive, and applicable to any crops
(Ding et al., 2001). They increase the AOX transcript level and reduces
the chilling incidence (Fung et al., 2014). When plants are treated with
SA and MeSA before chilling induces the HSPs biosynthesis and in­
creases the tolerance against chilling injury (Ding et al., 2008). Different
genetic studies show that many auxin-responsive genes are involved in
cold stress (Jain and Khurana, 2009). Likewise, PIN1 and PIN3, an auxin
transporter mediate root gravity response during the early phase is
inhibited by cold stress (Shibasaki et al., 2009). Zhao et al. (2009) in
tomato found a positive relationship between ethylene and freezing
tolerance from the negative effects of 1-methyl cyclopropene, which
inhibit ethylene biosynthesis on freezing tolerance. For freezing toler­
ance, the RARE COLD INDUCIBLE 1A (RCI1A) gene interacts with ACC
synthase. Dehydration-responsive element-binding proteins/C-repeat
binding factors (DREBs/CBFs) interact with dehydration-responsive
elements/C-repeat-elements (DRE/CRT) to execute a highly coordi­
nated transcriptional response to cold signals. C-repeat-binding factors
(CBF) cold response pathway is found in freezing-tolerant (FT) plants
(Skinner et al., 2005) and in freezing-sensitive (FS) plants (Zhang and
Huang, 2010).
3.4. Hormonal regulation of growth responses under salinity stress
condition
Soil salinity induced by excessive accumulation of sodium chloride
(NaCl) (Munns and Tester, 2008). The existence of a high amount of
non-essential soluble salts in irrigation water limits normal growth and
reproduction (Hussain et al., 2020). When salt content in irrigation
water crossed the threshold level may harm osmotic adjustment, ion
uptake, osmotic adjustment, photosynthesis, stomatal conductance,
protein synthesis, nucleic acid production, enzymatic activity, and
hormonal stability (Patane et al., 2013; Raga et al., 2016). Based on salt
stress tolerance potential, plants have been classified into two major
groups; (i) halophytes have the potential to survive and reproduce even
when salt concentration nearly 200-mM NaCl and (ii) glycophytes are
not able to survive under salt stress condition (Patane et al., 2013).
Glycophytes produce stress symptoms like decreased in total leaf area
and leaf growth, marginal and tip yellowing and tip scorching, marginal
and tip scorching of leaves, yellowing and bronzing, early leaf drop, twig
dieback, blackening, necrosis, and leaf burning (Bernstein et al., 2004).
Severe drought condition leads to access accumulation of salt in young
leaves can affect the normal photosynthesis mechanism by losing
photosynthesis pigments, and inhibition of Calvin cycle enzymes
(Acosta-Motos et al., 2017). Structural disorders were found in both
sensitive and tolerant genotypes, such as an increase in leaf thickness,
excessive chloride accumulation with reduced Mg2+, and loss of chlo­
rophyll content.
Osmotic regulation is one of the most important mechanisms to
avoid osmotic stress by accumulating metabolites like glycine, betaine,
proline, mannitol, and soluble sugars (Houimli, 2010). Accurate infor­
mation regarding environmental pressure, molecular influence, prote­
omics, and genotypic variations need to reduce salt stress in fruit crops
(Miranda et al., 2018). Morphologically some level of salinity stress can
be minimized in fruit crops by grafting of salt-sensitive improved high
yielding cultivars on salt tolerance rootstocks (Munns et al., 2006).
Lateral root formation in plants inhibited by high salinity or osmotic
stress condition (Galvan-Ampudia and Testerink, 2011). Whereas under
mid salinity stress, reduction in LR elongation and an increase in the
number of LRs observed. Besides, the reduction of LRs numbers
observed in auxin signaling mutants axr1, axr4, and tir1 completely
blocked auxin influx (Zolla et al., 2010). Even at low salt stress can
suppress auxin efflux carrier PIN2 and also decrease the PIN2: GFP
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concentration in root tips. It suggests that salt regulates both tran­
scription and posttranscriptional activities (Zhao et al., 2011). High salt
concentration causes rapid degradation of amyloplasts in root columella
cells thereby reduce the gravity response in roots (Sun et al., 2008). The
development of LRs by salt overly sensitive 3 (SOS3) under salinity stress
conditions has been reported by auxin redistribution activities (Zhao
et al., 2011). Mutants of SOS3 do not accumulate auxin in plant aerial
parts. The NAC transcription factors have a key role in plant growth and
stress responses. NAC1 and NAC2 proteins promote lateral root forma­
tion in Arabidopsis. AtNAC2 gene expression was induced by auxin,
ethylene, and ABA. Under salt stress condition auxin receptor mutants
tir1 suppressed. Nacl also downregulate TIR1 expression (Xie et al.,
2000)
When rootstocks of Microhyla zeylanica are used to cultivate mango
under 60-mM NaCl salinity condition, plants maintained a higher
photosynthetic rate with greater K+ /Na+ ratio in roots and lesser in
leaf/root Na+ (Schmutz, 2000). Whereas Pistacia Atlantica rootstock
gave the better result as compared to polycythemia vera rootstock when
used in pistachio plants under salinity stress (Tounsi et al., 2017). When
salt-tolerant (1103 Paulsen) and salt-sensitive (K 50-40) rootstocks of
grapes were evaluated, the study reveals that Cl has no direct relation­
ship with apoplastic tracer (Abbaspour et al., 2014).
External application of 30-mM/L calcium nitrate for citrus plants
significantly reduced Cl and Na+ uptake by the root system and also
enhanced the growth attributes and gaseous exchange mechanism
(Banuls et al., 1997). In olive plants, a reduction in stomatal conduc­
tance and photosynthesis observed when soil NaCl content crossed the
100 mM level (Tabatabaei, 2007). Jujube plants have more potential to
resist salinity compared to other cultivated fruit crops, in which two
wild Indian jujube (Ziziphus rotundifolia and Ziziphus nummularia) were
found to be more resistant to saline conditions due to higher stomatal
conductance, higher CO2 assimilation, higher hormonal regulation,
antioxidants accumulation and better nutrients translocation (Meena
et al., 2016) by accumulating high amount of Na+ in roots through
restricted translocation and high K+/Na+ ratio to keep ionic balance in
leaves (Mohammadkhani, 2018). Number fruit crops are identified
which can store excess K+ in leaf, stem and other tissues to mitigate the
negative effect of Salinity. The lowest Cl ion concentration and high K+
were recorded in lemon (Gimeno et al., 2009) and fig (Zarei et al., 2016)
plants when these plants are irrigated with water containing 50-mM
NaCl with 10-mM potassium nitrate. Arbuscular mycorrhiza like fungi
can increase the stress tolerance potential in trifoliate orange by accu­
mulating a low amount of Na+ and Ca2+ while higher K+, proline
content, and carbohydrate content.
Proline can help to store carbon and nitrogen and maintains the
structural integrity of cell membranes and proteins by purifying free
radicals (Jalili et al., 2009) and it maintains the buffering capacity of
cells against salinity (Heidari and Jamshid, 2010). Fozouni et al. (2012)
demonstrated that Vitis vinifera table grape significantly increases the
proline accumulation with increased salinity. But it harms photosyn­
thesis by degrading chlorophyll content (Behboudian et al., 1986). Due
to excess sodium content in saline soil affects the potassium uptake,
reduces the growth rate, delays flowering, and lowers the economic
yield. The optimum value for the soil K/Na is 2.5 for the banana if this
proportion exceeds by 8% can cause economic loss. Salt tolerant fruit
crops have succulent organs, salt exclusion, excretion of salts, and
accumulation of ions in the tonoplasts to maintain osmatic regulation.
Pomegranate tissues can accumulate sodium, chlorine, and potassium
ions without any stress symptoms in response to saline water irrigation
and it can tolerate salinity up to 40 mM NaCl but at higher NaCl con­
centration (>40mM) affects the main stem length, number of internodes
and the leaf surface area (Naeini et al., 2006) and decrease in soluble
sugar accumulation was observed (Naeini et al., 2004). Date palm plants
are considered as one of the exceptional halophytic plants and it can
survive under severe salt stress conditions. Ramoliya and Pandey (2003)
found that some datepalm cultivars adopted salt exclusion mechanisms
V. Rachappanavar et al.
and they can endure high soil salinity levels (up to 12.8 dS m-1) with no
visible effect on the seedlings (Greenway and Munns, 1983). Litchi trees
should not be irrigated with water with salt contains more than 0.5 dS
per m electric conductivity. If salinity level exceeds, symptoms like tips
and margins die were observed in old leaves (Babita and Kumar, 2008).
When chloride content in plants crossed the toxic level induce the
production of ethylene precursor ACC, leads to leaf senescence. In
counteract to ethylene activities, plants trigger the biosynthesis of ABA
to reduce ethylene production (Arbona et al., 2006). Silver thiosulfate
and phosgene synthetics are also used externally to inhibit the ethylene
lead senesce in salt-affected plants (Gómez-Cadenas et al., 2002). The
same result observed in a grapevine, where the extreme rise in ABA level
observed in resistant cultivars rootstock when planted in salt stress soil
(Upreti and Murti, 2010). Olive plants are grown under salinity, water
uptake, relative water uptake, and water potential drastically reduced
(Chartzoulakis, 2005; Yadav et al., 2011).
In sugarcane, salt tolerance induced by GA3 foliar application in
terms of enhancing nutrient uptake, morphological, and physiological
modification (Shomeili et al., 2011). GA3 application increase the
nutrient uptake, plant height, and yield in wheat (Ashraf and Shahbaz,
2003) and rice (Wen et al., 2011) when cultivated under saline condi­
tion. Different plants can tolerate salt stress with increasing GA con­
centration by more absorption of P and Ca2+ and less absorption of Na+
(Starck and Kozinska, 1980), stomatal resistance and enhanced plant
water use were observed at low salinity in tomato (Maggio et al., 2010)
and it enhances the catabolism of ABA (Gonai et al., 2004). The GA
metabolism and signaling play a major role in plant physiological
adaptation, low-temperature influences of bioactive GA as a result of an
increase in expression of three GA2ox genes Archard et al. (2012).
Rootstocks of grapes with high cytokinin can maintain a high K-Na ratio
and root-shoot dry mass ratio under salinity (Upreti and Murti, 2010). In
desert almond trees, cytokinin concentration peak in the morning time
and rapid decline in the afternoon based on the stomatal conductance
(Fusseder et al., 1992). Brault and Maldiney (1999) reported that cy­
tokinins act at the plasma membrane along with other signaling
compounds.
An important role of ethylene in salt tolerance by relating its ability
to retain K+ without decreasing Na in roots as well as in shoots (Yang
et al., 2013) and It stimulates H+-ATPase activity to modulate ion ho­
meostasis (Wang et al., 2009). Polyamine content in the grafted plant
increased against salt stress by Prompting superoxide dismutase,
peroxidase, and ascorbic peroxidase and glutathione reductase enzyme
activities. Upreti and Murti (2010) found a significant increase in
polyamines concentration intolerant grape rootstock roots by salinity
stress along with ABA and it helped to maintain the high root- shoot
biomass ratio and high K-Na ratio. Similarly, Anjum (2010) reported
that Cleopatra mandarin and citrus rootstock shows better growth and
chlorophyll efficiency at high spermine and carbohydrates with low
chloride ions in leaves and roots under salinity. Polyamines control the
Ca2+ allocation through regulating Ca2+ permeable channels and CAXs
by the prevention of Na+/K+ entry into the cytoplasm and enhance­
ment of Na+/K+ influx to the vacuole with suppression of Na+/K+
release from the vacuole. Polyamines regulate the physiology stomata
similar to ABA, by targeting KAT1-like inward K+ channel in guard cells
of stomata Liu et al. (2019).
Klingler et al. (2010) suggested that ABA-dependent amino oxidase
derived H2O2 affected stomatal physiology in grapevines. By genetic
modification or exogenous application of polyamines enhances salt
tolerance in grape (Farooq et al., 2009b). In Neo-Muscat grapes seedling
significantly accumulate putrescine during osmotic and salt stress by
temporary induction of arginine decarboxylase (pVvADC) gene followed
by spermine synthase (pVvSPMS) (Liu et al., 2012). The overexpression
of OsGSTU4, an auxin-responsive GST gene enhance the tolerance to
salinity and oxidative stress condition in Arabidopsis plants (Jain et al.,
2010).
13
Scientia Horticulturae 304 (2022) 111302
3.5. Hormonal regulation of growth responses under heavy metal stress
condition
No organic life can develop and persist without the involvement of
metal ions in this universe. The elementary constituents of living or­
ganisms may be classified as major, minor, and trace elements based on
their essentiality for normal biological activity. The essential microele­
ments are Cu, Zn, Fe, Mn, Mo, Ni, and Co whose uptake in excess to the
plant causes toxic effects (Monni et al., 2000; Vaculík et al. 2020). Fe, Cu
Cr, and Co are involved in a redox reaction, whereas Zn, Cd, Ni, Al, etc.
monitor the various enzymatic activities in plant cells. Transition metals
such as copper and iron have frequently unpaired electrons and are,
therefore, very good catalysts of oxygen reduction. Currently, the
contamination of the natural ecosystem by these heavy metals repre­
sents a global threat, endangering the agriculture ecosystem (Recatalá
et al., 2010). In small quantity (in the trace), these elements are essential
for normal plant growth and development and play a key role in
metabolism by activating different enzymatic activities (Roy and
Mcdonald, 2015). However, when the concentration of these metals
crossed threshold level cause a negative impact on physiology and
biochemistry of plants (Ivanov et al., 2016) and reduce the biomass
accumulation in plants as a result dwindle the photosynthesis rate
(Rodriguez et al., 2012), mineral nutrition uptake (Vernay et al., 2007)
and communication with water (Mukhopadhyay and Mondal, 2015).
Besides they can also cell toxicity by excess production of ROS, which
impaired antioxidant activities and causes severe oxidative stress (Rui
et al., 2016).
Previous research has shown that the application of plant growth-
promoting phytohormones can improve protection by counter­
balancing heavy metal toxicity in plants (Agami and Mohamed, 2013;
Masood et al., 2012). ABA concentration increase as plants is exposed to
heavy metals (Rauser and Dumbroff, 1981). Heavy metals like Zn, Al, Ni,
and Cd (Fediuc et al., 2005) have been revealed to raise in ABA level in
plants. Hsu and Kao (2007) reported that at high temperature
(30/35◦ C), ABA regulates and increase the resistance against Cd in rice
seedlings. It’s proven that ABA-mediated the Cd-precipitated stimula­
tion of O-acetylserine (OASTL), the enzyme responsible for cysteine
biosynthesis. The expression of ABA and drought resistance BjCdR9 (an
aldehyde dehydrogenase) and BjCd55 (RNA binding protein) genes
stimulated by Cd in Brassica Juncea supported the involvement of ABA
signal transduction and these genes denoting aquaporins PIP1 and PIP2
was found to be transcribed when exposed to Cd stress. The above
observation gave information that water stress is imposed by cd and that
ABA and Cd show the synergistic relationship (Fusco et al., 2005). When
plants are treated with ABA, both BjCd15 and TGA3 responded to ABA.
However, TGA shows more response to ABA as compared to BjCdR15
(Farinati et al., 2010).
Zhou et al. (2016) observed that Cd concentration increased in roots
and leaves at a higher dose of CdCl2 in JA- deficient tomato mutants.
The result shows that a lack of JA biosynthesis enhances the sensitivity
against Cd. According to Sirhindi et al. (2015), the exogenous applica­
tion of JA enhance the tolerance to NiCl2 in soybean seedlings and it
regulates the antioxidant activities to neutralize the heavy metal stress
at the cellular level in plants, while Azeem et al. (2018) revealed that
exogenous JA application increases the biomass production and protein
concentration in Ni-treated plants and minimize the CD accumulation
rate by enhancing osmotic and antioxidant activities, but also inhibiting
H2O2 and Malondialdehyde accumulation (Noriega et al., 2012). JA
inhibit the peroxidase activity by activating ascorbate or glutathione
antioxidant machinery. Such findings reports show that JA plant re­
sponses by inducing antioxidant system.
Although these research reports suggest that phytohormones
responsive genes are differentially expressed under various abiotic stress
conditions, further indicating/corroborating crosstalk between plant bio
stimulants and abiotic stress signalling.
V. Rachappanavar et al.
4. Conclusion and future prospectus
Environmental stresses can significantly reduce the yield potential
and shift in cultivation area may occur for commercially important fruit
crops by climate change. Now need to improve the biosynthesis and
utilization of phytohormones without any negative impact by genetic
modification in fruit crops to improve tolerance against abiotic stresses.
Recently, much attention has been given to the development of strate­
gies designed for breeding to alleviate the adverse effects of abiotic
stress by using different phytohormones, from the foregoing research
works discussion it is clear that plants utilize elaborate signaling path­
ways in response to various abiotic stresses. In addition to ROS, anti­
oxidants, and other biochemicals, plant hormones trigger specific signal
cascades upon abiotic perception. The vacillation in several key hor­
mone levels such as GA, ABA, AUX, BRs, SA and JA and crosstalk among
phytohormones occur as early response to stresses. Such crosstalk at the
hormonal level in plants helps to integrate various stress signaling
pathways. The enhanced level of tolerance to environmental stress at the
molecular level by adjusting growth factors is the key role to the plant
survival. These have been illustrated in the present review with appro­
priate research results drawn from the abiotic and biotic stress responses
in fruit crops.
Author contributions
Vinaykumar Rachappanavar wrote the first draft of this review
article, Arushi Padiyal contributed in Figures and tables and JK sharma
and SK Gupta provided valuable inputs, conceptualize and finalized the
draft.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgment
The authors are thankful to the MS Swaminathan School of Agri­
culture, Shoolini University, Solan and for providing required facility for
this work.
References
Abbaspour, N, Kaiser, B, Tyerman, S, 2014. Root apoplastic transport and water relations
cannot account for differences in Cl− transport and Cl− /NO−3 interactions of two
grapevine rootstocks differing in salt tolerance. Acta Physiol. Plant. 36, 687–698.
https://doi.org/10.1007/s11738-013-1447-y.
Abd El-Naby, S.K.M., Abdelkhalek, A., Baiea, M.H.M., Amin, O.A., 2020. Mitigation of
heat stress effects on Washington navel orange by using melatonin, gibberellin and
salicylic treatments. Plant Arch. 20 (1), 3523–3534.
Abe, H, Yamaguchi-Shinozaki, K, Urao, T, Iwasaki, T, Hosokawa, D, Shinozaki, K, 1997.
Role of arabidopsis MYC and MYB homologs in drought-and abscisic acid-regulated
gene expression. Plant Cell 9, 1859–1868. https://doi.org/10.1105/tpc.9.10.1859.
Abeles, FB, Morgan, PW, Saltveit, ME, 1992. Ethylene in Plant Biology, 2nd. Academic
Press, San Diego. ISBN 0-12-041451-1.
Achard, P, Genschik, P, 2009. Releasing the brakes of plant growth: how GAs shutdown
DELLA proteins. J. Exp. Bot. 60, 1085–1092. https://doi.org/10.1093/jxb/ern301.
Acharya, BR, Assmann, SM, 2009. Hormone interactions in stomatal function. Plant Mol.
Biol. 69, 451–462. https://doi.org/10.1007/s11103-008-9427-0.
Acosta-Motos, J.R., Hernández, J.A., Álvarez, S., Barba-Espín, G., Sánchez-Blanco, M.J.,
2017. The long-term resistance mechanisms, critical irrigation threshold and relief
capacity shown by Eugenia myrtifolia plants in response to saline reclaimed water.
Plant Physiol. Biochem. 1 (111), 244–256. https://doi.org/10.1016/j.
plaphy.2016.12.003.
Adato, I, Gazit, S, 1974. Water-deficit stress, ethylene production, and ripening in
avocado fruits. Plant Physiol. 53, 45–46. https://doi.org/10.1104/pp.53.1.45.
Adkins, SW, Samosir, YM, Ernawati, A, Godwin, ID, Drew, RA, 1997. Control of ethylene
and use of polyamines can optimise the conditions for somatic embryogenesis in
coconut (Cocos nucifera L.) and papaya (Carica papaya L.) >Proceedings of the
International Symposium on Biotechnology of Tropical and Subtropical Species. In: ,
461, pp. 459–466. https://doi.org/10.17660/ActaHortic.1998.461.54.
14
Scientia Horticulturae 304 (2022) 111302
Agami, RA, Mohamed, GF, 2013. Exogenous treatment with indole-3-acetic acid and
salicylic acid alleviates cadmium toxicity in wheat seedlings. Ecotoxicol. Environ.
Saf. 94, 164–171. https://doi.org/10.1016/j.ecoenv.2013.04.013.
Ahammed, GJ, Gantait, S, Mitra, M, Yang, Y, Li, X, 2020. Role of ethylene crosstalk in
seed germination and early seedling development: a review. Plant Physiol. Biochem.
151, 124–131. https://doi.org/10.1016/j.plaphy.2020.03.016. DOI: 10.1016/j.
plaphy.2020.03.016.
Ahammed, GJ, Choudhary, SP, Chen, S, Xia, X, Shi, K, Zhou, Y, Yu, J, 2013. Role of
brassinosteroids in alleviation of phenanthrene–cadmium co-contamination-induced
photosynthetic inhibition and oxidative stress in tomato. J. Exp. Bot. 64, 199–213.
https://doi.org/10.1093/jxb/ers323.
Ahmad, P, Prasad, MNV, 2012a. Abiotic Stress Responses in Plants: Metabolism,
Productivity and Sustainability. Springer Science Business Media, LLC, New York,
NY. https://doi.org/10.1007/978-1-4614-0634-1.
Ahmad, P, Prasad, MNV, 2012b. Environmental Adaptations and Stress Tolerance in
Plants in the Era of Climate Change. Springer Science Business Media, LLC, New
York, NY. https://doi.org/10.1007/978-1-4614-0815-4.
akbar Mozafari, A, Havas, F, Ghaderi, N, 2018. Application of iron nanoparticles and
salicylic acid in in vitro culture of strawberries (Fragaria× ananassa Duch.) to cope
with drought stress. Plant Cell Tissue Organ. Cult. 132, 511–523. https://doi.org/
10.1007/s11240-017-1347-8.
Alam, P, Albalawi, TH, Altalayan, FH, Bakht, MA, Ahanger, MA, Raja, V, et al., 2019. 24-
Epibrassinolide (EBR) confers tolerance against NaCl stress in soybean plants by up-
regulating antioxidant system, ascorbate-glutathione cycle, and glyoxalase system.
Biomolecules 9, 640. https://doi.org/10.3390/biom9110640.
Alferez, F, Sala, JM, Sanchez-Ballesta, MT, Mulas, M, Lafuente, MT, Zacarias, L, 2005.
A comparative study of the postharvest performance of an ABA-deficient mutant of
oranges: I. Physiological and quality aspects. Postharvest Biol. Technol. 37,
222–231. https://doi.org/10.1016/j.postharvbio.2005.05.010.
Ali, Q, Shahid, S, Nazar, N, Hussain, AI, Ali, S, Chatha, SAS, Perveen, R, Naseem, J,
Haider, MZ, Hussain, B, Hussain, SM, 2020. Use of Phytohormones in Conferring
Tolerance to Environmental Stress. In Plant Ecophysiology and Adaptation under
Climate Change: Mechanisms and Perspectives II. Springer, Singapore, pp. 245–355.
https://doi.org/10.1007/978-981-15-2172-0_11.
Allen, RG, Tresini, M, 2000. Oxidative stress and gene regulation. Free Radical Biol. Med.
28, 463–499. https://doi.org/10.1016/S0891-5849(99)00242-7.
Al-Saidi, M, Elagib, NA, 2017. Towards understanding the integrative approach of the
water, energy and food nexus. Sci. Total Environ. 574, 1131–1139. https://doi.org/
10.1016/j.scitotenv.2016.09.046.
Amri, E, Mohammadi, MJ, 2012. Effects of timing of drought stress on pomegranate
seedlings (Punica granatum L. cv〈 Atabaki〉) to exogenous spermidine and putrescine
polyamines. Afr. J. Microbiol. Res. 6, 5294–5300. https://doi.org/10.5897/
AJMR11.1355.
An, C, Mou, Z, 2011. Salicylic acid and its function in plant immunity F. J. Integr. Plant
Biol. 53, 412–428. https://doi.org/10.1111/j.1744-7909.2011.01043.x.
An, JP, Wang, XN, Yao, JF, Ren, YR, You, CX, Wang, XF, Hao, YJ, 2017. Apple MdMYC2
reduces aluminum stress tolerance by directly regulating MdERF3 gene. Plant Soil
418, 255–266. https://doi.org/10.1007/s11104-017-3297-7.
An, JP, Yao, JF, Xu, RR, You, CX, Wang, XF, Hao, YJ, 2018. An apple NAC transcription
factor enhances salt stress tolerance by modulating the ethylene response. Physiol.
Plant. 164, 279–289. https://doi.org/10.1111/ppl.12724.
Anjum, MA, 2010. Response of Cleopatra mandarin seedlings to a polyamine-
biosynthesis inhibitor under salt stress. Acta Physiol. Plant. 32, 951–959. https://
doi.org/10.1007/s11738-0-0483-0.
Arbona, V, Gómez-Cadenas, 2008. Hormonal modulation of citrus responses to flooding.
J. Plant Growth Regul. 27, 241–250. https://doi.org/10.1007/s00344-008-9051-x.
Arbona, V, López-Climent, MF, Mahouachi, J, Pérez-Clemente, RM, Abrams, SR, Gomez-
Cadenas, A, 2006. Use of persistent analogs of abscisic acid as palliatives against salt-
stress induced damage in citrus plants. J. Plant Growth Regul. 25, 1–9. https://doi.
org/10.1007/s00344-005-0038-6.
Arc, E, Sechet, J, Corbineau, F, Rajjou, L, Marion-Poll, A, 2013. ABA crosstalk with
ethylene and nitric oxide in seed dormancy and germination. Front. Plant Sci. 4
https://doi.org/10.3389/fpls.2013.00063 p.63.
Archard, GA, Earley, RL, Hanninen, AF, Braithwaite, VA, 2012. Correlated behaviour and
stress physiology in fish exposed to different levels of predation pressure. Funct.
Ecol. 26, 637–645. https://doi.org/10.1111/j.1365-2435.2012.01968.x.
Arif, Y, Sami, F, Siddiqui, H, Bajguz, A, Hayat, S, 2020. Salicylic acid in relation to other
phytohormones in plant: a study towards physiology and signal transduction under
challenging environment. Environ. Exp. Bot. (175), 104040 https://doi.org/
10.1111/j.1365-2435.2012.01968.x.
Aseri, GK, Jain, N, Panwar, J, Rao, AV, Meghwal, PR, 2008. Biofertilizers improve plant
growth, fruit yield, nutrition, metabolism and rhizosphere enzyme activities of
pomegranate (Punica granatum L.) in Indian Thar Desert. Sci. Hortic. 117, 130–135.
https://doi.org/10.1016/j.scienta.2008.03.014.
Ashraf, M, Shahbaz, M, 2003. Assessment of genotypic variation in salt tolerance of early
CIMMYT hexaploid wheat germplasm using photosynthetic capacity and water
relations as selection criteria. Photosynthetica 41, 273–280. https://doi.org/
10.1023/B:PHOT.0000011961.33120.b6.
Asins, MJ, 2002. Present and future of quantitative trait locus analysis in plant breeding.
Plant Breed. 121, 281–291. https://doi.org/10.1046/j.1439-0523.2002.730285.x.
Azevedo, EP, Kumar, V, 2012. Rheological, water uptake and controlled release
properties of a novel self-gelling aldehyde functionalized chitosan. Carbohydr.
Polym. 90, 894–900. https://doi.org/10.1016/j.carbpol.2012.06.017.
Azim, AA, Simon, KMR, Fino, D, Bochmann, G, 2018. The physiological effect of heavy
metals and volatile fatty acids on Methanococcus maripaludis S2. Biotechnol. Biofuels
11, 301. https://doi.org/10.1186/s13068-018-1302-x.
V. Rachappanavar et al.
Babita, S, Kumar, R, 2008. Feed your litchi orchard properly. Indian Hortic. 53, 18–20.
Bai, MY, Fan, M, Oh, E, Wang, ZY, 2012. A triple helix-loop-helix/basic helix-loop-helix
cascade controls cell elongation downstream of multiple hormonal and
environmental signaling pathways in Arabidopsis. Plant Cell 24, 4917–4929.
https://doi.org/10.1105/tpc.112.105163.
Bai, T, Li, C, Li, C, Liang, D, Ma, F, 2013. Contrasting hypoxia tolerance and adaptation in
Malus species is linked to differences in stomatal behavior and photosynthesis.
Physiol. Plant. 147, 514–523. https://doi.org/10.1111/j.1399-3054.2012.01683.x.
Bai, T, Li, C, Ma, F, Feng, F, Shu, H, 2010. Responses of growth and antioxidant system to
root-zone hypoxia stress in two Malus species. Plant Soil 327, 95–105. https://doi.
org/10.1007/s11104-009-0034-x.
Bajguz, A, 2000. Effect of brassinosteroids on nucleic acids and protein content in
cultured cells of Chlorella vulgaris. Plant Physiol. Biochem. 38, 209–215. https://doi.
org/10.1016/S0981-9428(00)00733-6.
Balcı G, Aras S, Keles, H (2021) Exogenous EBL (24-Epibrassinolide) Alleviate Cold
Damage in Strawberry. Erwerbs-Obstbau. pp. 1-6. 10.1007/s10341-021-00566-6.
Banerjee, A, Roychoudhury, A, 2016. Plant responses to light stress: oxidative damages,
photoprotection, and role of phytohormones. Plant Hormones Under Challenging
Environmental Factors. Springer, Dordrecht, pp. 181–213. https://doi.org/10.1007/
978-94-017-7758-2_8 pp.
Bañuls, J, Serna, MD, Legaz, F, Talon, M, Primo-Millo, E, 1997. Growth and gas exchange
parameters of Citrus plants stressed with different salts. J. Plant Physiol. 150,
194–199. https://doi.org/10.1016/S0176-1617(97)80202-7.
Bari, R, Jones, JD, 2009. Role of plant hormones in plant defence responses. Plant Mol.
Biol. 69, 473–488. https://doi.org/10.1007/s11103-008-9435-0.
Bartels, D, Sunkar, R, 2005. Drought and salt tolerance in plants. Crit. Rev. Plant Sci. 24,
23–58. https://doi.org/10.1080/07352680590910410.
Behboudian, M.H., Törökfalvy, E., Walker, R.R., 1986. Effects of salinity on ionic
content, water relations and gas exchange parameters in some citrus
scion—rootstock combinations. Sci. Hortic. 28 (1-2), 105–116. https://doi.org/
10.1016/0304-4238(86)90130-5.
Ben-Yehoshua, S., Aloni, B., 1974. Effect of water stress on ethylene production by
detached leaves of Valencia orange (Citrus sinensis Osbeck). Plant Physiol. 53 (6),
863–865. https://doi.org/10.1104/pp.53.6.863.
Bernstein, N., Meiri, A., Zilberstaine, M., 2004. Root growth of avocado is more sensitive
to salinity than shoot growth. J. Am. Soc. Hortic. Sci. 129 (2), 188–192. https://doi.
org/10.21273/JASHS.129.2.0188.
Bertini, I., 1997. Determinants of reduction potentials in metalloproteins. JBIC 2 (1),
108. -108.
Bhatnagar-Mathur, P., Devi, M.J., Vadez, V., Sharma, K.K., 2009. Differential
antioxidative responses in transgenic peanut bear no relationship to their superior
transpiration efficiency under drought stress. J. Plant Physiol. 166 (11), 1207–1217.
https://doi.org/10.1016/j.jplph.2009.01.001.
Bhatt, D., Nath, M., Sharma, M., Bhatt, M.D., Bisht, D.S. and Butani, N.V., 2020. Role of
growth regulators and phytohormones in overcoming environmental stress.
Protective Chemical Agents in the Amelioration of Plant Abiotic Stress: Biochemical
and Molecular Perspectives, pp.254-279. 10.1002/9781119552154.ch11.
Bidabadi, S.S., Ghobadi, C., Baninasab, B., 2012. Influence of salicylic acid on
morphological and physiological responses of banana (’Musa
acuminata’cv.’Berangan’, AAA) shoot tips to in vitro water stress induced by
polyethylene glycol. Plant Omics 5 (1), 33–39. https://doi.org/10.3316/
informit.007834090133454.
Biddington, N.L., Dearman, A.S., 1985. The effect of mechanically induced stress on the
growth of cauliflower, lettuce and celery seedlings. Ann. Bot. 55 (1), 109–119.
https://doi.org/10.1093/oxfordjournals.aob.a086869.
Boatwright, J.L., Pajerowska-Mukhtar, K., 2013. Salicylic acid: an old hormone up to
new tricks. Mol. Plant Pathol. 14 (6), 623–634. https://doi.org/10.1111/
mpp.12035.
Bouchabke, O., Chang, F., Simon, M., Voisin, R., Pelletier, G., Durand-Tardif, M., 2008.
Natural variation in Arabidopsis thaliana as a tool for highlighting differential
drought responses. PLoS One 3 (2). https://doi.org/10.1371/journal.pone.0001705.
Bouchabke-Coussa, O., Quashie, M.L., Seoane-Redondo, J., Fortabat, M.N., Gery, C.,
Yu, A., Linderme, D., Trouverie, J., Granier, F., Téoulé, E., Durand-Tardif, M., 2008.
ESKIMO1 is a key gene involved in water economy as well as cold acclimation and
salt tolerance. BMC Plant Biol. 8 (1), 125. https://doi.org/10.1186/1471-2229-8-
125.
Brault, M., Maldiney, R., 1999. Mechanisms of cytokinin action. Plant Physiol. Biochem.
37 (5), 403–412. https://doi.org/10.1016/j.ejar.2021.01.003.
Bray, E.A., 2002. Abscisic acid regulation of gene expression during water-deficit stress
in the era of the Arabidopsis genome. Plant Cell Environ. 25 (2), 153–161. https://
doi.org/10.1046/j.1365-3040.2002.00746.x.
Brodersen, P., Malinovsky, F.G., Hématy, K., Newman, M.A., Mundy, J., 2005. The role
of salicylic acid in the induction of cell death in Arabidopsis acd11. Plant Physiol.
138 (2), 1037–1045. https://doi.org/10.1104/pp.105.059303.
Burbidge, G., Hewitt, A., Narlikar, J.V., Gupta, P.D., 1990. Associations between quasi-
stellar objects and galaxies. Astrophys. J. Suppl. Ser. 74, 675–730.
Cao, S., Yang, Z., Cai, Y., Zheng, Y., 2011. Fatty acid composition and antioxidant system
in relation to susceptibility of loquat fruit to chilling injury. Food Chem. 127 (4),
1777–1783. https://doi.org/10.1016/j.foodchem.2011.02.059.
Chacko, E.K., 1986. Physiology of vegetative and reproductive growth in mango
(Mangifera indica L.) trees. In: Proceedings of the First Australian Mango Research
Workshop (Vol. 1, pp. 54-70). CSIRO, Cairns, QLD.
Chaitali, R., Sengupta, D.N., 2014. Effect of short term NaCl stress on cultivars of
S. lycopersicum: a comparative biochemical approach. J. Stress Physiol. Biochem. 10
(1).
15
Scientia Horticulturae 304 (2022) 111302
Chapman, E.J., Estelle, M., 2009. Mechanism of auxin-regulated gene expression in
plants. Annu. Rev. Genet. 43, 265–285.
Chartzoulakis, K.S., 2005. Salinity and olive: growth, salt tolerance, photosynthesis and
yield. Agric. Water Manag. 78 (1–2), 108–121. https://doi.org/10.1016/j.
agwat.2005.04.025.
Chen, M., Guo, H., Chen, S., Li, T., Li, M., Rashid, A., Xu, C., Wang, K., 2019. Methyl
jasmonate promotes phospholipid remodeling and jasmonic acid signaling to
alleviate chilling injury in peach fruit. J. Agric. Food Chem. 67 (35), 9958–9966.
https://doi.org/10.1021/acs.jafc.9b03853.
Chen, Y.E., Cui, J.M., Li, G.X., Yuan, M., Zhang, Z.W., Yuan, S., Zhang, H.Y., 2016. Effect
of salicylic acid on the antioxidant system and photosystem II in wheat seedlings.
Biologia plantarum 60 (1), 139–147. https://doi.org/10.1007/s10535-015-0564-4.
Cheong, Y.H., Chang, H.S., Gupta, R., Wang, X., Zhu, T., Luan, S., 2002. Transcriptional
profiling reveals novel interactions between wounding, pathogen, abiotic stress, and
hormonal responses in Arabidopsis. Plant Physiol. 129 (2), 661–677. https://doi.
org/10.1104/pp.002857.
Chernys, J.T., Zeevaart, J.A., 2000. Characterization of the 9-cis-epoxycarotenoid
dioxygenase gene family and the regulation of abscisic acid biosynthesis in avocado.
Plant Physiol. 124 (1), 343–354. https://doi.org/10.1104/pp.124.1.343.
Childs, A.C., Mehta, D.J., Gerner, E.W., 2003. Polyamine-dependent gene expression.
Cellular Mol. Life Sci. CMLS 60 (7), 1394–1406. https://doi.org/10.1007/s00018-
003-2332-4.
Chinnusamy, V., Schumaker, K., Zhu, J.K., 2004. Molecular genetic perspectives on
cross-talk and specificity in abiotic stress signalling in plants. J. Exp. Bot. 55 (395),
225–236. https://doi.org/10.1093/jxb/erh005.
Choi, J., Huh, S.U., Kojima, M., Sakakibara, H., Paek, K.H., Hwang, I., 2010. The
cytokinin-activated transcription factor ARR2 promotes plant immunity via TGA3/
NPR1-dependent salicylic acid signaling in Arabidopsis. Dev. Cell 19 (2), 284–295.
https://doi.org/10.1016/j.devcel.2010.07.011.
Choudhary, S.P., Yu, J.Q., Yamaguchi-Shinozaki, K., Shinozaki, K., Tran, L.S.P., 2012.
Benefits of brassinosteroid crosstalk. Trends Plant Sci. 17 (10), 594–605. https://doi.
org/10.1016/j.tplants.2012.05.012.
Choudhury, F.K., Rivero, R.M., Blumwald, E., Mittler, R., 2017. Reactive oxygen species,
abiotic stress and stress combination. Plant J. 90 (5), 856–867. https://doi.org/
10.1111/tpj.13299.
Claeys, H., Skirycz, A., Maleux, K., Inzé, D., 2012. DELLA signaling mediates stress-
induced cell differentiation in Arabidopsis leaves through modulation of anaphase-
promoting complex/cyclosome activity. Plant Physiol. 159 (2), 739–747. https://
doi.org/10.1104/pp.112.195032.
Clifford, S.C., Arndt, S.K., Popp, M., Jones, H.G., 2002. Mucilages and polysaccharides in
Ziziphus species (Rhamnaceae): localization, composition and physiological roles
during drought-stress. J. Exp. Bot. 53 (366), 131–138. https://doi.org/10.1093/
jexbot/53.366.131.
Colebrook, E.H., Thomas, S.G., Phillips, A.L., Hedden, P., 2014. The role of gibberellin
signalling in plant responses to abiotic stress. J. Exp. Biol. 217 (1), 67–75. https://
doi.org/10.1242/jeb.089938.
Colmer, T.D., Peeters, A.J.M., Wagemaker, C.A.M., Vriezen, W.H., Ammerlaan, A.,
Voesenek, L.A.C.J., 2004. Expression of α-expansin genes during root acclimations to
O 2 deficiency in Rumex palustris. Plant Mol. Biol. 56 (3), 423–437.
Conesa, M.R., De La Rosa, J.M., Domingo, R., Banon, S., Perez-Pastor, A., 2016. Changes
induced by water stress on water relations, stomatal behaviour and morphology of
table grapes (cv. Crimson seedless) grown in pots. Sci. Hortic. 202, 9–16. https://doi.
org/10.1016/j.scienta.2016.02.002.
Cushman, J.C., Bohnert, H.J., 2000. Genomic approaches to plant stress tolerance. Curr.
Opin. Plant Biol. 3 (2), 117–124. https://doi.org/10.1016/S1369-5266(99)00052-7.
Damour, G., Simonneau, T., Cochard, H., Urban, L., 2010. An overview of models of
stomatal conductance at the leaf level. Plant Cell Environ. 33 (9), 1419–1438.
https://doi.org/10.1111/j.1365-3040.2010.02181.x.
Depuydt, S., Hardtke, C.S., 2011. Hormone signalling crosstalk in plant growth
regulation. Curr. Biol. 21 (9), R365–R373. https://doi.org/10.1016/j.
cub.2011.03.013.
Dhanapal, A.P., Crisosto, C.H., 2013. Association genetics of chilling injury susceptibility
in peach (Prunus persica (L.) Batsch) across multiple years. 3 Biotech 3 (6), 481–490.
https://doi.org/10.1007/s13205-012-0109-x.
Ding, L., Li, Y., Wang, Y., Gao, L., Wang, M., Chaumont, F., Shen, Q., Guo, S., 2016. Root
ABA accumulation enhances rice seedling drought tolerance under ammonium
supply: interaction with aquaporins. Front. Plant Sci. 7, 1206. https://doi.org/
10.3389/fpls.2016.01206.
Ding, Q., Keller, J.N., 2001. Proteasome inhibition in oxidative stress neurotoxicity:
implications for heat shock proteins. J. Neurochem. 77 (4), 1010–1017. https://doi.
org/10.1046/j.1471-4159.2001.00302.x.
Ding, X., Cao, Y., Huang, L., Zhao, J., Xu, C., Li, X., Wang, S., 2008. Activation of the
indole-3-acetic acid–amido synthetase GH3-8 suppresses expansin expression and
promotes salicylate-and jasmonate-independent basal immunity in rice. Plant Cell 20
(1), 228–240. https://doi.org/10.1105/tpc.107.055657.
Dong, Q., Duan, D., Zheng, W., Huang, D., Wang, Q., Li, X., Mao, K., Ma, F., 2021.
MdVQ37 overexpression reduces basal thermotolerance in transgenic apple by
affecting transcription factor activity and salicylic acid homeostasis. Horticulture
Res. 8 (1), 1–16. https://doi.org/10.1038/s41438-021-00655-3.
Dos Reis, S.P., Lima, A.M., De Souza, C.R.B., 2012. Recent molecular advances on
downstream plant responses to abiotic stress. Int. J. Mol. Sci. 13 (7), 8628–8647.
https://doi.org/10.3390/ijms13078628.
Du, F., Ruan, G., Liu, H., 2012. Analytical methods for tracing plant hormones. Anal.
Bioanal.Chem. 403 (1), 55–74. https://doi.org/10.1007/s00216-011-5623-x.
V. Rachappanavar et al.
Fahad, S, Nie, L, Chen, Y, Wu, C, Xiong, D, Saud, S, Hongyan, L, Cui, K, Huang, J, 2015.
Crop plant hormones and environmental stress. Sustain Agric. Rev. 15, 371–400.
https://doi.org/10.1007/978-3-319-09132-7_10.
FAO, 2017. Fruit and vegetables for health initiative. http://www.fao.org/3/a-i6807e.
pdf. accessed on 26/9/2021 at 12:06.
Fariduddin, Q., Yusuf, M., Ahmad, I., Ahmad, A., 2014. Brassinosteroids and their role in
response of plants to abiotic stresses. Biol. Plant. 58 (1), 9–17. https://doi.org/
10.1007/s10535-013-0374-5.
Farinati, S., DalCorso, G., Varotto, S., Furini, A., 2010. The Brassica juncea BjCdR15, an
ortholog of Arabidopsis TGA3, is a regulator of cadmium uptake, transport and
accumulation in shoots and confers cadmium tolerance in transgenic plants. New
Phytol. 185 (4), 964–978. https://doi.org/10.1111/j.1469-8137.2009.03132.x.
Farooq, M., Wahid, A., Kobayashi, N., Fujita, D.B.S.M.A., Basra, S.M.A., 2009b. Plant
drought stress: effects, mechanisms and management. Sustainable Agriculture.
Springer, Dordrecht, pp. 153–188.
Fediuc, E., Lips, S.H., Erdei, L., 2005. O-acetylserine (thiol) lyase activity in Phragmites
and Typha plants under cadmium and NaCl stress conditions and the involvement of
ABA in the stress response. J. Plant Physiol. 162 (8), 865–872. https://doi.org/
10.1016/j.jplph.2004.11.015.
Feng, S., Martinez, C., Gusmaroli, G., Wang, Y., Zhou, J., Wang, F., Chen, L., Yu, L.,
Iglesias-Pedraz, J.M., Kircher, S., Schäfer, E., 2008. Coordinated regulation of
Arabidopsis thaliana development by light and gibberellins. Nature 451 (7177),
475–479. https://doi.org/10.1038/nature06448.
Fozouni, M., Abbaspour, N., Baneh, H.D., 2012. Leaf water potential, photosynthetic
pigments and compatible solutes alterations in four grape cultivars under salinity.
Vitis 51 (4), 147–152.
Fraire-Velazquez, S., Rodriguez-Guerra, R., Sanchez-Calderon, L., 2011. Abiotic and
biotic tress response crosstalk in plants. In: Shanker, A. (Ed.), Abiotic Stress Response
in lantsPhysiological, Biochemical and Genetic Perspectives. InTech Open Access
Company, Rijeka, Croatia, pp. 1–26.
Fu, J., Wu, H., Ma, S., Xiang, D., Liu, R., Xiong, L., 2017. OsJAZ1 attenuates drought
resistance by regulating JA and ABA signaling in rice. Front. Plant Sci. 8, 2108.
https://doi.org/10.3389/fpls.2017.02108.
Fujita, H., Morii, T., Fujishima, H., Sato, T., Shimizu, T., Hosoba, M., Tsukiyama, K.,
Narita, T., Takahashi, T., Drucker, D.J., Seino, Y., 2014. The protective roles of GLP-
1R signaling in diabetic nephropathy: possible mechanism and therapeutic potential.
Kidney Int. 85 (3), 579–589. https://doi.org/10.1038/ki.2013.427.
Fukao, T., Yeung, E., Bailey-Serres, J., 2011. The submergence tolerance regulator
SUB1A mediates crosstalk between submergence and drought tolerance in rice. Plant
Cell 23 (1), 412–427. https://doi.org/10.1105/tpc.110.080325.
Furr, J.R., Armstrong, W.W., 1975. Water and salinity problems of Abadan Island date
gardens. Ann. Date Growers Inst. 52, 14–17.
Fusco, N., Micheletto, L., Dal Corso, G., Borgato, L., Furini, A., 2005. Identification of
cadmium-regulated genes by cDNA-AFLP in the heavy metal accumulator Brassica
juncea L. J. Exp. Bot. 56 (421), 3017–3027. https://doi.org/10.1093/jxb/eri299.
Fusseder, A., Wartinger, A., Hartung, W., Schulze, E.D., Heilmeier, H., 1992. Cytokinins
in the xylem sap of desert-grown almond (Prunus dulcis†) trees: Daily courses and
their possible interactions with abscisic acid and leaf conductance. New Phytol. 122
(1), 45–52. https://doi.org/10.1111/j.1469-8137.1992.tb00051.x.
Galiba, G., Kocsy, G., Kaur-Sawhney, R., Sutka, J., Galston, A.W., 1993. Chromosomal
localization of osmotic and salt stress-induced differential alterations in polyamine
content in wheat. Plant Sci. 92 (2), 203–211. https://doi.org/10.1016/0168-9452
(93)90207-G.
Galvan-Ampudia, C.S., Testerink, C., 2011. Salt stress signals shape the plant root. Curr.
Opin. Plant Biol. 14 (3), 296–302. https://doi.org/10.1016/j.pbi.2011.03.019.
Gao, H., Zhang, Z., Lv, X., Cheng, N., Peng, B., Cao, W., 2016. Effect of 24-epibrassinolide
on chilling injury of peach fruit in relation to phenolic and proline metabolisms.
Postharvest Biol. Technol. 111, 390–397. https://doi.org/10.1016/j.
postharvbio.2015.07.031.
Garg, N., Mohanty, A., Lazarus, N., Schultz, L., Rozzi, T.R., Santhanam, S., Weiss, L.,
Snyder, J.L., Fedder, G.K., Jin, R., 2010. Robust gold nanoparticles stabilized by
trithiol for application in chemiresistive sensors. Nanotechnology 21 (40), 405501.
Ge Y.X., Zhang L.J., Li F.H., Chen Z.B., Wang C., Yao Y.C., Han Z.H., Zhang J., Shi Z.S.
Relationship between jasmonic acid accumulation and senescence in drought-stress.
Afr. J. Agric. Res. 2010 Aug 31;5(15):1978-83.
George, A.P., Nissen, R.J., 1992. Effects of water stress, nitrogen and paclobutrazol on
flowering, yield and fruit quality of the low-chill peach cultivar,‘Flordaprince’. Sci.
Hortic. 49 (3-4), 197–209. https://doi.org/10.1016/0304-4238(92)90157-8.
George, A.P., Broadley, R., Nissen, R.J., Hamill, S., 2004, September. Breeding and
selecting new varieties and rootstocks of custard apple (Annona spp. hybrids) in
subtropical Australia. In: Proceedings of the International Symposium on Harnessing
the Potential of Horticulture in the Asian-Pacific Region, 694, pp. 125–128. https://
doi.org/10.17660/ActaHortic.2005.694.18.
Ghorbani, B., Pakkish, Z., 2014. Brassinosteroid enhances cold stress tolerance of
Washington navel orange (Citrus sinensis L.) fruit by regulating antioxidant enzymes
during storage. Agriculturae Conspectus Scientificus 79 (2), 109–114. https://hrcak.
srce.hr/129092.
Gibbs, D.J., Conde, J.V., Berckhan, S., Prasad, G., Mendiondo, G.M., Holdsworth, M.J.,
2015. Group VII ethylene response factors coordinate oxygen and nitric oxide signal
transduction and stress responses in plants. Plant Physiol. 169 (1), 23–31. https://
doi.org/10.1104/pp.15.00338.
Gimeno, V., Syvertsen, J.P., Nieves, M., Simón, I., Martínez, V., García-Sánchez, F., 2009.
Additional nitrogen fertilization affects salt tolerance of lemon trees on different
rootstocks. Sci. Hortic. 121 (3), 298–305. https://doi.org/10.1016/j.
scienta.2009.02.019.
16
Scientia Horticulturae 304 (2022) 111302
Gómez-Cadenas, A., Pozo, O.J., García-Augustín, P., Sancho, J.V., 2002. Direct analysis
of abscisic acid in crude plant extracts by liquid chromatography—electrospray/
tandem mass spectrometry. Phytochem. Anal. 13 (4), 228–234. https://doi.org/
10.1002/pca.640.
Gómez-Cadenas, A., Tadeo, F.R., Talon, M., Primo-Millo, E., 1996. Leaf abscission
induced by ethylene in water-stressed intact seedlings of Cleopatra mandarin
requires previous abscisic acid accumulation in roots. Plant Physiol. 112 (1),
401–408. https://doi.org/10.1104/pp.112.1.401.
Gonai, T., Kawahara, S., Tougou, M., Satoh, S., Hashiba, T., Hirai, N., Kawaide, H.,
Kamiya, Y., Yoshioka, T., 2004. Abscisic acid in the thermoinhibition of lettuce seed
germination and enhancement of its catabolism by gibberellin. J. Exp. Bot. 55 (394),
111–118. https://doi.org/10.1093/jxb/erh023.
Gong, C., Wei, X., Wang, X., Wang, Y., Guo, G., Mao, Y., Luo, F., Qian, Z., 2010.
Biodegradable self-assembled PEG–PCL–PEG micelles for hydrophobic honokiol
delivery: I. Preparation and characterization. Nanotechnology 21 (21), 215103.
Gonzalez-Olmedo, J., Cordova, A., Aragon, C., Pina, D., Rivas, M., Rodriguez, R., 2004.
Tissue culture effect of an analogue of brassinosteroid on FHIA-18 plantlets exposed
to thermal stress. Infomusa 13, 18.
Granier, A., Reichstein, M., Bréda, N., Janssens, I.A., Falge, E., Ciais, P., Grünwald, T.,
Aubinet, M., Berbigier, P., Bernhofer, C., Buchmann, N., 2007. Evidence for soil
water control on carbon and water dynamics in European forests during the
extremely dry year: 2003. Agric. For. Meteorol. 143 (1-2), 123–145. https://doi.org/
10.1016/j.agrformet.2006.12.004.
Greenway, H., Munns, R., 1983. Interactions between growth, uptake of Cl− and Na+,
and water relations of plants in saline environments. II. Highly vacuolated cells.
Plant, Cell Environ. 6 (7), 575–589. https://doi.org/10.1111/j.1365-3040.1983.
tb01171.x.
Guda, C., Lee, S.B., Daniell, H., 2000. Stable expression of a biodegradable protein-based
polymer in tobacco chloroplasts. Plant Cell Rep. 19 (3), 257–262. https://doi.org/
10.1111/j.1365-3040.1983.tb01171.x.
Guo, T., Wang, N., Xue, Y., Guan, Q., van Nocker, S., Liu, C., Ma, F., 2019.
Overexpression of the RNA binding protein MhYTP1 in transgenic apple enhances
drought tolerance and WUE by improving ABA level under drought condition. Plant
Sci. 280, 397–407. https://doi.org/10.1016/j.plantsci.2018.11.018.
Gupta, A., Sinha, R., Fernandes, J.L., Abdelrahman, M., Burritt, D.J., Tran, L.S.P., 2020.
Phytohormones regulate convergent and divergent responses between individual
and combined drought and pathogen infection. Crit. Rev. Biotechnol. 40 (3),
320–340. https://doi.org/10.1080/07388551.2019.1710459.
Gururani, M.A., Mohanta, T.K., Bae, H., 2015. Current understanding of the interplay
between phytohormones and photosynthesis under environmental stress. Int. J. Mol.
Sci. 16 (8), 19055–19085. https://doi.org/10.3390/ijms160819055.
Hannah, M.A., Heyer, A.G., Hincha, D.K., 2005. A global survey of gene regulation
during cold acclimation in Arabidopsis thaliana. PLos Genet. 1 (2) https://doi.org/
10.1371/journal.pgen.0010026.
Harfouche, A., Meilan, R., Altman, A., 2014. Molecular and physiological responses to
abiotic stress in forest trees and their relevance to tree improvement. Tree Physiol.
34 (11), 1181–1198. https://doi.org/10.1093/treephys/tpu012.
Heidari, M., Jamshid, P., 2010. Interaction between salinity and potassium on grain
yield, carbohydrate content and nutrient uptake in pearl millet. J. Agric. Biol. Sci 5,
39–46.
Herbette, S., Roeckel-Drevet, P., Drevet, J.R., 2007. Seleno-independent glutathione
peroxidases: more than simple antioxidant scavengers. FEBS J. 274 (9), 2163–2180.
https://doi.org/10.1111/j.1742-4658.2007.05774.x.
Hirano, K., Kouketu, E., Katoh, H., Aya, K., Ueguchi-Tanaka, M., Matsuoka, M., 2012.
The suppressive function of the rice DELLA protein SLR1 is dependent on its
transcriptional activation activity. Plant J. 71 (3), 443–453. https://doi.org/
10.1111/j.1365-313X.2012.05000.x.
Hou, H., Lv, L., Huo, H., Dai, H., Zhang, Y., 2020. Genome-wide identification of the ABA
receptors genes and their response to abiotic stress in apple. Plants 9 (8), 1028.
https://doi.org/10.3390/plants9081028.
Houimli, S.I.M., Denden, M., Mouhandes, B.D., 2010. Effects of 24-epibrassinolide on
growth, chlorophyll, electrolyte leakage and proline by pepper plants under NaCl-
stress. Eurasian J. Biosci. 4 (1), 96–104. https://doi.org/10.5053/
ejobios.2010.4.0.12.
Hsu, Y.T., Kao, C.H., 2007. Toxicity in leaves of rice exposed to cadmium is due to
hydrogen peroxide accumulation. Plant Soil 298 (1-2), 231–241. https://doi.org/
10.1007/s11104-007-9357-7.
Hu, W., Zuo, J., Hou, X., Yan, Y., Wei, Y., Liu, J., Li, M., Xu, B., Jin, Z., 2015. The auxin
response factor gene family in banana: genome-wide identification and expression
analyses during development, ripening, and abiotic stress. Front. Plant Sci. 6, 742.
https://doi.org/10.1007/s11104-007-9357-7.
Hu, W.H., Xiao, Y.A., Zeng, J.J., Hu, X.H., 2010. Photosynthesis, respiration and
antioxidant enzymes in pepper leaves under drought and heat stresses. Biologia
Plantarum 54 (4), 761–765. https://doi.org/10.1007/s10535-010-0137-5.
Hu, Y.R., Ma, H., Zou, Z.Y., He, K., Xiao, Y.B., Wang, Y., Feng, M., Ye, X.L., Li, X.G., 2017.
Activation of Akt and JNK/Nrf2/NQO1 pathway contributes to the protective effect
of coptisine against AAPH-induced oxidative stress. Biomed. Pharmacother. 85,
313–322. https://doi.org/10.1016/j.biopha.2016.11.031.
Hu, Z., Weijian, L., Yali, F., Huiquan, L., 2018. Gibberellic acid enhances postharvest
toon sprout tolerance to chilling stress by increasing the antioxidant capacity during
the short-term cold storage. Sci. Hortic. 237, 184–191. https://doi.org/10.1016/j.
scienta.2018.04.018.
Huang, S., Huang, Q., Chang, J., Zhu, Y., Leng, G., Xing, L., 2015. Drought structure
based on a nonparametric multivariate standardized drought index across the Yellow
River basin, China. J. Hydrol. 530, 127–136. https://doi.org/10.1016/j.
jhydrol.2015.09.042.
V. Rachappanavar et al.
Hussain, MA, Fahad, S, Rahat, S, Muhammad, FJ, Muhammad, M, Qasid, A, et al., 2020a.
Multifunctional role of brassinosteroid and its analogues in plants. Plant Growth
Regul. https://doi.org/10.1007/s10725-020-00647-8.
Hyun, Y., Richter, R., Vincent, C., Martinez-Gallegos, R., Porri, A., Coupland, G., 2016.
Multi-layered regulation of SPL15 and cooperation with SOC1 integrate endogenous
flowering pathways at the Arabidopsis shoot meristem. Dev. Cell 37 (3), 254–266.
https://doi.org/10.1016/j.devcel.2016.04.001.
Iqbal, N., Umar, S., Khan, N.A., Khan, M.I.R., 2014. A new perspective of phytohormones
in salinity tolerance: regulation of proline metabolism. Environ. Exp. Bot. 100,
34–42. https://doi.org/10.1016/j.envexpbot.2013.12.006.
Issarakraisila, M., Ma, Q., Turner, D.W., 2007. Photosynthetic and growth responses of
juvenile Chinese kale (Brassica oleracea var. alboglabra) and Caisin (Brassica rapa
subsp. parachinensis) to waterlogging and water deficit. Sci. Hortic. 111 (2),
107–113. https://doi.org/10.1016/j.scienta.2006.10.017.
Iuchi, Y., Obora, Y., Ishii, Y., 2010. Iridium-catalyzed α-alkylation of acetates with
primary alcohols and diols. J. Am. Chem. Soc. 132 (8), 2536–2537. https://doi.org/
10.1021/ja9106989.
Ivanov, R.C., da Luz, C.A., Zorel Jr, H.E., Pereira Filho, J.I., 2016. Behavior of calcium
aluminate cement (CAC) in the presence of hexavalent chromium. Cement Concrete
Composites 73, 114–122. https://doi.org/10.1016/j.cemconcomp.2016.07.006.
Jackson, M.B., Young, S.F., Hall, K.C., 1988. Are roots a source of abscisic acid for the
shoots of flooded pea plants? J. Exp. Bot. 39 (12), 1631–1637. https://doi.org/
10.1093/jxb/39.12.1631.
Jacobsen, S.E., Olszewski, N.E., 1993. Mutations at the SPINDLY locus of Arabidopsis
alter gibberellin signal transduction. Plant Cell 5 (8), 887–896. https://doi.org/
10.1105/tpc.5.8.887.
Jain, M., Khurana, J.P., 2009. Transcript profiling reveals diverse roles of auxin-
responsive genes during reproductive development and abiotic stress in rice. FEBS J.
276 (11), 3148–3162. https://doi.org/10.1111/j.1742-4658.2009.07033.x.
Jain, R., Srivastava, S., Solomon, S., Shrivastava, A.K., Chandra, A., 2010. Impact of
excess zinc on growth parameters, cell division, nutrient accumulation,
photosynthetic pigments and oxidative stress of sugarcane (Saccharum spp.). Acta
Physiol. Plant. 32 (5), 979–986. https://doi.org/10.1007/s11738-010-0487-9.
Jalili, F., Khavazi, K., Pazira, E., Nejati, A., Rahmani, H.A., Sadaghiani, H.R.,
Miransari, M., 2009. Isolation and characterization of ACC deaminase-producing
fluorescent pseudomonads, to alleviate salinity stress on canola (Brassica napus L.)
growth. J. Plant Physiol. 166 (6), 667–674. https://doi.org/10.1016/j.
jplph.2008.08.004.
Jan, S, Noman, A, Kaya, C, Ashraf, M, Alyemeni, MN, Ahmad, P., 2020. 24-
Epibrassinolide alleviates the injurious effects of Cr (VI) toxicity in tomato plants:
insights into growth, physio-biochemical attributes, antioxidant activity and
regulation of Ascorbate–glutathione and Glyoxalase cycles. J. Plant Growth Regul.
1–18. https://doi.org/10.1007/s00344-020-10169-2.
Janeczko, A., 2011. The significance of ethanol as a hormone solvent in experiments on
the physiological activity of brassinosteroids. Brassinosteroids: A Class of Plant
Hormone. Springer, Dordrecht, pp. 361–374.
Jha, U.C., Nayyar, H., Siddique, K.H., 2021. Role of phytohormones in regulating heat
stress acclimation in agricultural crops. J. Plant Growth Regul. 1–24. https://doi.
org/10.1007/s00344-021-10362-x.
Jiang, J., Hou, R., Yang, N., Li, L., Deng, J., Qin, G., Ding, D., 2021. Physiological and
TMT-labeled proteomic analyses reveal important roles of sugar and secondary
metabolism in Citrus junos under cold stress. J. Proteomics, 104145. https://doi.
org/10.1016/j.jprot.2021.104145.
Jin, SH, Li, XQ, Wang, GG, Zhu, XT, 2015. Brassinosteroids alleviate hightemperature
injury in Ficus concinna seedlings via maintaining higher antioxidant defence and
glyoxalase systems. AoB plants 7 plv009. https://doi.org/10.1093/aobpla/plv009.
Jiroutova, P., Oklestkova, J., Strnad, M., 2018. Crosstalk between brassinosteroids and
ethylene during plant growth and under abiotic stress conditions. Int. J. Mol. Sci. 19
(10), 3283. https://doi.org/10.3390/ijms19103283.
Jogawat, A., Yadav, B., Lakra, N., Singh, A.K., Narayan, O.P., 2021. Crosstalk between
phytohormones and secondary metabolites in the drought stress tolerance of crop
plants: a review. Physiol. Plant. 172 (2), 1106–1132. https://doi.org/10.1111/
ppl.13328.
Jones, H.G., Tardieu, F., 1998. Modelling water relations of horticultural crops: a review.
Sci. Hortic. 74 (1-2), 21–46. https://doi.org/10.1016/S0304-4238(98)00081-8.
Juszczak, I., Bartels, D., 2017. LEA gene expression, RNA stability and pigment
accumulation in three closely related Linderniaceae species differing in desiccation
tolerance. Plant Sci. 255, 59–71. https://doi.org/10.1016/j.plantsci.2016.10.003.
Kang, H.K., Natelson, B.H., Mahan, C.M., Lee, K.Y., Murphy, F.M., 2003. Post-traumatic
stress disorder and chronic fatigue syndrome-like illness among Gulf War veterans: a
population-based survey of 30,000 veterans. Am. J. Epidemiol. 157 (2), 141–148.
https://doi.org/10.1093/aje/kwf187.
Kang, Y., Massagué, J., 2004. Epithelial-mesenchymal transitions: twist in development
and metastasis. Cell 118 (3), 277–279. https://doi.org/10.1016/j.cell.2004.07.011.
Karlidag, H., Yildirim, E., Turan, M., 2009. Salicylic acid ameliorates the adverse effect of
salt stress on strawberry. Scientia Agricola 66, 180–187.
Kende, Hans., 1993. Ethylene biosynthesis. Annu. Rev. Plant Biol. 44 (1), 283–307.
Khan, N., Bano, A., Ali, S., Babar, M.A., 2020b. Crosstalk amongst phytohormones from
planta and PGPR under biotic and abiotic stresses. Plant Growth Regul. 90 (2),
189–203. https://doi.org/10.1007/s10725-020-00571-x.
Khan, A., Bilal, S., Khan, A.L., Imran, M., Al-Harrasi, A., Al-Rawahi, A., Lee, I.J., 2020a.
Silicon-mediated alleviation of combined salinity and cadmium stress in date palm
(Phoenix dactylifera L.) by regulating physio-hormonal alteration. Ecotoxicol.
Environ. Saf. 188, 109885 https://doi.org/10.1016/j.ecoenv.2019.109885.
Khan, Hafiz Azhar Ali, Shad, Sarfraz Ali, Akram, Waseem, 2013. Resistance to new
chemical insecticides in the house fly, Musca domestica L., from dairies in Punjab,
17
Scientia Horticulturae 304 (2022) 111302
Pakistan. Parasitol. Res. 112 (5), 2049–2054. https://doi.org/10.1007/s00436-013-
3365-8.
Kim, J.M., To, T.K., Ishida, J., Matsui, A., Kimura, H., Seki, M., 2012. Transition of
chromatin status during the process of recovery from drought stress in Arabidopsis
thaliana. Plant Cell Physiol. 53 (5), 847–856. https://doi.org/10.1093/pcp/pcs053.
Kim, T.H., Böhmer, M., Hu, H., Nishimura, N., Schroeder, J.I., 2010. Guard cell signal
transduction network: advances in understanding abscisic acid, CO2, and Ca2+
signaling. Annu. Rev. Plant Biol. 61, 561–591. https://doi.org/10.1146/annurev.
arplant.52.1.627.
Kinoshita, N., Wang, H., Kasahara, H., Liu, J., MacPherson, C., Machida, Y., Kamiya, Y.,
Hannah, M.A., Chua, N.H., 2012. IAA-Ala Resistant3, an evolutionarily conserved
target of miR167, mediates Arabidopsis root architecture changes during high
osmotic stress. Plant Cell 24 (9), 3590–3602. https://doi.org/10.1105/
tpc.112.097006.
Klingler, J.P., Batelli, G., Zhu, J.K., 2010. ABA receptors: the START of a new paradigm
in phytohormone signalling. J. Exp. Bot. 61 (12), 3199–3210. https://doi.org/
10.1093/jxb/erq151.
Kodur, S., Tisdall, J.M., Tang, C., Walker, R.R., 2010. Accumulation of potassium in
grapevine rootstocks (Vitis) grafted to ‘Shiraz’ as affected by growth, root-traits and
transpiration. Vitis 49, 7–13.
Kolomeichuk, LV, Efimova, MV, Zlobin, IE, Kreslavski, VD, Ol’ga, KM, Kovtun, IS, et al.,
2020. 24-Epibrassinolide alleviates the toxic effects of NaCl on photosynthetic
processes in potato plants. Photosynth. Res. 1–13. https://doi.org/10.1007/s11120-
020-00708-z.
Kovtun, Y., Chiu, W.L., Zeng, W., Sheen, J., 1998. Suppression of auxin signal
transduction by a MAPK cascade in higher plants. Nature 395 (6703), 716–720.
https://doi.org/10.1038/27240.
Krishna, S., Miller, L.S., 2012. Host–pathogen interactions between the skin and
Staphylococcus aureus. Curr. Opin. Microbiol. 15 (1), 28–35. https://doi.org/
10.1016/j.mib.2011.11.003.
Kumari, S., Thakur, A., 2019. The effects of water stress and Brassinosteroid on apple
varieties. Int. J. Econ. Plants 6 (1), 1–6. https://doi.org/10.23910/IJEP/
2019.6.1.0278.
Kuromori, T., Fujita, M., Urano, K., Tanabata, T., Sugimoto, E., Shinozaki, K., 2016.
Overexpression of AtABCG25 enhances the abscisic acid signal in guard cells and
improves plant water use efficiency. Plant Sci. 251, 75–81. https://doi.org/10.1016/
j.plantsci.2016.02.019.
Larkindale, J., Vierling, E., 2008. Core genome responses involved in acclimation to high
temperature. Plant Physiol. 146 (2), 748–761.
Larson, K.D., Schaffer, B., Davies, F.S., 1991. Flooding, leaf gas exchange, and growth of
mango in containers. J. Am. Soc. Horticul. Sci. 116 (1), 156–160. https://doi.org/
10.21273/JASHS.116.1.156.
Lavenus, J., Goh, T., Roberts, I., Guyomarc’h, S., Lucas, M., De Smet, I., Fukaki, H.,
Beeckman, T., Bennett, M., Laplaze, L., 2013. Lateral root development in
Arabidopsis: fifty shades of auxin. Trends Plant Sci. 18 (8), 450–458. https://doi.
org/10.1016/j.tplants.2013.04.006.
León, P., Sheen, J., 2003. Sugar and hormone connections. Trends Plant Sci. 8 (3),
110–116. https://doi.org/10.1016/S1360-1385(03)00011-6.
Levitt, J., 1980. Responses of Plants to Environmental Stress, Volume 1: Chilling,
Freezing, and High Temperature Stresses. Academic Press.
Li, L., Lu, X., Ma, H., Lyu, D., 2017. Jasmonic acid regulates the ascorbate–glutathione
cycle in Malus baccata Borkh. roots under low root-zone temperature. Acta Physiol.
Plant. 39 (8), 1–11. https://doi.org/10.1007/s11738-017-2469-7.
Lim, S., Park, J., Lee, N., Jeong, J., Toh, S., Watanabe, A., Kim, J., Kang, H., Kim, D.H.,
Kawakami, N., Choi, G., 2013. ABA-INSENSITIVE3, ABA-INSENSITIVE5, and
DELLAs interact to activate the expression of SOMNUS and other high-temperature-
inducible genes in imbibed seeds in Arabidopsis. Plant Cell 25 (12), 4863–4878.
https://doi.org/10.1105/tpc.113.118604.
Liu, X.J., Dong, Y.H., Liu, X., You, C.X., Hao, Y.J., 2019. A C2-domain phospholipid-
binding protei MdCAIP1 positively regulates salt and osmotic stress tolerance in
apple. Plant Cell, Tissue Organ Culture (PCTOC) 138 (1), 29–39. https://doi.org/
10.1007/s11240-019-01600-y.
Liu, Y., Burgos, J.S., Deng, Y., Srivastava, R., Howell, S.H., Bassham, D.C., 2012.
Degradation of the endoplasmic reticulum by autophagy during endoplasmic
reticulum stress in Arabidopsis. Plant Cell 24 (11), 4635–4651. https://doi.org/
10.1105/tpc.112.101535.
Farooq, M.G., Liu, X.H. and Melville, I.D., International Business Machines Corp, 2009a.
Test structures for electrically detecting back end of the line failures and methods of
making and using the same. U.S. Patent 7, 622,737.
Llanes, A., Andrade, A., Masciarelli, O., Alemano, S., Luna, V., 2016. Drought and salinity
alter endogenous hormonal profiles at the seed germination phase. Seed Sci. Res. 26
(1), 1–13. https://doi.org/10.1017/S0960258515000331.
Lurie, S., Crisosto, C.H., 2005. Chilling injury in peach and nectarine. Postharvest Biol.
Technol. 37 (3), 195–208. https://doi.org/10.1016/j.postharvbio.2005.04.012.
Maggio, A., Barbieri, G., Raimondi, G., De Pascale, S., 2010. Contrasting effects of GA 3
treatments on tomato plants exposed to increasing salinity. J. Plant Growth Regul.
29 (1), 63–72. https://doi.org/10.1007/s00344-009-9114-7.
Mahajan, S., Tuteja, N., 2005. Cold, salinity and drought stresses: an overview. Arch.
Biochem. Biophys. 444 (2), 139–158. https://doi.org/10.1016/j.abb.2005.10.018.
Mahmood, M., Bidabadi, S.S., Ghobadi, C., Gray, D.J., 2012. Effect of methyl jasmonate
treatments on alleviation of polyethylene glycol-mediated water stress in banana
(Musa acuminata cv.‘Berangan’, AAA) shoot tip cultures. Plant Growth Regul. 68 (2),
161–169. https://doi.org/10.1007/s10725-012-9702-6.
Mahouachi, J., López-Climent, M.F., Gómez-Cadenas, A., 2014. Hormonal and
hydroxycinnamic acids profiles in banana leaves in response to various periods of
water stress. Sci. World J. 2014, 540962 https://doi.org/10.1155/2014/540962.
V. Rachappanavar et al.
Marcelis, L.F.M., Heuvelink, E., Goudriaan, J., 1998. Modelling biomass production and
yield of horticultural crops: a review. Sci. Hortic. 74 (1-2), 83–111. https://doi.org/
10.1016/S0304-4238(98)00083-1.
Masle, J., Gilmore, S.R., Farquhar, G.D., 2005. The ERECTA gene regulates plant
transpiration efficiency in Arabidopsis. Nature 436 (7052), 866–870. https://doi.
org/10.1038/nature03835.
Masood, A., Iqbal, N., Khan, N.A., 2012. Role of ethylene in alleviation of cadmium-
induced photosynthetic capacity inhibition by sulphur in mustard. Plant Cell
Environ. 35 (3), 524–533. https://doi.org/10.1111/j.1365-3040.2011.02432.x.
Medici, A., Laloi, M., Atanassova, R., 2014. Profiling of sugar transporter genes in
grapevine coping with water deficit. FEBS Lett. 588 (21), 3989–3997. https://doi.
org/10.1016/j.febslet.2014.09.016.
Meena, G.S., Singh, A.K., Borad, S., Panjagari, N.R., 2016. Effect of concentration,
homogenization and stabilizing salts on heat stability and rheological properties of
cow skim milk ultrafiltered retentate. J. Food Sci. Technol. 53 (11), 3960–3968.
https://doi.org/10.1007/s13197-016-2173-4.
Miranda, P.L., Cordeau, J.F., Ferreira, D., Jans, R., Morabito, R., 2018. A decomposition
heuristic for a rich production routing problem. Comput. Operat. Res. 98, 211–230.
https://doi.org/10.1016/j.cor.2018.05.004.
Mishra, A.K., Singh, V.P., 2011. Drought modeling–a review. J. Hydrol. 403 (1–2),
157–175. https://doi.org/10.1016/j.jhydrol.2011.03.049.
Mittler, R., 2006. Abiotic stress, the field environment and stress combination. Trends
Plant Sci. 11 (1), 15–19. https://doi.org/10.1016/j.tplants.2005.11.002.
Mohamed, H.I., Latif, H.H., 2017. Improvement of drought tolerance of soybean plants
by using methyl jasmonate. Physiol. Mol. Biol. Plants 23 (3), 545–556. https://doi.
org/10.1007/s12298-017-0451-x.
Mohammadkhani, N., Abbaspour, N., 2018. Absorption kinetics and efflux of chloride
and sodium in the roots of four grape genotypes (Vitis l.) differing in salt tolerance.
Iran. J. Sci. Technol. Trans. A 42 (4), 1779–1793. https://doi.org/10.1007/s40995-
017-0445-0.
Monni, S., Salemaa, M., Millar, N., 2000. The tolerance of Empetrum nigrum to copper and
nickel. Environ. Pollut. 109 (2), 221–229. https://doi.org/10.1016/S0269-7491(99)
00264-X.
Mubarik, M.S., Khan, S.H., Sajjad, M., Raza, A., Hafeez, M.B., Yasmeen, T., Rizwan, M.,
Ali, S., Arif, M.S., 2021. A manipulative interplay between positive and negative
regulators of phytohormones: a way forward for improving drought tolerance in
plants. Physiol. Plant. https://doi.org/10.1111/ppl.13325.
Mukhopadhyay, M., Mondal, T.K., 2015. Effect of zinc and boron on growth and water
relations of Camellia sinensis (L.) O. Kuntze cv. T-78. Natl. Acad. Sci. Lett. 38 (3),
283–286. https://doi.org/10.1007/s40009-015-0381-5.
Munns, R., Tester, M., 2008. Mechanisms of salinity tolerance. Annu. Rev. Plant Biol. 59,
651–681. https://doi.org/10.1146/annurev.arplant.59.032607.092911.
Munns, R., James, R.A., Läuchli, A., 2006. Approaches to increasing the salt tolerance of
wheat and other cereals. J. Exp. Bot. 57 (5), 1025–1043. https://doi.org/10.1093/
jxb/erj100.
Mutlu, S., Karadagoglu, O., Atici, O., Nalbantoglu, B., 2013. Protective role of salicylic
acid applied before cold stress on antioxidative system and protein patterns in barley
apoplast. Biol. Plant. 57, 507–513. https://doi.org/10.1007/s10535-013-0322-4,
10.1007/s10535-013-0322-4.
Naeini, M.R., Khoshgoftarmanesh, A.H., Lessani, H., Fallahi, E, 2004. Effects of sodium
chloride-induced salinity on mineral nutrients and soluble sugars in three
commercial cultivars of pomegranate. J. Plant Nutr. 27, 1319–1326. https://doi.org/
10.1081/PLN-200025832.
Naeini, M.R., Khoshgoftarmanesh, A.H., Fallahi, E., 2006. Partitioning of chlorine,
sodium, and potassium and shoot growth of three pomegranate cultivars under
different levels of salinity. J. Plant Nutr. 29 (10), 1835–1843. https://doi.org/
10.1080/01904160600899352.
Nakashima, K., Yamaguchi-Shinozaki, K., Shinozaki, K., 2014. The transcriptional
regulatory network in the drought response and its crosstalk in abiotic stress
responses including drought, cold, and heat. Front. Plant Sci. 5, 170. https://doi.org/
10.3389/fpls.2014.00170.
Nassar, A.H., Hashim, M.F., Hassan, N.S., Abo-Zaid, H., 2004. Effect of gamma
irradiation and phosphorus on growth and oil production of chamomile
(Chamomilla recutita L. Rauschert). Int. J. Agric. Biol. 6 (5), 776–780.
Navarro, A., Sánchez-Blanco, M.J., Bañon, S., 2007. Influence of paclobutrazol on water
consumption and plant performance of Arbutus unedo seedlings. Sci. Hortic. 111 (2),
133–139. https://doi.org/10.1016/j.scienta.2006.10.014.
Ng, L.M., Melcher, K., Teh, B.T., Xu, H.E., 2014. Abscisic acid perception and signaling:
structural mechanisms and applications. Acta Pharmacol. Sin. 35 (5), 567–584.
https://doi.org/10.1038/aps.2014.5.
Ning, J., Li, X., Hicks, L.M., Xiong, L., 2010. A Raf-like MAPKKK gene DSM1 mediates
drought resistance through reactive oxygen species scavenging in rice. Plant Physiol.
152 (2), 876–890. https://doi.org/10.1104/pp.109.149856.
Nishiyama, R., Watanabe, Y., Leyva-Gonzalez, M.A., Van Ha, C., Fujita, Y., Tanaka, M.,
Seki, M., Yamaguchi-Shinozaki, K., Shinozaki, K., Herrera-Estrella, L., Tran, L.S.P.,
2013. Arabidopsis AHP2, AHP3, and AHP5 histidine phosphotransfer proteins
function as redundant negative regulators of drought stress response. Proc. Natl.
Acad. Sci. 110 (12), 4840–4845. https://doi.org/10.1073/pnas.1302265110.
Noriega, G., Caggiano, E., Lecube, M.L., Santa Cruz, D., Batlle, A., Tomaro, M.,
Balestrasse, K.B., 2012. The role of salicylic acid in the prevention of oxidative stress
elicited by cadmium in soybean plants. Biometals 25 (6), 1155–1165. https://doi.
org/10.1007/s10534-012-9577-z.
Ogasawara, N., Hiramasu, T., Ishiyama, K., Fushimi, H., Suzuki, H., Takagi, H., 2001.
Effects of gibberellic acid and temperature on growth and root carbohydrates of
Delphinium seedlings. Plant Growth Regul. 33 (3), 181–187. https://doi.org/
10.1023/A:1017593815746.
18
Scientia Horticulturae 304 (2022) 111302
Ogawa, Atsushi, Kawashima, Choji, Yamauchi, Akira, 2005. Sugar accumulation along
the seminal root axis, as affected by osmotic stress in maize: a possible physiological
basis for plastic lateral root development. Plant Product.Sci. 8 (2), 173–180. https://
doi.org/10.1626/pps.8.173.
Okushima, Y., Fukaki, H., Onoda, M., Theologis, A., Tasaka, M., 2007. ARF7 and ARF19
regulate lateral root formation via direct activation of LBD/ASL genes in
Arabidopsis. Plant Cell 19 (1), 118–130. https://doi.org/10.1105/tpc.106.047761.
Ozturk, M., Turkyilmaz Unal, B., García-Caparrós, P., Khursheed, A., Gul, A.,
Hasanuzzaman, M., 2021. Osmoregulation and its actions during the drought stress
in plants. Physiol. Plant. 172 (2), 1321–1335. https://doi.org/10.1111/ppl.13297.
Pakkish, Z., Ghorbani, B., Najafzadeh, R., 2019. Fruit quality and shelf life improvement
of grape cv. Rish Baba using Brassinosteroid during cold storage. J. Food Measure.
Character. 13 (2), 967–975. https://doi.org/10.1007/s11694-018-0011-2.
Pandey, P., Irulappan, V., Bagavathiannan, M.V., Senthil-Kumar, M., 2017. Impact of
combined abiotic and biotic stresses on plant growth and avenues for crop
improvement by exploiting physio-morphological traits. Front. Plant Sci. 8, 537.
https://doi.org/10.3389/fpls.2017.00537.
Papdi, C., Pérez-Salamó, I., Joseph, M.P., Giuntoli, B., Bögre, L., Koncz, C., Szabados, L.,
2015. The low oxygen, oxidative and osmotic stress responses synergistically act
through the ethylene response factor VII genes RAP 2.12, RAP 2.2 and RAP 2.3. Plant
J. 82 (5), 772–784. https://doi.org/10.1111/tpj.12848.
Parajuli, R., Thoma, G., Matlock, M.D., 2019. Environmental sustainability of fruit and
vegetable production supply chains in the face of climate change: a review. Sci. Total
Environ. 650, 2863–2879. https://doi.org/10.1016/j.scitotenv.2018.10.019.
Park, J.E., Seo, P.J., Lee, A.K., Jung, J.H., Kim, Y.S., Park, C.M., 2007. An Arabidopsis
GH3 gene, encoding an auxin-conjugating enzyme, mediates phytochrome B-
regulated light signals in hypocotyl growth. Plant Cell Physiol. 48 (8), 1236–1241.
https://doi.org/10.1093/pcp/pcm086.
Pasternak, T., Potters, G., Caubergs, R., Jansen, M.A., 2005. Complementary interactions
between oxidative stress and auxins control plant growth responses at plant, organ,
and cellular level. J. Exp. Bot. 56 (418), 1991–2001. https://doi.org/10.1093/jxb/
eri196.
Patanè, C., Saita, A., Sortino, O., 2013. Comparative effects of salt and water stress on
seed germination and early embryo growth in two cultivars of sweet sorghum.
J. Agronomy Crop Sci. 199 (1), 30–37. https://doi.org/10.1111/j.1439-
037X.2012.00531.x.
Patel, N.M., Patel, D.K., Verma, L.R., Patel, M.M., 2010. Effect of cultural and chemical
treatments on fruit set and fruit yield of custard apple (Annona sqamosa Linn.) cv.
Sindhan. Asian J. Hortic. 5 (2), 498–502.
Pauwels, L., Goossens, A., 2011. The JAZ proteins: a crucial interface in the jasmonate
signaling cascade. Plant Cell 23, 3089–3100. https://doi.org/10.1105/
tpc.111.089300 [PMC free article] [PubMed] [Google Scholar].
Pedrosa, A.M., Martins, C.D.P.S., Goncalves, L.P., Costa, M.G.C., 2015. Late
embryogenesis abundant (LEA) constitutes a large and diverse family of proteins
involved in development and abiotic stress responses in sweet orange (Citrus sinensis
L. Osb.). PloS one 10 (12), e0145785.
Peleg, Z., Blumwald, E., 2011. Hormone balance and abiotic stress tolerance in crop
plants. Curr. Opin. Plant Biol. 14 (3), 290–295. https://doi.org/10.1016/j.
pbi.2011.02.001.
Perin, E.C., da Silva Messias, R., Borowski, J.M., Crizel, R.L., Schott, I.B., Carvalho, I.R.,
Rombaldi, C.V., Galli, V., 2019. ABA-dependent salt and drought stress improve
strawberry fruit quality. Food Chem. 271, 516–526. https://doi.org/10.1016/j.
foodchem.2018.07.213.
Proebsting Jr, E.L., Mills, H.H., 1978. A synoptic analysis of peach and cherry flower bud
hardiness [Elevation, soil type, cultural practices, cold resistance]. J. Am. Soc.
Horticultural Sci.
Qin, X., Zeevaart, J.A., 1999. The 9-cis-epoxycarotenoid cleavage reaction is the key
regulatory step of abscisic acid biosynthesis in water-stressed bean. Proc. Natl. Acad.
Sci. 96 (26), 15354–15361. https://doi.org/10.1073/pnas.96.26.15354.
Qin, X., Zeevaart, J.A., 2002. Overexpression of a 9-cis-epoxycarotenoid dioxygenase
gene in Nicotiana plumbaginifolia increases abscisic acid and phaseic acid levels and
enhances drought tolerance. Plant Physiol. 128 (2), 544–551. https://doi.org/
10.1104/pp.010663.
Raga, V., Intrigliolo, D.S., Bernet, G.P., Carbonell, E.A., Asins, M.J., 2016. Genetic
analysis of salt tolerance in a progeny derived from the citrus rootstocks Cleopatra
mandarin and trifoliate orange. Tree Genet. Genom. 12 (3), 34. https://doi.org/
10.1007/s11295-016-0991-1.
Ramoliya, P.J., Pandey, A.N., 2003. Effect of salinization of soil on emergence, growth
and survival of seedlings of Cordia rothii. Forest Ecol. Manag. 176 (1-3), 185–194.
https://doi.org/10.1016/j.foreco.2004.07.020.
Rangan, P., Subramani, R., Kumar, R., Singh, A.K., Singh, R., 2014. Recent advances in
polyamine metabolism and abiotic stress tolerance. Biomed. Res. Int. 2014 https://
doi.org/10.1155/2014/239621.
Rao, R., Li, Y., 2003. Management of flooding effects on growth of vegetable and selected
field crops. HortTechnology 13 (4), 610–616. https://doi.org/10.21273/
HORTTECH.13.4.0610.
Rauser, W.E., Dumbroff, E.B., 1981. Effects of excess cobalt, nickel and zinc on the water
relations of Phaseolus vulgaris. Environ. Exp. Bot. 21 (2), 249–255. https://doi.org/
10.1016/0098-8472(81)90032-0.
Ravi, I., Uma, S., Vaganan, M.M., Mustaffa, M.M., 2013. Phenotyping bananas for
drought resistance. Front. Physiol. 4, 9. https://doi.org/10.3389/fphys.2013.00009.
Recatalá, L., Sánchez, J., Arbelo, C., Sacristán, D., 2010. Testing the validity of a Cd soil
quality standard in representative Mediterranean agricultural soils under an
accumulator crop. Sci. Total Environ. 409 (1), 9–18. https://doi.org/10.1016/j.
scitotenv.2010.09.021.
V. Rachappanavar et al.
Reusche, M., Klásková, J., Thole, K., Truskina, J., Novák, O., Janz, D., Strnad, M.,
Spíchal, L., Lipka, V., Teichmann, T., 2013. Stabilization of cytokinin levels enhances
Arabidopsis resistance against Verticillium longisporum. Mol. Plant-Microbe Interact.
26 (8), 850–860. https://doi.org/10.1094/MPMI-12-12-0287-R.
Reyes-Díaz, M., Lobos, T., Cardemil, L., Nunes-Nesi, A., Retamales, J., Jaakola, L., et al.,
2016. Methyl Jasmonate: an alternative for improving the quality and health
properties of fresh fruits. Molecules 21, E567. https://doi.org/10.3390/
molecules21060567 [PMC free article] [PubMed] [Google Scholar].
Rhaman, M.S., Imran, S., Rauf, F., Khatun, M., Baskin, C.C., Murata, Y.,
Hasanuzzaman, M., 2021. Seed priming with phytohormones: an effective approach
for the mitigation of abiotic stress. Plants 10 (1), 37. https://doi.org/10.3390/
plants10010037.
Ristic, Z., Bukovnik, U., Prasad, P.V., West, M., 2008. A model for prediction of heat
stability of photosynthetic membranes. Crop Sci. 48 (4), 1513–1522. https://doi.
org/10.1007/s00018-003-2332-4.
Rodriguez, E., Santos, C., Azevedo, R., Moutinho-Pereira, J., Correia, C., Dias, M.C.,
2012. Chromium (VI) induces toxicity at different photosynthetic levels in pea. Plant
Physiol. Biochem. 53, 94–100. https://doi.org/10.1016/j.plaphy.2012.01.013.
Roghabadi, M.A., Pakkish, Z.A.H.R.A., 2014. Role of brassinosteroid on yield, fruit
quality and postharvest storage of’Tak Danehe Mashhad’sweet cherry (Prunus avium
L.). Agri. Commun. 2 (4), 49–56.
Roy, M., McDonald, L.M., 2015. Metal uptake in plants and health risk assessments in
metal-contaminated smelter soils. Land Degradat. Dev. 26 (8), 785–792. https://doi.
org/10.1002/ldr.2237.
Roychoudhury, A., Paul, S., Basu, S., 2013. Cross-talk between abscisic acid-dependent
and abscisic acid-independent pathways during abiotic stress. Plant Cell Rep. 32 (7),
985–1006. https://doi.org/10.1007/s00299-013-1414-5.
Ruan, J., Zhou, Y., Zhou, M., Yan, J., Khurshid, M., Weng, W., Cheng, J., Zhang, K., 2019.
Jasmonic acid signaling pathway in plants. Int. J. Mol. Sci. 20 (10), 2479. https://
doi.org/10.3390/ijms20102479.
Rui, J., Zhen, Y., 2016. An improved centroid localization algorithm based on iterative
computation for wireless sensor network. Acta Phys. Sin. 65 (3).
Sadeghi, F., Shekafandeh, A., 2014. Effect of 24-epibrassinolide on salinity-induced
changes in loquat (Eriobotrya japonica Lindl). J. Appl. Bot. Food Qual. 87 https://
doi.org/10.5073/JABFQ.2014.087.026.
Saradhi, P.P., Suzuki, I., Katoh, A., Sakamoto, A., Sharmila, P., Shi, D.J., Murata, N.,
2000. Protection against the photo-induced inactivation of the photosystem II
complex by abscisic acid. Plant Cell Environ. 23 (7), 711–718. https://doi.org/
10.1046/j.1365-3040.2000.00579.x.
Satisha, J., Prakash, G.S., Murti, G.S.R., Upreti, K.K., 2005. Response of grape genotypes
to water deficit: root, shoot, growth and endogenous hormones. Indian J. Plant
Physiol. 10 (3), 225.
Savchenko, T., Kolla, V.A., Wang, C.Q., Nasafi, Z., Hicks, D.R., Phadungchob, B.,
Chehab, W.E., Brandizzi, F., Froehlich, J., Dehesh, K., 2014. Functional convergence
of oxylipin and abscisic acid pathways controls stomatal closure in response to
drought. Plant Physiol. 164 (3), 1151–1160. https://doi.org/10.1104/
pp.113.234310.
Save, R., Biel, C., Domingo, R., Ruiz-Sánchez, M.C., Torrecillas, A., 1995. Some
physiological and morphological characteristics of citrus plants for drought
resistance. Plant Sci. 110 (2), 167–172. https://doi.org/10.1016/0168-9452(95)
04202-6.
Schaffer, S.W., Azuma, J., Madura, J.D., 1995. Mechanisms underlying taurine-mediated
alterations in membrane function. Amino Acids 8 (3), 231–246. https://doi.org/
10.1007/BF00806821.
Schmutz, U., 2000. Effect of salt stress (NaCl) on whole plant CO2 gas exchange in
mango. Acta Hortic 509, 269–276. https://doi.org/10.17660/
ActaHortic.2000.509.29.
Scholefield, P.B., Oag, D.R., Sedgley, M., 1986. The relationship between vegetative and
reproductive development in the mango in northern Australia. Aust. J. Agric. Res. 37
(4), 425–433. https://doi.org/10.1071/AR9860425.
Seidel, C., Walz, A., Park, S., Cohen, J.D., Ludwig-Müller, J., 2006. Indole-3-acetic acid
protein conjugates: novel players in auxin homeostasis. Plant Biol. 8 (03), 340–345.
https://doi.org/10.1055/s-2006-923802.
Sengupta, S., Patra, B., Ray, S., Majumder, A.L., 2008. Inositol methyl tranferase from a
halophytic wild rice, Porteresia coarctata Roxb. (Tateoka): regulation of pinitol
synthesis under abiotic stress. Plant Cell Environ. 31 (10), 1442–1459. https://doi.
org/10.1111/j.1365-3040.2008.01850.x.
Seo, J.S., Joo, J., Kim, M.J., Kim, Y.K., Nahm, B.H., Song, S.I., Cheong, J.J., Lee, J.S.,
Kim, J.K., Choi, Y.D., 2011. OsbHLH148, a basic helix-loop-helix protein, interacts
with OsJAZ proteins in a jasmonate signaling pathway leading to drought tolerance
in rice. Plant J. 65 (6), 907–921. https://doi.org/10.1111/j.1365-313X.2010.04477.
x.
Serra, I., Strever, A., Myburgh, P.A., Deloire, A., 2014. the interaction between rootstocks
and cultivars (V itis vinifera L.) to enhance drought tolerance in grapevine. Aust. J.
Grape Wine Res. 20 (1), 1–14. https://doi.org/10.1111/ajgw.12054.
Sessa, G., Meller, Y., Fluhr, R., 1995. A GCC element and a G-box motif participate in
ethylene-induced expression of the PRB-1b gene. Plant Mol. Biol. 28 (1), 145–153.
https://doi.org/10.1007/BF00042046.
Shannon, M.C., Grieve, C.M., 1998. Tolerance of vegetable crops to salinity. Sci. Hortic.
78 (1-4), 5–38. https://doi.org/10.1016/S0304-4238(98)00189-7.
Sharma, A, Kumar, V, Thukral, AK, Bhardwaj, R., 2017. 24-epibrassinolide restores the
synthesis of proteins and amino acids in Brassica juncea L. leaves under imidacloprid
stress. J. Hortic. Res. 25 (2), 85–90.
Shen, C., Bai, Y., Wang, S., Zhang, S., Wu, Y., Chen, M., Jiang, D., Qi, Y., 2010.
Expression profile of PIN, AUX/LAX and PGP auxin transporter gene families in
19
Scientia Horticulturae 304 (2022) 111302
Sorghum bicolor under phytohormone and abiotic stress. FEBS J. 277 (14),
2954–2969. https://doi.org/10.1111/j.1742-4658.2010.07706.x.
Shen, X., Guo, X., Zhao, D., Zhang, Q., Jiang, Y., Wang, Y., Peng, X., Wei, Y., Zhai, Z.,
Zhao, W., Li, T., 2017. Cloning and expression profiling of the PacSnRK2 and
PacPP2C gene families during fruit development, ABA treatment, and dehydration
stress in sweet cherry. Plant Physiol. Biochem. 119, 275–285. https://doi.org/
10.1016/j.plaphy.2017.08.025.
Shi, H., Chen, L., Ye, T., Liu, X., Ding, K., Chan, Z, 2014. Modulation of auxin content in
Arabidopsis confers improved drought stress resistance. Plant Physiol. Biochem. 82,
209–217. https://doi.org/10.1016/j.plaphy.2014.06.008.
Shibasaki, K., Uemura, M., Tsurumi, S., Rahman, A., 2009. Auxin response in Arabidopsis
under cold stress: underlying molecular mechanisms. Plant Cell 21 (12), 3823–3838.
https://doi.org/10.1105/tpc.109.069906.
Shinozaki, K., Yamaguchi-Shinozaki, K., 2007. Gene networks involved in drought stress
response and tolerance. J. Exp. Bot. 58 (2), 221–227. https://doi.org/10.1093/jxb/
erl164.
Shkolnik-Inbar, D., Bar-Zvi, D., 2010. ABI4 mediates abscisic acid and cytokinin
inhibition of lateral root formation by reducing polar auxin transport in Arabidopsis.
Plant Cell 22 (11), 3560–3573. https://doi.org/10.1105/tpc.110.074641.
Shomeili, M., Nabipour, M., Meskarbashee, M., Memari, H.R., 2011. Effects of gibberellic
acid on sugarcane plants exposed to salinity under a hydroponic system. Afr. J. Plant
Sci. 5 (10), 609–616. https://doi.org/10.5897/AJPS.9000093.
Siddiqui, H, Ahmed, KBM, Hayat, S., 2018. Comparative effect of 28-homobrassinolide
and 24-epibrassinolide on the performance of different components influencing the
photosynthetic machinery in Brassica juncea L. Plant Physiol. Biochem. 129,
198–212. https://doi.org/10.1016/j.plaphy.2018.05.027.
Sinha, B.K., Watson, D.C., 2007. Stress, coping and psychological illness: a cross-cultural
study. Int. J. Stress Manage. 14 (4), 386. https://psycnet.apa.org/doi/10.1037/107
2-5245.14.4.386.
Sirhindi, G., Mir, M.A., Sharma, P., Gill, S.S., Kaur, H., Mushtaq, R., 2015. Modulatory
role of jasmonic acid on photosynthetic pigments, antioxidants and stress markers of
Glycine max L. under nickel stress. Physiol. Mol, Biol. Plants 21 (4), 559–565.
https://doi.org/10.1007/s12298-015-0320-4.
Skinner, J., Staiger, D., 2005. Technology adoption from hybrid corn to beta blockers
(No. w11251). Natl. Bureau Econ. Res.
Spoel, S.H., Koornneef, A., Claessens, S.M., Korzelius, J.P., Van Pelt, J.A., Mueller, M.J.,
Buchala, A.J., Métraux, J.P., Brown, R., Kazan, K., Van Loon, L.C., 2003. NPR1
modulates cross-talk between salicylate-and jasmonate-dependent defense pathways
through a novel function in the cytosol. Plant Cell 15 (3), 760–770. https://doi.org/
10.1105/tpc.009159.
Starck, Z., Kozińska, M., 1980. Effect of phytohormones on absorption and distribution of
ions in salt-stressed bean plants. Acta Societatis Botanicorum Poloniae 49 (1-2),
111–125. https://doi.org/10.5586/asbp.1980.010.
Stern, R.A., Naor, A., Bar, N., Gazit, S., Bravdo, B.A., 2003. Xylem-sap zeatin-riboside and
dihydrozeatin-riboside levels in relation to plant and soil water status and flowering
in ‘Mauritius’ lychee. Sci. Hortic. 98 (3), 285–291. https://doi.org/10.1016/S0304-
4238(02)00229-7.
Stoll, G., Jander, S., Schroeter, M., 2000. Cytokines in CNS disorders: neurotoxicity
versus neuroprotection. Advances in Dementia Research. Springer, Vienna,
pp. 81–89. https://doi.org/10.1007/978-3-7091-6781-6_11.
Su, Q., Zheng, X., Tian, Y., Wang, C., 2020. Exogenous brassinolide alleviates salt stress
in Malus hupehensis Rehd. by regulating the transcription of NHX-Type Na+ (K+)/H
+ antiporters. Front. Plant Sci. 11, 38. https://doi.org/10.3389/fpls.2020.00038.
Sulochanamma, B.N., Yellamanda Reddy, T., Subbi Reddy, G., 2005. Effect of basin and
drip irrigation on growth, yield and water use efficiency in pomegranate cv. Ganesh.
Acta horticulturae. https://doi.org/10.17660/ActaHortic.2005.696.48.
Sun, F., Zhang, W., Hu, H., Li, B., Wang, Y., Zhao, Y., Li, K., Liu, M., Li, X., 2008. Salt
modulates gravity signaling pathway to regulate growth direction of primary roots in
Arabidopsis. Plant Physiol. 146 (1), 178–188. https://doi.org/10.1104/
pp.107.109413.
Suzuki, N., Rivero, R.M., Shulaev, V., Blumwald, E., Mittler, R., 2014. Abiotic and biotic
stress combinations. New Phytol. 203 (1), 32–43. https://doi.org/10.1111/
nph.12797.
Swietlik, D., Miller, S.S., 1983. The effect of paclobutrazol on growth and response to
water stress of apple seedlings. J. Am. Soc. Hortic. Sci. 108 (6), 1076–1080.
Tabatabaei, S.J., 2007. Salinity stress and olive: an overview. Plant Stress 1 (1), 105–112.
Tan, B.C., Schwartz, S.H., Zeevaart, J.A., McCarty, D.R., 1997. Genetic control of abscisic
acid biosynthesis in maize. Proc. Natl. Acad. Sci. 94 (22), 12235–12240. https://doi.
org/10.1073/pnas.94.22.12235.
Tang, Y., Bao, X., Zhi, Y., Wu, Q., Yin, X., Zeng, L., Li, J., Zhang, J., He, W., Liu, W.,
Wang, Q., 2019. Overexpression of a MYB family gene, OsMYB6, increases drought
and salinity stress tolerance in transgenic rice. Front. Plant Sci. 10, 168. https://doi.
org/10.3389/fpls.2019.00168.
Thakur, P., Kumar, S., Malik, J.A., Berger, J.D., Nayyar, H., 2010. Cold stress effects on
reproductive development in grain crops: an overview. Environ. Exp. Bot. 67 (3),
429–443. https://doi.org/10.1016/j.envexpbot.2009.09.004.
Thompson, A.J., Mulholland, B.J., Jackson, A.C., McKee, J.M., Hilton, H.W., Symonds, R.
C., Sonneveld, T., Burbidge, A., Stevenson, P., Taylor, I.B., 2007. Regulation and
manipulation of ABA biosynthesis in roots. Plant, Cell Environ. 30 (1), 67–78.
https://doi.org/10.1111/j.1365-3040.2006.01606.x.
Torres, C.A., Sepulveda, A., Gonzalez-Talice, J., Yuri, J.A., Razmilic, I., 2013. Fruit water
relations and osmoregulation on apples (Malus domestica Borkh.) with different sun
exposures and sun-injury levels on the tree. Sci. Hortic. 161, 143–152. https://doi.
org/10.1111/j.1365-3040.2006.01606.x.
Torrey, J.G., 1976. Root hormones and plant growth. Ann. Rev. Plant Physiol. 27 (1),
435–459.
V. Rachappanavar et al.
Tounsi, S., Feki, K., Hmidi, D., Masmoudi, K., Brini, F., 2017. Salt stress reveals
differential physiological, biochemical and molecular responses in T. monococcum
and T. durum wheat genotypes. Physiol. Mol. Biol. Plants 23 (3), 517–528. https://
doi.org/10.1007/s12298-017-0457-4.
Tran, L.S., Urao, T., Qin, F., Maruyama, K., Kakimoto, T., Yamaguchi-Shinozaki, K.,
2007. Functional analysis of AHK1/ATHK1 and cytokinin receptor histidine kinases
in response to abscisic acid, drought, and salt stress in Arabidopsis. Proc. Natl. Acad.
Sci. U. S. A. 104 (51), 20623–20628. https://doi.org/10.1073/pnas.0706547105.
Tuberosa, R., Salvi, S., 2006. Genomics-based approaches to improve drought tolerance
of crops. Trends Plant Sci. 11 (8), 405–412. https://doi.org/10.1016/j.
tplants.2006.06.003.
Upreti, G.C., Jensen, K., Munday, R., Duganzich, D.M., Vishwanath, R., Smith, J.F., 1998.
Studies on aromatic amino acid oxidase activity in ram spermatozoa: role of
pyruvate as an antioxidant. Anim. Reprod. Sci. 51 (4), 275–287. https://doi.org/
10.1016/S0378-4320(98)00082-7.
Upreti, K.K., Murti, G.S.R., 2010. Response of grape rootstocks to salinity: changes in
root growth, polyamines and abscisic acid. Biol. Plant. 54 (4), 730–734. https://doi.
org/10.1007/s10535-010-0130-z.
Upreti, K.K., Murti, G.S.R., Bhatt, R.A., 2000. Water stress-induced changes in the levels
of ethylene, ACC and ACC-oxidase activity in french bean. Indian J. Plant Physiol. 5
(4), 369–373.
Vaculík, M., Lukačová, Z., Bokor, B., Martinka, M., Tripathi, D.K., Lux, A., 2020.
Alleviation mechanisms of metal (loid) stress in plants by silicon: a review. J. Exp.
Bot. 71 (21), 6744–6757. https://doi.org/10.1093/jxb/eraa288.
Vardhini, B.V., Anuradha, S., Rao, S.S.R., 2006. Brassinosteroids-New class of plant
hormone with potential to improve crop productivity. Indian J. Plant Physiol. 11 (1),
1.
Verma, S.P., 2020. Trace Element Geochemistry. Road from Geochemistry to
Geochemometrics. Springer, Singapore, pp. 201–225. https://doi.org/10.1007/978-
981-13-9278-8_3.
Vernay, P., Gauthier-Moussard, C., Hitmi, A., 2007. Interaction of bioaccumulation of
heavy metal chromium with water relation, mineral nutrition and photosynthesis in
developed leaves of Lolium perenne L. Chemosphere 68 (8), 1563–1575. https://doi.
org/10.1016/j.chemosphere.2007.02.052.
Vinocur, B., Altman, A., 2005. Recent advances in engineering plant tolerance to abiotic
stress: achievements and limitations. Curr. Opin. Biotechnol. 16 (2), 123–132.
https://doi.org/10.1016/j.copbio.2005.02.001.
Wang, D., Pajerowska-Mukhtar, K., Culler, A.H., Dong, X., 2007a. Salicylic acid inhibits
pathogen growth in plants through repression of the auxin signaling pathway. Curr.
Biol. 17 (20), 1784–1790. https://doi.org/10.1016/j.cub.2007.09.025.
Wang, H., Liang, X., Wan, Q., Wang, X., Bi, Y., 2009. Ethylene and nitric oxide are
involved in maintaining ion homeostasis in Arabidopsis callus under salt stress.
Planta 230 (2), 293–307. https://doi.org/10.1007/s00425-009-0946-y.
Wang, J., Xia, H., Lin, L.J., He, H., Liang, D., Lv, X.L., 2016. Exogenous abscisic acid
increases resistances against abiotic stress and improve fruit quality of grape. JAPS
26 (5).
Wang, L., Chen, S., Kong, W., Li, S., Archbold, D.D., 2006. Salicylic acid pretreatment
alleviates chilling injury and affects the antioxidant system and heat shock proteins
of peaches during cold storage. Postharvest Biol. Technol. 41 (3), 244–251. https://
doi.org/10.1016/j.postharvbio.2006.04.010.
Wang, L.J., Li, S.H., 2006. Salicylic acid-induced heat or cold tolerance in relation to Ca2
+ homeostasis and antioxidant systems in young grape plants. Plant Sci. 170 (4),
685–694. https://doi.org/10.1016/j.plantsci.2005.09.005.
Wang, L.J., Fan, L., Loescher, W., Duan, W., Liu, G.J., Cheng, J.S., Luo, H.B., Li, S.H.,
2010. Salicylic acid alleviates decreases in photosynthesis under heat stress and
accelerates recovery in grapevine leaves. BMC Plant Biol. 10 (1), 1–10. https://doi.
org/10.1186/1471-2229-10-34.
Wang, D., Pei, K., Fu, Y., Sun, Z., Li, S., Liu, H., Tang, K., Han, B., Tao, Y., 2007b.
Genome-wide analysis of the auxin response factors (ARF) gene family in rice (Oryza
sativa). Gene 394 (1–2), 13–24. https://doi.org/10.1016/j.gene.2007.01.006.
Wang, S.Y., Steffens, G.L., 1985. Effect of paclobutrazol on water stress-induced ethylene
biosynthesis and polyamine accumulation in apple seedling leaves. Phytochemistry
24 (10), 2185–2190. https://doi.org/10.1016/S0031-9422(00)83007-1.
Wang, X., Chen, X., Wang, Q., Chen, M., Liu, X., Gao, D., Li, D., Li, L., 2019. MdBZR1 and
MdBZR1-2like transcription factors improves salt tolerance by regulating gibberellin
biosynthesis in apple. Front. Plant Sci. 10, 1473. https://doi.org/10.3389/
fpls.2019.01473.
Wang, Y., Jiang, H., Mao, Z., Liu, W., Jiang, S., Xu, H., Su, M., Zhang, J., Wang, N.,
Zhang, Z., Chen, X., 2021. Ethylene increases the cold tolerance of apple via the
MdERF1B–MdCIbHLH1 regulatory module. Plant J. 106 (2), 379–393. https://doi.
org/10.1111/tpj.15170.
Wani, R.A., Malik, T.H., Malik, A.R., Baba, J.A., Dar, N.A., 2014. Studies on apple seed
germination and survival of seedlings as affected by gibberellic acid under cold arid
conditions. Int. J. Sci. Technol. Res. 3 (3), 2010–2016.
Wani, S.H., Kumar, V., Shriram, V., Sah, S.K., 2016. Phytohormones and their metabolic
engineering for abiotic stress tolerance in crop plants. Crop J. 4 (3), 162–176.
https://doi.org/10.1016/j.cj.2016.01.010.
Wartinger, A., Heilmeier, H., Hartung, W., Schulze, E.D., 1990. Daily and seasonal
courses of leaf conductance and abscisic acid in the xylem sap of almond trees
[Prunus dulcis (Miller) DA Webb] under desert conditions. New Phytol. 116 (4),
581–587. https://doi.org/10.1111/j.1469-8137.1990.tb00542.x.
Watkinson, J.I., Sioson, A.A., Vasquez-Robinet, C., Shukla, M., Kumar, D., Ellis, M.,
Heath, L.S., Ramakrishnan, N., Chevone, B., Watson, L.T., van Zyl, L., 2003.
Photosynthetic acclimation is reflected in specific patterns of gene expression in
drought-stressed loblolly pine. Plant Physiol. 133 (4), 1702–1716. https://doi.org/
10.1104/pp.103.026914.
20
Scientia Horticulturae 304 (2022) 111302
WATTS, S., Rodriguez, J.L., EVANS, S.E., Davies, W.J., 1981. Root and shoot growth of
plants treated with abscisic acid. Ann. Bot. 47 (5), 595–602. https://doi.org/
10.1093/oxfordjournals.aob.a086056.
Weiner, J.J., Peterson, F.C., Volkman, B.F., Cutler, S.R., 2010. Structural and functional
insights into core ABA signaling. Curr. Opin. Plant Biol. 13 (5), 495–502. https://doi.
org/10.1016/j.pbi.2010.09.007.
Wen, X.P., Ban, Y., Inoue, H., Matsuda, N., Kita, M., Moriguchi, T., 2011. Antisense
inhibition of a spermidine synthase gene highlights the role of polyamines for stress
alleviation in pear shoots subjected to salinity and cadmium. Environ. Exp. Bot. 72
(2), 157–166. https://doi.org/10.1016/j.envexpbot.2011.03.001.
Wien, H.C., 1997. The Physiology of Vegetable Crops. Cab International.
Wohlbach, D.J., Quirino, B.F., Sussman, M.R., 2008. Analysis of the Arabidopsis histidine
kinase ATHK1 reveals a connection between vegetative osmotic stress sensing and
seed maturation. Plant Cell 20 (4), 1101–1117. https://doi.org/10.1105/
tpc.107.055871.
Wolters, H., Jürgens, G., 2009. Survival of the flexible: hormonal growth control and
adaptation in plant development. Nat. Rev. Genet. 10 (5), 305–317. https://doi.org/
10.1105/tpc.107.055871.
Wright, A.D., Sampson, M.B., Neuffer, M.G., Michalczuk, L., Slovin, J.P., Cohen, J.D.,
1991. Indole-3-acetic acid biosynthesis in the mutant maize orange pericarp, a
tryptophan auxotroph. Science 254 (5034), 998–1000. https://doi.org/10.1126/
science.254.5034.998.
Wu, K., Zhang, L., Zhou, C., Yu, C.W., Chaikam, V., 2008. HDA6 is required for jasmonate
response, senescence and flowering in Arabidopsis. J. Exp. Bot. 59 (2), 225–234.
https://doi.org/10.1093/jxb/erm300.
Xi, Z, Wang, Z, Fang, Y, Hu, Z, Hu, Y, Deng, M, Zhang, Z, 2013. Effects of 24-epibras­
sinolide on antioxidation defense and osmoregulation systems of young grapevines
(V. vinifera L.) under chilling stress. Plant Growth Regul. 71, 57–65. https://doi.org/
10.1007/s10725-013-9809-4.
Xi, Zhu-mei, Zhang, Zhen-wen, Huo, Shan-shan, Luan, Li-ying, Gao, Xiang, Ma, Li-na,
Fang, Yu-lin, 2013. Regulating the secondary metabolism in grape berry using
exogenous 24-epibrassinolide for enhanced phenolics content and antioxidant
capacity. Food Chem. 141 (3), 3056–3065. https://doi.org/10.1016/j.
foodchem.2013.05.137.
Xia, X.J., Wang, Y.J., Zhou, Y.H., Tao, Y., Mao, W.H., Shi, K., Asami, T., Chen, Z., Yu, J.
Q., 2009. Reactive oxygen species are involved in brassinosteroid-induced stress
tolerance in cucumber. Plant Physiol. 150 (2), 801–814. https://doi.org/10.1104/
pp.109.138230.
Xie, J.H., Zapata-Arias, F.J., Shen, M., Afza, R., 2000. Salinity tolerant performance and
genetic diversity of four rice varieties. Euphytica 116 (2), 105–110. https://doi.org/
10.1023/A:1004041900101.
Xin, Z., Browse, J., 2000. Cold comfort farm: the acclimation of plants to freezing
temperatures. Plant Cell Environ. 23 (9), 893–902. https://doi.org/10.1046/j.1365-
3040.2000.00611.x.
Xiong, L., Zhu, J.K., 2003. Regulation of abscisic acid biosynthesis. Plant Physiol. 133
(1), 29–36. https://doi.org/10.1104/pp.103.025395.
Xiong, L., Gong, Z., Rock, C.D., Subramanian, S., Guo, Y., Xu, W., Galbraith, D., Zhu, J.K.,
2001. Modulation of abscisic acid signal transduction and biosynthesis by an Sm-like
protein in Arabidopsis. Dev. Cell 1 (6), 771–781. https://doi.org/10.1016/S1534-
5807(01)00087-9.
Xu, H., Song, J., Luo, H., Zhang, Y., Li, Q., Zhu, Y., Xu, J., Li, Y., Song, C., Wang, B.,
Sun, W., 2016. Analysis of the genome sequence of the medicinal plant Salvia
miltiorrhiza. Mol. Plant 9 (6), 949–952. https://doi.org/10.1016/2Fj.
molp.2016.03.010.
Yadav, B., Jogawat, A., Gnanasekaran, P., Kumari, P., Lakra, N., Lal, S.K., Pawar, J.,
Narayan, O.P., 2020. An overview of recent advancement in phytohormones-
mediated stress management and drought tolerance in crop plants. Plant Gene,
100264. https://doi.org/10.1016/2Fj.molp.2016.03.010.
Yadav, S.S., Redden, R.J., Hatfield, J.L., Lotze-Campen, H., Hall, A.J., 2011. Crop
Adaptation to Climate Change. John Wiley & Sons.
Yang, J., Duan, G., Li, C., Liu, L., Han, G., Zhang, Y., Wang, C., 2019. The crosstalks
between jasmonic acid and other plant hormone signaling highlight the involvement
of jasmonic acid as a core component in plant response to biotic and abiotic stresses.
Front. Plant Sci. 10 https://doi.org/10.3389/fpls.2019.01349.
Yang, W., Liu, X.D., Chi, X.J., Wu, C.A., Li, Y.Z., Song, L.L., Liu, X.M., Wang, Y.F.,
Wang, F.W., Zhang, C., Liu, Y., 2011. Dwarf apple MbDREB1 enhances plant
tolerance to low temperature, drought, and salt stress via both ABA-dependent and
ABA-independent pathways. Planta 233 (2), 219–229. https://doi.org/10.1007/
s00425-010-1279-6.
Yang, Y., Yang, F., Li, X., Shi, R., Lu, J., 2013. Signal regulation of proline metabolism in
callus of the halophyte Nitraria tangutorum Bobr. grown under salinity stress. Plant
Cell Tissue Organ. Cult. 112 (1), 33–42. https://doi.org/10.1007/s11240-012-0209-
7.
Yang, Y.Y., Zheng, P.F., Ren, Y.R., Yao, Y.X., You, C.X., Wang, X.F., Hao, Y.J., 2021.
Apple MdSAT1 encodes a bHLHm1 transcription factor involved in salinity and
drought responses. Planta 253 (2), 1–13. https://doi.org/10.1007/s00425-020-
03528-6.
Yanping, L., Zhenhai, H., Xuefeng, X., 2004. Accumulation of jasmonic acid in apple
seedlings under water stress. Acta Hortic. Sin. 31 (1), 16.
Yao, Y., Graziano, D.J., Riddle, M., Cresko, J., Masanet, E., 2016. Prospective energy
analysis of emerging technology options for the United States ethylene industry. Ind.
Eng. Chem. Res. 55 (12), 3493–3505. https://doi.org/10.1021/acs.iecr.5b03413.
Ying, Y., Yue, Y., Huang, X., Wang, H., Mei, L., Yu, W., Zheng, B., Wu, J., 2013. Salicylic
acid induces physiological and biochemical changes in three Red bayberry (Myric
rubra) genotypes under water stress. Plant Growth Regul. 71 (2), 181–189. https://
doi.org/10.1007/s10.
V. Rachappanavar et al.
Yiu, J.C., Liu, C.W., Fang, D.Y.T., Lai, Y.S., 2009. Waterlogging tolerance of Welsh onion
(Allium fistulosum L.) enhanced by exogenous spermidine and spermine. Plant
Physiol. Biochem. 47 (8), 710–716. https://doi.org/10.1016/j.plaphy.2009.03.007.
Yoshida, T., Fujita, Y., Sayama, H., Kidokoro, S., Maruyama, K., Mizoi, J., Shinozaki, K.,
Yamaguchi-Shinozaki, K., 2010. AREB1, AREB2, and ABF3 are master transcription
factors that cooperatively regulate ABRE-dependent ABA signaling involved in
drought stress tolerance and require ABA for full activation. Plant J. 61 (4),
672–685. https://doi.org/10.1111/j.1365-313X.2009.04092.x.
Zaharah, S.S., Razi, I.M., 2009. Growth, stomata aperture, biochemical changes and
branch anatomy in mango (Mangifera indica) cv. Chokanan in response to root
restriction and water stress. Sci. Hortic. 123 (1), 58–67. https://doi.org/10.1016/j.
scienta.2009.07.022.
Chapter 8 Zaid, A., de Wet, P.F., 2002. Pollination and bunch management. In: Zaid, A
(Ed.), Date Palm Cultivation. FAO, plant production and protection, p. 156.
Zarei, A., Trobacher, C.P., Shelp, B.J., 2016. Arabidopsis aldehyde dehydrogenase 10
family members confer salt tolerance through putrescine-derived 4-aminobutyrate
(GABA) production. Sci. Rep. 6, 35115. https://doi.org/10.1038/srep35115.
Zhang, Y., Chen, K., Zhang, S., Ferguson, I., 2003. The role of salicylic acid in postharvest
ripening of kiwifruit. Postharvest Biol. Technol. 28, 67–74. https://doi.org/
10.1016/S0925-5214(02)00172-2.
Zhang, W., Wang, Z., Song, J., Yue, S., Yang, H., 2019. Cd2+ uptake inhibited by
MhNCED3 from Malus hupehensis alleviates Cd-induced cell death. Environ. Exp.
Bot. 166, 103802 https://doi.org/10.1016/j.envexpbot.2019.103802.
Zhang, Y., Xiong, Y., Yarbrough, W.G., 1998. ARF promotes MDM2 degradation and
stabilizes p53: ARF-INK4a locus deletion impairs both the Rb and p53 tumor
suppression pathways. Cell 92 (6), 725–734. https://doi.org/10.1016/S0092-8674
(00)81401-4.
Zhang, Z., Huang, R., 2010. Enhanced tolerance to freezing in tobacco and tomato
overexpressing transcription factor TERF2/LeERF2 is modulated by ethylene
biosynthesis. Plant Mol. Biol. 73 (3), 241–249. https://doi.org/10.1007/s11103-
010-9609-4.
Zhao, H., Wu, Q.Q., Cao, L.F., Qing, H.Y., Zhang, C., Chen, Y.H., Wang, H., Liu, R.R.,
Xu, D.X., 2014. Melatonin inhibits endoplasmic reticulum stress and epithelial-
21
Scientia Horticulturae 304 (2022) 111302
mesenchymal transition during bleomycin-induced pulmonary fibrosis in mice. PLoS
One 9 (5). https://doi.org/10.1371/journal.pone.0097266.
Zhao, J., Gao, Y., Zhang, Z., Chen, T., Guo, W., Zhang, T., 2013a. A receptor-like kinase
gene (GbRLK) from Gossypium barbadense enhances salinity and drought-stress
tolerance in Arabidopsis. BMC Plant Biol. 13 (1), 110. https://doi.org/10.1186/
1471-2229-13-110.
Zhao, R., Satpradit, O., Rijnaarts, H.H.M., Biesheuvel, P.M., Van der Wal, A., 2013b.
Optimization of salt adsorption rate in membrane capacitive deionization. Water
Res. 47 (5), 1941–1952. https://doi.org/10.1016/j.watres.2013.01.025.
Zhao, X.Q., Bai, F.W., 2009. Mechanisms of yeast stress tolerance and its manipulation
for efficient fuel ethanol production. J. Biotechnol. 144 (1), 23–30. https://doi.org/
10.1016/j.jbiotec.2009.05.001.
Zhao, Y., 2010. Auxin biosynthesis and its role in plant development. Annu. Rev. Plant
Biol. 61, 49–64. https://doi.org/10.1146/annurev-arplant-042809-112308.
Zhao, W., Wang, Y., Liu, S., Pan, M., Yang, J., Chen, S., 2013c. Denitrification activities
and N2O production under salt stress with varying COD/N ratios and terminal
electron acceptors. Chem. Eng. J. 215, 252–260. https://doi.org/10.1016/j.
cej.2012.10.084.
Zhao, Y., Wang, R., Tu, K., Liu, K., 2011. Efficacy of preharvest spraying with Pichia
guilliermondii on postharvest decay and quality of cherry tomato fruit during
storage. Afr. J. Biotechnol. 10 (47), 9623–9630. https://doi.org/10.5897/
AJB11.870.
Zhou, W., Li, C., Sun, C., Yang, X., 2016. Simultaneously determination of trace Cd2+ and
Pb2+ based on l-cysteine/graphene modified glassy carbon electrode. Food Chem.
192, 351–357. https://doi.org/10.1016/j.foodchem.2015.07.042.
Zhou, Y., Yu, B., Levon, K., 2005. Potentiometric sensor for dipicolinic acid. Biosens.
Bioelectron. 20 (9), 1851–1855. https://doi.org/10.1016/j.bios.2004.05.005.
Zhu, Z., An, F., Feng, Y., Li, P., Xue, L., Mu, A., Jiang, Z., Kim, J.M., To, T.K., Li, W.,
Zhang, X., 2011. Derepression of ethylene-stabilized transcription factors (EIN3/
EIL1) mediates jasmonate and ethylene signaling synergy in Arabidopsis. Proc. Natl.
Acad. Sci. 108 (30), 12539–12544. https://doi.org/10.1073/pnas.1103959108.
Zolla, G., Heimer, Y.M., Barak, S., 2010. Mild salinity stimulates a stress-induced
morphogenic response in Arabidopsis thaliana roots. J. Exp. Bot. 61 (1), 211–224.
https://doi.org/10.1093/jxb/erp290.