See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/358873344

Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops

Book · February 2022

DOI: 10.5772/intechopen.102033

CITATIONS READS

6 510

9 authors, including:

M.N. Arun
Directorate of Rice Research
18 PUBLICATIONS   14 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Nutrient Use Efficiency in RICE View project

Irrigation and Water Use Efficiency in Rice View project

All content following this page was uploaded by M.N. Arun on 11 April 2022.

The user has requested enhancement of the downloaded file.

 We are IntechOpen,
the world’s leading publisher of
Open Access books
Built by scientists, for scientists

5,700
Open access books available
139,000
International authors and editors
175M
Downloads

Our authors are among the

154
Countries delivered to
TOP 1%
most cited scientists
12.2%
Contributors from top 500 universities

Selection of our books indexed in the Book Citation Index

in Web of Science™ Core Collection (BKCI)

Interested in publishing with us?

Contact [email protected]
Numbers displayed above are based on latest data collected.
For more information visit www.intechopen.com
Chapter

Seed Priming: The Way Forward to

Mitigate Abiotic Stress in Crops
Melekote Nagabhushan Arun, Shibara Shankara Hebbar,
Bhanuprakash, Thulasiram Senthivel, Anil Kumar Nair,
Guntupalli Padmavathi, Pratima Pandey and Aarti Singh

Abstract

Abiotic stress is a major threat to the farming community, biasing the crop
productivity in arid and semi-arid regions of the world. The seed is an important
component of agriculture, contributing significantly to the booming production
of food and feed crops across the different agro-ecological regions of the world
with constant challenges with reference to production, storage, and quality control.
Germination, plant growth, and development via non-normal physiological pro-
cesses are detrimentally affected by stress. Seed priming is an alternative, low cost,
and feasible technique, which can improve various abiotic stress tolerances through
enhanced and advanced seed production. Seed priming is a process that involves
imbibing seed with a restricted amount of water to allow sufficient hydration and
advancement of metabolic processes but preventing germination. The beneficial
influence of priming on the germination performance of diverse species is attributed
to the induction of biochemical mechanisms of cell repair: the resumption of meta-
bolic activity that can re-impose cellular integrity, through the synthesis of nucleic
acids (DNA and RNA) and proteins and the improvement of the antioxidant defense
system metabolic damage incurred by dry seed and thus fortifying the metabolic
machinery of the seed. With this background, this chapter highlights the morpho-
logical, physiological, biochemical, and molecular responses of seed priming and
recent advances in priming methods as a tool to combat abiotic stress in crop plants.

Keywords: abiotic stress, seed priming, crop establishment, physiological changes,

biochemical changes

1. Introduction

Crop plants are subjected to multiple abiotic stresses during their life span
that significantly reduces productivity and foreshadow global food security. The
improvement of crop plants through direct selection-based conventional breeding
for drought and salinity stress appears to be quite complex [1]. Abiotic stresses such
as salinity, drought, flooding, heat, cold, freezing, excess light, UV radiation, and
heavy metal toxicity have a significant impact on seed which reduce germination
rate, seedling growth and yield with significant variations from crop to crop world-
wide [2]. Water stress is a key agronomic problem globally and is one of the most
important factors reducing crop productivity. Drought is one of the most important

1
Plant Stress Physiology - Perspectives in Agriculture

environmental component limiting plant growth and productivity. The advance-

ment of crop plants through direct selection-based conventional breeding for
drought and salinity stress appears to be quite complex. In agriculture, alternative
innovative technologies such as plant tissue culture, seed priming and genetic engi-
neering could play a major role in combating abiotic stresses for increasing yield.
The quality of seed in the present time can be accomplished by various means where
the basic and applied knowledge of plant physiology, genetics and seed technology
all are integrated to improve the criteria of quality of seeds. Seed germination is a
dynamic and complex stage of plant ontogeny, with several interactive metabolic
processes changing from a storage phase to a mobilization phase [3]. The multitude
of simultaneous metabolic processes taking place in germinating seed makes it
difficult to explorte events related to the initiation of the germination process. The
catabolic processes in the storage tissues can confound measurements and inter-
pretation of anabolic and growth processes occurring in the developing embryo in
the entire seed. The time from sowing to the seedling establishment is of consider-
able importance in crop production and has major impacts on plant growth, final
yield and post-harvest seed quality [4]. Efficient seed germination is important for
agriculture. Successful foundation of early seedling indeed requires an accelerated
and uniform emergence and root growth. In nature, plants often face the chal-
lenge of severe environmental conditions, which include various biotic and abiotic
stresses that exert unfavorable effects on plant growth and development causing
considerable losses in crop productivity. Drought, salinity and extreme temperature
cause osmotic stress on crop plants which generate an imbalance at cellular, molecu-
lar and physiological levels which ultimately lead to plant death [5]. Abiotic stress
causes many physiological and biochemical changes in the seedlings, which include
the generation of reactive oxygen species (ROS), leading to membrane injury and
cell leakage and destruction of photosynthetic components [6]. Seed priming is a
simple, safe, cost effective, and effective approach for enhancement of seed germi-
nation, early seedling growth, and yield under stressed and non-stressed conditions
[7]. Seed priming is a form of seed preparation in which seeds are pre-soaked before
planting with a certain solution that allows partial hydration but will not germinate
and redried to original moisture content [8].

2. Abiotic stress and its effect on crop growth and development

Abiotic stresses are closely related individually or cohesive; they cause physical,
morphological, biochemical and cellular changes that adversely affect plant growth
and productivity and ultimately yield. Heat, drought, salinity and cold are major
abiotic pressures that cause severe cell damage to a wide variety of plants, including
crop plants (Figure 1). Water stress or drought is a major challenge for agricultural
production worldwide. Excessive wilting causes a change in the ratio of membrane
lipids and this may result in increased electrolyte leakage [9]. Drought is known as
a prime abiotic factor that limits plant growth and production in arid and semi-arid
regions and is the most significant factor in world security and sustainability in
agricultural production. Drought slows growth, induces stomatal closure and there-
fore reduces photosynthesis, growth and yield in a number of plant species [10].
Water stress has been revealed to be one of the factors limiting the productivity of
cowpea as it affects not only the production of the grain, but also the whole process
of growth of all organs of the plant and its metabolism [11]. Water stress limits
the size of individual leaves and leaf number. Physiological disorders occur during
seed germination under abiotic stress is a decrease in water uptake by the seed due
to the low water potential of the germination medium. Slow or abnormal growth

2
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

Figure 1.
Direct and indirect causes of low productivity of field and horticultural crops.

and emergence result in fewer and smaller plants, which are more susceptible to
various abiotic stresses [12]. In addition to causing various structural changes at
different levels of organization in the seed, slow rate of imbibitions may lead to a
series of metabolic changes, including up-regulation or down-regulation of enzyme
activities, perturbance in the mobility of inorganic nutrients to developing tissues,
disturbance in N metabolism, imbalances in the levels of plant growth regulators,
reduction in hydrolysis and utilization of food reserves and accumulation of com-
patible osmotic such as soluble sugars, free proline and soluble proteins [13]. Stress
processes may lead to poor or complete lack of germination under abiotic condi-
tions. Salinity is also considered as substantial abiotic stress and significant factor
affecting crop production globally and especially in arid and semi-arid regions [14].
The salinity of soil reduces water availability of plant root via negative (low)
osmotic potential as well as decrease of germination dynamics of plant seeds by
ionic toxicity of Na+ and Cl− [15]. Of the extensive forms of abiotic stress, heat stress
has an independent mode of action on the physiology and metabolism of plant cells.
Due to high temperatures, various physiological injuries have been observed such
as scorching of leaves and stems, leaf abscission and senescence and root and shoot
growth inhibition or fruit damage, which as a result lead to decreased plant produc-
tivity [16]. The plant growth is reduced by affecting the shoot, net assimilation rates
and finally the total dry weight of the plant due to high temperature [17]. The stress
is extensively prominent on reproductive development than on vegetative growth,
and the sudden reduction in yield with temperature is mainly associated with
pollen infertility in many crop species under effects of high-temperature [18]. Heat
stress which causes multifarious, and often adverse, alterations in plant growth,
development, physiological processes, and yield is one of the major consequences

3
Plant Stress Physiology - Perspectives in Agriculture

of high-temperature stress [19]. Due to stress limitation of available technology, we

should think of various alternate technologies such as priming, mutagenesis, and
tissue culture for combating abiotic stresses. In crop species, seed germination and
early development of seedling are the most sensitive stages to salinity stress. Salinity
delays the onset, reduce the rate and increase the dispersion of germination phenol-
ogy, leading to a reduction in plant growth and final crop yield. Thus, if the effect
of abiotic stress can be mitigated at the early stages, the chance of establishing a
successful crop under stress will be improved [20].

3. Seed priming and its importance

Priming involves prior exposure to elicitors which brings a cellular state that
arrest the harmful effects of abiotic stress, and plants raised after priming are more
tolerant of abiotic stress. Seed dormancy is an emerging issue related to germination
that is common to many plant species. It is a practice that allows a species to estimate
the germination period in a population. Some species use a environmental indicator
(such as drought, rain or temperatures) to integrate germination of many seeds at a
particular time of year. Temperature, humidity, air and light conditions are the main
factors for seed germination. The minimum temperature is the minimal temperature
at which a seed germinates effectively and the maximum is the highest temperature
at which seeds can germinate [21]. At optimal temperatures, germination is rapid and
uniform but with meager change in this temperature can damage seeds or make them
go into the dormancy condition. Seeds need accurate moisture to initiate internal
processes leading up to germination. Osmotic adjustment or priming of seeds before
sowing is known as a potent way to increase germination and emergence rate in some
species with incremental seed development [22]. Inadequate seed germination and
subsequently poor field establishment are a common occurrence at adverse condi-
tions of the environment. Seed germination and early seedling growth are the most
sensitive stages of water limitation and the water deficit may impede the onset and
reduce the rate and uniformity of germination, leading to poor crop accomplishment
and yield in several crops. Seed priming is a water-based approach with low external
water potential that restrict hydration (controlled hydration of seed) and permit
metabolic processes necessary for enhancing germination rate and seed quality by
managing the seed moisture content and temperature in which the seed is taken
through the initial biochemical processes within the initial stages of germination but
preventing the seed transition towards full germination [23]. This will assure better
field emergence and disease resistance under various adverse conditions. The purpose
of priming is to reduce the germination time and improve stand and germination
percentage under unfavorable environmental conditions. Primed seeds are used
instantly but may be dried and stored for a short time for later use. Primed seeds
attain the potential to rapidly imbibe and revive seed metabolism thus enhancing the
germination rate [24]. These attributes have practical agronomic implications notably
under adverse germination conditions. Pre-treatment using a priming induced
stimulus like sodium nitroprusside, hydrogen peroxide, melatonin and polyamines
results in enhanced cell tolerance and amelioration of stress-induced plant growth
inhibition [25]. Therefore, the beneficial effects of priming may be more evident
in under favorable rather than unfavorable conditions [26]. Primed seeds generally
exhibit an increase germination rate, greater germination uniformity, and, at times,
greater total germination percentage. Abiotic stresses produce oxidative stress and
activate similar cell signaling pathways and cellular responses [27]. But, seed priming
seems to activate these signaling pathways during the early stages of growth, result-
ing in faster defense responses [28]. The abiotic stress tolerance generated by seed

4
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

priming is accorded via the synchronization of divergent physiological, biochemical,

systemic, cellular and molecular modulations [29]. The purpose of seed germination
is to reduce the germination period and to protect the seed from environmental stress
during the critical stage of seedling growth to integrate the growth leading to uniform
establishment and improved yields. It minimizes the effect of salinity on the morpho-
logical parameter of the plants. One of the priming methods called osmopriming is a
commercially available method of improving seed germination and strength. It con-
trols seed imbibition to initiate the germination process followed by the seed drying
up to its original weight. Various methods of seed planting such as hormonal priming
and support for chemo priming in improving the process of germination, flowering
and growth of plants are done for commercial purposes on the farm or on the farm.
Planting seed on a farm requires electricity, high-tech seed harvesting, seed harden-
ing or seed-drying process is accessible to farmers to help them with cool farming and
horticulture [30]. The effective use of a seed management method depends on the
type of test, method of application, crop selection, early performance of the plant,
chemical selection, concentration, duration of treatment and the purpose of the
application. Priming method in seed management techniques is proven very essential
factor for enhancing quality issues, germination rate and establishment. Priming can
interfere with some of the degenerative effects of aging, leading to improved seed
performance [31]. It has shown an immense effect of priming to activate different
processes related to cell cycle and to induce to the synthesis of nuclear DNA in radial
tip cells [32]. Seed priming had the most beneficial effect on leaf area indicating the
advantage of rapid seedling emergence [33]. The reason attributed may be due to cell
division, cell number due to multiplication in various plant tissues, auxin multiplica-
tion, cell wall plasticity and permeability of cell membrane, increases photosynthesis,
cell enlargement and rapid cell wall elongation [34].

4. Factors affecting seed priming process

Seed priming is controlled by many factors such as aeration, light, temperature,

concentration of priming solution, time (duration), and seed quality.

4.1 Aeration

Aeration is considered an important step to assist seed respiration, seed viability

and contributes to synchronize the emergence [35] Germinating seeds respire very
actively and need sufficient oxygen. The consequence of aeration varies according
to species: in onion aeration of the PEG solution increased the germination process
compared to non-aerated treatment [36]. No difference was noticed in the germina-
tion of lettuce seeds between aerated and non-aerated K3PO4 priming [37].

4.2 Duration

Duration (maximum length of time) of priming is one of the key factors for seed
priming. Seed priming for 7 h and 14 h is sufficient to augment seed and seedling
vigor, stand establishment, and grain yield contrary to soaking the seeds for 21 h
for optimizing duration of hydro priming in green gram [38]. Seeds primed for 12 h
took significantly fewer days to emerge and reach maturity when compared to the
untreated dry seed, whereas 36 h primed seeds showed poor germination and 48 h
primed seeds inhibited germination. This inhibition may be attributed to the pro-
longed period of priming that led to excess water in the seeds and greater reduction
in the O2 availability to the embryo [39].

5
Plant Stress Physiology - Perspectives in Agriculture

4.3 Temperature

The lower temperatures during priming seem to result in slower imbibition

of seeds, increasing the duration of phase II of the triphasic pattern of imbibi-
tions [40]. This would allow the activation mechanisms to repair the membrane
systems and prevent destruction caused by rapid imbibition. Hydro priming at
15°C increased synchronization of germination and speed of seedling emergence in
Solanum lycocarpum [41]. Seed priming at 15°C showed good vigor, mean germina-
tion time and growth response in french bean compared to seed primed at 30°C
which inhibits germination [42]. Seed priming at low temperature showed the
beneficial effects on germination index, speed of germination, length of radical
and plumule, fresh weight of seedling and seedling vigor index compared to high
temperature in wheat [43]. Priming was effective in reducing the time for 50% ger-
mination, mean germination time and increase germination percent and seedling
vigor index when primed with potassium di-hydrogen phosphate at low tempera-
ture in sunflower [44]. GA3 (20 ppm) primed seeds at low temperature showed
significantly higher germination, root length, shoot length and seedling dry weight
over higher temperature in soybean [45]. Seed priming at 15°C showed better
percent increase in germination, mean germination time than at higher temperature
at 30°C. Seed priming at low temperature associated with a buildup of nucleic acids
and protein synthesis and membrane repair [46].

4.4 Concentration

Seedling growth was not proper and subsequent growth of seedling was
arrested probably due damage of cell organelles due to higher concentration
when seed primed with ethrel. Increasing the concentration of priming solu-
tion from 10−3 M to 10−1 M at constant temperature decreased the germination
to the extent of 21% and 56% when papaya seeds were treated with oxalic acid
and mannitol [47]. Soaking of wheat seeds in GA3 at low concentration not only
enhances the speed of germination but also increase the length of radical as
well as plumule [48]. Osmopriming with PEG at low concentration in spinach
improved the final germination percentage, germination rate and uniformity
[49]. Seed priming at lower concentration of growth regulators favors the
increase enzymatic activity which leads to the favorable environment for the
germination. Growth regulators at higher concentration inhibit the seed germi-
nation which might have been due to solute leakage and lipid per oxidation which
limit the necessary material for germination and seedling growth [50]. Tomato
seeds osmo-primed with PEG 6000 at low concentration improved mean ger-
mination time, seedling emergence percentage and cell membrane stability over
higher concentration [51]. Seed priming with 1% sodium molybdate reduced the
seed germination due to toxic effect on physiological and biochemical processes
within the cell [52]. In case of seed priming with higher concentration of ammo-
nium molybdate and magnesium nitrate solution germination was absent. The
reason attributed due to higher concentration of chemical which cause detrimen-
tal effect on cellular mechanism and mitochondrial; membrane in seed [46].

4.5 Light

Light effect changed widely according to crop species. Illumination during seed
priming of celery seeds reduced dormancy [53]. The best germination in lettuce was
obtained with seeds primed in the dark [37]. Light played a vital role in maximizing
seed germination with guava seeds primed at 12 h of light and 12 h of dark [54].

6
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

5. Seed priming and physiological changes during priming

Seed treatment technology is an important link between seed producers and

crop production. Seed priming is the technology which is a novel concept of treating
seeds using various solvents including water which activates physiological processes
of seeds. When dry seeds are immersed in water, water absorption occurs in three
stages [55]. Stage I is imbibition where there is a swift initial water uptake due to
the seed’s low water potential. Proteins are synthesized using existing mRNA and
DNA and mitochondria are repaired during stage I phase. In stage II, there is a steady
increase in seed water content, but physiological activities associated with germina-
tion are initiated, including synthesis of proteins by translation of new mRNAs and
new mitochondria. There is a swift uptake of water in stage I where the process of
germination is completed culminating in radicle emergence Stages I and II are the
foundations of successful seed priming where the seed is brought to a seed moisture
content that is just short of radicle protrusion [56]. The pattern of water uptake
during priming is identical to that during slow germination and controlled. Seed
hydration triggers germination via three stages: imbibition, lag phase and radicle
protrusion through the testa [57]. Seed requires oxygen, water, and a suitable tem-
perature for germination. The time from sowing to the seedling establishment is of
considerable importance in crop production and has a major impact on plant growth,
yield and post-harvest seed quality [4]. During germination process of orthodox
seeds three distinct phases is manifested where in (1) Phase I: seed hydration process
related to passive imbibition of dry tissues associated with water movement prelimi-
nary occurring in the apoplastic spaces; (2) Phase II: activation phase associated with
the rejuvenate of metabolic activities and repairing processes at the cell level; and (3)
Phase III: initiation of developing processes associated to cell elongation and leading
to radical protrusion. Phases I and III both entail an increase in the water content
while hydration remains stable during Phase II. Before the conclusion of Phase II, it
is considered that germination remains a reversible process: the seeds may be dried
again and remain functioning during storage and able to subsequently re-initiate
germination under propitious condition. Water-based seed priming is elucidated as
a pre-sowing dressing that partially hydrates seeds without allowing emergence [58].
Different treatments may indeed be applied during the reversible stage of germina-
tion (point 3). They broadly differ according to the osmotic potential of the prim-
ing solution, the duration, the external temperature, and the existence of specific
chemical compounds. The efficient treatments trigger metabolic processes activated
during phase II of germination, which are then transitory stopped before a loss of
desiccation occurs [59]. Priming is a technique that allows controlled seed hydration
to trigger pre-germinative metabolism but does not allow the seed for the transition
towards full germination. In the case of primed seed hydration treatment allows
regulated imbibition and induction of the pre-germinative metabolism (“activa-
tion”), but radicle emergence is prevented, represented by the extended second
phase. Final phase (phase III) represents the germination and post-germination
phase which is again similar in the case of non-primed seeds. In the case of primed
hydration treatment seeds allow regulated imbibition and the introduction of pre-
germinative metabolism (“initiation”), but the emergence of radicle is inhibited,
represented by an extended second phase. The final stage (phase III) represents
germination and the post-emergence phase which is similar in the case of unprimed
seeds. During Phase I Imbibition there is a rapid absorption of water due to the
lower seed potential than outside. Initially there is water movement in apoplastic
spaces, proteins are synthesized from existing mRNAs and DNA and mitochondria
are repaired during Phase I. In phase II, there is the activation of metabolic and
repairing activity along with the synthesis of proteins by translation of new mRNAs

7
Plant Stress Physiology - Perspectives in Agriculture

of new mitochondria, where as phase III is associated with regaining capacity of

rapid water uptake and initiation of growing processes linked with cell elonga-
tion that leads to radical protrusion. Priming allows a seed to hydrate up till a seed
moisture content involving the entire phase I and before the end of phase II when
the germination remains a reversible process just short of the radical protrusion [56].
Thus priming activates ‘pre-germinative metabolism’ that included a wide range
of physiological functioning. This activates DNA repair pathways, ROS scavenging
systems (that impart for seed repair response) and also helps in preserving genome
integrity [60]. Priming solutions can be supplemented with hormones or beneficial
micro-organisms. The seed may be dried back for storage, distribution and planting.
Priming can induce the germination by improving the speed and synchronization
of seed germination [61]; it can improve seed vigor which require very short or no
activation time during germination. The advanced germination status of primed
seeds contributes to increased germination under stressful conditions [62]. Besides it
also facilitates the initiation of many germination-related activities such as enhanced
energy metabolism, early reserve mobilization, embryo expansion and endosperm
weakening [31]. Priming also enhances the specific stress-responsive systems which
include induced accumulation of LEA and heat shock proteins [63], activation of
catalase and other antioxidant scavenging enzymes [64] and up-regulation of genes
encoding peroxiredoxin [65].

6. Biochemical and molecular basis of stress tolerance

The type of test, method of application, selection of crop, initial performance

of the crop, selection of chemical, duration of treatment, its concentration, and
the purpose of implication helps in successful application of seed management
technique. Priming method in seed management techniques is established, which
is very important factor for enhancing quality issues, germination rate and estab-
lishment. Priming can improve some of the aging-induced deteriorative events,
resulting in improved seed performance [31]. It has shown an immense effect to
activate different processes related to cell cycle and to induce synthesis of nuclear
DNA in radial tip cells in tomato [32]. Prolonged storage of seeds resulted in a
decrease in protein content which led to an increase in oxidation of amino acids,
due to increased respiratory function and progression in the process of deteriorat-
ing stored seed. Seed degradation results in loss of membrane integrity, changes in
enzymatic functions and reduction of protein and nucleic acid synthesis and lesions
in DNA [66]. Priming with NaCl and 30% PEG for 24 h of rice seed initiated in
increase in the activity of superoxide dismutase (SOD) and peroxidise (POD) which
enhance the intensity of respiration of plant and cause an increase in vigourity in
germination [67]. Priming is also thought to increase the activity of many enzymes
involved in metabolism of carbohydrates (α- and β-amylases), proteins (proteases)
and lipids mobilization (iso citratelyase) that are implicated in the stored reserves
mobilization [68]. These enzymes are vital in the breakdown of macromolecules
for the development and growth of the embryo that ultimately result in early and
higher seedling emergence [69]. There are reports that priming facilitates the repair
of chromosomal damage [70], permits early DNA replication and repair, increases
RNA and de novo protein synthesis and reduces the leakage of metabolites [24].
Thus, total seed protein, peroxidases, polyphenol oxidases, RNA and de novo
protein synthesis were enhanced significantly by seed priming. Among the vari-
ous processes of priming, osmopriming could enhance rapid seed germination by
reducing mechanical hindrance on the germinating embryo. The pre-treatment
of seeds with priming agents facilitates the active absorption of ionic molecules

8
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

with greater ATP availability and repair of deteriorated seed parts for reducing
leakage of metabolites leading to faster embryo development [71]. It also, reflected
in greater cellular membrane integrity, counteraction of lipid per oxidation,
and free radical chain reaction often are found to be directly correlated with the
maintenance of viability and reduce moisture uptake by hydrated-dehydrated seed
[72], repair of biochemical lesions by the cellular enzymatic repair system [73] and
metabolic removal of toxic substances [74], counteraction of free radical and lipid
peroxidation reactions [75], biochemical changes like enzyme activation [76], and
improvement of germination rate particularly in old seeds [77]. Priming provides a
‘head-start’ of seed transition from quiescent to germinating state, thus increasing
the potential to germinate. Seed priming thrust abiotic stress on seeds that represses
radicle protrusion but stimulates stress-responsive elements [78].

7. Reversal of seed deterioration by priming

Seed deterioration is defined as the loss of seed viability and vigor due to
aging effects and adverse environmental factors distinctly higher temperature,
relative air humidity and oxygen/carbon-dioxide ratio [66]. Seed deterioration
is associated with several cellular, metabolic and chemical alterations includ-
ing lipid per oxidation, membrane disruption, and DNA damage, impairment
of RNA and protein synthesis and causes several detrimental effects on seeds
[79]. The cause of seed deterioration is damage to cellular membranes and other
sub cellular components by harmful free radicals generated by peroxidation of
unsaturated and polyunsaturated membrane fatty acids. Seed storage causes a
decrease in the protein content which may be related to oxidation of the amino
acids due to the increase in the respiratory activity and advance in the deteriora-
tion process of the stored seeds [80]. Poor storage conditions may accelerate seed
deterioration of seeds [81]. As seed deterioration increases, seed performance
progressively decreases. Plants that originated from deteriorated seed can reduce
growth rate. The aging of seeds, during long term storage deteriorated their vital
status which was expressed in change in their moisture content, decreasing of
their sowing qualities and development of weaker seedlings with higher water
content [82]. The main mechanism for aging of seed is associated with increased
peroxidation of lipid membranes [65]. Priming can reverse some of the aging
induced deteriorative factors and thus improve seed performance [31]. The ben-
eficial effects of priming are associated with the repair and building up of nucleic
acid, increased synthesis of proteins as well as the repair of both mitochondria
and membranes [24]. Priming for 24 h with GA3 and ammonium molybdate in
aged seeds showed increase enzyme activity restored almost entire protein profile
and esterase and peroxidase isozyme profile as it allowed repair system to combat
sub-cellular damage and activated synthesis of enzymes and protein [83]. Under
invigorated, metabolic repair processes in deteriorated seeds occur before onset
of seed germination process [84]. Seed priming is more useful for enhancing
germination of low-quality seed lots than higher-quality ones which indicates
that repair of aging is one of the primary advantages [85]. Significant changes in
enzyme activities were observed in primed seeds compared to un-primed seeds.
Desiccation and storage of seeds has been suggested to result in progressive loss
of integrity of the membrane components of the seeds, which in turn bring about
to seed deterioration as measured by loss of seed vigor and viability. Maintenance
of the integrity of DNA by repairing the damages incurred naturally is crucial
for generating error free template for transcription and replication with fidel-
ity. The damage to DNA which accumulates during the seed aging is repaired by

9
Plant Stress Physiology - Perspectives in Agriculture

aerated hydration [86]. During imbibition prior to germination the integrity of

cell membranes need to be re-established. Rapid imbibitions by the seed at this
time probably reverse the damage and cell will attain maximum vigor by repair
mechanism. It is thought that hydro priming initiates an oxidative stress, which
generates reactive oxygen species, and catalase is synthesized to minimize cell
damage. In addition to catalase, levels of superoxide dismutase, another essential
enzyme quenching free radicals also increases during priming. Increased levels of
these free radical scavenging enzymes due to the oxidative stress during priming
might also protect the cell against membrane damage due to lipid peroxidation
occurring naturally [68]. Priming with GA3 and ammonium molybdate allowed
repair system to combat sub-cellular damage activated enzyme synthesis due to
accelerated aging. The changes in the activation of the enzymes, upon priming
advocate that mobilization of storage material may be responsible for increased
germination and vigor in primed seeds when compared to unprimed aged
seeds [87].

8. Effect of priming on reserve mobilization and management of

oxidative status

It is proposed that germination-related processes such as respiration, energy

metabolism, and initial reserve mobilization also occur during priming. Higher
respiratory activity is required to cover energy pool for speed up germination.
Increased respiratory activity has been associated with pre-sowing treatments.
Seed priming increased the respiratory activity of seeds and reduced the oxygen-
time constant and increased the standard deviation of germination responses [88].
During seed germination, storage proteins, which provide a source of reduced
nitrogen and inorganic minerals, need to be mobilized to support seedling growth
[89]. Soluble protein content increased in pepper seeds after 12 days of prim-
ing in –1.34 MPa NaCl solution [90]. Pepper seedlings developed from primed
seed had improved soluble protein [90]. Osmo-priming induced accumulation
of stress proteins, such as late embryogenesis abundant (LEA) proteins and heat
shock proteins (HSP) [91]. Management of oxidative status is also an important
part of primed seed physiology. Priming activates the response of the antioxidant
system and modifies the prepared seeds for potential stresses [92]. In the early
stages of seed intake and germination, the production of reactive oxygen species
(ROS) is primarily due to the respiratory activity of mitochondria, β-oxidative
pathway activity, and enzymes such as NADPH oxidase, extra-cellular peroxidase,
and oxalate oxidase [93]. Antioxidants, by breaking down high ROS during early
endocytosis, play an important role in ensuring successful germination, especially
under stressful conditions [94]. Seed priming in tomato seeds revealed enhanced
activity of antioxidant enzymes such as ascorbate peroxidase, catalase, peroxidase,
glutathione reductase and superoxide dismutase [95]. Free radical scavenging
enzymes such as catalase and super dismutase are synthesized during hydro prim-
ing to defend the cell from damage due to lipid peroxidation, which occurs due to
the oxidative stress induced by hydro priming. Priming synchronizes all the cells
of the germinating embryo in the G2 phase of the cell cycle so that upon further
imbibitions, cell division proceeds uniformly in all the cells ensuring uniform
development of all parts of the seedlings. Seed priming of pepper (Capsicum
annuum L.) conducted under temperature stress (low 15° C and high 35°C) for
two consecutive runs revealed enhanced germination even in stressful conditions.
Priming was found to enhance repair of membranes, the activities of hydrolytic
enzymes, and antioxidant system. However, it was noticed that priming decreased

10
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

sucrose content, whereas the fatty acid composition remained unchanged and
increased enzymatic activity of catalase which was enhanced significantly in
pepper seeds [96]. Osmopriming with –1.5 MPa PEG 6000 for 6 days of aged seeds
of sweet pepper resulted in an improved germination with decreased levels of
malondialdehyde (MDA)and total antioxidant activity, total ascorbate, de-hydro
ascorbate, and catalase activity in primed seeds enhanced the defense mechanism
in protecting the cell membrane damage from reactive oxygen species [29]. Nano
priming augmented the performance of seeds by enhancing α-amylase activity,
increasing soluble sugar content to support early seedling growth, up-regulating
the expression of aquaporin gene in germinating seeds, increased stress tolerance
through lower ROS production and creation of nano pores for enhancing water
uptake in crops in field [74]. The main obstacle to the practical application of
primed seeds is storage and viability. This barrier can be overcome by knowing the
genes/markers associated with seed germination and the identified markers can be
used to assess the effect of priming on germination efficiency and seed vigor [97].
Genes/markers involved in rice seed priming were identified by comparing differ-
ential proteins between the dry and imbibed seed using two-dimensional electro-
phoresis [98]. Improved germination was reported in polyethylene glycol-6000
(PEG-6000) primed rapeseeds which, on germination, resulted in differential
expression of 952 genes and 75 proteins [99].

9. Methods of seed priming and role in improving crop productivity

Seed priming technique such as hydro priming, halo priming, chemical prim-
ing, osmopriming, hormone priming, solid matrix priming and nutrient priming
are extensively used in crop plants for many environmental stresses. Seed priming
increases germination and growth especially under environmental constraints.
However, the degree of efficacy of different priming agents varies with plant species
and diverse environmental conditions [100]. Different seed priming methods
employed to mitigate stress and salt tolerance as reported by many researchers are
shown in Tables 1-5.

S. no Priming agent Crop Attributed improved References

1. Water (12 h) Rice Accelerated germination, early emergence time, [101]
increased initial growth after emergence, increased
dry root weight and dry matter productivity
Wheat Higher germination percent, increased water use [102]
efficiency, homogeneity of seedling emergence,
increase growth and yield parameters. Aged seeds
when primed with water improved induced increase
in enzyme activity, improved germination and
seedling characteristics

Pearl millet Increased crop emergence and crop yield in arid zone [103]

Chickpea Improved membrane integrity and electrical [104]

conductivity of seed leachate. Increased germination
indices and seedling growth

Sunflower Increased germination percentage, seed vigor index [105]

and seedling growth rate. Reduced time to 50%
seedling emergence

Table 1.
Hydropriming technique and their effectiveness in improving growth of various crops under adverse condition.

11
Plant Stress Physiology - Perspectives in Agriculture

S. no Priming agent Crop Concentration Attributed improved References

1. PEG wheat −1.0 MPa Seed germination and grain yield [106]

Rape seed −0.5 MPa Percentage of germination under [107]
saline soil, seedling length, and
increased metabolic process in
seeds

Chickpea −0.5 MPa Seed germination, yield and [108]

improvement in seed quality
attributes

2. Mannitol Broad bean 1% Percentage of germination, higher [109]

seedling length and seed vigor
index

Rice 1% Enhanced germination [110]

percentage, growth of seedlings
and plant survival under salt stress

3. Sorbitol Wheat 1% Improved seed vigor, plant [111]

morphology and upregulation of
plant growth regulator

Table 2.
Osmoconditioning techniques and their effectiveness in improving growth of various crops under adverse
condition.

S. no Priming agent Crop Attributed improved References

1. Gibberellic acid Rice Improve crop emergence, crop establishment and [112]
(50 ppm) yield in direct seeded rice
2. Cytokinins Soybean Hasten seed germination rate and seedling [113]
(100 ppm) development. Improving root length and enhanced
nutrient uptake and water use efficiency

3. IAA (20 mg L−1) Cotton Improves the germination, root length, seedling [114]
height and seedling growth, biomass and leaf
photosynthesis capacity and yield

4. Auxin (50 ppm) Wheat Increased grain filling rate and grain yield and [115]
positive effect of photosynthesis
5. Salicyclic acid Sesame Increased germination percent and seedling length [116]
(0.9%) and vigorous growth, reduced germination time

Table 3.
Priming with plant growth regulators and their effectiveness in improving growth of various crops under
adverse condition.

9.1 Hydropriming

Soaking seeds with water overnight and then drying before sowing markedly
improved seedling emergence, plant growth establishment, vigor and final yield in
field crops [12]. Slow and inconsistent germination of seeds has prompted the need
for water-based seed priming. Hydro priming is a very sustainable, cost-effective
and environmentally friendly technique, mainly involving soaking the seeds in
water for a predetermined time and then drying them back to their initial moisture
level [89]. The process of seed germination occurs in three phases, viz., rapid water
uptake or imbibition (phase I), lag or plateau phase (phase II), and protrusion of
seminal root and resumption of growth (phase III) [55]. Hydro priming reduces the

12
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

S. no Priming agent Crop Attributed improved References

1. Potassium nitrate Rice Improve crop emergence, crop establishment [117]

(2%) (50 ppm) and yield in DSR

2. CaCl2(50 mM) Sorghum Increased germination rate, root and shoot [112]

length under salt stress condition

3. KH2PO4 (1%) Maize Increased field emergence, plant height number [26]
of leaves and seed yield

4. Mg(NO3)2 + ZnSO4 Wheat Increased plant height, number of leaves, leaf [118]

(2%) area and chlorophyll content and increased yield
under drought stress
5. Ammonium Common Improved germination percent, net CO2 [119]
molybdate (0.1%) Bean assimilation rate, chlorophyll content and
increased grain yield

6. KCl (1%) Green Enhancing crop stand and increasing yield [120]
gram under drought condition

Table 4.
Nutrient priming techniques and their effectiveness in improving growth of various crops under adverse
condition.

S. no Priming agent Crop Attributed improved References

1. Rhizobium Green gram Increased germination percentage, [121]
sp + Trichoderma viride synchronized seed germination, growth
and yield components.

2. Bacillus amyloliquefacians Rice Enhanced activities of peroxidase and [122]

polyphenol oxidase in seedlings. Improved
germination percentage, increased leaf area

3. Bacillus sp. (MGW9) Maize Improved the germination energy, seedling [123]
length, relative water content, field
seedling emergence and seedling growth
4. Trichoderma asperelllum Wheat Plant growth promoting activities, [124]
(24 h) uniformity in seed emergence, good
seedling vigor and establishment under
stress conditions
5. Azotobacter chroococuum Chickpea Implant plant growth, and dry weight and [125]
yield

6. Azospirillum lipoferum Barley Tolerance to stress and improved plant [126]

growth and productivity

Table 5.
Biopriming priming techniques and their effectiveness in improving growth of various crops under adverse
condition.

lag period ensures rapid and uniform germination for good stand establishment
[127]. Controlled seed hydration as a pre-sowing strategy triggers pre-germination
metabolic activities in the form of cellular physiological, biochemical, and molecular
changes [93]. Ameliorated germination of hydro primed seeds is a repercussion
of stimulation of enzymes (amylase, protease, phosphatase, lipase, etc.), ATP
production, RNA and protein synthesis, DNA replication, detoxification of ROS
and lowering of lipid per oxidation by antioxidant enzymes [superoxide dismutase
(SOD), catalase (CAT), peroxidase (POD), and glutathione reductase (GPx)],
accumulation of germination enhancing metabolites (proline, soluble sugars, etc.),
higher utilization of seed reserves (proteins, carbohydrates, lipids, and phosphorus

13
Plant Stress Physiology - Perspectives in Agriculture

compounds), and supplementary metabolic repairing mechanisms. The major

limitation associated with hydro priming is uncontrolled water uptake which result
in unsynchronized germination [12]. Plants produced from hydro-primed seeds had
substantial water uptake which is positively associated with seedling growth. Hydro-
priming is a simple method to improve abiotic stress tolerance and improve germina-
tion percentage in cauliflower [28]. Seed soaking in water improved germination,
seedling establishment and yield in wheat crop under contrasting environmental
conditions [128]. Hydro-priming of rice seeds improved the germination rate, speed
and uniformity even under less than optimum field condition in upland conditions
[129]. Hydropriming of soybean seeds for 12 h was effective to increase number of
pods, grain yield and biological yield under drought stress conditions [130]. Hydro-
priming for 48 h of naturally aged rice seeds were more efficient in seed germina-
tion, emergence and seeding vigor under direct seeded rice [131].

9.2 Osmoconditioning

Osmopriming is known as osmotic priming, osmotic conditioning or osmocon-

ditioning. It is a pre-sowing treatment in an osmotic solution that allows seeds to
imbibe water to proceed to the first stage of germination but prevent radicle protru-
sion through the seed coat [132]. In this methodology, seeds are soaked in osmotic
solutions of organic compounds such as polyethylene glycol, mannitol, glycerol,
and sorbitol having low water potential so as to regulate the water uptake by seeds
and allow the pre-germinative metabolic events to continue, but restrict the seminal
root protrusion [20]. Osmohardening with PEG improved seed quality of maize
and leading to early germination and better performance under field conditions
[133]. Polyethylene glycol (PEG) as an inert material which can prevent embryo
toxicity problem during priming. The large size of PEG molecule also prevents
its penetration into seed tissues, avoiding lowering the osmotic potential [134].
Seeds primed with PEG were effective in improving seed germination and seedling
establishment of sorghum under unfavorable soil moisture conditions. Seed prim-
ing with PEG reduced lipid peroxidation and stabilized cell membrane, resulting
in elevation of stress tolerance under drought environment [135]. Osmopriming
or Osmoconditioning is the seed soaking in solutions with low water potential. In
osmopriming, degree and rate of imbibitions is restricted through the exposure of
seeds to low external water potential. Osmopriming can maintain the integrity of
plasma membrane and gives better germination percentage [28].
Osmopriming of rice seeds enhanced starch to improve sugar availability
in embryo and produced strong seedling growth. Osmopriming economically,
technically and methodologically is more challenging as it yields easier and faster
results than water conservation systems. Osmopriming of spinach seeds with PEG
increased germination percentage, stimulated anti-oxidant defense systems and
thereby induced tolerance to spinach plants. Osmopriming has advantages which
include rapid and uniform germination and emergence, improved seedling growth
and better stand establishment under water stress condition [64]. Osmopriming in
adequate concentration of PEG improved seedling growth and germination in rice
[136]. Seed osmopriming with lower dose of PEG addressed the inhibitory effects
of salinity on green gram plants in terms of greater values for osmolytes accumu-
lation, chlorophyll content and better antioxidant defense system and osmotic
adjustment [137]. Priming of french bean seeds with PEG 0.1Mpa enhanced
germination, emergence time and seedling vigor index [138]. Osmopriming with
mannitol mitigated the inhibitory effects of salinity and drought on plant growth in
chickpea. Plant produced from seeds soaked in different concentration of mannitol
(2–4%) improved biomass and length of shoot and roots under saline conditions

14
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

[139]. Plants from seeds primed with mannitol had higher activities of antioxidant
enzymes and minimal electrolyte leakage and malondialdehyde contents [140].
Osmoconditioning of cucumber (Cucumis sativus L.) seed with 0.7 M mannitol
improved the rate of germination at 25°C and 15°C in water. Osmoconditioning
stimulated the rate of radical extension, seedling emergence and expansion of the
cotyledons and first leaf of cucumber [51].

9.3 Priming with plant growth regulators

Presoaking seeds with optimal concentrations of plant growth regulators has

shown to effectively improve germination as well as growth and yield performances
of various crop species crops under environmental stress conditions [141]. Growth
regulators normally used for seed priming include auxins (IAA, IBA, and NAA), gib-
berellins (GA), kinetin, salicyclic acid, abscisic acid, ethylene and ascorbic acids. The
use of plant hormones and other plant growth regulators as seed pre-sowing treatment
can improve plant growth under stressful conditions [28]. Pre-soaking GA3 in guava
seeds at 100 ppm at a temperature range of 32°C/20°C resulted in a significant increase
in germination to 84–88% compared to unprimed seeds [54]. Seed priming with GA3
(100 ppm) for 24 h at low concentration and priming temperature at 15°C in cowpea
increased the leaf area index, relative growth rate, crop growth rate and net assimila-
tion rate under limited soil water conditions [142]. Rye seeds soaked with gibberellic
acid increased germination percentage under water stress conditions [143].
Seed priming with ethylene minimizes the effect of high temperature on lettuce
seed germination [144]. The pre-treated wheat seeds with salicylic acid improved seed
germination, rate of germination and total chlorophyll content significantly under
salinity levels [145]. Phytohormonal priming can augment seed germination through
enhancing some enzymes such as amylase activities and protease that hydrolyzed
starch and protein molecules into simple forms available for the embryo to germinate
[146]. GA3 priming has been found to improve seed germination, possibly as a result
of nutrients stored inside the seeds, and to make embryos available during germina-
tion. Seed endosperm is found in the embryo through the action of other hydrolase
enzymes [147]. It is very important that GA3 promotes the synthesis and production
of hydrolases, especially α-amylase, which leads to seed germination. Seed priming
is controlled through suppression effects of excess ABA on the expansion of embryo
organs caused by inhibition of GA3 effects on the growth of radical and hypocotyl
[148]. Seeds primed with ascorbic acid improved emergence, growth and yield
of maize under water deficit [149]. Seed priming with gibberellic acid induced an
increase in grain yield of wheat plants, modulation of ion uptake and partitioning and
hormone homeostasis under saline conditions [150].

9.4 Nutrient priming

Nutrient priming or nutripriming means soaking of seeds in nutrient solution of a

specific concentration, for a certain period of time or duration prior to sowing [151].
Seed priming with nutrients (macro or micro) can increase seed nutrient content and
improve seed quality for better germination, seedling establishment, plant growth,
nutrient uptake and water use efficiency of several crop species. Nutrient priming is
one of the methods of priming practices that includes salts like ammonium molybdate,
Mg (NO3)2, CaCl2, CaSO4, KBr, MgSO4, KH2PO4, ZnSO4, KNO3, sodium molybdate,
KCl and NaCl in such a way that pre-emergence metabolic functions begin to prevent
major outbreaks followed by seed drying at the initial humidity level [24]. In this way,
the seeds are immersed in various salts that promote the germination process and
subsequent emergence of seedlings even under adverse environmental conditions.

15
Plant Stress Physiology - Perspectives in Agriculture

Salt priming of hot pepper seed induced salinity tolerance at seedling stage, wherein
seed priming improved significantly the germination percentage and vigor index,
plumule and radical length and dry weight of seedling as compared to the non-primed
seeds (control) [110]. Nutrient priming in 3% KNO3 solution for 40 h at normal room
temperature increased speed of emergence, seedling vigor index, root length and shoot
length over hydro priming, and control in pepper [152]. Pre-sowing seed treatment
with ammonium molybdate (10−3 M) enhanced germination, improved vigor and
growth of root system, increased drought tolerance which helped in higher nutrient
uptake in cowpea crop under limited soil moisture [46]. Pepper seeds primed in 1%
KNO3 recorded the highest germination percentage as compared to non-priming
[153]. Seed priming with CaCl2 (2%) and sodium molybdate (100 ppm) increased
the harvest index over dry seed under drought stress situation [154]. Nutrient prim-
ing is a simple and low cost agro-technique and found suitable to be recommended
to the farmers owing to better synchrony of emergence and crop stand under various
conditions of environment [7]. Seed ripening with CaCl2 has been very successful in
implanting a high salt tolerance to maize with an improved percentage of germina-
tion and biomass of plants. Plants grown from extracted seeds also raised the cellular
levels of Ca2+, K+, and Na+. Chloride content was important for maize plants raised
from seeds incorporated into NaCl and KCl [155]. Effect of seed priming with KNO3
and urea increased the seedling growth, germination percentage, germination rate
and proline and protein content in maize hybrids under severe and moderate salt and
drought stress [156]. Seed priming with ammonium molybdate (10−3 M) improved
germination, stimulated growth, seed yield, biological yield and water use efficiency
in cowpea under limited water supply conditions in cowpea [157]. Seed priming of
chickpea seeds in a 0.05% solution of zinc sulphate (ZnSO4) has been found quite
effective to exhibit 19% higher seed yield and 29% more Zn concentration in seeds over
that of non-primed seeds [158]. Seed priming with potassium nitrate (0.5%) recorded
higher emergence, shoot length, shoot fresh weight, maximum root length and root
fresh weight of dry direct-seeded rice compared with non-primed seeds [159]. Seed
priming for pepper crop with osmotic solution KCl (10 mM) for 36 h improved the
plant biomass, number of leaves per plant, shoot and root length, leaf area and carot-
enoid content under saline stress [90]. Seed priming sorghum seeds with 50 mM Ca Cl2
enhanced the germination potential, germination rate, germination index, vigor index,
root and shoot length, root and shoot fresh weight and root and shoot dry weight under
salt stress condition [26].

9.5 Biopriming

Bio-priming of seeds has diverse process to stimulate morphogenesis and

plant immunity, viz., production of phytohormones, induced expression of plant
growth-promoting genes, mycoparasitism, increased nutrient status into the plant,
antibiosis, trigger phenolic production, activation of antioxidant production, and
systemic defense activation. Biopriming plays an important role in improving seed
viability, germination, uniformity in emergence, plant vigor, growth and yield [160].
Biopriming agents comprehend plant growth promoting microorganisms (PGPM),
Plant growth promoting bacteria (PGPB), plant growth promoting fungi (PGPF)
and plant growth cyanobacteria (PGPC). PGPC are responsible for enhancing the
crop growth through nitrogen fixation and release of metabolites, improving soil
fertility by soil aggregation and enhancing water holding capacity [161]. Rhizoshere
microbes play a very crucial role in enhanced uptake of three essential nutrients N, P
and K [162]. Application of Trichoderma sp. through seed biopriming enhanced the
enzyme activity through release of metabolites in maize plants [163]. Application
of Trichoderma harzianum to cucumber seeds as aqueous slurry and incubated

16
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

this mixture for 4 days at 20°C increased seedling emergence [164]. Slurry coating
of non-primed cucumber (C. sativus L.) seeds with Trichoderma harzianum and
Trichoderma viridae or combination of both reduced percentage of damping-off dis-
ease and increased the final emergence percentage up to 58.10% and greater seedling
fresh weight [165]. Biopriming is recently used as an alternative method for control-
ling many seed- and soil borne pathogens [166]. Combined effect of Pseudomonas
fluorescens and Trichoderma harzianum as seed biopriming resulted in significant
growth of pepper seedlings [167]. Among abiotic stress amelioration by bio priming,
Trichoderma spp. has been used in controlling salinity and drought stress in maize
and wheat which exhibited better physiological and morphological parameters when
compared to untreated control [168]. Biopriming with the biofungicide and clove
oil 0.06% or 0.1% was an effective seed treatment to improve the vigor and relative
speed of germination in hot pepper seeds [169]. Seed priming with Rhizobium +
Pseudomonas at 10% for 12 h recorded significantly higher germination percent
and speed of germination, and seedling vigor in chickpea [170]. Application of
Pseudomonas aureofaciens through drum priming system enhanced the stand estab-
lishment in tomato [171]. The results showed that seed inoculation with plant growth
promoting rhizobacteria had significantly effects on grain yield, grain 1000 weight,
number of grains per plant, plant height and all of grain filling parameters such as
grain filling period, rate and effective grain filling period inlentil [172]. The tech-
nique of biopriming to document using two strains including Azospirillum brasilense
and Bacillus amyloliquefaciens increased drought tolerance in wheat plants through
regulation of genes related to stress. Biopriming, an amalgamation of seed priming
with application of plant beneficial fungi and bacteria, can remarkably improve
seed germination and emergence, seedling establishment, crop growth, and yield
parameters under normal and stress conditions [173]. Thallasso bacillus denorans
and Oceano bacillus kapialis isolates from salt mine showing halophillic behavior
enhanced the growth of fine rice variety under varying salinity concentrations and
exhibited improvement in morphological and physiological parameters after 15
and 28 days, respectively, when applied through biopriming [174]. Bioprimping of
Medicago truncatula seeds with Bacillus spp. Improved seed germination and seedling
biomass and at the molecular level reflected in the up regulation of genes involved in
DNA damage repair and antioxidant defense [88].

9.6 Solid matrix priming

In solid matrix priming (SMP) or matrix conditioning, solid or semi solid

medium is used as a substitute in place of liquid medium. This technique is accom-
plished by mixing seeds with a solid or semi solid medium and specified amount
of water. In solid matrix priming, a small quantity of seed and solid particles are
used. During solid matrix priming, water is slowly delivered to seeds and thus, slow
or controlled imbibition occurs, allowing cell repair mechanisms to function [28].
Predominant solid matrices are exfoliated vermiculate, expanded calcined clay, bitu-
minous coal, sodium polypropionate gel or synthetic calcium silicate. Solid matrix
priming using saw dust, ground charcoal, green gram seeds responded favorably to
shorten incubation periods. The longer incubation periods and higher water levels
were harmful to the seeds because they encouraged fungal growth [175].

10. Conclusion and future perspective

Seed priming emerges as a reassuring technology for combating abiotic stress

in crops and alleviating the detrimental effects of abiotic stress without much

17
Plant Stress Physiology - Perspectives in Agriculture

influencing its fitness. Seed priming technique is innovative, cheap and simple to
apply at farmer’s field conditions. Oxidative stress, temperature extremes, salin-
ity, and drought are associated and frequently induce similar type of damage.
Seed priming stimulates signaling pathways earlier and enhances plant defense
responses. Experimental results reveal that improved germination and vigorous
growth of seedlings occur in early seed by combining stored nutrients and using
genes responsible for the synthesis of essential enzymes. Priming is also capable of
repairing damage that occurs inside the seed. Seed priming effects on early stage of
germination, and it modulates the DNA replication, transcription, and translation.
Storage and short shelf life of the primed seeds are a limitation of this technology.
There is a need to standardize suitable priming methods in different crops to com-
bat abiotic stress sustainably. Seed priming may indeed be considered as a valuable
strategy to improve stand establishment under detrimental agro-climatic condi-
tions (rainfed, dry farming and dry land farming regions) with enhanced yield,
increased tolerance to stress situations, enhanced crop competitiveness against
weeds, increased resistance against diseases and increased water use efficiency. In
an outline, seed priming acts as an important criterion for the induction of toler-
ance in plants against a wide range of abiotic stresses. However, more investigation
will be needed in unraveling the mechanism of plant growth regulators and their
substitutes, especially with stress-responsive genes.

Author details

Melekote Nagabhushan Arun1*, Shibara Shankara Hebbar2, Bhanuprakash2,

Thulasiram Senthivel3, Anil Kumar Nair2, Guntupalli Padmavathi4,
Pratima Pandey5 and Aarti Singh1

1 Department of Agronomy, Indian Council of Agricultural Research-Indian

Institute of Rice Research (ICAR–IIRR), Hyderabad, India

2 Department of Vegetable Crop, ICAR-Indian Institute of Horticultural Research,

Bangalore, India

3 Department of Agriculture and Animal Husbandry, Gandhigram Rural Institute,

Dindigul, India

4 Department of Plant Breeding, ICAR-Indian Institute of Rice Research,

Hyderabad, India

5 Department of Seed Science, Nepal Agricultural Research Council, Kathmandu,

Nepal

*Address all correspondence to: [email protected]

© 2022 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms
of the Creative Commons Attribution License (http://creativecommons.org/licenses/
by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,
provided the original work is properly cited.

18
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

References

[1] Kooyers NJ. The evolution of drought emergence, vigor and antioxidant
escape and avoidance in natural activities of maize. African Journal of
herbaceous populations. Plant Science. Biotechnology. 2012;11:1127-1137
2015;234:155-162
[9] Liu Y, Li P, Xu GC, Xiao L, Li ZB.
[2] Zhao TJ, Liu Y, Yan YB, Feng F, Growth, morphological and
Liu WQ, Zhou HM. Identification of the physiological responses to drought stress
amino acids crucial for the activities of in Bothrochloa ischaemum. Frontiers in
drought responsive element binding Plant Science. 2017;8:230
factors (DREBs) of Brassica napus. FEBS
Letters. 2007;581:3044-3050 [10] Alzahrani FO. Metabolic of osmo
protectants to elucidate the mechanism
[3] Bewley JD, Black M. Seeds: of salt stress tolerance in crop plants.
Physiology of Development and Planta. 2021;253:1-17
Germination. 2nd ed. New York: Plenum
Press; 1994. DOI: 10.1007/978-1-4899- [11] Blum A. Osmotic adjustment is a
1002-8 prime drought stress adaptive engine in
support of plant production: Osmotic
[4] Zhang Y, Pan J, Huang X, Gua D, adjustment and plant production. Plant
Lou H, Hou Z, et al. Differential effects Cell Environment. 2017;40:4-10
of a post-anthersis heat stress on wheat
[12] Harris D, Joshi A, Khan PA,
(Triticum aestivum L.) grain proteomic
Gothkar P, Sodhi PS. On-farm seed
determine iTRAQ. Science Reporter.
priming in semi-arid agriculture
2017;7:3468
development and evaluation in maize,
rice and chickpea in India using
[5] Hamidi H, Safarnejad A. Effect of
participatory methods. Experimental
drought stress on alfalfa cultivars
Agriculture. 1999;35:15-29
(Medicago sativa L.) in germination
stage. American-Eurasian Journal of [13] Ashraf M, Afaf YR, Qureshi MS,
Agricultural and Environmental Sarwar G, Naqvi MH. Salinity induced
Sciences. 2010;8(6):705-709 changes in amylase and protease
activities and associated metabolism in
[6] Jisha KC, Puthur JT. Halopriming of
cotton varieties during germination and
seeds imparts tolerance to NaCl and early seedling growth stages. Acta
PEG induced stress in Vigna radiata (L.) Physiologiae Plantarum. 2002;24:37-44
Wilczek varieties. Physiology and
Molecular Biology of Plants. 2014;20(3): [14] Khajeh-Hosseini M, Powell AA,
303-312 Bimgham IJ. The interaction between
salinity stress and seed vigour during
[7] Sedghi M, Nemati A, Amanpour- germination of soybean seeds. Seed
Balaneji B, Gholipouri A. Influence of Science and Technology. 2003;31:715-725
different priming materials on
germination and seedling establishment [15] Hussian S, Zhang J, Zhong C, Zhu L,
of milk thistle (Silybum marianum) Cao X, Yu S. Effect of salt stress on rice
under salinity stress. World Applied growth, development characteristics
Sciences Journal. 2010;11(5):604-609 and the regulating ways: A review.
Journal of Integrative Agriculture.
[8] Ahmad I, Khaliq T, Ahmad A, 2017;16(11):2357-2374
Basra SMA, Hasnain Z, Ali A. Effect of
seed priming with ascorbic acid, [16] Vollenweider P, Gunthardt-Goerg MS.
salicylic acid and hydrogen peroxide on Diagnosis of abiotic and biotic stress

19
Plant Stress Physiology - Perspectives in Agriculture

factors using the visible symptoms in Sheffield Academic Press, Sheffield,

foliage. Environmental Pollution. UK; 2000. pp. 287-325
2005;137:455-465
[25] Savvides A, Ali S, Tester M,
[17] Wahid A, Gelani S, Ashraf M, Fotopoulas V. Chemical priming of
Foolad MR. Heat tolerance in plants: An plants against multiple abiotic stresses:
overview. Environmental and Mission possible? Trends in Plant
Experimental Botany. 2007;61(3): Science. 2016;21:329-340
199-223
[26] Chen X, Zhang R, Xing Y, Jiang B,
[18] Zinn KE, Tunc-Ozdemir M, Li B, Xu X. The efficacy of different
Harper JF. Temperature stress and plant seed priming agents for promoting
sexual reproduction: Uncovering the sorghum germination under salt stress.
weakest links. Journal of Experimental PLoS One. 2021;16(1):e0245505.
Botany. 2010;61:1959-1968 DOI: 10.1371/journal.pone.0245505

[19] Hasanuzzaman M, Nahar K,

[27] Fujita M, Fujita Y, Noutoshi Y,
Fujita M, Ahmad P, Chandna R, Takahashi F, Narusaka Y,
Prasad M, et al. Enhancing plant Yamaguchi-Shinozaki K, et al. Crosstalk
productivity under salt stress: Relevance between abiotic and biotic stress
of poly-omics. In: Salt Stress in Plants. responses: A current view from the
New York, NY: Springer; 2013. pp. points of convergence in the stress
113-156. DOI: 10.1007/978-1- signaling networks. Current Opinion
4614-6108-1_6 Plant Biology. 2006;9:436-442
[20] Ashraf M, Foolad MR. Pre-sowing
[28] Jisha KC, Vijayakumari K, Puthur JT.
seed treatment—A shotgun approach to
Seed priming for abiotic stress tolerance:
improve germination, plant growth, and
An overview. Acta Physiologiae
crop yield under saline and non-saline
Plantarum. 2013;35:1381-1396
conditions. Advance of Agronomy.
2005;88:223-276
[29] Siri B, Vichitphan K, Kaewnaree P,
[21] Nleya T, Ball RA, Vandenberg A.
Vichitphan S, Klanrit P. Improvement of
Germination of common bean under quality, membrane integrity and
constant and alternating cool antioxidant systems in sweet pepper
temperatures. Canadian Journal of Plant (Capsicum annuum Linn) seeds affected
Sciences. 2005;85(3):577-585 by osmopriming. Australian Journal of
Crop Science. 2013;7(13):2068-2073
[22] Sivritepe HO. The effects of osmotic
conditioning treatments on salt tolerance [30] Harris D, Raghuwanshi BS,
of Onion seeds. In: 3rd National Gangwar JS, Singh SC, Joshi KD,
Symposium on Vegetable Production; Rashid A, et al. Participatory evaluation
Isparta, Turkey. 2000. pp. 475-481 by farmers of on-farm seed priming in
wheat in India. Nepal and Pakistan.
[23] Bradford KJ. Manipulation of seed Experimental Agriculture. 2001;37:
water relations via osmotic priming to 403-415
improve germination under stress
conditions. HortScience. 1986;21: [31] Khan FA, Maqbool R, Narayan S.
1105-1112 Reversal of age-induced seed
deterioration through priming in
[24] McDonald MB. In: Bewley MJD, vegetable crops—A review. Advance in
editor. Seed Priming, Black, Seed Plants and Agriculture Research.
Technology and Its Biological Basis. 2016;4(6):403-411

20
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

[32] Sano N, Seo M. Cell cycle inhibitors [41] Anese S, Da Silva EAA, Davide AC,
improve seed storability after priming Rocha Faria JM, Soares GCM,
treatments. Journal of Plant Research. Matos ACB, et al. Seed priming
2019;132:263-271 improves endosperm weakening,
germination, and subsequent seedling
[33] Arun MN, Bhanuprakash K, development of Solanum lycopersicum St.
Shankara Hebbar S, Senthivel T. Effect Hil. Seed Science and Technology.
of seed priming with various chemicals 2011;39:125-139
on germination and vigour in cowpea
(Vigna unguiculata (L.) Walp). [42] Sarika G, Basavaraju GV,
Progressive Research—An International Bhanuprakash K, Chaanakeshava V,
Journal. 2016;11:2113-2117 Paramesh R, Radha BN. Investigation on
seed viability and vigour of aged seed by
[34] Salisbury FB, Ross CW. Plant priming in french bean. Vegetable
Physiology. Belmont, CA: Wadsworth; Science. 2013;40:169-173
1992. pp. 357-407, 531-548
[43] Yari L, Aghaalikhani M, Khazaei F.
[35] Hussain A, Rizwan M, Ali Q, Ali S. Effect of seed priming duration and
Seed priming with silicon nano particles temperature on seed germination
improved the biomass and yield while behavior of bread wheat (Triticum
reduced the oxidative stress and aestivum L.). ARPN Journal of
cadmium concentration in wheat grains. Agricultural and Biological Science.
Environmental Science and Pollution 2010;5(1):5-8
Research. 2019;26(8):7579-7588
[44] Kausar M, Mahmood T, Basra SMA,
[36] Heydecker W, Coolbear P. Seed Arshad M. Invigoration of low vigor
treatments for improved performance; sunflower hybrids by seed priming.
survey and attempted prognosis. Seed International Journal of Agriculture and
Science and Technology. 1977;5:353-425 Biology. 2009;11:521-528

[37] Cantliffe DJ. Priming of lettuce seed [45] Assefa MK, Hunje R.
for early and uniform emergence under Standardization of seed priming
conditions of environmental stress. Acta duration in soybean [Glycine max (L.)
Horticulturae. 1981;122:29-38 Merill]. Seed Research. 2011;39:1-4

[38] Ghassemi-Golezani K, Jabbarpour S, [46] Arun MN, ShankaraHebbar S,

Zehtab-Salmasi S, Mohammadi A. Bhanuprakash K, Senthivel T. Effect of
Response of winter rapeseed (Brassica seed priming on phenology, growth and
napus L.) cultivars to salt priming of yield of vegetable cowpea (Vigna
seeds. African Journal of Agricultural unguiculata (L.) Walp) under different
Research. 2010;5:1089-1094 intensities of moisture stress.
Progressive Research—An International
[39] Rajpar I, Kanif YM, Memon AA. Journal. 2016;11:2118-2122
Effect of seed priming on growth and
yield ofwheat (Triticum aestivum L.) [47] Bhanuprakash K, Yogeesha HS,
under non-saline conditions. Arun MN, Naik LB. Effect of priming
International Journal of Agricultural and ageing on seed germination and
Research. 2006;1:259-264 vigour in papaya (Carica papaya).
Indian Journal of Agricultural Sciences.
[40] Anju BR, Sheeja R. Seed priming: 2009;79(4):295-297
An approach towards agricultural
sustainability. Journal of Applied and [48] Parashar A, Verma SK. Effect of
Natural science. 2019;11(1):227-234 pre-sowing seed soaking in gibberellic

21
Plant Stress Physiology - Perspectives in Agriculture

acid, duration of soaking temperatures [55] Bewley JD. Seed germination and
and their interaction on seed dormancy. The Plant Cell. 1997;9(7):
germination and early seedling growth 1055-1066
of wheat under saline conditions. Plant
Physiology and Biochemistry. 1988; [56] Bray CM. Biochemical processes
15(2):189-197 during the osmopriming of seeds. In:
Kigel J, Galili G, editors. Seed
[49] Chen K. Antioxidants and dehydrin Development and Germination. Marcel
metabolism associated with Dekker: New York; 1995. pp. 767-789
osmopriming-enhanced stress tolerance
of germinating spinach (Spinacia [57] Ibrahim EA. Seed priming to
oleracea L. cv. Bloomsdale) seeds. In: alleviate salinity stress in germinating
Graduate Theses and Dissertations, seeds. Journal of Plant Physiology.
Paper 10471. USA: Iowa State 2016;192:38-46
University; 2011. DOI:10.31274/
ETD-180810-355 [58] Chen K, Arora R. Dynamics of the
antioxidant system during seed
[50] Barthwal P, Prabha D. Effect of osmopriming, post priming
ageing and hormonal priming on germination, and seedling establishment
different physiological attributes on in spinach (Spinacia oleracea). Plant
French beans (Phaseolus vulgaris). Science. 2011;180:212-220
Advances in Agriculture and Natural
[59] Paparella S, Araújo SS, Rossi G,
Sciences for Sustainable Agriculture.
Wijayasinghe M, Carbonera D,
2018;5:76-79
Balestrazzi A. Seed priming: State of the
art and new prospective. Plant Cell
[51] Amooaghaie R, Vaviland M. The
Reports. 2015;34:1281-1293
combined effect of gibberallic acid
andlong time osmoprimng on seed [60] Bradford KJ, Bewley JD. Seeds:
germination and subsequent seedling Biology, technology and role in
growth of Klussia odoratissima Mozaff. agriculture. In: Chrispeels MJ,
African Journal of Biotechnology. Sadava DE, editors. Plants, Genes and
2011;10(66):14873-14880 Crop Biotechnology. 2nd ed. Boston:
Jones and Bartlett; 2002. pp. 210-239
[52] Renugadevi J, Natarajan N,
Srimathi P. Efficacy of botanicals in [61] Bose B, Tandon A. Effect of
improving the seeds and seedling magnesium nitrate on metabolism in
quality characteristics of cluster bean. germinating maize seeds. Indian Journal
Legume Research—An International of Plant Physiology. 1991;34:69-71
Journal. 2008;31:164-168
[62] Pandita VK, Anand A, Nagarajan S.
[53] Khan FA, Maqbool R, Narayan S, Enhancement of seed germination in
Bhat SA, Narayan R, Khan FU. Reversal hot pepper following presowing
of age-induced seed deterioration treatments. Seed Science and
through priming in vegetable crops—A Technology. 2007;35:282-290
review. Advances in Plants and
Agriculture Research. 2016;4(6):403-410 [63] Catusse J, Meinhard J, Job C,
Strub JM, Fischer U, Pestova E, et al.
[54] Bhanuprakash K, Yogeesha HS, Proteomics reveals potential biomarkers
Vasugi C, Arun MN, Naik LB. Effect of of seed vigor in sugar beet. Proteomics.
pre-soaking treatments and temperature 2011;11:1569-1580
on seed germination of guava (Psidium
guajava L.). Seed Science and [64] Chen Y, Cui J, Li G, Yuan M,
Technology. 2008;36:792-794 Zhang Z, Yuan S, et al. Effect of salicylic

22
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

acid on the antioxidant system and [73] Villiers TA, Edgcumbe DJ. On the
photosystem II in wheat seedlings. cause of seed determination in dry
Biology of Plant. 2016;60:139-147 storage. Seed Science and Technology.
1975;3:761-774
[65] Li X, Cai J, Liu F, Dai T, Cao W,
Jiang D. Cold priming drives the sub- [74] Panda D, Mondal S. Seed
cellular antioxidant systems to protect enhancement for sustainable agriculture:
photosynthetic electron transport against An overview of recent trends. Plant
subsequent low temperature stress in Archives. 2020;20(1):2320-2332
winter wheat. Plant Physiology and
Biochemistry. 2014;82:34-43 [75] Ali M, Hayat M, Ahmad H, Ghani MI,
Amin B, Atif MJ, et al. Priming of
[66] McDonald MB. Seed deterioration: Solanum melongena L. seeds enhancing
Physiology, repair and assessment. Seed germinatiuon, alters antioxidant
Science and Technology. 1999;27:177-237 enzymes, modulated ROS and improves
early seedling growth: Indicating aqueous
[67] Sen A, Jos TP. Influence of different
garlic extract as seed-priming bio-
seed priming technique on oxidative
stimulant for egg plant production.
and antioxidative responses during the
Applied Sciences. 2019;9:1-18
germination of Oryza sativa varieties.
Physiology and Molecular Biology of
[76] Sananda M, Bose B. Kinetics studies
Plants. 2020;26(3):551-556
on α-amylase extracted from
germinating wheat endosperm of
[68] Varier A, Vari AK, Dadlani M. The
primed and non-primed seeds. Indian
sub cellular basis of seed priming.
Journal of Agricultural Biochemistry.
Current Science. 2010;99(4):450-456
2012;25:137-141
[69] Farooq M, Basra SMA, Hussain M,
Rehman H, Saleem BA. Incorporation of [77] Lee SS, Kim JH, Hong SB, Yun SH,
polyamines in the priming media Park EH. Priming effect of rice seeds on
enhances the germination and early seedling establishment under adverse
seedling growth in hybrid sunflower soil conditions. Korean Journal of Crop
(Helianthus annuus L). International Science. 1998;43:194-198
Journal of Agriculture and Biology.
2007;9:868-872 [78] Chen K, Arora R. Priming memory
invokes seed stress-tolerance.
[70] Sivritepe HO, Dourado AM. The Environmental and Experimental
effect of priming treatments on the Botany. 2013;94:33-45
viability and accumulation of
chromosomal damage in aged pea seeds. [79] Jyoti MCP. Seed deterioration: A
Annals of Botany. 1995;75:165-171 review. International journal of life
science botany and pharmaceutical.
[71] Dahal P, Bradford KJ, Jones RA. Research. 2013;(3):374-385
Effects of priming and endosperm
integrity on seed germination rates of [80] Manonmani V, Ameer Junaithal
tomato genotypes. II. Germination at Begum M, Jayanthi M. Halo priming of
reduced water potential. Journal of seeds. Research Journal of Seed Science.
Experimental Botany. 1990;41: 2014;7:1-13
1441-1453
[81] Georghiou K, Thanos CA, Passam HC.
[72] Dollypan BRN. Mid-storage and Osmoconditioning as a means of
pre-sowing seed treatments for lettuce counteracting the ageing of pepper seeds
and carrot. Scientia Horticulturae. during high-temperature storage. Annals
1985;33:1026-1027 of Botany. 1987;60:279-285

23
Plant Stress Physiology - Perspectives in Agriculture

[82] Panayotov N, Aladjadjiyan A. Effect approaches for an old empirica

of long-term storage of pepper technique, new challenges in seed
(Capsicum annuum L.) seeds on their biology. In: Susana Araújo S,
viability measured by selected Balestrazzi A, editors. Basic and
thermodynamıc parameters. Acta Translational Research Driving Seed
Scientiarum Polonorum Hortorum Technology. Open, Rijeka: In Tech;
Cultus. 2014;13(2):151-162 2016. pp. 1-47

[83] Arun MN, Shankara Hebbar S, [90] Aloui H, Eymen EM, Cherif H. Seed
Bhanuprakash K, Senthivel T, Nair AK, priming to improve seedling growth of
Pratima PD. Biochemical investigation pepper cultivars exposed to salt
on vigour enhancement in fresh and concentrations. International Journal of
aged seeds upon seed priming in cowpea Vegetable Science. 2017;23:489-507
(Vigna unguiculata (L.) Walp). Legume
Research—An International Journal. [91] Gallardo K, Job C, Groot SPC,
2021;44(12):1497-1505. DOI: Puype M, Demol H, Vandekerekhove J,
10.18805/LR-4476 et al. Proteomics of Arabidopsis seed
germination and priming. In:
[84] Srinivasan K, Jain SK, Saxena S, Nicholas G, editor. The Biology of Seeds:
Radhamani J, Uprety M. Seed priming Recent Advances. CABI: Cambridge;
and fortifcation. Seed Research. 2002. pp. 199-209
2009;37(1&2):1-13
[92] Kibinza S, Bazin J, Bailly C,
Farrant JM, Corbineau F,
[85] Ermiş S, Kara F, Özden E, Demir I.
El-Maarouf-Bouteau H. Catalase is a key
Solid matrix priming of cabbage seed
enzyme in seed recovery from ageing
lots: Repair of ageing and increasing
during priming. Plant Science.
seed quality. Journal of Agricultural
2011;181:309-315
Sciences. 2016;22:588-595
[93] Wojtyla Ł, Lechowska K, Kubala S,
[86] Thornton JM, Collins ARS, Garnczarska M. Different modes of
Powell AA. The effect of aerated hydrogen peroxide action during seed
hydration on DNA synthesis in embryos germination. Frontiers in Plant Science.
of Brassica oleracea L. Seed Science 2016;7:66. DOI: 10.3389/fpls.2016.
Research. 1993;3:195-199 00066
[87] Arun MN, Bhanuprakash K, [94] Bailly C. Active oxygen species and
Shankara Hebbar S, Senthivel T. Effect antioxidants in seed biology. Seed
of seed priming on biochemical Science Research. 2004;14:93-107
parameters and seed germination in
cowpea (Vigna unguiculata (L.). Walp). [95] Coolbear P, Slater RJ, Bryant JA.
Legume Research—An International Changes in nucleic acid levels associated
Journal. 2017;40(3):562-570 with improved germination
performance of tomato seeds after
[88] Chiari F, Ajay S, Anjali S, Alma S, low-temperature pre-sowing treatments.
Prasad V, Anca M. Hydropriming and Annals of Botany. 1990;65:187-195
biopriming improve Medicago truncatula
seed germination and upgrade DNA [96] Kaya G, Demir I, Tekin A, Yasar F,
repair and antioxidant genes. Genes. Demir K. The effect of priming
2020;11:1-15 application on germination, fatty acids,
sugar content and enzyme activity at the
[89] Lutts S, Benincasa P, Wojtyla L, temperature of stress of pepper seeds.
Kubala S, Pace R, Lechowska K, et al. Journal of Agricultural Science.
Seed priming: New comprehensive 2010;16:9-16

24
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

[97] Job D, Capron I, Job C, Dacher F, on germination indices, seeding growth

Corbineau F, Coˆme D. Identification of and enzymatic activities of chickpea
germination-specific protein markers and (Cicer arietinum L.) seed. International
their use in seed priming technology. In: Journal of Ecology and Environmental
Black M, Bradford KJ, Va’zquez-Ramos J, Sciences. 2021;3(1):405-410
editors. Seed Biology: Advances and
Applications. Wallingford, UK: CAB [105] Catiempo RL, Photchanachai S,
International; 2000. pp. 449-459 Bayogan ERV, Chalermchai W. Impact
of hydropriming on germination and
[98] Cheng J, Wang L, Zeng P, He Y, seedling establishment of sunflower
Zhou R, Zhang H, et al. Identification of seeds at elevated temperature. Plant Soil
genes involved in rice seed priming in and Environment. 2021;67:491-498
the early imbibition stage. Plant Biology.
2017;19:61-69 [106] Elien Lemmens, Lomme JD, Neils
De Brier, Wannes L DeMan, Marice De
[99] Kubala S, Garnczarska M, Proft, Els Primsen, Jan AD. The impact
Wojtyla L, Clippe A, Kosmala A, of hydropriming and Osmopriming on
Zmienko A, et al. Deciphering priming- seedling characteristics, plant hormone
induced improvement of rapeseed concentrations, activity of selected
(Brassica napus L.) germination through hydrolytic enzymes and cell wall and
an integrated transcriptomic and phytate hydrolysis in sprouted wheat.
proteomic approach. Plant Science. ACS Omega 2019;4:22089-22100.
2015;231:94-113 Available from: http//pubsacs.org/
journal/acsdf
[100] Iqbal M, Ashraf M. Seed treatment
with auxins modulates growth and ion [107] Pirosteh-Anosheh H, Hashemi SE.
partitioning in salt-stressed wheat Priming, a promising practical approach
plants. Journal of Integrative Plant to improve seed germination and plant
Biology. 2007;49:1003-1015 growth in saline soils. Asian Journal of
Agriculture and Food Sciences.
[101] Nakao Y, Asea G, Yoshino M, 2020;8(1):6-10
Kojima N, Hanada H, Miyamoto K, et al.
Development of hydropriming [108] Sumar Varshini P, Bayyapu
techniques for sowing seeds of upland Reddy K, Radhika K, SaidaNaik V. Effect
rice in Uganda. American Journal of of concentration and duration of
Plant Sciences. 2018;9:2170-2182 osmopriming on germination and vigour
of aged seed of chickpea. International
[102] Bhusal D, Thakur DP. Seed Journal of Current Microbiology and
hydropriming technique in cereal crops: Applied Sciences. 2018;7(10):2410-2421
A review. Reviews in Food and
Agriculture. 2020;1(2):85-88 [109] Cokkizgin A, Girgel U,
Cokkizgin H. Mannitol effects on
[103] Elisabeth Zida P, James Neya B, germination of broad bean (Vicia faba
RomainSoalla W, Sereme P. Ole Sogaard L.) seeds. Forestry Research and
Lund. Hydropriming of pearl millet in Engineering International Journal.
northern and central Burkino Faso 2019;3(1):20-22
applying six hours of soaking and
overnight drying of seeds. African [110] PiyadaTheerakulpisut NK,
Journal of Agricultural Research. BunikaPanwang S. Priming alleviated
2017;12(149):3441-3446 salt stress effects on rice seedlings by
improving Na+/K+ and maintaining
[104] Sharma R, OmvatiVerma S. membrane integrity. International
Response to pre-sowing seed treatment Journal of Plant Biology. 2016;7:53-58

25
Plant Stress Physiology - Perspectives in Agriculture

[111] Shelar A, Singh AV, Maharjan RS, Microbial Inoculants in Sustainable

Laux P, Luch A, Gemmati D, et al. Agricultural Productivity: Vol. 1:
Sustainable agriculture through Research Perspectives. New Delhi:
multidisciplinary seed nanopriming: Springer; 2016. pp. 211-228
Prospects of opportunities and
challenges. Cell. 2021;10:6-22. [118] Sathiyanarayanan G, Suvarna G,
DOI: 10.3390/cells10092428 Baradhan G, Prakash G. Effect of
various seed halo-priming treatments
[112] Dhillon BS, Kumar V, Sagwal P, on seed yield and quality in maize. Plant
Kaur N, Singh Mangat G, Singh S. Seed Archives. 2019;19(1):377-383
priming with potassium nitrate and
gibberellic acid enhances the [119] Singhal RK, Pandey S, Bose B. Seed
performance of dry direct seeded Rice priming with Mg(NO3)2 and ZnSO4 salts
(Oryza sativa L.) in North-Western triggers physio-biochemical and
India. Agronomy. 2021;11:849. antioxidant defense to induce water
DOI: 10.3390/agronomy11050849 stress adaptation in wheat (Triticum
aestivum L.). Plant Stress. 2021;2:1-12.
[113] Mangena P. Effect of hormonal seed DOI: 10.1016/j.stress2021.10003
priming on germination, growth, yield
and biomass allocation in soybean grown [120] Majda C, Khalid D, Aziz A,
under induced drought stress. Indian Rachid B, Badr A-S, Lotfi A, et al.
Journal of Agricultural Research. Nutrient-priming as an efficient means
2020;54:441. DOI: 10.18805/IJARe.A-441 to improve the agronomic performance
of molybdenum in common bean
[114] Zhao T, Ding X, Xiao Q, Han Y, (Phaseolus vulgaris L.). Science of the
Zhu S, Chen J. IAA improves the Total Environment. 2019;661:654-663
germination and seedling growth in
cotton via regularing the endogenous [121] Krishnaprabu S. Response of green
phytohormones and enhancing the gram to pre-sowing seed priming
sucrose metabolism. Industrial Crops chemicals. International Journal of Pure
and Products. 2020;1555:1-9. and Applied Biosciences. 2018;6(6):
DOI: 10.1016/j.indcrop.2020.112788 455-458

[115] Khanmdi N, Nabipour M, [122] Bisht N, Mishra SK, Chauhan PS.

Roshanfekr H, Rahmana A. Effect of Bacillus amyloliquifaciens inoculation
seed priming and application and alters physiology of rice (Oryza sativa L.
application of cytokinin and auxin on var.IR36) through modulating
growth and grain yield of wheat under carbohydrate metabolism to mitigate
Alvay climatic condition. Iranian Journal stress induced by nutrient starvation.
of Crop Science. 2019;21(1):31-44 International Journal of Biological
Macromolecules. 2020;15(143):935-951
[116] Ahmad F, Iqbal S, Khan MR,
Abbas MW, Ahmad J, Nawaz H, et al. [123] Li H, Yue H, Li L. Seed
Influence of seed priming with salicylic biostimulant Bacillus sp. MGW8
acid on germination and early growth of improves the salt tolerance of maize
sesame. Pure and Applied Biology. during seed germination. AMB Express.
2019;8(2):1206-1213 2021;11:74. DOI: 10.1186/s13568-021-
012371
[117] Prasad SR, Kamble UR,
Sripathy KV, Udaya Bhaskar K, [124] Mahmood A, Kataoka R. Potential
Singh DP. Seed bio-primingfor biotic of biopriming in enhancing crop
and abiotic stress management. In: productivity and stress tolerance. In:
Singh DP, Singh HB, Prabha R, editors. Rakshit A, Singh H, editors. Advances in

26
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

Seed Priming Singapore. 2018. pp. [133] Jalal R, Khan AZ, Anwar S,
127-145. DOI: 10.1007/978-981-13- Ahmad J, Safia B, Ahmad F, et al.
0032-5_9 Influence of different pre-sowing
invigoration techniques on early growth
[125] Sumbul A, Ansari RA, Rizvi R, of different maize hybrids. International
Mahmood I. A potential bio-fertilizer Journal of Biosciences;15(2):370-379
for soil and plant health management.
Saudi Journal of Biological Sciences. [134] Brocklehurst PA, Dearman J.
2020;27(12):3634-3640 Interaction between seed priming
treatments and nine seed lots of carrot,
[126] Fukami J, Cerezini P, Hungria M. celery and onion. II. Seedling emergence
Azospirillium: Benefits that go far and plant growth. Annals of Applied
beyond biological nitrogen fixation. Biology. 2008;102:583-593
AMB Express. 2018;8:73.
DOI: 10.10.11186/s13568-018-0608 [135] Zhang F, Yu J, Johnston CR,
Wang Y, Zhu K, Lu F, et al. Seed priming
[127] Ahammad KU, Rahman MM, with polyethylene glycol induces
Ali MR. Effect of hydro priming method physiological changes in
on maize (Zea mays) seedling emergence. sorghum(Sorghum bicolour L. Moench)
Bangladesh Journal of Agricultural seedlings under suboptimal soil
Research. 2014;39(1):43-150 moisture environments. PLoS One.
2015;10(10):e0140620. DOI: 10.1371/
[128] Patra SS, Mehera B, Rout S, journal.pone.0140620
Tomar SS, Singh M, Kumar R. Effect of
hydro-priming and different sowing [136] Sun L, Zhou Y, Wang C, Xiao M,
dates on growth and yield attributes of Tao Y, Xu W. Screening and identification
wheat (Triticum aestivum L.). Journal of of sorghum cultivars for salinity tolerance
Applied and Natural Science. during germination. Scientia Agricultura
2016;8(2):971-980 Sinica. 2012;45(9):1714-1722
[129] Nakao Y, Asea G, Minoru Y, [137] Saha P, Chatterjee P, Biswas AK.
Nobuki K, Hiroyuki H, Kisho M, et al. NaCl pretreatment alleviates salt stress
Development of hydro priming by enhancement of antioxidant defense
techniques for sowing seeds of upland system and osmolyte accumulation in
rice in Uganda. American Journal of mungbean (Vigna radiate (L.) Wilczek).
Plant Sciences. 2018;9(11):2170-2182 Indian Journal of Experimental Biology.
2010;48:593-600
[130] Langeroodi ARS, Noora R. Seed
priming improves the germination and [138] Bhanuprakash K, Yogeesha HS,
field performance of soybean under Arun MN, Naik LB. Studies on seed
drought stress. The Journal of Animal quality in relation to ageing and priming
and Plant Sciences. 2017;27(5):1611-1621 in french beans cv.Arka komal. In:
Proceeding of the Fourth International
[131] Banjobpudsa S, Sripichitt A, Food Legume Research Conference—IV
Sarutayophat T. The effect of pre- New Delhi India. 2005. pp. 771-776
sowing treatments on germination and
vigour of upland rice (Oryza sativa L.). [139] Kaur D, Grewal S, Kaur J, Singh S.
International Journal of Agricultural Differential proline metabolism in
Technology. 2017;13:1343-1353 vegetative and reproductive tissues
determine drought tolerance in chickpea.
[132] Heydecker W, Higgins J, Biology of Plant. 2017;61(2):359-366
Gulliver RL. Accelerated germination by
osmotic seed treatment. Nature. [140] Matias JR, Ribeiro RC, Arago CA,
1973;5427:42-44 Araujo GGL, Dantas BF. Physiological

27
Plant Stress Physiology - Perspectives in Agriculture

changes in osmo and hydro-primed max L.) seeds under lead (Pb) stress.
cucumber seeds germinated in biosaline Bioscience Research. 2017;14(1):42-56
water. Journal of Seed Science.
2015;37(1):7-15 [148] Voegele A, Linkies A, Muller K,
Leubner-Metzger G. Members of the
[141] Bahrani A, Pourreza J. Gibberellin GIBBERELLIN receptor gene
acid and salicylic acid effects on seed family GID1 (GIBBERELLIN
germination and seedlings growth of INSENSITIVEDWARF1) play distinct
wheat under salt stress condition. World roles during Lepidium sativum and
Applied Science Journal. 2012;18(5): Arabidopsis thaliana seed germination.
633-641 Journal of Experimental Botany.
2011;155:1851-1870
[142] Arun MN, Shankara Hebbar S,
Bhanuprakash K, Senthivel T, Nair AK,
[149] Farooq M, Hussain M, Wakeel A,
Pratima PD. Influence of seed priming
and different irrigation levels on growth Kadambot HMS. Salt stress in maize:
parameters of cowpea (Vigna Effects, resistance mechanisms and
unguiculata (L.) Walp). Legume management—A review. Agronomy for
Research—An International Journal. Sustainable Development.
2018;43(1):99-104 2015;35:461-448

[143] Ansari O, Azadi MS, [150] Iqbal M, Raja NI, Yasmeen F,

Sharif-Zadeh F, Younesi E. Effect of Hussain M, Ejaj M, Shah MA. Impacts of
hormone priming on germination heat stress on wheat: A critical review.
characteristics and enzyme activity of Advances in Crop Sciences and
mountain rye (Secale montanum) seeds Technology. 2017;5(1):01-09
under drought stress conditions. Journal
of Stress Physiology and Biochemistry. [151] Shivay YS, Singh U, Prasad R,
2013;9:61-71 Kaur R. Agronomic interventions for
micronutrient biofortification of pulses.
[144] Nascimento WM, Cantliffe DJ, Indian Journal of Agronomy.
Huber DJ. Ethylene evolution and 2016;61(4th IAC Special Issue):161-172
endo-beta-mannanase activity during
lettuce seed germination at high [152] Maiti R, Rajkurmar D, Jagan M,
temperature. Scientia Agricola. Pramanik K, Vidasagar P. Effect of seed
2004;61(2):156-163 priming on seedling vigour and yield of
tomato and chilli. International Journal
[145] Azeem M, Qasim M, Abbas MW, of Bio-resource and Stress Management.
Tayyab SB, Yousuf M, Ali H. Salicyclic 2013;4(2):119-125
acid seed priming modulates some
biochemical parameters to improve [153] Dutta SK, Singh AR, Boopathi T,
germination and seedling growth of salt Singh SB, Singh MC, Malsawmzuali.
stressed wheat (Triticum aestivum L.). Effects of priming on germination and
Pakistan Journal of Botany. 2019;51(2):1-7 seedling vigour of bird’s eye chilli
(Capsicum frutescens L.) seeds collected
[146] Miransari M, Smith DL. Plant from eastern Himalayan region of India.
hormones and seed germination. The. Bioscan. 2015;10(1):279-284
Environmental and Experimental
Botany. 2014;99:110-121 [154] Manjunath BL, Dhanoji MM.
Effect of seed hardening with chemicals
[147] Abu-Muriefah SS. Phytohormonal on drought tolerance traits and yield in
priming improves germination and chickpea (Cicer arietinum L.). Journal of
antioxidant enzymes of soybean (glycine Agricultural Science. 2011;3(3):186-189

28
Seed Priming: The Way Forward to Mitigate Abiotic Stress in Crops
DOI: http://dx.doi.org/10.5772/intechopen.102033

[155] Ashraf M, Rauf H. Inducing salt mediated plant defense against

tolerance in maize (Zea mays L.) phytopathogens: Readdressing for
through seedpriming with chloride salts: enhancing efficacy. Soil Biology and
Growth and ion transport at early Biochemistry. 2015;87:25-33
growth stages. Acta Physiologiae
Plantarum. 2001;23(4):407-414 [163] Lopez LVP, Rodríguez AR,
Coronado MES, Hernandez PEM,
[156] Anosheh HP, Sadeghi H, Emam Y. Segovia AO. Effects of hydro priming
Chemical priming with urea and KNO3 treatments on the invigoration of aged
enhances maize hybrids (Zea mays L.) Dodonaea viscosa seeds and water-
seed viability under abiotic stress. holding polymer on the improvement of
Journal of Crop Science and seedling growth in a lava field.
Biotechnology. 2011;14(4):289-295 Restoration Ecology. 2016;24(1):61-70

[157] Arun MN, Shankara Hebbar S, [164] Desmukh AJ, Jaiman RS,
Bhanuprakash K, Senthivel T. Seed Bambharolia RP, Vijay AP. Seed
priming improved irrigation water use biopriming—A review. International
efficiency, yield and yield components of Journal of Economic Plants.
summer cowpea under limited water 2020;7(1):38-43
conditions. Legume Research—An
International Journal. 2017;40(5):864-871 [165] Pill WG, Collins CM, Goldberger B,
Gregory N. Responses of nonprime or
[158] Harris D, Rashid A, Miraj G, primedseeds of “Marketmore 76”
Arif M, Yunas M. ‘On-farm’ seed cucumber (Cucumis sativus L.) slurry
priming with zinc in chickpea and coated with Trichoderma species to
wheat in Pakistan. Plant and Soil. planting in growth media infested with
2008;306:3-10 Pythium aphanidermatum. Scientia
Horticulturae. 2009;121:54-62
[159] Zheng M, Tao Y, Hussain S, Jiang Q,
Peng S, Huang J, et al. Priming in dry [166] Reddy PP. Bio-priming of seeds. In:
direct seeded rice: Consequences for Recent Advances in Crop Protection.
emergence seedling growth and New Delhi: Springer; 2012. p. 83
associated metabolic events under
drought stress. Plant Growth Regulation. [167] Kumar VV. Plant growth-
2016;78(2):167-178. DOI: 10.1007/ promoting microorganisms: Interaction
s10725-015-0083-5 with plants and soil. In: Hakeem KR,
Akhtar MS, Abdullah SNA, editors.
[160] Devi K, Barua PK, Barua M. Plant, Soil and Microbes:Volume 1:
Integrated effect of pre-sowing seed Implications in Crop Science. Springer,
treatment, sowing windows and seasons Cham; 2016. pp. 1-16
on seed yield and quality of green gram.
Legume Research. 2021;44(8):956-961 [168] Pehlivan N, Yesilyurt AM,
Durmus N, Karaoglu SA. Trichodermalixii
[161] Shariatmadari Z, Riahi H, Seyed ID11Dseed biopriming mitigates dose
Hashtroudi M, Ghassempour A, dependent salt toxicity in maize. Acta
Aghashariatmadary Z. Plant growth Physiologiae Plantarum. 2017;39:79.
promoting cyanobacteria and their DOI: 10.1007/s11738-017-2375-z
distribution in terrestrial habitats of
Iran. Soil Science and Plant Nutrition. [169] Ilyas S, Asie KV, Sutariati GAK. Bio
2013;59:535-547 matriconditioning or biopriming with
biofungicides or biological agents
[162] Sarma BK, Yadav SK, Singh S, applied on hot pepper (Capsicum
Singh HB. Microbial consortium annuum L.) seeds reduced seed borne

29
 Plant Stress Physiology - Perspectives in Agriculture

Colletotrichum capsici and increased seed

quality and yield. Acta Horticulturae.
2015;1105:89-96

[170] Vishwas S, Chaurasia AK, Bara BM,

Debnath A, Parihar NN, Brunda K, et al.
Effect of priming on germination and
seedling establishment of chickpea
(Cicer arietinum L.) seeds. Journal of
Pharmacognosy and Phytochemistry.
2017;6(4):72-74

[171] Prachi S, Jyoti S, Shatrupa R,

Rahul SR, Anukool V, Rakesh KS, et al.
Seed biopriming with antagonistic
microbes and ascorbic acid and induce
resistance in tomato against Fusarium sp
wilt. Microbiological Research.
2020;237:1-13

[172] Abadeh MR, Sharifi RS, Imani A.

Influence of nitrogen and seed
biopriming with plant growth
promoting rhizobacter (PGPR) on yield
and agronomic characteristics of red
lenthil. Journal of Applied
Environmental and Biological Sciences.
2013;3(11):117-123

[173] Kasim WA, Osman ME, Omar MN,

Abd El-Daim IA, Bejai S, Meijer J.
Control of drought stress in wheat using
plant-growth-promoting bacteria.
Journal of Plant Growth Regulation.
2013;32:122-130

[174] Shah G, Jan M, Afrien M, Anees M,

Rehman S, Daud MK, et al. Halophillic
bacteria mediated phyto remediation of
salt-affected soils cultivated with rice.
Journal of Geochemical exploration.
2017;174:59-65

[175] Sen SK, Mandal P. Solid matrix

priming with chitosan enhances seed
germination and seedling vigouration in
mung bean under salinity stress. Journal
of Central European Agriculture.
2016;17(3):749-762

30

View publication stats