International Journal of Biological Macromolecules 234 (2023) 123708

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International Journal of Biological Macromolecules

journal homepage: www.elsevier.com/locate/ijbiomac

Review

Use of whey protein as a natural polymer for the encapsulation of plant

biocontrol bacteria: A review
Roohallah Saberi Riseh a, *, Mozhgan Gholizadeh Vazvani a, Mohadeseh Hassanisaadi a,
Vijay Kumar Thakur b, c, d, John F. Kennedy e, *
a
Department of Plant Protection, Faculty of Agriculture, Vali-e-Asr University of Rafsanjan, 7718897111 Rafsanjan, Iran
b
Biorefining and Advanced Materials Research Center, Scotland’s Rural College (SRUC), Edinburgh EH9 3JG, UK
c
School of Engineering, University of Petroleum & Energy Studies (UPES), Dehradun 248007, India
d
Centre for Research and Development, Chandigarh University, Mohali 140413, Punjab, India
e
Chembiotech Laboratories Ltd, WR15 8FF Tenbury Wells, United Kingdom

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

Keywords: Climate changes, drought, the salinity of water and soil, the emergence of new breeds of pests and pathogens, the
Whey protein industrialization of countries, and environmental contamination are among the factors limiting the production of
Encapsulation agricultural products. The use of chemicals (in the form of fertilizers, pesticides and fungicides) to enhance
Natural polymers
products against biotic and abiotic stresses has limitations. To eliminate the effects of agricultural chemicals,
Plant growth-promoting rhizobacteria
Sustainable agriculture
synthetic agrochemicals should be replaced with natural substances and useful microorganisms. To be more
Biocontrol effective and efficient, plant biocontrol bacteria need a coating layer around themselves to protect them from
adverse conditions. Whey protein, a valuable by-product of the cheese industry, is one of the important natural
polymers. Due to its high protein content, safety, and biodegradability, whey can have many applications in
agriculture and encapsulation of bacteria to resist pests and plant diseases. This compound is a rich source of
amino acids that can activate plant defense systems and defense enzymes. Considering the amazing potentialities
of formulation whey protein, this review attends to the efficiency of whey protein as coating layers on fruit and
vegetables and in the packaging system to increase the shelf life of agricultural products against phytopathogens.

1. Introduction emission of greenhouse gases (such as carbon dioxide, methane, and

Soil and water salinity, drought, heavy metals, pests, and pathogens
are among the issues that the agricultural industry faces. These factors
can affect the quantity and quality of agricultural products. Living
plants, as sessile beings, are exposed to biotic and abiotic stresses.
Climate change is the most serious because it affects agriculture by
reducing the amount of water available to the plant and causing drought
and salinity stress. Scientists in NASA’s Goddard Institute for Space
Studies (GISS) have reported that the average global temperature since
1880 has increased by about 1 ◦ C [1]. The following reasons cause
climate changes; 1) Fossil fuel burning, 2) Agriculture through the
nitrous oxide), and 3) Toxic and residual effects of synthetic chemicals
(commercial fertilizers) [2,3].
Plant pathogens with an extremely high rate of evolution are
spreading new races daily, and this issue is not suitable for agriculture.
Using different agrochemicals to control plant pathogens and insect
pests leads to environmental contamination (due to environmental
toxicity and persistence) [4,5]. Agricultural chemicals, with their effects
on soil, water, air, and other living organisms, can cause the reduction of
biodiversity and the destruction of non-target organisms [4,6]. Contin­
uous use of chemical pesticides creates chemical resistance against
fungal pathogens. Moreover, they are also harmful to humans and the

Abbreviations: GISS, Goddard Institute for Space Studies; PGPRs, Plant growth promoter rhizobacteria; VOCs, Volatile organic compounds; CE, Circular economy;
WP, Whey protein; β-LG, β-lactoglobulin; α-LA, α-lactalbumin; BSA, Bovine serum albumin; WPC, Whey protein concentrate; WPI, Whey protein isolate; PGPB, Plant
growth-promoting bacteria; IAA, Indoleacetic acid; ABA, Abscisic acid; ACC, 1-aminocyclopropane-1-carboxylate; HCN, Hydrogen cyanide; EAAs, Essential amino
acids; BCAAs, Branched-chain amino acids; pI, Isoelectric point; DA, Degree of acetylation; GA, Gum Arabic; CMC, Carboxymethyl cellulose; PB, Peanut butter; CNT,
Carbon nanotubes; PDA, Polydopamine; ROS, Reactive oxygen species.
* Corresponding authors.
E-mail addresses: r.saberi@vru.ac.ir (R. Saberi Riseh), vijay.thakur@sruc.ac.uk (V.K. Thakur), jfk@chembiotech.co.uk (J.F. Kennedy).

https://doi.org/10.1016/j.ijbiomac.2023.123708
Received 29 November 2022; Received in revised form 7 February 2023; Accepted 11 February 2023
Available online 19 February 2023
0141-8130/© 2023 Elsevier B.V. All rights reserved.
R. Saberi Riseh et al.
Fig. 1. Harmful effects of agrochemicals and poisons on living organisms.
environment [7,8]. These materials’ effects on living organisms are
depicted in Fig. 1. The entry of chemicals into nature directly and
indirectly leads to environmental contamination and a negative impact
on humans and other living organisms. The roots and rhizosphere absorb
chemical materials from fertilizers and herbicides. The output of these
materials has a negative effect on the genetic and metabolic structure of
the plant and beneficial soil microorganisms. The second harmful output
is the use of chemicals is an increase in greenhouse gases in the form of
O3, N2O, CH4, and CO2. The addition of these greenhouse gases to the
atmosphere leads to global warming. Another output of chemical ma­
terials is the impact on underground water and absorption by aquatic
organisms.
In addition, feeding animals from plants that have been treated with
chemicals also indirectly affects livestock and livestock products. As the
fourth output, we can mention the human use of the plant, aquatic, and
livestock products. Human life in this environment leads to respiratory
and skin diseases. In addition, human nutrition from these products
leads to changes in the content of the genome in the form of mutations in
the genetic material, oxidative stress and various diseases and disorders.
In general, chemicals have a momentary effect on controlling pests,
diseases, and increasing product yield. This momentary effect will have
no output other than negatively impacting the environment and living
microorganisms.
Generally, plant pathogens can damage agricultural products in two
stages; 1) The pre-harvest stage and 2) The post-harvest stage [9].
Fungal pathogens are one of the most important limiting factors for
economically important crops in field, greenhouse, or post-harvest
conditions [7]. Pathogens that cause postharvest damage to citrus and
other subtropical fruit crops (apples, grapes, and strawberries) include
Botrytis, Penicillium, Mucor, Alternaria, Colletotrichum, Lasiodiplodia,
Rhizopus, Fusarium, and Aspergillus [10]. Mycotoxins are toxic organic
compounds produced by the secondary metabolism of fungi. Citrus
fruits, grapes, apples, and strawberries are the products infected with
pathogens during the harvesting and post-harvest stages [11,12].
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International Journal of Biological Macromolecules 234 (2023) 123708
Studies have shown that antagonistic microorganisms effectively
control the disease post-harvesting citrus fruits, apples, grapes, and
strawberries with different application methods. Control of post-harvest
diseases using plant growth promoter rhizobacteria (PGPRs) is done in
two ways: (1) by direct application of natural epiphytic antagonists on
the surface of fruits and vegetables, (2) by indirect use of selective mi­
crobial antagonists that control postharvest diseases [7]. Biocontrol
bacteria are highly effective as biological control agents in managing
plant diseases (pre-harvest and post-harvest). In most plant species,
rhizosphere bacteria can be isolated from the surface of plant tissues,
soil, and roots. These factors lead to resistance induction and colonizing
plant tissues to develop resistance against pathogens [7,13]. Sustainable
agriculture refers to all processes in which healthy agricultural products
are produced by relying on non-chemical and natural factors. Sustain­
able agriculture emphasizes reducing the use of chemical agents and
increasing the use of natural factors in crop management.
Biocontrol bacteria are one of the factors considered in sustainable
agriculture. They have advantages that can be used in sustainable
agriculture, including The presence of chitinase enzyme in some of Ba­
cillus and Pseudomonas genera (as antifungal or antimicrobial). Extra­
cellular enzyme activity, the presence of different antibiotics,
production of secondary metabolites including phenazine-1-carboxylic
acid and 2, 4-diacetyl phloroglucinol, polysaccharides, and the pro­
duction of volatile organic compounds (VOCs) [7,14–16]. With all the
advantages that bacteria have, using PGPRs alone is not desirable in
biological control and does not have high efficiency [7]. As it was said at
the beginning, severe climate changes, drought, and water and soil
salinity are limiting factors that agriculture faces. PGPR as a living
agent, cannot last in such conditions and may have a momentary effect
on the quality and quantity of products and does not have a long-term
effect. Therefore, researchers should look for a way to increase the ef­
ficiency of biological agents.
In today’s world, the Circular economy (CE) is a promising strategy
to deal with the spread of pests and diseases, climate change, increasing
R. Saberi Riseh et al.
Fig. 2. CE and effects of bacteria encapsulation into WP.
protection, increasing food, and supporting sustainable and regenerative
agriculture [17]. This strategy can save and positively exploit natural
and renewable resources, reduce the negative effects of using agricul­
tural inputs, and improve economic performance (increase production)
[17–19]. The goals of the CE are reducing waste and making the best use
of it by using economic procedures to increase its value and return it to
the agrifood chain [20]. Creating alternative technological solutions to
create sustainable agriculture and a CE helps increase agricultural pro­
duction. In the meantime, developing materials based on natural poly­
mers in the form of capsules and carriers with useful microorganisms can
be a natural solution for problems related to agriculture. Controlled
agrochemical release, growth promotion, soil conditioning, and soil
condition detection are sustainable agricultural practices. Natural
polymers provided a better solution for the sustained release of agro­
chemicals economically and the CE [21].
CE and sustainable agriculture are important issues that should be
considered today. Considering biodegradable and environmentally
friendly materials is one of the important issues of CE and sustainable
agriculture. It has been reported that until 2050, agricultural production
should increase by 70 % [17,22]. Increasing agricultural production
should ensure a balance between production (exploitation of nature)
and nature protection to manage sustainable and long-term natural re­
sources [23,24].
Encapsulating bacterial cells to provide an environment that ensures
the possibility of protection and slow release of bacteria is essential. This
technique causes less exposure to adverse environmental conditions,
especially drought stress, for the bacterial cells. In addition, the selection
of encapsulation technique and wall type for encapsulation to create an
effective encapsulation, controlled release protection, and bacterial ac­
tion is of utmost importance. Using plant biocontrol bacteria in formu­
lation (solid or liquid) is a reliable and smart alternative to reduce the
use of chemical fertilizers. The selection of a bacterial encapsulation
strategy to develop the final bio product will depend on the balance
between stability, efficacy, economic feasibility, and ease of use to
generate a more sustainable and eco-friendly agricultural practice [25].
It has been reported that using natural polymers such as gum Arabic,
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International Journal of Biological Macromolecules 234 (2023) 123708
cassava starch, whey protein (WP), beeswax, and chitosan can activate
plant defense responses and reduce diseases. Using gum Arabic in
strawberries reduced the activity of Bacillus cinerea [26–29]. Some types
of natural polymers such as brewery sludge, powdered brewery, spent
grain or hydrolyzed spent brewing yeast, skim milk with whey
concentrate, bean gum, xanthan gum, and gum Arabic can be used as the
encapsulation of bacteria. Using these low-cost and natural materials
can act as the carbon source for the biocontrol bacteria. Also, due to
their biocompatibility with the environment, these materials can play an
important role in sustainable agriculture and CE [30]. Milk proteins are
caseins (about 80 % of total protein content) and whey (about 20 % of
total protein content). Caseins are similar in structure and found in milk
as nanoparticles (micelle). Casein micelles (300–600 nm) are composed
of several thousand molecules bonded via calcium phosphate [31]. Also,
the proteins of β-lactoglobulin (β-LG), α-lactalbumin (α-LA), bovine
serum albumin (BSA), immunoglobulins, protease peptone, lactoferrin,
lactollin, glycoproteins, lactoperoxidase, and transferrin make up about
1 % of cow’s milk [32–34]. Whey is a byproduct of cheese and casein in
the dairy industry. After the casein curd is separated from the milk, the
remaining watery liquid is called whey. Whey derivatives include whey
protein concentrate (WPC) and whey protein isolate (WPI). WPC is
divided into two types, with a protein percentage of 35–34 % (WPC35)
and 80 % (WPC80). WPC35 is used in producing yogurt, processed
cheese, and milk powder and various bakery applications due to its
thickening properties and nutritional benefits, combining the effects of
protein, lactose, and minerals. WPC80 is used in gelation and emulsifi­
cation applications due to its low carbohydrate content, high gel
strength, and high water-binding ability. WPI contains 90 % protein.
Due to its high protein content, WPI acts as a water-binding, gelling,
emulsifying, and foaming agent [35]. Proteins and lactose in whey could
form a matrix embedding the bacterial cells and protecting them against
several detrimental stresses during freezing and drying, such as the
formation of intracellular ice. This is because milk proteins provide
bacterial and carrier cell proteins through the reaction between amino
groups [36,37]. Whey is a low-cost compound that can be used as a
nutrient and a protective agent in biomass production and formulation
R. Saberi Riseh et al.
[25]. Fig. 2 depicts the use of WP as a natural polymer to encapsulate
biocontrol bacteria. This figure shows the benefits of using this method
in the CE of sustainable agriculture. Whey is used as the most important
encapsulants and preservatives for PGPRs. Due to its physicochemical
ability, this protein can enclose and release bioactive compounds in a
controlled manner. Whey protein has a dual role, which means that due
to its nutritional values, it protects the microorganism and also leads to
the protection of the product and the health of consumers [38].
Following the interactions that occur between whey-PGPR-plant, a se­
ries of defense reactions occur. Whey as a natural polymer with a protein
structure enters the long-term protection of bacteria, bacteria and whey
lead to the activation of antioxidant and defense reactions in the plant.
This interaction has several important outputs, including: improvement
of plant growth, producing more milk in animals, resistance to iotic and
abiotic stresses, increase of benefical microorganisms, improvement of
soil properties, and increase in plant nutrients.
Agriculture researchers must look for a way to deal with the increase
in the world population and the growing need for food. The production
of resistant cultivars and using natural and synthetic inducers are
important factors that should be paid special attention to in this regard.
Natural inducers are more popular than synthetic inducers due to their
biodegradability, biocompatibility properties, and return to the food
cycle.
In addition, environmental issues and consumer demand for sus­
tainable, healthy, and safe foods have led industrialists to focus on
developing edible packaging materials, especially whey protein (as an
edible, biodegradable, and environmentally friendly alternative
biopolymer). Edible films and coatings provide significant potential as a
functional barrier between food and the environment that ensures food
safety and quality. Edible whey film is a dry polymer network with a
three-dimensional gel-type structure. Films or coatings made of whey
proteins colorless, odorless, flexible, and transparent with excellent
mechanical and inhibitory properties (compared to polysaccharide and
other protein polymers), high water vapor permeability, low tensile
strength, and excellent oxygen permeability compared to other are
protein films [39].
In this review, the encapsulation of bacteria in WP coating for plant
protection against plant pathogens as well as the use of whey coating in
the postharvest process for long-term storage of fruits and vegetables,
have been discussed.
2. Sustainable agriculture and CE
The production of greenhouse gases, agricultural pesticides, and
non-renewable materials causes the annual production of waste in
agriculture [20,40]. On the other hand, climate changes, change in
global precipitation patterns, reductions in crop yields, and proliferation
of weeds and pests on agricultural land has greatly affected agricultural
systems [20,41]. CE is a technology and a profitable business practice
that uses agricultural wastes, by-products, and co-products. The goal of
this system is economic improvement and environmental sustainability.
This system should ensure that these materials have no environmental
impact and reduce agricultural waste production [20,42–44]. The CE
should become a model as a sustainable path for the development of
every country. This system is a road to achieve harmonious de­
velopments of the economy and the environment. In other words, the CE
uses agricultural resources and recycled usage [45]. Among the main
features of the CE, we can mention the following: 1) Economic devel­
opment, 2) Environmental protection, 3) Saving, recycling, and reusing
resources.
CE has requirements that should be taken into account [45]. These
essentials include different industries, internal and overall circulation
dimensions. Internal circulation means the development of ecological,
industrial, and sustainable agriculture. In the overall circulation, green
consumption and resource recycling should be developed. Eventually,
the positive interaction of these two components will expand together in
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International Journal of Biological Macromolecules 234 (2023) 123708
various industries such as agriculture and animal husbandry [45].
Research has shown that the implementation of the CE in the agricul­
tural sector can be summarized in the following four cases [46]: 1)
Minimizing the use of limited resources, 2) Using regenerative re­
sources, 3) Preventing natural resources from leaving the system, 4)
Reusing and recycling of resources.
3. Factors affecting sustainable agriculture and CE
In order to realize sustainable agriculture, CE, and ecological
development, the agricultural economy includes a chain of agricultural
procedures, such as cultivation, processing, distribution, and consump­
tion [47]. Different factors threaten the agricultural industry, such as
limited resources, a rapidly increasing human population (9.6 billion by
2050), and biotic and abiotic stresses [48,49]. Therefore, we should
search to develop sustainable agriculture and reduce poverty and hun­
ger in the world. Using chemical fertilizers and poisons is not ideal for
this purpose because they are unstable and have harmful environmental
effects (such as groundwater pollution, water eutrophication, soil
quality degradation, and air pollution) [48–50]. In sustainable agricul­
ture, nano fertilizers can be used to increase fertilizer efficiency [49].
Researchers have presented the advantages and disadvantages of using
nano and chemical fertilizers. Using chemical fertilizer inputs can cause
limited efficiency of nutrients, environmental restrictions, cost increases
due to over-application of chemical fertilizers, and reduced profit for
growers. However, the use of nano fertilizers has the following advan­
tages, 1) decreasing volatilization and environmental hazards, 2)
Increasing crop productivity and enhancing seed germination, 3)
Enhancing seedling growth and photosynthesis, 4) Improving nitrogen
metabolism, protein, and carbohydrate synthesis, 5) Improving stress
tolerance [49,51–56].
The application of nanotechnology in agriculture with nanoparticles
can increase the efficiency of using agricultural inputs [57,58]. For this
purpose, nano-plant growth promoters, nano-pesticides, nano-fertil­
izers, nano-herbicides, and agrochemical encapsulated nano-carrier
systems have been developed in sustainable agriculture [59–61]. The
use of natural polymers as coating for plant growth-promoting rhizo­
bacteria has a high potential in CE and sustainable agriculture. These
two components alone and in combination with each other have ad­
vantages and applications. For example, the reduced use of chemical
fertilizers with increased applications of PGPR has high efficiency in
decreasing the pressure on the environment caused by current agro­
nomical practices [62]. Encapsulation of bacteria is necessary for
providing a structure that protects the bacteria and keeps their function
constant. This technique has the following advantages: stabilizing bac­
teria cells, providing less exposure to abiotic and biotic stresses, and
enhancing viability and stability in the production, storage, and
handling processes [30,63]. On the other, hand, it is necessary to pay
attention to a natural and environmentally friendly structure as a
capsule with biodegradability and food quality for the bacteria. Natural
polymers are economical, rapidly available, non-toxic, biodegradable,
and biocompatible [64]. The combination of natural polymer-PGPR is a
promising option available in CE and sustainable agriculture [64]. In
other words, the combination of PGPR (as a plant growth stimulator and
resistance factor against biotic and abiotic stresses) with natural poly­
mers that have biodegradable properties, can be used as a promising
option in the sustainable agriculture sector. Encapsulating bacteria in
natural polymer coatings that are compatible with the environment and
biodegradable can be used in recycling materials and returning these
natural polymers to nature due to their rich compounds can be consid­
ered in the circular economy.
4. Biocontrol bacteria
Phytopathogens cause a wide range of harms to plants, including 1)
Changes in the physiological response of plants and disrupting defense
R. Saberi Riseh et al.
Table 1
The number of application plant biocontrol bacteria in sustainable agriculture.
Species
Azospirillum lipoferum
Azospirillum lipoferum, Bacillus
megaterium and Rhizobium
radiobacter
Azospirillum lipoferum,
Pseudomonas striata,
Mythylobacterium spp., and
Bacillus circulans
Bacillus subtilis
Bacillus velezensis
Pseudomonas
Pseudomonas sp.
Pseudomonas putida
Pseudomonas aeruginosa
pathways, 2) Disruption in the synthesis of secondary metabolites and
organic molecules, and 3) Reduction in yield and quality of crops.
Effective use of plant growth-promoting bacteria (PGPB) is a valuable
Goal
Combined use of phosphogypsum and Azospirillum
lipoferum to improve soil properties and increase
crop yield to improve the negative effects of
salinity on plants
Promot growth, enhance yield and increase
nutrient uptake
Combined use of bacteria to manage diseases and
sucking pests in cotton
Investigate Bacillus’s ability to suppress Rhizoctonia
solani in tomatoes (damping-off disease)
The identification of the specific secondary
metabolite gene clusters and their role in the
suppression of pathogens and plant growth
promotion
In vitro and in vivo screening of Pseudomonas
strains against Ralstonia solanacearum
Application of pseudomonas for sustainable
agriculture and investigation of their potential to
increase crop production in fields contaminated
with heavy metals
Efficacy to control common bean rust disease under
greenhouse conditions.
Investigating the antifungal effects of bacteria
against Fusarium oxysporum, Botrytis cinerea,
Botryosphaeria dothidea, and Fusarium fujikuroi
Fig. 3. The different encapsulation methods and the general structure of the microcapsule.
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International Journal of Biological Macromolecules 234 (2023) 123708
Host Result Ref.
Increase in chlorophyll, carotenoids, antioxidant enzymes,
microbial communities, soil enzymes activity, and nutrient
Zea mays L. contents [82]
The increased soil quality, nutrient content, stimulation of
plant growth, and yield
Glycine soja [83]
Increase in seed cotton yield, using these bio inoculants to the
economic management of major diseases and sucking pest
Gossypium complex in cotton
spp.
Solanum Bacillus was recovered from the soil as spores at the end of the
lycopersicum experimental period
– Producing the range of secondary metabolites, such as cyclic [84]
lipopeptides (surfactin, fengycin, bacillibactin, and
bacilycin), and polyketides (difficidin, macrolactin, and
bacillaene).
Solanum Reducing the disease incidence, producing 2,4-diacetyl [85]
lycopersicum phloroglucinol, pyoluteorin, and orfamides A and B
secondary metabolite, and inhibiting the growth of the
disease agent
Oryza sativa IAA production, nitrogen fixation, and phosphate [86]
solubilization under Cd stress; the colonization of root; has
biological control potential against plant disease (the
production of chitinase, β-1,3-glucanase, and protease).
Phaseolus Increasing the activity of enzymes (chitinase, lipase, and [87]
vulgaris protease) and reducing the severity of the disease
Lilium davidii Increase plant growth. The presence of secondary metabolite [88]
compounds in bacteria includes quinolones; 3,9-dimethox­
ypterocarpan; cascaroside B; dehydroabietylamine;
epiandrosterone; nocodazole; oxolinic acid; pyochelin;
rhodotulic acid; 9,12-octadecadienoic acid; di-peptides; tri-
peptides; ursodiol, and venlafaxine and the effect on the
growth of bulbs of Lilium davidii and inhibiting the growth of
fungi.
alternative to minimizing chemicals [61].
Sustainable agriculture production can be done using biocontrol
bacteria. PGPR or biocontrol bacteria increase plant growth in three
R. Saberi Riseh et al.
Fig. 4. Different conditions and the mechanism of delivery in the encapsulation process.
different ways: 1) By synthesizing growth-promoting hormones that are
essential for plants, 2) By facilitating the uptake of nutrients from the
soil and providing them to the plants, and 3) By reducing or preventing
plant diseases [65,66]. Biocontrol bacteria with auxin production
(indoleacetic acid (IAA), abscisic acid (ABA), gibberellic acid, and cy­
tokinins); can produce 1-aminocyclopropane-1-carboxylate (ACC)
deaminase, siderophore, antibiotics, and oxidative stress and lead to
root length increase, growth, and resistance to biotic and abiotic stresses
[63,67].
Plant biocontrol bacteria have several important advantages,
including 1) Improving plant nutrient acquisition, 2) Inducing tolerance
to be biotic and abiotic stresses in the plant, and 3) Developing plant
growth [68–71]. These bacteria are bio-stimulant and biocontrol agents
in sustainable agriculture [71].
A study by Sehrawat et al. [4] showed that hydrogen cyanide (HCN)
producing bacteria effectively inhibiting the growth of pathogenic fungi,
weeds, and insects. HCN, the secondary metabolite, inhibits the activity
of mitochondrial cytochrome carbon oxidase and leads to the disruption
of the activity of pathogens [4]. Therefore, these bacteria are considered
as bio-pesticides and are eco-friendly for sustainable agriculture. Apart
from controlling pests and plant diseases by producing IAA, antibiotics,
and fluorescent insecticidal toxins, ACC, HCN utilizes deaminase and
solubilizes phosphate and leads to the stimulation of plant growth [4].
HCN is reported in some fluorescent pseudomonas and some members
of the genera Aeromonas, Chromobacterium, Burkholderia, certain
rhizobia, cyanobacteria, and Bacillus sp. [4,72–79]. Due to their potential
for growth promotion, these bacteria are used as biological fertilizers or
biocontrol agents to enhance crop production in sustainable agriculture
[1,80,81]. Applying beneficial microorganisms (as mentioned above) is
the best idea for achieving sustainable agriculture and CE. Table 1 shows
studies on biocontrol bacteria use in biotic and abiotic stresses and their
related effects on sustainable agriculture.
4.1. Encapsulation and the release mechanism of the encapsulated
bacteria
Encapsulation is a process in which active compounds and, generally
microorganisms are stabilized through materials capable of maintaining
their chemical, physical and biological properties. In general, in
encapsulation, we come across two terms: 1- active substance or core (as
materials covered, for example, bacterial cells), 2- carrier material or
shell. The core and shell structure can be in different forms such as a
sphere, microcapsule, microbead, single core, multi-core, matrix, and
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International Journal of Biological Macromolecules 234 (2023) 123708
multi-shell [89]. The carrier material for PGPR encapsulation acts as a
protector. The carrier should be non-toxic, chemically stable, readily
available, and inexpensive. The carrier must be able to retain moisture,
lead to cell viability during storage, and be able to lead to the effective
transfer of bacterial cells to the intended target (effective release) [90].
Fig. 3 shows the different encapsulation methods and the general
structure of the microcapsule.
Encapsulation of biological control bacteria in microcapsules is done
to increase stability, slow release, and overcome the limitation of
chemicals in nature. An encapsulation method is necessary to obtain an
optimal formulation to preserve biological agents against biotic and
abiotic stresses [91–97]. Carbohydrates are the main wall materials for
encapsulation. On the other hand, proteins and other polymers have
many advantages that could be used in capsule production alone or
together (combined with carbohydrates) [98]. Using natural polymers
with a high percentage of protein is very useful for creating a healthy
environment for encapsulated bacteria for the intended purposes
[99–102].
Stabilizing beneficial microorganisms inside biodegradable poly­
mers in the form of encapsulation is a promising biological formula and
method that leads to increased plant growth, maximum cell viability and
survival, and increased colonization of plant rhizosphere and roots [90].
Biological encapsulation is divided into three types of macro, micro, and
nano, depending on the size of the produced particles, and is generally
summarized in the following three steps: 1- Microorganism absorption
and mixing in a polymer matrix; 2- stirring liquid mechanics and for­
mation of solid particles; 3- The particles formed in the previous step are
subjected to polymerization and physicochemical stabilization of the
formed particles [90,103]. Encapsulation stabilizes cells, limits exposure
to abiotic and biotic stress, improves their bioactivity and stability in
production, safe storage, and handling, and provides additional pro­
tection during rehydration. In general, the advantages of encapsulation-
based formulations: 1) increased efficiency due to increased encapsu­
lated surface area; 2), improved systemic activity due to small size, and
greater mobility of particles 3) reduced toxicity compared to fungicides,
pesticides, herbicides, and chemical fertilizers [104]. Since most of the
natural polymers used in the encapsulation of bacteria are hydrophilic,
with the increase of water absorption in the capsule’s environment, the
pores are increased, and the process of releasing the bacteria is done
quickly. It has also been reported to control the proliferation of bio­
logical bacteria by adding biodegradable additives to the matrix
[105,106]. The metabolites secreted from bacteria (such as auxin,
siderophore, chitinase, protease, ACC bind with the protein compounds
R. Saberi Riseh et al. International Journal of Biological Macromolecules 234 (2023) 123708

Fig. 5. The origin of natural polymers and the location of whey protein.

in natural polymers and follow the interaction between the substances.,
shock is created in the bacteria, and it is released from the capsule. The
release of bacteria from the capsule (long or short time) is different
under different conditions.
The purpose of delivery system encapsulation in the agricultural
sector is to provide safe products that do not degrade over a period of
time and have maximum efficiency and activity. Encapsulation should
be designed in such a way that it has different reactions in different
conditions and situations [104]. Fig. 4 shows the different conditions
and the mechanism of action in the encapsulation process.

Fig. 6. The structure and features of β-lactoglobulin.

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R. Saberi Riseh et al. International Journal of Biological Macromolecules 234 (2023) 123708

Fig. 7. The structure and features of α-lactalbumin.

5. Natural polymers products or industrial by-products, is necessary to ensure the efficient

In the agricultural sector, >70 % of the water resources are used for
the cultivation of crops and food production; this issue has become a
concern due to the severe lack of water and decreased rainfall [107]. The
use of superabsorbent hydrogels, which are derived from agricultural
use of water and soil stability. These hydrogels have several advantages,
including controlled humidification of the environment, increased
water-holding capacity, increased plant growth, and sustained release of
fertilizers [108–112]. A polymer is a macromolecule combined with
repeating structural units connected by covalent chemical bands.

Fig. 8. The structure and features of bovine serum albumin.

8
R. Saberi Riseh et al.
Fig. 9. The structure and features of immunoglobulins.
Natural polymers are preferable in terms of being economical, rapidly
available, non-toxic, biodegradable, biocompatible, and cost-effective
[64].
These characteristics make natural polymers a promising option in
CE and sustainable agriculture. Synthetic polymers have become a major
concern due to their environmental effects. On the contrary, natural
polymers as environmentally friendly agents can be an alternative op­
tion [113]. Natural polymers play different roles in different industries
as binders, matrix formers or drug release modifiers, film coating for­
mers, thickeners or viscosity enhancers, stabilizers, disintegrants, sol­
ubilizers, emulsifiers, suspending agents, gelling agents and
bioadhesives [64]. Natural polymers such as carbohydrates, lipids or
proteins are often used to manufacture micro- or nano-capsules
[114,115]. Natural polymers protect bacterial cells in different ways,
such as nutrient provision, improvement of physical characteristics, and
osmoprotectant [116,117]. These polymers create a protective micro­
environment for bacteria by limiting heat transfer, having high water
activity, and protecting the bacterial environment from dryness [25].
Low-cost additives derived from living organisms and nature can greatly
help the development of sustainable agriculture. A list of natural poly­
mers and their origin is shown in Fig. 5.
Whey is one of the industrial by-products that, as a soil softener and
high content of organic compounds, has the potential to act as a nutrient
to enhance the growth, yield, and quality of crops [112,118]. This
product can also be used for the encapsulation of bacteria. Following
paragraphs are presented a body of research in this regard.
5.1. WP, structure and function
Cow’s milk consists of two types of proteins. These two proteins are
called whey (about 20 % of total protein content) and casein as the
dominant milk protein (about 80 % of total protein content) [32–34]. In
general, there are four main classes of proteins in whey: (1) β-lg, (2) α-la,
(3) BSA, and (4) Ig. These proteins have a spherical structure and are
sensitive to denaturation and aggregation caused by heat and high
pressure. β-lg is a globular, acid-stable protein containing 162 amino
9
International Journal of Biological Macromolecules 234 (2023) 123708
acids. α-la is a globular protein containing 123 amino acids. This protein
plays an important role in lactose biosynthesis. BSA has 582 amino
acids, and its biological function is related to the transport of fatty acids
through membranes. Due to its low concentration (5 % of whey protein
and 1 % of total milk protein), BSA does not greatly affect functional
properties. It is a complex heterogeneous mixture of large glycoproteins
with the antibody activity. It constitutes approximately 2 % of total milk
protein and 10 % of whey protein [35]. The structure of these proteins is
shown in Figs. 6, 7, 8, and 9 shows the four main classes of proteins in
whey.
WPs are separated from the casein during cheese making and are
purified to different concentrations depending on the final composition.
Whey is one of the by-products and the original one obtained from
cheese. About 9 l of whey are normally obtained from each kilogram of
cheese [25]. In addition to the high nutritional value of whey, its
amphipathic structure and good film formation can be mentioned [119].
Whey and proteins derived from it are widely used in many products,
including bioethanol, single-cell protein, bioplastics, antimicrobial
peptides, enzymes, biogas, and other organic compounds [25,120]. WP
concentrate is produced by drying whey and includes 50 % lactose, 10 %
proteins, 2 % fat, 5 % minerals, and a broad range of vitamins, cofactors,
and mineral salts [121,122]. Ultrafiltration, diafiltration, electrodial­
ysis, gel filtration, ion-exchange chromatography, and reverse osmosis
processes are methods for WP recovery. The most common shapes of WP
are WPC and WPI. WPC is produced through ultrafiltration (selective
removal of high molecular weight materials by choosing a permeable
membrane) and by separating lactose and minerals from the protein.
This composition has the lowest amount of fats, cholesterol, and
bioactive compounds. WPI is prepared by WPC processing using the ion-
exchange chromatographic method [39,123,124]. WP is complete and
has high quality. This protein includes essential amino acids (EAAs) and
branched-chain amino acids (BCAAs), which are very important for
tissue growth and repair [34]. The following factors are among the
important advantages of WP: protein synthesis, muscle building, glucose
metabolism, High protein concentration compared to soybean, meat,
and wheat, fatigue symptoms reduction, and immunity enhancement
R. Saberi Riseh et al. International Journal of Biological Macromolecules 234 (2023) 123708

[34,125]. WPs in pH 4.6 and 20 ◦ C will be obtained from milk during 2 Table 2
steps: 1) Acid whey: the addition of acid and precipitation of caseins, Other properties of natural polymers.
and 2) Sweet whey: after the deletion of caseins and from rennet coag­ Polymer Properties Reference
ulation [31,34]. WP provides the nutritional requirements of entrapped
Alginate linear heteropolysaccharid;. The combination [99,127,141–146]
microorganisms [30]. WP is used in two ways: 1) WP edible film is a of β-D-mannuronic acid and α-L-guluronic
polymer network and three-dimensional gel-type structure that is acid leads to subunits binding and gel
applicable as a wrapping on the food products, and 2) Immersion of food formation. Alginate microcapsules have a
products into film-forming solution and air drying, which is called WP cracked and porous structure that reduces
their mechanical and chemical stability. The
coating. WP edible and WP coating are flexible, colorless, odorless, and viscosity of sodium alginate increases with
transparent; they also have amphiphilic nature, conformation, dena­ the number of monomer units
turation, and electrostatic charges [39,126]. (macromolecule length). The disadvantages
Studies of interactions among whey proteins have shown that lac­ of alginate beads are that they are
incompatible with alkaline conditions. The
toferrin can interact with both β-Lg and BSA at room temperature. It has
defects of this polymer can be solved by
also been shown that, by increasing the amount of lactoferrin in whey, it mixing alginate with different biopolymers
is possible to enhance the gel strength of whey protein gels. The whey and covering the capsules with other
protein interactions between lactoferrin and β-Lg/BSA are ionic. Thus, polymers such as gelatin, chitosan, and whey
their large difference in isoelectric point (pI) for lactoferrin (8.7), β-Lg protein. Due to its porous structure and
solubility, sodium alginate has a weak
(4.8), and BSA (5.6), enables them to exist with the opposite net charge protective ability, and the addition of whey
at the same pH. However, some selectivity exists in the ionic interaction, protein isolate can reduce the release rate of
as lactoferrin does not interact with α-La (pI = 4.7). Of the four major cells.
whey proteins (β-Lg, α-La, BSA, and lactoferrin), the structure of β-Lg is Gelatin Gelatin contains high levels of glycine, [99,115,131,147–149]
hydroxyproline, and proline and can be used
dominated by β-strands, whereas α-La, BSA, and lactoferrin are mainly
alone or in combination with other
composed of α-helices [127]. β-Lg can bind to hydrophobic and ingredients for PGPR encapsulation. The
amphiphilic compounds such as flavor compounds, vitamins, fatty acids, application of gelatin is limited due to its low
and polyphenols. Interactions between β-Lg and bio actives are mainly network stability. However, its physical
caused by hydrophobic bonds, although hydrogen bonds are also properties can be modified through the
addition of cross-linking agents. This
involved in the binding of polyphenols and fatty acids. It has been
substance is amphoteric in nature. Therefore,
suggested that β-Lg binds hydrophobic compounds preferentially in the it is a suitable polymer for mixing with
internal calyx. The interaction parameter (localization and number of anionic carbohydrates such as carboxymethyl
binding sites on the protein and correlation) depends on the ligand’s cellulose, alginate and whey protein. Because
gelatin is neutral, it is the most suitable option
chemical nature, the substrate’s physico-chemical conditions and the
for reaction with gellan gum.
protein’s structure. Binding stability of curcumin to plant β-Lg beyond Chitosan Chitosan is chemically composed of repeating [150–156]
that of heat-denatured protein [128]. units of D-glucosamine and N-acetyl-D-
glucosamine. Application in encapsulation
5.2. Comparison of whey protein to other biopolymers technology, controlled release coatings, drug
delivery system, nanofiltration and tissue
engineering; Chitosan is soluble in neutral
As mentioned, there are natural polymers with unique properties and acidic environments, but the viscosity
that can be used in the encapsulation process depending on different and solubility of the solution depends on the
purposes and availability and have different efficiencies. degree of acetylation and the length of the
chains. Adding polymers such as alginate,
The unique functional and nutritional properties of whey proteins
xanthanan, and whey protein forms ionic
make them useful food ingredients for various applications. The nutri­ hydrogels, which are used in various sciences
tional quality of whey proteins and the essential amino acids of this such as agriculture, food, and medicine due to
protein is high. The emulsifying, foaming and gelling properties of whey their biodegradability and biocompatibility.
protein are the basis of its widespread use as a functional ingredient in Starch The chemical structure of starch is α-D-glucose [96,105,157–159]
compound units connected by glycosidic
foods [129]. In the following, we will explain the capabilities and bonds. Starch is insoluble and dense, and its
characteristics of several other natural polymers. hydrate forms only in water at normal
Alginates are natural marine polysaccharides extracted from temperatures. By heating at 80 degrees
seaweed. Depending on the origin of alginate, three types of blocks may Celsius, the structure of this polymer is
destroyed. Starch and gelatin are effective in
be found: homopolymeric M blocks (M-M), homopolymeric G blocks (G-
protecting beneficial microbes in acidic
G) and alternating heteropolymeric MG blocks (M-G). The composition conditions. Starch-alginate mixture is a very
and structure of the block are strongly related to the functional prop­ effective compound for encapsulating and
erties of alginate molecules in an encapsulation matrix. By dissolving protecting Raoultella planticola. Tensile
alginate in water, a viscous solution is obtained, the viscosity of which strength, and percent strain at failure in corn
starch-whey mixture increased due to
increases with the number of monomer units, and its solubility is also interaction between proteins.
affected by pH (at pH < 3, it precipitates as alginic acid) [130].
Gelatin is a heterogeneous mixture of single or multi-stranded
polypeptides and has between 300 and 4000 amino acid units. Gelatin [130]. The basis of choosing a polymer for encapsulation is its high
is a form of hydrolyzed collagen obtained from skin, boiled crushed functional properties and high protein, which can provide a suitable
bones, connective tissues, organs, and some intestines of animals. There environment for the microorganism so that the microorganism and the
are two types of gelatin depending on the type of treatment to obtain the polymer interact with each other to have maximum efficiency.
gelatin: type A gelatin (raw materials treated with acid) and type B Recently, research has shown that the composition of amino acids,
gelatin (alkali treatment). Gelatin is soluble in water, has a high vis­ especially hydrophobic amino acids (determining the physical, chemi­
cosity, and forms a thermal hydrogel that turns into a liquid when cal, and functional properties of gelatin) extracted from poultry is higher
heated and solidifies when cooled again. Gelatin gels are formed only in than gelatin obtained from mammals and fish [115]. Also, in a study, an
a certain temperature range. It melts at high temperatures and crystal­ increase in the viability of probiotic bacteria was observed in gelatins
lizes at low temperatures. Chromium salts are needed to stabilize the gel synthesized from duck feet [131]. Another natural polymer is chitosan.

10
R. Saberi Riseh et al.
Fig. 10. PGPB-WP encapsulation and application in plant and its result.
Chitosan is a deacetylated derivative of chitin found in the shells of
crustaceans, fungi, insects, and molluscs. This polymer is a linear poly­
saccharide that can be considered a copolymer consisting of β-(1, 4)-
linked D-glucosamine and N-acetyl-D-glucosamine. The functional
properties of chitosan are determined by the molecular weight, as well
as by the degree of acetylation (DA), which indicates the ratio of N-
acetyl-D-glucosamine units with respect to the total number of units
[158]. Chitosan is soluble in acidic to neutral environments, but the
solubility and viscosity of the solution depends on the chain length and
DA. Chitosan is a positively charged polymer and forms ionic hydrogels
by adding anions such as pentasodium tripolyphosphate (TPP) and also
by interacting with negatively charged polymers (alginate or xanthan)
[130,132]. Table 2 shows other characteristics of natural polymers.
Due to its high nutritional value, functional activities, and afford­
ability, whey is commonly used for various food applications (dairies,
bakeries, food products, beverages, nutritional supplements or func­
tional foods). This composition is a substitute for other proteins and can
improve the functional properties of food products. This composition
has wide compatibility with the human body. So that the addition of
hydrolyzates of whey in drinks increases sensory and physicochemical
properties such as taste, smell, low precipitation and storage stability,
this composition has functional properties such as antioxidant, antimi­
crobial, and antihypertensive consumption of whey hydrolysates pro­
duces high protein synthesis in the body [133–136]. Whey protein
hydrolysates are important in food processing due to their technological
properties, including oil and water retention, emulsifying capacity,
foaming capacity, and solubility. They can promote the formation of
volatile compounds in food products regardless of being added in small
amounts [136–140]. Whey presents an important content of proteins,
group B vitamins, minerals, and lactose [136]. In general, whey protein
has important advantages [127]:
1- WP is used in the food industry to increase protein content, replace
fat, increase storage capacity, and improve texture.
2- A strong biopolymer network is formed by heating concentrated WP,
which includes covalent and non-covalent intermolecular bonds.
3- It works in the form of hydrogel and as a carrier of biologically active
compounds.
These things have made whey superior compared to other polymers.
The triple advantages of this protein for different purposes have
turned it into a polymer with different efficiencies, which is of interest in
the circular economy. Whey protein in sustainable agriculture alone, in
combination with other polymers and in the encapsulation of bacteria in
the fields of disease resistance, food packaging, the post-harvest process
of fruits and vegetables, and resistance to abiotic stresses are of interest
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International Journal of Biological Macromolecules 234 (2023) 123708
to researchers, has been placed and has shown high efficiency.
5.3. The ability of WP as a carrier in the encapsulation of PGPR
Three important features that whey proteins have and can be effec­
tive in hydrogel and microcapsule synthesis are:: 1- As surface agents in
the formation and stabilization of emulsions containing hydrophobic
bioactive substances, 2- Formation of covalent complexes or electro­
statics with the desired molecules and trapping bioactive substances
through gel formation, 3- self-assembly or co-assembly to form super­
structures that allow the encapsulation and transportation of a variety of
small molecules [128]. Carriers increase the survival and shelf life of
bacteria by protecting them from desiccation, and also provide a suit­
able microenvironment for rapid growth after release. The successful
encapsulation of PGPR is done by integrating physical, chemical, and
biological parameters, which leads to an increase in the population of
the target organism and the long-term survival of microorganisms over
time in suboptimal conditions. The material used as a carrier must have
high water retention capacity, non-toxicity, sufficient adhesion, and
desired acidity [160]. As mentioned in the previous sections, whey
protein has advantages over other polymers. In fact, this protein can
provide small peptides, amino acids, and growth factors that positively
stimulate bacteria’s growth [161]. Studies have shown that the highest
growth of Lactobacillus rhamnosus bacteria has been observed in the
medium containing whey due to low molecular weight peptides. It has
also been reported that the lactose in whey can further increase the cell
viability of microorganisms [161,162]. It has also been reported that
using whey protein as a wall material for bacteria can maintain their
viability for 3 months of storage at 20 ◦ C [163]. Therefore, it seems that
WP as a protector has positive effects on the viability of PGPRs and can
be used as a suitable carrier in the encapsulation of bacteria.
6. Encapsulation with WP
Encapsulation uses natural polymers as core materials to protect the
stability of sensitive active components against temperature, light, pH,
oxygen, moisture, and metal ion. WP, as a protein polymer, is used alone
or combined with other carbohydrate polymers (such as starch, chito­
san, sucrose, and dextrin) and plant water-soluble gums (gum Arabic,
carrageenan, xanthan gum, and sodium alginate) as wall material in
microencapsulation [164].
WPs are valuable by-products of the cheese industry. Due to their
gelling and emulsification properties, these proteins are used in food and
non-food applications, such as agriculture. The main factor of whey
gelation is the presence of b-Lactoglobulin (BLG) protein [165]. In a
study, Bacillus velezensis was capsulated with alginate combined by WP,
R. Saberi Riseh et al.
zedo, mastic, and tragacanth gums. Silica and titania nanoparticles coat
these microcapsules in two-layer and multilayer capsules [61]. In order
to provide a suitable formula for biofertilizers based on biodegradable
compounds, including whey, a survey was conducted. Results revealed
that the mixture of alginate-WP concentrate-zedo offered the highest
encapsulation efficiency. That is because of a multilayer coating around
the bacterial cells. X-ray diffraction (XRD) and Fourier Transform
Infrared Spectrometer (FTIR) showed that alginate, WP, and zedo were
mixed properly and confirmed the compatibility of these compounds
with each other and with bacterial cells [61]. In another study, WPI and
Gum Arabic (GA) were used to encapsulate the probiotic bacterium
Lactobacillus plantarum. For microcapsules production, spray-drying and
freeze-drying methods were used [166]. Lactose acts as a potential
protector in whey for its binding capacity to water and reduces the
formation of ice crystals during the storage of liquid formulations at low
temperatures. Lactose is known to stabilize cell membranes and preserve
the structure and function of proteins [167]. In addition, the presence of
eight hydroxyl groups in lactose molecules protects bacterial cells
against free radicals produced during storage [25,168]. The encapsu­
lation of biocontrol bacteria within a biodegradable matrix of protein
such as WP and poly-Lysine polysaccharide or other polymers protects
these bacteria from biotic and abiotic stress factors, leads to better dis­
ease control and promotes shelf life and persistence in the environment
[169,170]. Fig. 10 depicts essential factors of sustainable agriculture
and CE. A WP as a natural polymer coating derived from nature and
plant growth-promoting bacteria (PGPB) can guarantee the necessary
conditions for plant survival in adverse biological and non-biological
conditions.
6.1. The encapsulation of WP-biocontrol bacteria for resistance to plant
pathogens
Preserving the efficacy of plant probiotic bacteria, such as viability,
activity, survivability, and controlled release in the soil is a major
challenge for effectively controlling plant pathogens. Accordingly, to
determine the best encapsulation method for Pseudomonas bacteria,
alginate (Alg) was combined with WPC, carboxymethyl cellulose (CMC),
and peanut butter (PB) [100]. These microcapsules were investigated to
control Rhizoctonia solani disease in potato plants under greenhouse
conditions [100]. Encapsulation efficiency for Alg-WPC was about
84.23 %. Spherical, smooth, and compact shapes with no pores were
recorded for Alg-WPC. This microcapsule (Alg-WPC) could reduce the
severity of the disease by about 90 % and improve growth in greenhouse
conditions [100]. Due to many amino acids, WP is an important source
for plant growth, metabolism, and synthesis of defense proteins against
pathogens [100,171,172]. Another investigation tested different
methods for microencapsulation and control of Rhizoctonia solani in
potatoes. These methods include extrusion, emulsification, and spray
drying. For the microencapsulation of biocontrol bacteria, alginate was
used from WP [145]. Alginate–WP microcapsules were able to reduce
the Severity of Rhizoctonia disease (up to 70 %). The presence of whey
as a substance rich in amino acids and carbon nanotubes (CNT) to in­
crease rooting and proliferation can be used to resist induction and
reduce physiological complications. This technique increases the effi­
ciency and persistence of probiotic bacteria, gradual and controlled
release of them, maintains their biological control properties, enhances
plant growth, and reduces pathogens’ negative effects on the plant. In
other words, this formulation (ALG-WP-CNT) can be used as an ideal
biological fertilizer in agriculture [145]. Moradi Pour et al. evaluated
the efficiency of Bacillus velezensis (as a Biological Control Agent) [61].
This research used alginate with WP, zedo, mastic, and tragacanth gums
as microcapsules. Silica and Titania nanoparticles in two and multilayer
assemblies were used in these microcapsules. These microcapsules were
used to control the take-all disease.
This research obtained multilayer microcapsules with Alg-WPC-ZG
and Alg-MG-ZG [61]. Amorphous structure in Alg-WPC microcapsules
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International Journal of Biological Macromolecules 234 (2023) 123708
can be due to electrostatic interaction between the carboxyl group (in
alginate) and an amino group (in WP). Trapped cells are released more
easily in this structure of their high solubility [61,173]. Alg-WPC-ZG
formulation (in multilayer form) with TiO2/SiO2 nanoparticles with a
high percentage of encapsulation was more capable of controlling take-
all disease in laboratory conditions. These microcapsules protect the
bacteria against environmental factors and biotic stresses [61]. Another
experiment was conducted to biological control bacteria with natural
and synthetic polymers against Clavibacter michiganensis subsp. nebras­
kensis in maize seeds. This study showed that whey as a coating would
stabilize bacteriophages as a biological control agent. Whey can act as a
substitute for traditional antibiotics in agriculture [174].
6.2. The encapsulation of WP-bacteriophage for resistance to plant
pathogens
Bacteriophages as natural biopesticides are another alternative to
chemical pesticides. However, they are unstable against UV rays and
drying. Therefore, a new formulation has been used to increase the
stability of bacteriophages against UV rays and e bacterial contamina­
tion (Escherichia coli (E. coli)) on plants [175]. In research, the in­
vestigators obtained a mixture of microscale polydopamine (PDA)
particles with WPI–glycerol formulation. This formulation was con­
ducted to improve the phage stability, enhance the antimicrobial ac­
tivity of phages, and prevent bacterial contaminations on the leaf
surface in the presence of UV-A [175]. The antibacterial activity of
phage is increased in this formulation (PDA-WPI-glycerol). This
formulation kept the phage alive during radiation exposure, which is
probably due to tribute to interactions of PDA (polydopamine) with the
bacterial cell surface and self-polymerization of PDA onto cells that may
influence cell viability [175–177]. Phage stability and infectivity are
modulated by 1) Protein-protein interactions, 2) Protein-DNA in­
teractions, and 3) DNA stability. These interactions include electrostatic
and non-electrostatic interactions. These interactions can be influenced
by oxidative stress generated by UV radiations and thermal and desic­
cating factors commonly encountered on a leaf substrate [175]. In
summary, PDA–WPI-Glycerol had these advantages, including
improving phage stability, enhancing phage antimicrobial activities,
and preventing bacterial contamination in field simulated conditions.
The PDA-WPI-glycerol formulation provided protection activity for
phages in agricultural practices [175]. In order to investigate the pro­
duction of protease from the Bacillus thuringiensis-whey combination, an
experiment was conducted [178]. A combination of whey+skim
milk+NaCl+CaCl2 achieved the maximum protease production.
B. thuringiensis has a high ability to produce extracellular protease.
Combined with the high functional properties of whey, this feature can
be highly effective in reducing environmental pollution. In another
research, encapsulated bacteriophages into WP films and glycerol were
used to develop pathogen-specific antimicrobial storage and release.
This study was performed to develop pathogen-specific antimicrobial
packaging materials using WP isolate and bacteriophage [179]. The
phage was highly stable in the whey coating [179]. One of the unique
properties of WP that makes phages stable is that it prevents the pene­
tration of oxygen, which can limit oxidative damage and stabilization of
viral particles with proteins and organic substances in phage. In addi­
tion, glycerol in the WPI-phage coating helps stabilize the viral capsid
after drying the WPI films [179–182]. The growth inhibition method by
measuring antimicrobial activity showed that phage-WPI could effec­
tively reduce the rate of E. coli growth [179].
7. The application and mechanisms of WP in the postharvest of
fruits and vegetables
Vegetables and fruits are essential for a balanced diet, vigor, and
body health with cellular detoxification and cell regeneration. In the
fruit ripening process, a series of complex reactions occur, including
R. Saberi Riseh et al.
ethylene production, an increase in photosynthesis rate, a change in
sugar content, synthesis of aromatic compounds, and many other
physiological and biochemical reactions. Following these changes,
reactive oxygen species produced by mitochondrial electron transfer can
cause cell damage and reduce the quality of fruits and vegetables
[183–185].
Nowadays, decreasing the quality of fruits and vegetables is one of
the important issues that researchers are facing, and they are looking for
a way to increase the shelf life of fruits and vegetables. Dehydration and
yellowing in vegetables due to the breakdown of chlorophyll reduce the
postharvest quality of agricultural products [186,187]. Using natural
polymers as a coating for fruits can greatly increase their quality. These
edible films and coatings reduce the diminution in quality attributes by
setting up a semipermeable safety barrier around the vegetable and
fruits [188]. WP in the coating can impart nutritional value and have
antioxidant properties [189]. The ability to synthesize antioxidants re­
duces oxidative damage caused by reactive oxygen species (ROS). Whey
is a rich food source of protein, and studies have shown that this com­
pound has free radical scavenging activity. Proteins in whey probably
act as an inducing agent that stimulates defense activities in plants,
fruits, and vegetables. Stimulating defense activities leads to greater
host resistance against plant pathogens and increases fruit and vegeta­
bles’ shelf life and marketability. WP can act as an intermediate and
biodegradable agent for the microencapsulation of fruits and vegetables
to neutralize the harmful effects of oxidative stress by increasing the
expression of defense enzymes, defense signals, and physiological
activities.
WP, a protein-rich composition with different mechanisms, increases
the quality and quantity of fruits and vegetables in the post-harvest
stages.
As explained, whey can have antibacterial and antifungal activity as
a strong antioxidant and can be effective as an effective agent in the
post-harvest processes of fruits and vegetables. Due to its cysteine-rich
structure, studies have shown that whey can cause the synthesis of
glutathione, a strong intracellular antioxidant. This antioxidant consists
of glycine, glutamate, and cysteine. The cysteine compound in gluta­
thione contains a sulfhydryl group that can act as an active reducer in
preventing oxidation and tissue damage [190]. Studies have shown that
N-acetylcysteine and glutathione as anti-browning agents can inhibit
microbial growth in freshly cut fruits. Also, an increase in the content of
vitamin C and polyphenols has been reported in freshly coated sliced
pears [191].
7.1. WP-based coating as edible coating for preserving fruits and
vegetables against phytopathogens
Since fruits and vegetables are subject to a series of complex physi­
ological processes after harvest, these reactions cause bacterial and
fungal infections on the products and reduce marketability. Therefore,
environmentally friendly polymers with potential protection against
pathogens can be used as a product coating. Due to the creation of a
semipermeable barrier against gases and moisture, edible coating on
products (fruits and vegetables) increases mechanical quality, prevent­
ing aging, maintains the structure of coated products, and subsequently
increases their storage time [192]. As an edible coating rich in nutrients,
whey can help to increase the shelf life of the taste and aroma of fruits
and vegetables and maintain their quality and moisture. Studies have
been conducted concerning the use of WP to increase the shelf life of
agricultural products. Among the advantages of edible coatings formu­
lated with different materials, the following can be mentioned: 1- direct
inhibitory effects on pathogens causing important postharvest diseases;
2- Preventing the evaporation of water, gas, and solutes; 3- increase
quality, storability, and safety of fruits [193]. The efficacy of edible
coatings to control postharvest infections in fruits may be enhanced by
incorporating specific antimicrobials [193]. The effect of WPC-mango
peel extract (MPE) formulation was evaluated on broccoli’s
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International Journal of Biological Macromolecules 234 (2023) 123708
physiochemical-microbial properties and shelf life [187]. The results
revealed that coated broccoli showed less weight loss. Preservation of
color and reduction of yellowness of florets and fungi were observed in
broccoli coated with WPC-mango peel extract (MPE). WPC alone, or in
combination with MPE, seemed to have a favorable impact on the
quality retention of minimally processed broccoli by delaying the yel­
lowing processes and changes in their weight loss, respiration rate,
bioactive compounds, and total antioxidants [187]. A study was con­
ducted to investigate the effect of WPI, lemon, and lemongrass on color,
hardness, polyphenol content, flavonoids, structure, and sensory attri­
butes in pears (storage temperature: 4 ◦ C). WPI as coating reduced
browning. The content of polyphenols and flavonoids in pears decreased
during 28 days of storage [194]. It seems that whey deactivates the
pathogens which cause softening and browning in fruits and makes the
fruit firm by preventing the penetration of the pathogen into the fruit
tissue. WPC-apple pomace extract (WPC-APE) increased the useful life of
apples. WPC-APE acted as an antimicrobial cover against pathogens. In
addition, the lowest amount of browning and moisture loss was
observed in freshly cut fruits [195].
7.2. The use of additives containing whey to enrich coatings for better
control of fruits and vegetables against plant pathogens
Polymers’ edible films or coatings act as a barrier on the surface of
fruits and vegetables and lead to higher moisture and water retention;
they create favorable micro-environments by optimizing the concen­
tration of gases and delaying ripening [185]. These edible films and
coatings are enriched with health-promoting compounds, creating a
semipermeable safety barrier. This barrier guarantees the freshness and
shelf life of the products [188,196]. Using edible coatings in combina­
tion with chemical additives, natural plant extracts, and different nano-
technological techniques increases the shelf life and quality of food
[197]. The effect of an edible coating containing WPC-rosemary essen­
tial oil (WPC-REO) was evaluated on extending the shelf life of spinach.
This formulation prevented the loss of weight, color, and pH. Further­
more maintained the chlorophyll content of the leaves. Lactoferrin is one
of the proteins in whey. This protein has these features: 1- Iron chelation
to dispossess microorganisms. 2- Destruct the lipopolysaccharides of the
outer membrane of gram-negative bacteria. On the other hand, the
interaction between essential oils and biomembranes of microorganisms
can lead to cell disruption, leakage of cell content, and cell death. In
essential oils, phenolic compounds (such as thymol and carvacrol) have
antimicrobial activities. Calcium is another beneficial factor in effective
coating formulation with whey. Calcium plays an important role in
managing different physiological procedures in plants during post and
pre-harvest treatments [198,199].
Another study evaluated the effect of the edible coating process and
irradiation treatment of Strawberry fragaria spp. on storage-keeping
quality. In this study, coating formulations were based on WP, CMC,
and glycerol combined with Calcium caseinate. CaCl2 and a mixture of
polysaccharides were also added to the caseinate–whey coating formu­
lation and compared for their efficiency in delaying the apparition of
molds in strawberries. Covered strawberries were treated with gamma-
radiation and were stored in a refrigerator [200]. Adding salt (CaCl2)
and a pectin-agar mixture enhanced the coating efficiency by delaying
mold growth on strawberries. Coating with milk-protein formulations or
the gamma-irradiation treatment effectively reduces mold infections
and thus extends the shelf life of fresh strawberries Fragaria spp. when
stored at 4 ± 1 ◦ C [200]. It seems that gamma-irradiation through cross-
links leads to increased protein accumulation. Increasing water vapor
resistance and delaying mold growth due to creating more cross-links in
WPs through disulfide bonds and hydrophobic boundaries are the ad­
vantages of using gamma-irradiation in whey formulation [200–205].
Research has proved that calcium in mixed-protein coatings containing
CaCl2 leads to electrostatic interactions between carboxylic groups
adjacent to polypeptide chains. These interactions lead to an increase in
R. Saberi Riseh et al. International Journal of Biological Macromolecules 234 (2023) 123708

Fig. 11. The use of whey protein as coating fruits against phytopathogens.

Fig. 12. Whey protein for encapsulating biocontrol bacteria to use in the packaging system.

the cohesion of hydrogels. The mutual combination of calcium ions and
gamma-irradiation increases the formation of B-tyrosine during mutual
reactions and enhances the properties of the coating formulation
[200,205]. CaCl2 could be used as a natural biological control agent for
disease control of fresh fruits and vegetables [200]. Using WP with other
activators, including essential oils and calcium, and activator methods,
such as radiation therapy, can increase the effectiveness of the coating
formulation. Fig. 11 shows the use of whey in coating different products.
In this figure, the role of whey as a mediating agent for the protection of
products and its application and the activation of defense pathways is
shown in comparison to cases with no coating.
7.3. The application of WP for food sustainable packaging against
pathogens
Food sustainable packaging means that food can be safely trans­
ported over long distances, protecting it from external factors, viral,
bacterial, and fungal contamination [39,206]. Using natural resources
polymers due to the properties of antimicrobial, antioxidant, O2 and
CO2 absorbers, CO2 emitters, humidity regulators, and flavor releases
can impact the quality of packaging and replace traditional packaging.
Due to mechanical-optical-physical properties, protein is an excellent
candidate for biodegradable films and coating production. Trans­
parency, flexibility, and resistance in proteins can be positive options for

14
R. Saberi Riseh et al. International Journal of Biological Macromolecules 234 (2023) 123708

Table 3 considering the mechanisms of nitrogen fixation, increasing plant

Other studies on the application of whey in the protection of crops and plants in growth, synthesis of primary and secondary metabolites, stimulating the
the composition of activators and bio activators. plant’s defense system, etc. Using bacteria in a polymer coating in­
Research Result Ref. creases biocontrol bacteria effectiveness and controlled release. In other
Microencapsulation conjugated High stability against oxidation (at [209]
words, microencapsulation or encapsulation of bacteria in polymer
linoleic acid (CLA) using WPC least 60 days) coatings is done to increase viability and delivery to the target areas.
Inhibiting TYLCV1 on tomato plants Preventing the virus from [210] Whey, a natural polymer that is a by-product of milk, is rich in protein
by WP (native and esterified) multiplying and amino acids. Other research has shown that WPs have been
In greenhouse conditions, the negative correlation between the [211]
considered an excellent delivery material for bacterial encapsulation
evaluation of WP against powdery severity of powdery mildew and
mildew (Podosphaera xanthii) of whey concentration, due to their high nutritional value and ability to bind to various com­
cucumber and zucchini squash. control of powdery mildew in pounds, gelation, emulsifying properties, and inhibitory effects. Whey
cucumber and zucchini can be used as an intermediary agent in the encapsulation of bacteria
Whey as replacements for synthetic Collapse and damage the hyphae of [212] and as the main agent for postharvest agricultural products preserving
fungicides and sulfur in the E. necator, free radical production,
control of powdery mildew in and control of powdery mildew.
fruits and vegetables, and increasing their shelf life. Considering the fact
Grapevine. that this compound has biocompatibility and gelling properties, it can be
The effect of WPs on the antioxidant Interaction of antimicrobial effects [213] used in agricultural programs as a biological and environmentally
and antimicrobial capacity of in the presence of WPs with friendly fertilizer, which in addition to protecting plants against biotic
different types of green tea polyphenol content
and abiotic stresses, leads to soil enrichment and increasing
Evaluation of inhibitory effects of Increase in level defense enzymes [214]
methylated α-lactalbumin, ascorbate peroxidase, microorganisms.
methylated β-lactoglobulin, dehydroascorbate reductase,
methylated lactoferrin against glutathione reductase, glutathione CRediT authorship contribution statement
tobacco mosaic virus in tobacco peroxidase, guaiacol-dependent
seedlings. peroxidases, and defense-related
enzymes phenylalanine ammonia-
Roohallah Saberi Riseh: Supervision, Project administration, Re­
lyase, and lipooxygenase. view and editing, Correspondences. Mozhgan Gholizadeh Vazvani:
The use of whey for control reduce the Severity of powdery [215] Writing-original draft, Investigation. Mohadeseh Hassanisaadi:
Grapevine powdery mildew mildew Writing-original draft, Investigation, Review & editing, Conceptualiza­
(Erysiphe necator) under field
tion, Validation. Vijay Kumar Thakur: Investigation, Validation,
conditions
Evaluation of the potential of whey reduce the severity of powdery [216] Visualization. John F. Kennedy: Review & editing, Investigation,
to reduce powdery mildew of mildew Validation, Visualization, Correspondence.
Grapevine under field conditions
The evaluation of WPI with onion The increase in growth traits, total [217]
Funding
extract powder for induction of antioxidant status, and vitamin C
systemic resistance against contents
PMMoV2 in pepper plants This research received no external funding.
The use of stable antimicrobial Oregano or clove oil in WPI had the [218]
edible films based on WPI-EO3 most inhibitory properties of
Institutional review board statement
against the different bacteria. microbial growth.
Antimicrobial activity and barrier Effective activity against S. aureus [219]
properties of WPI, glycerol, lemon, and E. coli, remarkable responses Not applicable.
and bergamot essential oils in against water vapor and oxygen
coating edible films permeability in films
Informed consent statement
1: tomato yellow leaf curl virus; 2: Pepper mild mottle virus; 3: essential oils
Not applicable.
efficient use in different industries. Proteins can also be a barrier against
external factors such as aromas and organic compounds [207,208] [39]. Declaration of competing interest
In addition to protecting various products in the pre-harvest and post­
harvest stages, whey can be effectively used as food packaging. Since The authors declare that they have no known competing financial
this study is focused on the application of whey on different agricultural interests or personal relationships that could have appeared to influence
products, providing details on this issue which is out of our discussion the work reported in this paper.
has been avoided. Fig. 12 depicts food packaging materials enriched
with plant biocontrol bacteria encapsulated into WP to extend shelf life Data availability
against pathogens. Table 3 shows other studies on the application of WP
(WPI and WPC) in the protection of crops and plants in the composition No data was used for the research described in the article.
of activators and bio activators.
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