J Americ Oil Chem Soc - 2022 - Hao - Recent Developments and Prospects in The Extraction Composition Stability Food

15589331, 2022, 8, Downloaded from https://aocs.onlinelibrary.wiley.com/doi/10.1002/aocs.12618 by New York University, Wiley Online Library on [06/03/2023].
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Received: 31 July 2021 Revised: 7 May 2022 Accepted: 21 May 2022
DOI: 10.1002/aocs.12618
REVIEW
Recent developments and prospects in the extraction,

composition, stability, food applications, and in vitro
digestion of plant oil bodies
Jia Hao | Xiaoyu Li | Qiuyu Wang | Wenwen Lv | Wenguan Zhang |

Duoxia Xu
Beijing Advanced Innovation Center for Food Abstract

Nutrition and Human Health (BTBU), School
of Food and Health, Beijing Engineering and
Oil bodies (OBs) are micron- or submicron-sized sub-organelles widely found
Technology Research Center of Food in plants seeds and nuts. The structure of OBs is composed of a core of neutral
Additives, Beijing Higher Institution lipids covered by a phospholipid-protein layer, which ensures the stability of
Engineering Research Center of Food
Additives and Ingredients, Beijing Key
OBs under extreme environmental conditions and further protects core lipids
Laboratory of Flavor Chemistry, Beijing as energy reserves. As naturally pre-emulsified oil-in-water emulsions, OBs
Laboratory for Food Quality and Safety, have been gradually applied to replace synthetically engineered oil droplets. In
Beijing Technology and Business University,
Beijing, China
this paper, the recent research on the extraction, composition, stability, food
applications, in vitro digestion, and future perspectives of plant OBs were
Correspondence reviewed. Recent studies have focused on the large-scale extraction tech-
Duoxia Xu, Beijing Technology and Business niques and OBs surface protein identification and function. Electrostatic depo-
University, No. 11, Fucheng Road, Beijing
sition and viscosity effect of polysaccharides significantly improve the stability
100048, China.
Email: xdxbtbu@126.com of OBs emulsions. OBs have broad application prospects in replacing milk fat
droplets, as fat replacers and delivery vehicles, and fabricating emulsion-gels,
Funding information oleogels or edible films. Notably, oleosins present a potential allergen risk. OBs
National Natural Science Foundation of China, are digested slowly, resulting in increased satiety and effectively helping to
Grant/Award Number: 32072216
reduce calorie intake, but this can negatively affect the bioaccessibility of nutri-
tional compounds in OBs. When OB as a delivery vehicle, its natural structure
may be disrupted, whereas OBs-based delivery system facilitates the bio-
accessibility of hydrophobic active compounds.
KEYWORDS
composition, extraction, food applications, in vitro digestion, oil bodies, stability
INTRODUCTION have isolated more than 20 kinds of seed OBs in the

range of 0.5–2.0 μm (Table 1a), including peanut,
Organisms store lipids in sub-organelles as energy sunflower and walnut with high oil content and soy-
reserves. These micron- or submicron-sized sub- bean, corn, germ and bran with low oil content
organelles mostly can be found in plants seeds and (Huang, 2018). The small size of OBs provides a
nuts called oil bodies (OBs) (Huang, 1996). Since maximal surface area per unit triacylglycerol (TAG)
1970, scientists have reported on the extraction, for lipase binding during germination (Tzen
structure, stability and application of OBs from vari- et al., 1993). OBs are extracted from plant materials
ous sources. OBs either intracellularly or in isolated using aqueous media and the difference of specific
preparations have exhibited unique physical and gravity and solubility between OBs and other compo-
chemical stabilities due to the presence of a surface nents. The isolated OBs droplets can be dispersed in
membrane composed of phospholipids (PLs) and aqueous environments to form natural emulsions. As
proteins preventing their aggregation or coalescence a naturally pre-emulsified oil-in-water emulsion, OB
and protecting core lipids from extreme environmen- can be widely applied in food production, which is
tal conditions (Nikiforidis et al., 2014). Researchers beneficial to the low energy consumption and it has
J Am Oil Chem Soc. 2022;99:635–653. wileyonlinelibrary.com/journal/aocs © 2022 AOCS. 635

15589331, 2022, 8, Downloaded from https://aocs.onlinelibrary.wiley.com/doi/10.1002/aocs.12618 by New York University, Wiley Online Library on [06/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
636 JOURNAL OF THE AMERICAN OIL CHEMISTS’ SOCIETY
T A B L E 1 Average diameters and the contents of basic composition (a), fatty acids composition (b), and PLs composition (c) of OBs isolated
from various plants (Cao et al., 2015; Dave et al., 2019; Ding et al., 2019; Furse et al., 2013; Gallier et al., 2013; Iwanaga et al., 2007; Lan
et al., 2020; Liu et al., 2022; Loman et al., 2018; Mantzouridou et al., 2019; Millichip et al., 1996; Nantiyakul et al., 2013; Niu et al., 2021; Payne
et al., 2014; Trombetta et al., 2020; Tzen et al., 1993; White et al., 2009; Zhao et al., 2016; Zhou et al., 2019)
(a) Average diameters and the contents of Basic composition

Materials Average diameter (μm) Neutral lipids (%) Proteins (%) PLs (%)
Soybean 0.25–1.75 88.51–93.72 5.21–9.72 0.80

Peanut 1.00–2.00 75.99–98.17 0.94–9.20 0.45–0.82
Rapeseed 0.65 94.21 3.46 1.97
Flaxseed 1.34–1.50 97.65 1.34 0.90
Sesame 2.00 97.37 0.59 0.57
Sunflower 2.30 96.00–98.63 1.06 0.31
Safflower 0.28–1.40 92.60 7.30 -
Almond 0.34–2.62 10.20 4.30 -
Coconut 9.30 20.10 2.50 -
Oat 1.07 - 1.40–23.00 -
Rice bran 0.50–5.80 - 1.20 -
Maize germ 0.30–1.45 - 1.43 0.91
Echium plantagineum - - 0.90–2.40 -
(b) Fatty acids composition

Materials fatty acids (%) Soybean Peanut Sunflower Almond Coconut Maize germ
Linoleic acid (18:2) 55.00 35.01 28.00–29.00 24.34 1.10 56.30

Oleic acid (18:1) 24.00 40.70 54.00–56.00 66.58 4.10 29.70
Palmitic acid (16:0) 10.00 14.08 8.00–9.00 5.83 7.30 13.00
Stearic acid (18:0) 4.00 4.21 7.00–9.00 1.62 2.70 1.50
Arachidic acid (20:0) - 1.55 - 0.07 - -
Arachidonic acid (20:1) - 0.73 - - - -
Behenic acid (22:0) - 2.42 - - - -
Tetracosanoic acid (24:0) - 1.30 - - - -
(c) PLs composition

Materials PLs (%) PC PE PI PS PA
Peanut 50.60–61.60 5.00–16.76 8.40–23.04 10.80–25.00 5.33–10.00

Rapeseed 59.90 5.90 14.00 20.20 -
Flaxseed 57.20 2.80 6.90 33.10 -
Sesame 41.20 15.80 20.90 22.10 -
Sunflower 79.00–81.00 13.00 5.00 -
Almond 31.90 6.63 15.80 - 39.30
Maize germ 64.10 8.10 7.60 20.20 -
Echium plantagineum 52.40–55.10 11.70–13.20 4.00–5.20 30.70–32.60 -
excellent stability and emulsifying performance discussed (Figure 1). The paper reported the future
(Nikiforidis, 2019). research direction and development potential of
This paper reviewed the research advances of OBs natural OBs.
derived from plants. It provided a general description of
OBs composition, structure and the methods to main-
tain the stability of OBs emulsions. The technologies Oil bodies extraction
for large-scale and complete extraction of OBs were
highlighted. In addition, the food application and in vitro The microstructure of plant cells was observed, and
digestion behavior based on OBs emulsions were different sizes of abundant natural OBs were found
JOURNAL OF THE AMERICAN OIL CHEMISTS’ SOCIETY 637
F I G U R E 1 An overview of OBs
including extraction, composition and
structure, stability, food applications,
in vitro digestion, and future
perspectives. OBs, oil bodies
F I G U R E 2 Microstructure of plants cells: TEM images of rice bran (a) (unpublished data), Cryo-SEM images of hazelnut after soaking for 48
h (b), and CLSM images of peanut (c). OBs, oil bodies; PBs, protein bodies; CW, cell wall. Source: (b): Reproduced with permission from
Capuano et al. (2018) and copyright Royal Society of Chemistry. (c): Reproduced with permission from Zaaboul et al. (2018) and copyright John
Wiley and Sons
(Figure 2). Starches and proteins were distributed Aqueous extraction

around the OBs, and each region was separated by the
cell wall to maintain the integrity of the OBs. Therefore, The aqueous extraction of OBs requires soaking seeds
destruction of cell walls is the key to extracting natural in an aqueous medium, then blending or pressing to
OBs. Due to the hydrophilic surface of OB, it can be disrupt cell walls and release intracellular materials.
dispersed into an aqueous phase to form a uniform Urea, sucrose, deionized water, salt, alkali and buffer
emulsion (Nikiforidis, 2019). Based on this characteris- solution (including Tris–HCl and PBS) are often used
tic, the isolation process of OBs can be divided into four as the grinding medium for aqueous extraction. In
steps: (1) the structure of plant cell wall is destroyed by 1992, 0.6 M sucrose solution was first used to extract
mechanical crushing, and the OB is separated from plant OBs including maize, rice, soybean, rapeseed,
protein body and starch grain to release in an aqueous jojoba, wheat bran, flax, sesame, tripsacum, teosinte,
medium; (2) the slurry is filtered to produce a “milk”, yucca, mustard, jojoba, sunflower, peanut, palm, castor
which is a suspension of OBs in an aqueous medium bean, Brazil nut and oat bran, the grinding medium also
with a small number of seed particles; (3) OB is con- contained 1 mM EDTA, 10 mM KCl, 1 mM MgCl2, 2 mM
centrated into cream by centrifugation of the “milk” to DTT, and 0.15 M Tricine buffer (Tzen et al., 1992,
obtain crude OB; (4) the crude OB is washed to remove 1993). Since 1996, the extraction medium was simpli-
the impurities on the surface and obtain pure OBs fied (Chuang et al., 1996; Tzen et al., 1997). Lacey
(Nikiforidis et al., 2014) (Figure 3). Current extraction et al. (1998) found that OBs extracted by 9 M urea were
methods of OBs mainly include aqueous extraction and not affected by exogenous proteins, and integral
enzyme assisted extraction based on the different prin- oleosin (the dominant OBs surface proteins) could be
ciples of disrupting the cell wall. completely preserved. Urea can disrupt non-covalent
F I G U R E 3 Process involved in
the extraction of OBs using aqueous
extraction and enzyme assisted
extraction and impact factors (a), OBs
extraction schematic diagram (b), and
extraction process of safflower OBs
(c). OBs, oil bodies; PBs, protein
bodies. Source: (c): Reproduced with
permission from Lan et al. (2020) and
copyright Elsevier
bonds of proteins allowing the removal of passively Enzyme assisted extraction

associated proteins from the OBs surface (Millichip
et al., 1996). However, urea is not a food grade mate- Plant cell walls are composed of cellulose, hemicellu-
rial, thus limiting its use. At present, deionized water loses, lignin and pectin. Thus, in addition to mechanical
and buffer solution are often selected as extraction methods, the use of carbohydrases such as cellulase,
media (Iwanaga et al., 2007; Lan et al., 2020; hemicellulose, pectinase, xylanase, and β-glucanase
Nikiforidis & Kiosseoglou, 2009), while, OBs still con- may also destroy the cell wall. The high specificity of
tain some intracellular substances such as exogenous enzymes greatly limits the degree of hydrolysis due to
proteins and phytochemicals, and endogenous proteins the diversity of cell wall components and raw materials.
are also vulnerable to destruction. Therefore, De In this case, the combination of multiple single carbohy-
Chirico et al. (2018) pointed out that the purity of rape- drase can be considered. Protease is not considered in
seed OBs extracted with sodium bicarbonate solution the extraction process. Although the recovery of OBs
(0.1 M) was the same as that preparation with 9 M urea, can be improved, the associated proteins may be des-
and the physical stability of OBs was improved. The troyed into small peptides through hydrolysis. Kapchie
result provides a new method for the aqueous extrac- et al. (2008) compared the efficiency by mixing Multifect
tion of intact and pure OBs. Pectinase FE (Pectinase, cellulase, and hemicellulase
complex), Cellulase A (Cellulase, β-glucanase, Pretreatment

hemicellulase, and xylanase complex) and Multifect CX
13 L (β-glucanase complex) extraction and aqueous The pretreatment of plant materials usually involves
extraction of soybean OBs and found that the total soy- grinding and soaking. Grinding provides better expo-
bean oil recovered from OBs extracted by the enzy- sure of OBs to the water as a result of cells structure
matic method was 36.42%–63.61% and the yield of rupture, and soaking allows water molecules to pene-
soybean oil in OBs could reach 84.65% after three suc- trate into the network of cells for more efficient extrac-
cessive extractions of the residue. However, the total tion. For seed raw materials, mechanical grinding is
soybean oil recovered from OBs was only 28.65%– usually carried out in the extraction medium after
34.28% by a conventional method. Xu et al. (2021) soaking for 8–72 h. For processing by-products such as
studied the structure, physical properties and chemical wheat bran and rice bran, after sieving and removing
compositions of rice bran OBs extracted by plant impurities can be soaked or not for a short time,
exacted enzyme, xylanase and their mixture, and followed by mixed with the extraction medium for
results showed that the yield of OBs obtained by the homogenous extraction of OBs (Lan et al., 2020;
mixture of xylanase and plant extracted enzyme was Nantiyakul et al., 2013; Tzen & Huang, 1992; White
76.95%. Extracting OBs from plant raw materials is a et al., 2006).
relatively new research field. It is necessary to invest in At present, pretreatment methods such as mechani-
developing special equipment and consider the actual cal crushing, extrusion, high pressure, and ultrasound
production conditions to achieve industrial production. have been successfully used for aqueous oil extraction.
Towa et al. (2011) conducted a pilot scale equipment It is worth noting that these pretreatment methods have
by which the yield of OBs isolated from soybean was the potential to improve the OBs extraction yield
93.40% compared to a laboratory-scale process. (Koubaa et al., 2016; Mat Yusoff et al., 2015). The
crushing degree of raw materials directly affects the
extraction yield of OBs. The size of raw material cells
Comparison of aqueous extraction and should be considered during mechanical crushing.
enzyme assisted extraction When the particle size of the material is smaller than
the cell size, the cell wall can be destroyed and the con-
The selection of OBs extraction method mainly tact area between the OBs cells and the extraction
depends on the extraction yield, complexity, produc- medium be enlarged. While excessive crushing may
tion cost, environmental friendliness, and safety. destroy the complete structure of the OBs. Nikiforidis
Aqueous extraction is a traditional method widely and Kiosseoglou (2009) showed that the extraction
applied for extraction of OBs, which has the advan- yield of OBs was the highest at any pH and number of
tages of no chemical pollution and low-energy con- extraction steps applied when the corn particle size
sumption, however, there are certain limitations that it was less than 0.8 mm. However, there was a possibility
requires large amounts of solvents, and thus makes that a single OB merged into a larger droplet due to the
up-scaling difficult. Although the enzymatic extraction destructive of mechanical force.
of OBs requires a long reaction time, and the enzyme The extrusion process increases the pressure gen-
has strict requirements on temperature and pH, the erated by compression to exceed the maximum shear
extraction efficiency of OBs is higher. There are major stress that the plant cell wall can withstand, causing the
problems that must be solved to develop special cell wall to rupture (Li et al., 2020; Peng et al., 2021). It
enzymes, reduce the dosage and cost of enzymes in is not conducive to the solvent percolation for the fine
the industrialization of enzymatic extraction of OBs powder raw materials. Extrusion can also reshape the
technology. Furthermore, the extraction procedures material into porous expanded particles, which
require simple equipment that can be operated safely increases percolation rate of the solvent and hence bet-
to facilitate industrial production and commercial appli- ter efficiency of OBs extraction (Liu et al., 2020).
cation of OBs. Romero-Guzma n et al. (2020) used a twin-screw press
to extract OBs at pH 7.0, and showed that the yield
could reach 90%. Twin-screw press is a promising
THE MAIN INFLUENCING FACTORS OF alternative to scale up the aqueous extraction of OBs
EXTRACTION PROCESS and the water usage is significantly reduced.
It has been reported that the high pressure-
In order to further improve the OBs yield, reduce the processing transmitted isostatic pressure (100–400
cost and obtain high quality and purity OBs, the key MPa) to plant materials can be useful for destroying
parameters in the extraction process can be controlled. cells and increasing the solubility of bioactive com-
Generally, there are many factors affecting the extrac- pounds (Bueno et al., 2020; Ninčevic Grassino
tion procedures including pretreatment, pH, purity, et al., 2020). However, the structure of the product is
material to liquid ratio and centrifugal force. changed at a pressure of around 600 MPa, resulting in
a decrease in extractability (Butz et al., 2002; Dobrinčic pH 8.0 and 2S albumins were removed at pH 11.0. At pH
et al., 2020). Kapchie et al. (2008) demonstrated that 11.0, highly purified OBs were obtained from jicama, sun-
the highest OBs yield of 71.39% was obtained with the flower, peanut, and sesame, whereas there were still
ultrahigh pressurizations of soybeans flours at 200 MPa enzymes remained in the castor bean and rapeseed
for 5 min at 25 C, while, the OBs yield was 21.82% with OBs. Therefore, the extraction pH can be selected
material pressurized at 500 MPa. according to the properties of OBs. High protein-
Ultrasound induced impacts that can be attributed containing OBs can be extracted at pH close to neutral,
to the cavitation phenomenon referring to bubble forma- while, highly purified OBs can be obtained at high-alkaline
tion, growth and implosion during the propagation of pH. It is clear that as the extraction pH increases, the
the ultrasonic wave into medium. The bubble implosion exogenous protein is gradually reduced, however, it is not
creates a hot spot with a temperature of up to 5000 K sure whether oleosins can also be removed by alkaline
and a pressure of 5000 atm (Khadhraoui et al., 2021). pH (Chen & Ono, 2010). Cao et al. (2015) confirmed that
This may result in effects on cell walls, enhancing the alkaline pH not only removed contaminated proteins but
penetration of the solvent into the internal structure and also oleosins, and more and more oleosins were
facilitating the release of the target compound. In addi- removed with increasing alkaline pH.
tion, the strong shear forces and turbulence generated The content and types of bioactive components in
by the propagation of the ultrasonic waves appear to the OBs may be affected by pH. Zaaboul et al. (2018)
further accelerate the exchange between the raw mate- first investigated the main compounds of peanut OBs
rial and extraction solvent (Chemat et al., 2017; Soria & and found that δ-tocopherol content increased when pH
Villamiel, 2010). Loman et al. (2018) found that recovery got increased, conversely, α-tocopherol con-
1.5 W/ml pulsed ultrasound treatment for 5 min every tent decreased. This was because α-tocopherol bound
3 h could significantly improve the performance and to PL-protein membrane, whereas the δ-tocopherol
separation of OBs from protein during the enzyme was more bonded with the TAG matrix, so it was more
processing. It was studied that ultrasonic pretreatment resistant to protein removal with alkaline pH. However,
would increase the OBs yield, but the yield showed a Chen et al. (2014) showed that the pH did not affect the
downward trend after a certain time, because the ultra- tocopherol content and isoforms profile in
sonic energy was not uniformly distributed in the pre- soybean OBs.
treatment process (Albu et al., 2004; Kapchie Research showed that pH also affected the rheolog-
et al., 2008; Toma et al., 2001). ical properties of soybean OBs, emulsions with 40%
OBs content (pH 6.8–9.5) could form gels, for solid-type
OBs products, pH 9.5 should be selected by consider-
pH ing the influence of thermal treatment on shelf life; for
liquid-type OBs products, pH 11.0 should be selected
pH is an important parameter in the OBs extraction pro- (Zhao et al., 2016).
cess. The isoelectric point of the OBs is between 4.0
and 6.0 due to the existence of proteins on the surface
(Tzen et al., 1993; Wang et al., 2019). Therefore, it is Purified OBs
not considered that the extraction pH is lower than 6.0.
At this point, the natural and independent OBs in the It is necessary to determine whether pure OBs or a mix-
cells may aggregate after separation. The pH value of ture of OBs and seed storage proteins is needed when
the medium when extracting OBs is usually between extracting OBs. For some applications, especially
6.5 and 11.0. In this pH range, the surface of OBs foods, such as salad dressings, where proteins have
maintains a negative charge, and electrostatic repul- been used to modulate the macro properties of the sys-
sion and steric hindrance effect exist in the OBs, which tem, so mixtures containing OBs and storage proteins
allow OBs distribute independently and maintain natu- are beneficial (Nikiforidis et al., 2012; Karefyllakis
ral structure. et al., 2019). However, in other applications, pure OBs
It is known that the pH value of the extraction buffer may be required. To obtain pure OBs, several cleaning
not only has a direct influence on the natural structure steps are required including urea washing, sucrose
of OBs, but also has a great influence on the protein washing, deionized water washing, salt washing, alkali
composition of the extracted OBs, which affects the washing and buffer solution (including Tris–HCl and
OBs properties, and then impacts the OBs utilization. PBS) washing. Generally, the washing medium is simi-
Zhao et al. (2016) studied jicama, sunflower, peanut, lar to the extraction medium, while the medium used in
castor bean, rapeseed, and sesame to explore the extracting crude OBs and obtaining pure OBs can be
effects of pH (6.5–11.0) on protein compositions of different. It was reported that OBs washed by 9 M urea
OBs. The results showed that there were many extrin- were significantly enriched in lipids and low in proteins
sic proteins (globulins and 2S albumins) presented in compared with unwashed, water-washed, and salt-
pH 6.5-extracted OBs. Globulins were mostly removed at washed OBs and washing significantly reduced the
total phenolic content of the oat OBs but significantly storage form, a small amount of diglycerides, free fatty
increased concentrations of vitamin E (White acids, and Vitamin E, etc (Abdullah & Zhang, 2020).
et al., 2006). For soybean OBs from different varieties, The neutral lipids of the OBs account for about 92%–
the contents of vitamin E and total phenolics were 98%, the contents of PLs and proteins are about 0.6%–
decreased by urea washing (Fisk & Gray, 2011). Mur- 2.0% and 0.6%–3.0%, respectively (Table 1a) (Ding
phy and Cummins (1989) explored the influence of et al., 2019; Tzen et al., 1993).
washing times on OBs and found that the composition
of OBs was not affected by washing times, but multiple
washing times could greatly reduce the final production Neutral lipids
of OBs. The OBs endogenous protein was tightly
bound to the surface of OBs and could not be removed The lipid composition of OBs was examined by chro-
by washing and pure OBs fraction could be obtained by matography and the content of TAGs was found to be
two floatation steps at most. the highest (Abdullah & Zhang, 2020). Linoleic acid,
oleic acid, palmitic acid, and stearic acid are the main
fatty acids in soybean OBs, peanut OBs, sunflower
Other influencing factors OBs, almond OBs, coconut OBs, and maize germ OBs
(Table 1b). TAGs cannot only be as energy reserves
The key factors affecting OBs extraction also include for germination and post germinative growth of the
material to liquid ratio, centrifugal force, incubation time, seedlings, but also are important for cell division and
temperature, stirring speed. The viscosity of the system expansion, membrane lipid remodeling and organ for-
is determined by the ratio of the raw material to extrac- mation (Yang & Benning, 2017). If the seeds have been
tion medium. When the viscosity is too low, it may result stored for a long time, the hydrolysis of TAGs and PLs
in less force borne by OBs during the recovery process by internal or external lipase or nonspecific acyl hydro-
and reduce the collision and damage of OBs particles. lases may lead to the increase of minor lipids, espe-
However, low-water content and high viscosity of the cially free fatty acids, causing rancidity and quality
system result in low yield of the OBs. It had been con- degradation (Huang, 1992).
firmed that the OB particle size distribution presented a
wide distribution at a high solid-phase loading ratio, and
droplet aggregation was observed in the optical image, Phospholipids
while, in a more diluted seed grinding system, the OB
was smaller and integrity (De Chirico et al., 2018). The The electron microscope results showed that the
centrifugal force can affect the size distribution of the core of OBs was an electronically opaque TAGs
OBs. Zhang et al. (2017) studied the impact of centrifu- matrix surrounded by one electron-dense layer
gal forces ranging from 5000 RCF to 20,000 RCF on (Figure 4b). This single electron-dense layer was
OBs yield and found that small OBs could be emulsified determined a “half-unit” PLs membrane, in which the
by low-centrifugal force, while OBs trended to coalesce hydrophobic tail faced inward to interact with the
by high-centrifugal force. Some researchers modify the TAGs, and the hydrophilic head was exposed to the
traditional extraction method by giving the raw materials cytoplasm (Tzen & Huang, 1992). Typically, PLs
a certain incubation time and temperature under a stir- accounted for more than 80% of the surface area of
ring speed (Nantiyakul et al., 2012; Niu et al., 2021). the OBs, and the thickness is 0.9–2.5 nm (Purkrtova
These operations affect proteins denaturation, OBs dis- et al., 2008; Yang et al., 2020). The dense crystalline
persion and aggregation, and extraction yield. For enzy- phase of the PL membrane results in a more thermo-
matic extraction, the amount of added enzyme is also an dynamic stability structure than other artificial emul-
important factor. Generally, there are many factors sions (Zhang et al., 2022). It was determined by TLC
affecting the yield of OBs during the extraction process, that the major PLs in Echium plantagineum OBs
while systematic and statistical optimization of the signif- were phosphatidylcholine (PC; 52.4%), followed by
icant factors has not been attempted to obtain the opti- phosphatidylserine (PS; 32.6%), phosphatidyletha-
mal extraction conditions and the ideal yields. nolamine (PE; 12.3%) and phosphatidylinositol (PI;
4.2%) (Payne et al., 2014). 31P NMR measured the
highest content of PC (50.6%) in peanut OBs,
COMPOSITION AND STRUCTURE OF OBs followed by PI (13.6%), PE (12.7%), PS (10.8%) and
phosphatidic acid (PA; 10.0%) (Niu et al., 2021). PC
The composition of the OBs conforms to a formula has a strong hydrophobic interaction with the OBs
describing a spherical particle surrounded by a shell of membrane proteins, therefore, the OBs are much
a monolayer of PLs embedded with proteins more stable. Tzen et al. (1993) analyzed the PLs in
(Figure 4a,c). The spherical particle is a neutral lipid the OBs from seven kinds of oilseeds found that they
core consisting of TAGs as a rich source of oil in contained substantial amounts of the uncommon
F I G U R E 4 The simulation
diagram of OBs structure (a), electron
micrograph of monolayer PLs
membrane of OBs isolated from
peanuts (b), and microstructure of
OBs in rice bran (c) (unpublished
data). (c): CLSM images (A–D),
phospholipid membrane is stained
with Rd-DHPE (purple, A); triglyceride
core is stained with Nile red (green, B);
proteins are stained with fast green
FCF (red, C); combination of
triglyceride core and proteins is shown
in (D), Cryo-SEM image (E) and TEM
image (F). Source: (b): Reproduced
with permission from Yatsu and Jacks
(1972) and copyright Oxford University
Press.
negatively charged PS and PI (Table 1c). It has been in maintaining OBs stability, since proteins provide not
proposed that these negatively charged PLs can only amphiphilic surfaces but also steric hindrances. At
interact with the basic amino acid residues of the present, the research mainly focuses on the identifica-
oleosins on the surface of the PLs layer. The fatty tion and function of OB surface proteins.
acid composition of the PLs in the OBs is determined
to be high in saturated fatty acids, which contributed
to the anchorage of the oleosins and thus to the sta- Identification of OB surface proteins
bility of the OBs (Furse et al., 2013; Katavic
et al., 2006; Payne et al., 2014). The dominant OBs-associated proteins are called
oleosins (Figure 5a). Oleosin has a low-molecular mass
of 15–26 kDa with three structural regions. Two amphi-
Proteins philic N- and C-terminal regions moieties locate at the
surface of the OBs, with the positively charged residues
It was studied that OBs isolated from plants maintained interacting with the negative charge of the phosphate
their discreteness, but coalescence was observed after molecule towards the lumen of the organelle. The nega-
only 1 min of trypsin treatment, OBs could be seen as tively charged residues facing the cytosol make the OBs
much larger particles after 6 h of treatment. The enzy- surface negatively charged and generate electrostatic
matic activity of trypsin on the oleosins on the surface repulsion to maintain the stability of the OBs, also can
of the OBs resulted in the production of smaller mainly prevent external phospholipase from acting on the PLs
of 8 kD, such polypeptides represented the central membrane (Napier et al., 1996). A central hydrophobic
hydrophobic domain of the oleosins which would not be region of about 70 residues formed by two approxi-
accessible to the external trypsin. Phospholipase A2 or mately 11 nm long antiparallel strands connected by a
C exerted its activity on OBs only after the exposed “proline knot” is inserted in the acyl moieties of PLs and
portion of oleosins had been removed by trypsin (Tzen the TAGs matrix and forms a hairpin-like structure
et al., 1992). This confirms the importance of proteins (Napier et al., 1996; Tzen et al., 1992).
F I G U R E 5 The basic structures of three major surface proteins on the plant OBs: oleosin (a), caleosin (b) and steroleosin (c).Source: (a):
Reproduced with permission from Yang et al. (2020) and copyright Elsevier. (b): Reproduced with permission from Frandsen et al. (2001) and
copyright John Wiley and Sons. (c): Reproduced with permission from Lin et al. (2002) and copyright Oxford university press
Caleosin is another group of protein on the OBs sur- storage proteins such as gij31432342, gij24899397,
face, with a molecular mass of 25–35 kDa (Figure 5b). and gij12039336 are also identified in rice bran OBs
It has the same molecular structure as oleosin, includ- (Xu et al., 2021).
ing a hydrophilic N-terminal group, a hydrophilic C-
terminal region, and a central hydrophobic region
inserted into the core of TAGs (Næsted et al., 2000). Functions of oleosin
Caleosin is characterized by a more hydrophilic N-
terminal segment as it contains a Ca2+ binding region Oleosins are thought to be the most important inter-
(Chen et al., 1999). It has been reported that caleosin face protein, covering the whole surface of the OBs.
may be involved in OBs fusion, membrane-fission There are some researches on the function of
and/or fusion and lipid intracellular trafficking and oleosin. The biological function of oleosin in stabiliz-
metabolism (Frandsen et al., 2001; Næsted ing OBs makes it an excellent emulsifier for the pro-
et al., 2000). duction of food and cosmetics. Nikiforidis et al.
Steroleosin is a comparatively bigger protein with (2013) used interfacial rheometer to study the foam
molecular mass more than 35 kDa (Figure 5c). It was stabilization capacity of oleosin extracted from maize
first identified as a minor protein in sesame OBs, con- germ at the air–water interface and found that
taining a hydrophobic anchoring segment at the same oleosin formed a strong elastic film at the air–water
length as caleosin, followed by a sterol-binding dehy- interface, which implied that oleosin could be a good
drogenase domain, which could anchor the soluble ste- foaming agent. However, oleosin has strong hydro-
rol binding dehydrogenase domain to the surface of the phobicity and low-extraction rate limited its applica-
OBs through the N-terminal hydrophobic fragment (Lin tion (Rayner, 2015). At present, the methods used in
et al., 2002). oleosin extraction include precipitation with organic
Tnani et al. (2011) identified new proteins in maize solvent, PCR technology, E. coli production and
embryos OBs besides oleosin, caleosin and ultrasound (Sun et al., 2022).
steroleosin, such as karyopherin-beta and a stress- A novel technology named oleosin-fusion technol-
induced membrane pore protein were involved in mem- ogy has been developed in which exogenous pro-
brane transport. Jolivet et al. (2004) found that oleosins teins are fused to the N- or C-terminus of oleosin in
accounted for 79% of OBs proteins in Arabidopsis OBs surface and expressed under the control of an
thaliana ecotype WS, and the 18.5 kDa oleosin was the oleosin gene promoter or seed-specific promoter for
most abundant. Meanwhile, they found a probable the production of recombinant proteins (Chiang
aquaporin and a glycosylphosphatidylinositol-anchored et al., 2005; Yang et al., 2018). There have been
protein, which had never been found in plant OBs, with- many reports on oleosin-fusion technology being
out known functions. Plant nsLTP, a soluble protein used with recombinant proteins and some fusion pro-
with a molecular mass of less than 20 kDa, presents in teins have reached commercial production standards
rice bran OBs, which contributes to the transfer of fatty (Capuano et al., 2007; Yang et al., 2015). For exam-
acids, PLs, glycolipids and steroids between mem- ple, recombinant human precursor insulin or human
branes, and plays a key role in the process of plant epidermal growth factor (HEGF) fused with oleosin
resistance to pathogens. Embryo-specific protein has been successfully expressed in Arabidopsis
belonging to plant antimicrobial peptides family and (Bhatla et al., 2010).
Bioactive components polysaccharide and the system changes from positive

to negative. Soybean OBs suspension containing 1 wt
OBs may contain a small amount of bioactive compo- % soybean OBs and 0.06 wt% beet pectin was pre-
nents that are contribute to their chemical stability, such pared at pH 4.5, and laccase was added to promote
as tocopherols, phytosterols, essential amino acids, crosslinking and electrostatic deposition of beet pectin
and isoflavones (Gallier et al., 2012; White et al., 2006; on OBs surface, which significantly improved soybean
Zaaboul et al., 2018). OBs from different sources may OBs stability at pH ranging from 3.0 to 7.0, NaCl addi-
contain special bioactive components that give them tion (0–500 mM), and freeze–thaw cycle (20 C for 22
unique functions, such as soybean isoflavones in soy- h; 40 C for 2 h) (Chen et al., 2010). It had been shown
bean OBs, and oryzanol in rice bran OBs have antioxi- that carrageenan (0.05–0.5 wt%) could be adsorbed
dant properties (Fisk & Gray, 2011; Nantiyakul stably on the surface of soybean OBs emulsions in an
et al., 2012). acidic environment (pH 3–5) (Wu et al., 2011;
Wu et al., 2012).
The stability of emulsion depends on the interaction
Stability of OBs of dispersed particles, and the collision of particles cau-
sed by Brownian motion, which is affected by the vis-
As a natural emulsion system, OBs can deliver stable cosity of solution (Lan et al., 2020). According to
pre-emulsified oil into appropriate food systems to Stokes formula, the increase of viscosity can increase
obtain products. It can reduce the need to extract and the drag force exerted on the droplets and reduce their
purify the oil using organic solvents and then emulsify it creaming rates in the emulsion (Zhao, 2008). Zhang
using a homogenizer, thereby achieving more sustain- et al. (2019) found that low concentrations of sodium
able and environmentally friendly processing opera- alginate (0.30–0.45 wt%) could stabilize the 1 wt% pea-
tions. The utilization of OBs in food products requires a nut, sesame, and rapeseed OBs emulsions below the
thorough understanding of their functional properties isoelectric point of the OBs through electrostatic inter-
under complex environmental conditions. During food action. While high concentrations of sodium alginate
processing, storage, transport and utilization, the pH (1.2–1.5 wt%) improved the OBs emulsions stability
value of the system may change. For taste, preserva- through the viscosity effect at pH 7.0. In a few cases,
tion and modification of physicochemical properties, complexation may occur even if the protein and poly-
salt is added to the food. Many emulsion-based food saccharide carry the same negative charge. The
products may undergo various kinds of thermal treat- reports ascribe this phenomenon to the existence of
ments such as pasteurization, sterilization, temperature small “charge patches” onto the protein, which may
fluctuations, baking, cooking, freezing and thawing pro- carry the local positive charges to bind with the anionic
cess. Studies have shown that the rheology and stabil- polysaccharide (Yang et al., 2021).
ity of OBs suspensions are susceptible to the effects of Ding et al. (2019) successfully and effectively encap-
pH and salt concentration, resulting in reduced electro- sulated soybean OBs by using maltodextrin (MD)–
static repulsion between OBs and instability such as chitosan (CS)–Epigallocatechin-3-gallate (EGCG) cova-
flocculation (White et al., 2008). Therefore, to utilize lent conjugates (CSEG) as coating material and apply-
OBs commercially in food systems, the stability of OBs ing spray drying technology, and the oxidative stability
must be enhanced to adjust to environmental changes and thermal stability of OBs microparticles were
in terms of pH, ionic strength and thermal treatment. improved and significantly reduced the amount of oil
It has been noted that OBs emulsions rich in exoge- released during the whole digestion process. It demon-
nous proteins have much higher stabilities against coa- strates that this novel spray drying encapsulating OBs
lescence after long-term aging than purified OBs technology has the potential to overcome stability
emulsions (Nikiforidis & Kiosseoglou, 2010). Many defects.
researchers have attempted to enhance the stability of
OBs by adding polysaccharides to OBs emulsions and
controlling the pH to produce electrostatic deposition FOOD APPLICATIONS OF OBs
and steric repulsion effects (Iwanaga et al., 2008).
Anionic polysaccharides are the most selected, mainly OBs replace milk fat droplets
including pectin, carrageenan, xanthan gum, and gum
arabic. The surface of the OBs has a negative charge OBs in plant-based milks
because of the existence of interfacial protein. There-
fore, after adding anionic polysaccharides into the OB Consumer demand for cow’s milk alternatives has
emulsions, it is necessary to adjust the pH less than increased as a result of lactose intolerance of some
the isoelectric point of OBs (pH 3.0–4.0), so that the people and the demand for vegetarianism and health
surface of the OBs is positively charged, resulting in (Aydar et al., 2020). Plant-based milks are often per-
electrostatic interaction between the OBs and anionic ceived as healthy possibly due to they contain dietary
fiber, vitamins, minerals, and antioxidants (Jeske et al., 2016; Selani et al., 2016). However, the direct
et al., 2018). Generally, plant-based milk is an emulsi- incorporation of plant-derived oil into meat product for-
fied system composed of OBs, solid particles and mulations may lead to a soft and sticky texture and
starch granules all dispersed in an aqueous phase, reduce stability against oxidation (Bolumar et al., 2015;
where the milk fat droplets are replaced by OBs and Nacak et al., 2021). Studies had shown that the alter-
the presence of water-soluble proteins on the surface ations in meat products could be alleviated by utilizing
of the OBs improves the texture, stability and nutrition oil in their pre-emulsified form (Serdarog lu et al., 2016;
of plant-based milks (Huang, 2018; Zaaboul Utama et al., 2019).
et al., 2019). Ultrafiltration is commonly used to pro- OB has been positioned as a “clean label” ingredi-
duce an OBs-based emulsion with a lipid content simi- ent as a natural pre-emulsified oil-in-water emulsion
lar to cow milk and rich in polyunsaturated and (Asioli et al., 2017). In addition, it has good thermal and
monounsaturated fatty acids to obtain plant-based oxidative stability, which may help inhibit lipid oxidation
milks and the plant-based milks can maintain its aroma in meat products (Sukhotu et al., 2014; Zaaboul
characteristics and bioactive constituents upon heating et al., 2018). These advantages make it as an alterna-
and exhibit appreciable microbial, physical and oxidative to animal fat. Bibat et al. (2022) found that
tive storage stabilities (Naziri et al., 2017). replacing pork fat with rapeseed OBs could produce a
Shakerardekani et al. (2013) prepared pistachio milk by softer, less gummy and less chewy meat system, and
using pistachio OB as a natural pre-emulsified emul- improved the emulsification and oxidation stability of
sion and 5.0% sugar, 0.02% vanilla. meat paste.
OBs in yogurt OBs-based delivery system

Fat globules play an important role in the production of Emulsions-based delivery system in food industry is
yogurt. The acidification of milk induces the formation widely recognized to be a promising method of encap-
of continuous gel networks of aggregated protein mole- sulating and delivering bioactive compounds in product
cules where fat globules are incorporated manufacturing and storage processes for preventing
(Horne, 1999). The strength of the gel depends on the chemical degradation and increasing bioavailability
size of the fat globules and the extent of their surface (McClements, 2015). Due to the unique PL-protein
interactions with the gel network (Kirimlidou membrane structure and ability of the hydrophobic core
et al., 2017). The physical properties and microstruc- to solubilize non-polar bioactive agents, OBs can be
ture of the gel are reflected in the texture and sensory used as a new carrier for transporting bioactive compo-
characteristics of yogurt (Öztürk et al., 2018). Man- nents (Zheng et al., 2019). The challenge, however, is
tzouridou et al. (2019) replaced the milk fat globules in to load the hydrophobic bioactives into pre-
yogurt with maize germ OBs, which had a good acidifi- existing OBs.
cation kinetic pattern, forming a liquid structure acidic Studies have confirmed that soybean OB are natu-
gel and the rheology and texture are similar to yogurt ral and environment-friendly emulsifying agents, which
prepared with whole cow milk. The milk fat globules in can be used as substitutes for emulsifiers such as low-
yogurt prepared from whole cow milk as fillers of the molecular-weight one and animal proteins (Ishii
milk protein matrix aiding in the formation of the gel et al., 2017). A new type of oil-in-water emulsion loaded
structure. For yogurt-like products, small-sized OBs with hydrophobic active substances can be formed by
acted as fillers and might interact with the milk protein applying the traditional homogeneous emulsification
network by skim milk incorporation and thus as rein- method using OBs as the aqueous phase and dispers-
forcing material of the gel structure (Matsakidou ing the hydrophobic active substances into external oil
et al., 2013). phase. As shown in Figure 6, there are three possibili-
ties about the behavior of OBs (sunflower OBs as an
example) at the oil–water interface during emulsifica-
OBs as fat replacers in meat products tion: (a) the oil–water interface is completely stabilized
by storage proteins; (b) large-sized OBs (>1–5 μm)
Excessive intake of processed meat products has decomposition, PL-protein membrane recombination
potential health risks as they commonly contain a con- stabilization interface; (c) small-sized OBs (<1 μm) and
siderable amount of animal fat, which is known to be storage protein stabilize the interface together. In this
high in saturated fat and cholesterol (Zhu et al., 2020). case, OBs structure is intact and the interface stability
Substituting plant-derived oil for animal fat has been mechanism is similar to the Pickering emulsion
addressed as a feasible strategy to achieve healthier (Karefyllakis et al., 2019). Pickering emulsion is a
meat products by increasing the ratio of monounsatu- mechanism of stabilization of a membrane interface by
rated fatty acid (MUFA) and polyunsaturated fatty acid solid particles that do not deform upon absorption like
(PUFA) to saturated fatty acid (SFA) (Alejandre proteins do. As it can be seen in Figure 6c, these small-
F I G U R E 6 Possible behavior of
OBs at the oil–water interface:
Storage protein plays a dominant role
(a), large size OB decomposition, PL-
protein membrane recombination
(b) and small size OB has an intact
structure together with storage protein
to stabilize the oil–water interface (c).
sized OBs particles seem to widely absorb at the O/W most recently used as the matrix (Bi et al., 2021;
interface without rupturing, and maintain their spherical Dickinson, 2012; Fontes-Candia et al., 2020; Hu
shape and act as Pickering stabilizers. Obviously, the et al., 2021; Li et al., 2020; Saavedra Isusi et al., 2020).
mechanism based on OBs as bioactive substance Emulsion-gels have broad application prospects in food
delivery vehicles obtained by conventional emulsifica- industry such as fat reduction, probiotics release and fla-
tion methods is complex and requires in-depth analysis. vor control due to their diversity in structure and compo-
It is well known that PLs and proteins can be used as sition (Lin et al., 2020). OBs can replace the traditional
natural emulsifiers. Previous researchers have loaded oil/fat droplets in the gel due to their nutritive value and
bioactive components and pharmaceutical ingredients natural emulsification properties (Nikiforidis et al., 2014).
(such as DHA and curcumin) into reconstituted OBs However, to the best of our knowledge, there are very
formed by homogenization of an oil and aqueous phase few studies on the emulsion-gels filled with OBs.
together in the presence of seed proteins and PLs Nikiforidis and Scholten (2015) only used natural OBs to
(Chang et al., 2013; Chen et al., 2020). prepare a high internal phase emulsion gel with volume
In order to maintain the structural integrity of natural fractions of 0.91 and its shear elastic modulus was
OBs, pH-driven and ultrasonic loading methods are between 102 and 105 Pa. Kirimlidou et al. (2017) rev-
developed. Zheng et al. (2019) dissolved curcumin crys- ealed that by modulating the pH of the OBs-gelatin mix-
tals in a strong alkaline solution (pH >8) to deprotonate ture at 5.8–6.2, it allowed gelatin molecules (positively
the hydroxyl group, resulting in an increase in negative charged with an isoelectric point of around 8.1) to inter-
charge and hydrophilicity of curcumin. This solution was act electrostatically with the surface proteins of the OBs
mixed with an acidified OBs suspension and the final pH (negatively charged) so that a large number of dispersed
of the mixed system was around neutral or less, which OBs could be acted as positive filler materials of the gel-
caused the curcumin in the aqueous phase to become atin protein matrix network, enhancing the network and
non-polar and move into the hydrophobic interior of the improving the gel strength of the composite gels.
OBs. Sun et al. (2022) successfully prepared a stable Yang et al. (2020) constructed an emulsion-gel with soy-
curcumin-loaded OBs emulsion by ultrasound-assisted bean OBs as filling oil/fat droplets and κ-carrageenan as
treatment. This stability was attributed to the pressure gel matrix by modulating the electrostatic interaction
and shear force generated by ultrasound treatment between the matrix and the proteins on the surface of
made the OBs emulsion particles smaller, effectively OBs at different pH, and found that at pH 4.0, the elec-
increased the amount of free foreign protein adsorbed at trostatic interaction between oppositely charged
the surface of the soybean OBs and ultimately resulting κ-carrageenan and protein-coated OBs facilitated gela-
in a higher concentration of the formed interfacial protein, tion of κ-carrageenan; at pH 7.0, the electrostatic repul-
which promoted the curcumin-coated oil bounded to the sion between κ-carrageenan and OBs delayed gel
hydrophobic region of the OBs protein through hydro- formation. OBs emulsion-gels exhibited better lubrication
phobic interactions. properties and an ultralow boundary friction coefficient
(μ) was achieved, which was significant to study the oral
processing of OB emulsion-gels when it was used in
OBs-based emulsion-gels semi-solid food.
Emulsion-gel is a typical semi-solid food system

consisting of gel matrix filled with oil/fat droplets OBs-based oleogels
(Dickinson, 2012). Proteins such as gelatin, soy protein,
casein and polysaccharides such as starch, carra- Oleogels are low in saturated fat content but similar to
geenan, pectin, alginate, and flaxseed gum are the solid lipid particles, which can serve as a healthy
alternative to solid fats while maintaining the food sen- reported that the initial OB film was less elastic and
sory properties (Hwang, 2020; Okuro et al., 2020). easy to tear, and Tris–HCl plasticizer, Ca2+/Mg2+
Recent studies have shown that oleogels can be used crosslinking agent and carboxymethyl cellulose could
in chocolate, cakes, cookies, ice cream, and meat effectively modify the mechanical properties of OBs
products (Martins et al., 2020). Edible oleogels are membranes, making them had a wide range of tensile
composed of an edible liquid phase (vegetable oil) that strength and elongation properties (Wang, 2004). To
is entrapped by a structural agent (structurant or avoid the homogenization, small sized natural OBs also
oleogelator) network, leading to the formation of a can be added to the initial biopolymer solution to
three-dimensional gel system (Tavernier et al., 2017). replace vegetable oil. The interaction between the sur-
Proteins, polysaccharides and their combinations face of maize germ OBs and caseinate molecules
are the most common high molecular weight resulted in the effective binding of the OBs to the pro-
oleogelators, which can form three-dimensional poly- tein matrix, and then the composite film with milky white
meric systems through physical interactions (Wijaya appearance, strong hydrophobicity, relative viscosity
et al., 2019). Most of the proteins and polysaccharides and flexibility was prepared. The fact that mechanical
used to prepare oleogels are hydrophilic polymers. and optical characteristics of the composite films mar-
Therefore, the gelators need to be dispersed into water ked alterations upon storage due to water uptake or
and adsorbed at the air–water or oil–water interface to OBs movement (Matsakidou et al., 2013, 2018). The
form hydrocolloids, and then carries out the dehydration preparation of biopolymer-based films involves two
step to build a three-dimensional network, which can steps: (i) preparation of film-forming dispersions and
entrap liquid oil within the matrix to form oleogels addition of plasticizers such as glycerine, propylene
(Ashok & Dewettinck, 2016). OBs can provide an excel- glycol, sorbitol to hinder the interaction between poly-
lent medium for forming soft solids from liquid oils as mers and improve the thermo-plasticity of polymer,
they are already in the form of encapsulated oil droplets (ii) the film is formed by casting method also known as
in surface proteins. Mert and Vilgis (2021) adopted solvent casting method. First, the solution is poured in
xanthan gum and pectin to stabilize natural OBs suspen- the mold, and then dried to form the film (Suhag
sion based on electrostatic deposition, and converted it et al., 2020).
into soft solid oleogel structure after drying. Emulsion-
template method, an indirect method of oil structuring, is
applied in this study. This method first fabricates an oil- In vitro digestion of OBs
in-water emulsion with proteins as stabilizers and then
uses polysaccharides to resist unstable forces, such as The digestive behavior of OBs emulsions is similar to
droplets coalescing and emulsifying in the process of that of protein-stabilized emulsions, with flocculation of
dehydration (Bascuas et al., 2021). Then the emulsion is the OBs occurring due to the proteolysis by pepsin
dried to remove water, resulting in a high internal phase under gastric conditions. It had been reported that
emulsion (HIPE), which is used as a template to be under intestinal conditions, hydrolysis of TAGs led to
sheared to create protein/polysaccharide oleogels the spontaneous formation of a new type of multiple
(Li et al., 2022; Urbankova et al., 2021). emulsions in which there was a liquid-crystalline phase
of bile salts and lipolytic products at the surface of the
oil droplets and some bile salt crystals at the surface of
Edible films the inner water-protein-rich droplets (Gallier
et al., 2013). It is presumed that the interaction between
Edible films based on biodegradable materials (biopoly- the PUFA-rich fatty acid structure of some OBs and bile
mer matrix such as polysaccharides, proteins, lipids, salts may be the cause of “spontaneous” formation of
and composite materials) have potential uses in food multiphase emulsions.
packaging and as carriers of active compounds like In general, lipids digestion in OBs is slow and the
antioxidants and antimicrobials (Jeevahan et al., 2020). digestive efficiency may be affected by the following
However, the water barrier property of water-soluble three aspects (Wang et al., 2020). First, the OBs mem-
hydrocolloid edible films is poor. It had been reported brane has a negative impact on the digestion efficiency.
that vegetable oil was added into the biopolymer film Second, oleosins are partially broken down during
matrix to reduce the water vapor permeability gastrointestinal digestion and the rearrangement of pro-
(Jeevahan & Chandrasekaran, 2019). The biopolymer- tein profile (from high- and low-MW proteins after gas-
oil mixture needs to be homogenized to obtain a uni- tric digestion to higher concentrations of low-MW
form distribution with small-sized droplets, thus increas- proteins after intestinal digestion) may limit the lipids
ing the tortuosity factor and improving the water barrier bioaccessibility. Third, long chain fatty acids, the main
performance of the film (Vargas et al., 2011). OB is kind lipolytic products, accumulate at the surface of the
of a natural composite film dispersion contains both OBs limited the activity of pancreatic lipase (Gallier
natural PL-protein membrane and oil, but it had been et al., 2013; Gallier & Singh, 2012; Trombetta
et al., 2020). Additionally, in the complex food matrix, a OBs extraction process, such as pretreatment, mate-
protective layer is formed around OBs under the influ- rial to liquid ratio and centrifugal force have a direct
ence of macromolecules such as protein and polysac- impact on the OBs extraction yield, while pH and purity
charide, which promote the flocculation of OBs droplets affect the protein composition and complete structure
and inhibit the ability of pepsin and lipase, so as of OBs. The basic structure of OBs has been deter-
to reduce the absorption rate of lipids and related mined, but the content of components in OBs from dif-
lipophilic compounds (Wu et al., 2012). It had been ferent sources may be different. As a key component
reported that the bioaccessibility of important nutri- of OBs, oleosin is an excellent emulsifier, foaming
tional compounds (α-tocopherol and fatty acids) in agent and vector of recombinant protein. The stability
OBs was significantly lower than that in artificial emul- of OBs in food system can be improved to adapt to
sions emulsified with Tween 20 and whey protein iso- environmental changes by using low concentration
late (White et al., 2009). As a carrier for the delivery of polysaccharide electrostatic deposition method, high
lipophilic bioactive compounds, in the process of concentration polysaccharide viscosity effect or spray
digestion, OBs show the ability to improve the bio- drying encapsulation technology. As a pre-emulsified
accessibility of nutrients. Zheng et al. (2019) found oil-in-water (O/W) emulsion, OB can replace milk fat
that the bioaccessibility of curcumin in the curcumin droplets in dairy products or as a fat substitute applied
nanocrystals, curcumin-loaded nanoemulsions and in meat products. Delivery system, emulsion-gels,
curcumin-loaded soybean OBs was significantly oleogels, or edible films based on OBs also have been
higher than that of curcumin crystals, and the bio- reported. It is worth noting that the oleosins in pea-
accessibility of curcumin in the three delivery systems nuts, sesame and hazelnuts have been registered as
was: curcumin-loaded nanoemulsions > curcumin- allergens (Jappe & Schwager, 2017). The slow diges-
loaded soybean OBs > curcumin nanocrystals. Zhang tion rate of OBs leads to increased satiation and effec-
et al. (2022) reported that during lipid digestion in the tively helps reduce calorie intake. The bioaccessibility
small intestinal fluids, the generated free fatty acids of important nutritional compounds in OBs
and monoglycerides combined with the bile salts and (α-tocopherol and fatty acids) is not high, while the
PLs to form mixed micelles with a higher solubilization delivery system based on OBs is beneficial to the bio-
capacity for curcumin, contributing to its accessibility of hydrophobic active compounds. Over-
bioaccessibility. all, there are relatively studies on plant OBs, which
The slower digestion of OBs affects gastrointestinal provide a research basis for the development of OBs
tract physiology and may result in increasing satiety, from different sources.
effectively helping to reduce calorie intake, but the bio-
accessibility of important nutritional compounds
(α-tocopherol and fatty acids) in OBs is not high. The FUTURE PERSPECTIVES
delivery system based on OBs emulsion is beneficial to
the bioaccessibility of hydrophobic active compounds, It is necessary to design a process with low cost, sus-
though the rate of lipid digestion is slower and no rela- tainability, high yield and complete structure of OBs.
tionship between them during digestion has been There has been a tendency to develop alternative
reported. Meanwhile, the bioaccessibility of hydropho- extraction processes including enzyme assisted extrac-
bic active compounds also depends on the structure of tion and supercritical fluid extraction. Complete OBs
the compound loaded-OBs, which is complex and extraction can be realized under the condition of ensur-
uncertain. ing that the OBs membrane is not destroyed by screen-
ing specific enzyme and adjusting operating
parameters. At the same time, high pressure, ultra-
CONCLUSIONS sonic, microwave, steam flash and other auxiliary
processing technology can be developed to achieve
OB is an organelle composed of PLs-protein layer cov- efficient extraction. The special structure of OBs can be
ering central lipid. This special structure ensures the used as a reference for the development of synthetic oil
stability of the lipid used as an energy source under droplets. It is also worth exploring the potential function-
extreme environmental conditions. In order to better ality of OBs and individual OBs components. The appli-
study this spontaneous and natural oil–water mixture to cation of OBs in food will remain a trendy subject for
prevent oil oxidation system, scientists begin to sepa- the future. Future research should have full understand-
rate and extract the OBs. It is found that OB can form a ing on the impact of the composition and structure of
natural pre-emulsified oil-in-water (O/W) emulsion in OBs-based application system on its bioaccessibility,
vitro. bioavailability and functionality. The potential allergenic
At present, there are three main OBs extraction behavior and toxicity of OBs also cannot be ignored
methods: deionized water extraction, buffer extraction and in vitro and in vivo studies are needed to take care
and enzymatic extraction. The key parameters in the of the associated side effects.
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15589331, 2022, 8, Downloaded from https://aocs.onlinelibrary.wiley.com/doi/10.1002/aocs.12618 by New York University, Wiley Online Library on [06/03/2023].

Recent developments and prospects in the extraction,

Jia Hao | Xiaoyu Li | Qiuyu Wang | Wenwen Lv | Wenguan Zhang |

Beijing Advanced Innovation Center for Food Abstract

INTRODUCTION have isolated more than 20 kinds of seed OBs in the

J Am Oil Chem Soc. 2022;99:635–653. wileyonlinelibrary.com/journal/aocs © 2022 AOCS. 635

(a) Average diameters and the contents of Basic composition

Soybean 0.25–1.75 88.51–93.72 5.21–9.72 0.80

(b) Fatty acids composition

Linoleic acid (18:2) 55.00 35.01 28.00–29.00 24.34 1.10 56.30

(c) PLs composition

Peanut 50.60–61.60 5.00–16.76 8.40–23.04 10.80–25.00 5.33–10.00

(Figure 2). Starches and proteins were distributed Aqueous extraction

bonds of proteins allowing the removal of passively Enzyme assisted extraction

complex), Cellulase A (Cellulase, β-glucanase, Pretreatment

Bioactive components polysaccharide and the system changes from positive

OBs in yogurt OBs-based delivery system

Emulsion-gel is a typical semi-solid food system

A UT H O R C ON T R IB UT IO NS Bolumar T, Toepfl S, Heinz V. Fat reduction and replacement in dry-

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