Review

Received: 5 March 2019 Revised: 27 June 2019 Accepted article published: 1 July 2019 Published online in Wiley Online Library: 4 September 2019

(wileyonlinelibrary.com) DOI 10.1002/jsfa.9905

Effect of low-temperature preservation

on quality changes in Pacific white shrimp,
Litopenaeus vannamei: a review
Chuang Pan, Shengjun Chen, Shuxian Hao and Xianqing Yang*

Abstract
Shrimp has been widely accepted as an excellent resource for white meat due to its high-protein and low-fat content, especially
low cholesterol. However, shrimps are highly perishable during preservation and retailing procedures due to the activities
of enzymatic proteolysis, lipid oxidation, and microbial degradation. With increasing knowledge of and demands for safety,
nutrition, and freshness of shrimp products, energy efficient, quality, maintained, and sustainable preservation technologies
are needed. Low-temperature preservation, a practical processing method for improving the shelf life of food products,
is widely used in the aquatic industry. This review focuses on the effects of low-temperature preservation on the quality
changes in Litopenaeus vannamei. It considers physicochemical properties, sensory evaluation, melanosis assessment, and
microbiological analysis. The perspectives of non-protein-based techniques on quality analysis of shrimps during preservation
are also discussed.
© 2019 Society of Chemical Industry

Keywords: Litopenaeus vannamei; preservation; quality; spoilage; temperature

INTRODUCTION and activities inhibition.13 Temperature control is well known to

The increasing world population and awareness of food freshness,
nutrition, and safety have contributed to an increased consump-
tion of fresh foods, especially in aquatic products.1–3 Fresh aquatic
products are highly perishable due to the existence of large num-
ber of nutrients like proteins, functional peptides, unsaturated fats,
and minerals.1,4–6 Enzymatic proteolysis plays an important role in
the quality and preservation of aquatic products. It was reported
that the quality loss in muscle food products, especially in aquatic
products, was primarily correlated with the activities of endoge-
nous enzymes during storage.7–10 Lipid oxidation, known as fat
rancidity, occurred. In this process, the unsaturated fatty acids
degrade into aldehydes, ketones, and lower fatty acids,11,12 which
can bring about a reduction in the nutritive properties and qual-
ity deterioration in fresh aquatic products.9,13–16 As a result of their
high unsaturated fatty acid content, aquatic products are more
susceptible to lipid oxidation than other muscle foods, like pork,
beef, and lamb.2,9,12 Furthermore, lipid oxidation products may also
react with proteins, phospholipids, and even DNA inside the mus-
cles, with adverse health effects for human beings and unfavor-
able sensory assessments.9,11 In addition to the above two factors,
microbial growth is another important parameter for the deter-
mination of shelf life and the quality of aquatic foods.17 It was
reported that about 25–30% of the gross primary agricultural and
fishery products are lost, and this loss is correlated with microbial
activities.18–20 Moreover, autolysis products from protein and lipid
will be used as substrates for bacteria growth, thus accelerating the
spoilage of fresh aquatic foods.21–24
To maintain quality and improve the shelf life of aquatic prod-
ucts, different preservative strategies have been employed for stor-
be an essential aspect of the food supply chain, and low storage
temperature is used to increase the shelf life of aquatic products
by reducing the rates of enzymatic autolysis, lipid oxidation, and
microbial degradation for decades.25–28 The most common meth-
ods used for low-temperature preservation include refrigerated
storage between 0 to 4 ∘ C, frozen storage at −18 to −40 ∘ C, and
the super-chilled range of −1 to −4 ∘ C.29–31 At low temperatures,
microbial activities are inhibited, and most bacteria are unable
to grow.32,33 Chemical and enzymatic reactions are also retarded,
especially the degradation of ATP and its related compounds.24,34,35
Crustaceans are cultured widely around the world, with 27%
of shrimps cultured in China.36 Crustacean aquaculture has been
dominated by Pacific white shrimp (Litopenaeus vannamei).37,38
Recently, with rapid economic growth and increasing awareness
of food nutrition and safety in China, the demand for high-quality
aquatic products has been increasing. It is a significant volume
and value of the high quality aquatic market in future China.36
The challenge for the aquatic food industry, which includes the
shrimp industry, is to develop an optimal preservation method
to maintain the freshness, quality, and safety of its products. The
∗ Correspondence to: XQ Yang, Key Laboratory of Aquatic Product Processing,
Ministry of Agriculture and Rural Affairs, National R&D Center for Aquatic Prod-
uct Processing, South China Sea Fisheries Research Institute, Chinese Academy
of Fishery Sciences, Guangzhou 510300, China. E-mail: yxqgd@163.com
Key Laboratory of Aquatic Product Processing, Ministry of Agriculture and Rural
Affairs, National R&D Center for Aquatic Product Processing, South China Sea
Fisheries Research Institute, Chinese Academy of Fishery Sciences, Guangzhou,
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age and distribution, like temperature control, moisture control, China

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Figure 1. ATP-related compounds and their degradation pathways in fish, shellfish, and mollusk.42
effects of low-temperature preservation on quality changes in
L. vannamei are therefore considered in this review for a better
understanding of the preservation technology on shrimp prod-
ucts, considering physicochemical properties, and using sensory
evaluation, melanosis assessment, and microbiological analysis.
Meanwhile, the perspectives of non-protein-based analysis tech-
niques and real-time polymerase chain reaction (RT-PCR) on the
determination of the quality of shrimps during preservation are
also discussed.
PHYSICOCHEMICAL ANALYSIS
Chemical indexes
K value
The K value is a freshness indicator, which is calculated based on
the quantification of adenosine triphosphate (ATP) and its related
compounds, including adenosine diphosphate (ADP), adenosine
monophosphate (AMP), inosine 5′ monophosphate (IMP), inosine
(HxR) and hypoxanthine (Hx) (Fig. 1).39–42 A higher K value indicates
a higher rate of ATP degradation, reflecting a greater loss of
freshness in aquatic products.35,42 It can be calculated as in the
following equation:
HxR + Hx
K(%) = × 100%
ATP + ADP + AMP + IMP + HxR + Hx
It is well known that the degradation of ATP in fish and shell-
fish after death is followed by ATP → ADP → AMP → IMP→HxR →
Hx → uric acid.42 In fresh fish muscle, the IMP usually accumulates
due to the slow degradation speed of IMP to HxR. However, in crus-
taceans, AMP tended to accumulate because of the low activity of
AMP deaminase (Fig. 1).42,43 In general, a K value of 20% or lower
suggests a good quality aquatic product, while 60% has been rec-
ognized as the rejection point.44
Huang et al. (2016) compared the change in the K values of
white shrimp during storage at 25 ∘ C and at 4 ∘ C. The K value
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with an initial value of 2.5% increased significantly when stored at
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25 ∘ C preservation and reached a final value of 46.8% after 24 h.
This value was reached at 14 days of storage at 4 ∘ C, indicating
that a low preservation temperature can effectively reduce the
degradation of ATP-related compounds in white shrimp muscle.45
After the death of the shrimp, glycogen anaerobically converted
to lactic acid due to the anaerobic metabolism that took over in
the shrimp muscle, resulting in the reduction of the pH in the
muscle, which could activate the ATPase.34,46 Similar results were
also obtained in banana shrimp, Japanese spiny lobster, kuruma
shrimp, and tanner crab.43,47,48 Mendes et al. (2002) mentioned that
deep-water pink shrimp (Parapenaeus longirostris) obtained from
the Portuguese coast had an initial K value of 9% and reached 40%
at 10 days of storage under 2 ∘ C.49 However, Nirmal and Benjakul
(2009) reported that the L. vannamei treated with catechin had
the lower rate of increase in the K value, which finally reached
32% under the same preservation condition.5 Furthermore, the
loss of ATPase activity showed a significant correlation with the
concentration of catechin.5 Zou et al. (2010) demonstrated that
the K value of L. vannamei under frozen storage increased from
5.62% for the fresh sample to 7.18% after 10 days of preservation
at −18 ∘ C and this value showed no significant change in the
next 30 days of preservation.50 However, long-term frozen storage
will increase the drip loss to the samples due to the damage of
shrimp muscle structure induced by ice crystals.50,51 These results
indicated that a low preservation temperature could effectively
reduce the degradation of ATP and its related compounds in
shrimps. It was explained that the temperature correlated with a
reduction in the activity of endogenous enzymes in muscle when
preserved at low-temperature conditions. A lower temperature
can retard the degradation of ATP-related compounds but it will
lead to a severe drip loss in the final products, which will affect the
texture assessment and sensory evaluation.
Total volatile basic nitrogen (TVB-N)
Total volatile basic nitrogen is a widely used index of the dete-
rioration of aquatic products, resulting from the degradation of
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nitrogenous compounds in muscle like proteins, peptides, amino
acids, and nucleotides during storage.52,53 Resulting from the
degradation of nitrogenous compounds including dimethylamine
(DMA), trimethylamine (TMA), ammonia and other volatile basic
nitrogenous compounds in muscle during storage.52,53 Huang
et al. (2016) reported that the formation of TVB-N in L. van-
namei increased significantly from 4.2 mg/100 g (fresh) to a final
49.0 mg/100 mg after 24 h of storage at 25 ∘ C. While the increas-
ing tendency of TVB-N was effectively limited when the temper-
ature was decreased to 4 ∘ C, it eventually reached 52.8 mg/100 g
after 14 days of storage at 4 ∘ C.45 The formation of other volatile
basic nitrogenous compounds was also slowed at low preserva-
tion temperatures. The TMA, ammonia, and urea content increased
to 14.4 mg/100 g, 21.8 mg/100 g and 6.0 mg/100 g, respectively,
when stored at 25 ∘ C for 24 h.45 However, the TMA content was
50% lower after 14 days of storage at 4 ∘ C compared with 25 ∘ C.
For ammonia and urea, it took 7 and 10 days to reach the same
value during storage at 4 ∘ C.45 Zhang et al. (2015) investigated the
changes in the TVB-N value of L. vannamei stored under slurry ice
conditions and revealed that the TVB-N value of samples treated
with slurry ice remained at 19.38 mg/100 g after 16 days of chilled
storage (icing storage).54 These results indicated that the TVB-N
value could be limited effectively by simply lowering the storage
temperature. It has also been reported that the TVB-N content
could be successfully controlled by low-temperature storage com-
bined with other techniques.55–62 Wang et al. (2016) demonstrated
the effect of modified atmosphere on the quality of shelf life of
L. vannamei under controlled freezing-point storage at −0.8 ∘ C.55
With an increase in storage time, atmospheres with higher CO2
concentrations notably inhibited the increase of TVB-N contents.
Shrimp samples with 100%CO2 and 67%CO2 /17%O2 /17%N2 had
the least changes in TVB-N content during storage experiments.55
However, the samples packaged without CO2 showed the steep-
est change in TVB-N content due to the activity of spoilage bac-
teria and endogenous enzymes, which resulted in the formation
of compounds including ammonia and primary, secondary and
tertiary amines, imparting characteristic off flavors to shrimps.56,57
Moreover, low-temperature storage combined with edible coating
technologies also showed significant limitations on the formation
of TVB-N.58–62
pH
The changes in pH were consistent with the results of TVB-N con-
tent due to the accumulation of basic compounds induced by the
activity of bacteria or enzymes.63,64 The increase in pH reflected
the production of alkaline bacterial metabolites in spoiling shrimp
muscle.65 The pH of the fresh L. vannamei ranged from 7.20 to
7.33 and the shrimps were of good quality before the pH reached
7.7.4,58,66,67 The pH of L. vannamei increased to 7.6 at the seventh
day of storage under 4 ∘ C,45 7.48 at the ninth day during chilled
storage,48 and 7.6 at the eighth day of storage at −0.8 ∘ C.55 More-
over, the increase in the pH in shrimp during storage could be
inhibited significantly by low-temperature combined with a coat-
ing strategy. Mu et al. (2012) investigated the effect of different
concentrations of cinnamaldehyde essential oil on the shelf-life of
L. vannamei stored at 4 ∘ C. The spoilage pH value was detected at
the tenth day of storage with 1 g/kg cinnamaldehyde, while the
samples treated with 5 g/kg cinnamaldehyde only reached 7.43
after 11 days of storage, which is almost the same as the pH in
fresh state.64 Similar results have also been found in other edi-
ble coating treatments like green curry paste marinated shrimp,68
chitosan coating combined with pomegranate peel extract,59
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ascorbic acid combined with green tea extract,69 caprylic acid coat-
ing treatment,60 chitosan-gelatin coating treatment,62 grape-seed
extract coating experiment,4 and basil seed gum coating com-
bined with thymol.70 Interestingly, the increasing of pH could
not be inhibited by some coating treaments. Huang et al. (2012)
demonstrated that the pH of shrimp samples treated with chitosan
and O-carboxymethyl chitosan showed no significant differences.
They suggested that the thick crust may prevent the penetration of
marinated solution into the flesh.58 Basiri et al. (2015) revealed that
the shrimp only coated with pomegranate peel extract showed a
higher pH value than the non-coating group, and there were no
significant differences among coating concentrations.61
Thiobarbituric acid reactive substances (TBARS)
Lipid peroxidation, corresponding to oxidative deterioration of
polyunsaturated fatty acids in muscle, leads to the production
of off flavors and off odors, thereby shortening the shelf life of
seafood.54 Thiobarbituric acid reactive substances, the products
of secondary lipid oxidation, have a close correlation with sen-
sory assessment in aquatic products.71,72 It has been reported that
low-temperature treatment is an effective way of reducing lipid
formation in aquatic products, like sardines (Sardina pilchardus),71
farmed turbot (Psetta maxima),73 and European hake (Merluccius
merluccius).74 Zhang et al. (2015) revealed that slurry ice particles
showed significant inhibition of lipid oxidation compared with
flake ice particles due to the round shape of slurry ice particles,
which could make contact with the samples directly and keep the
desired low temperature during storage.54 In addition to the ice
shape, some edible coating materials have also been found to pos-
sess antioxidant ability against lipid oxidation. Khazaei et al. (2017)
suggested that the shrimp samples coated with basil seed gum
and thymol had greater stability towards lipid oxidation.70 Shrimp
samples treated with green tea extract showed lower TBARS con-
tent after 12 days of iced storage.69 Basiri et al. (2015) revealed
that the pomegranate peel extract (PPE) possesses a markedly
antioxidant capacity against superoxide anion, hydroxyl, and per-
oxyl radicals.61 Shrimp samples treated with PPE showed about
30% lower on TBARS content when compared with the controls
after 10 days of storage. In the meantime, the coating concentra-
tions of PPE exhibited no significant effect on the inhibition of lipid
peroxidation.61 Similar results were also found in a shrimp storage
experiment in which shrimp was coated with catechin, which has
been reported to have strong antioxidant activity including radi-
cal scavenging activity.75,76 However, the well-known oxygen scav-
enger, ascorbic acid, showed no significant effect on antioxidative
activity during the low-temperature storage of L. vannamei.69,77
Proteins
Shrimp muscle protein mainly consists of myofibrillar and sar-
coplasmic proteins, which account for 60 to 70% and 20 to 30%,
respectively.78,79 Zhang et al. (2015) revealed that the deterio-
ration of myofibrillar protein in L. vannamei muscle could be
reduced effectively by low-temperature storage.54 The decreases
in muscle myofibrillar and intramuscular tissue have been reported
to be correlated with proteolytic enzymes like cathepsins80 and
calcium-dependent proteases.81 The release of these enzymes
may cause an acceleration of myofibrillar protein degradation,
which results in the deterioration of shrimp muscle.82 At of
low-temperature process, a large number of small ice crystals were
formed in muscle nuclei, which could prevent the irreversible
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destruction of the myofibrils from the formation of large ice
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crystals. Hence, the quality and shelf-life of aquatic product could
be maintained.54 Shaban et al. (1987) reported quality changes
in kuruma shrimp when stored at −20 ∘ C for 1 to 3 months.83
They found that 1) the contents of sarcoplasmic proteins remained
unchanged, 2) the myofibrillar proteins decreased, 3) the con-
tents of alkali-soluble proteins increased.83 However, Ando et al.
(2004) demonstrated that no changes could be detected in these
water-soluble proteins when stored at both 5 ∘ C and − 2 ∘ C for
3 days.79 Investigations of changes in shrimp muscle proteins dur-
ing low-temperature storage are relatively scarce and further stud-
ies are needed on this aspect.
Melanosis
Melanosis is characterized by black spot formation during the
post-harvest of crustaceans, such as shrimp.4 It has been explained
as the polymerization of phenolin to an insoluble black pigment,
melanin, which was mainly initiated by the action of polyphenol
oxidase (PPO).84 This change decreases the quality and marketabil-
ity of the shrimp during storage and it was difficult to suppress
by simply lowering the storage temperature.85,86 Yuan et al. (2016)
reported the effect of chitosan coating combined with green tea
extract on the melanosis of L. vannamei during icing storage.66
Melanosis formation increased during the storage in all the treat-
ment conditions; however, it was significantly inhibited and visual
quality was significantly improved in coating groups compared
with the control.66 Many natural extracts or compounds, such as
catechin, ferulic acid, extracts from pomegranate, green tea and
grape seed could also inhibit the melanogenesis of shrimps dur-
ing low-temperature storage.4,59,61,69 Polyphenol oxidase is the key
enzyme for the synthesis of melanin and some phenolic com-
pounds could inhibit PPO activity by interacting with the active
site of the enzyme.86 The phenolic compounds could also interact
with protein or enzymes through a hydrogen bond or hydrophobic
interaction.87 Moreover, it was reported that chitosan could delay
the appearance of black spots in Pandlus borealis due to its chelat-
ing action and coating-induced oxygen exclusion, which could
prevent the enzyme activity of PPO.88
Physical indexes
Color
Color is one of the most important visual properties for consumers
in seafood, especially in shrimps.89–91 Normally, changes in shrimp
color during different preservation conditions were quantified by
determination of L* (lightness / darkness), a* (redness / green-
ness) and b* (yellowness / blueness). Zhang et al. (2015) reported
that the L* values of raw L. vannamei tended to decrease and b*
values tended to increase after 16 days of icing storage.54 Com-
pared with flaked-iced shrimps, slurry ice-treated samples exhib-
ited higher L* values and lower b* values, which mean that the
slurry ice treatment could improve the color properties of shrimps
during storage.54 The raw crustacean body including shrimp and
crab turns blackish during low-temperature storage. This has been
reported to be correlated with melanin, which was generated
by the action of phenol / polyphenol oxidase on tyrosine or its
derivatives.92,93 Lipid oxidation, which degrades carotenoid pig-
ments, is another important factor in shrimp color change.93,94
Furthermore, color changes on shrimp were also considered as
a result of microbiological actions under different storage con-
ditions and packaging techniques.95 It has also been reported
that color changes in shrimp could be effectively retarded by low
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temperatures combined with an edible coating treatment.59,60,62,70
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Khazaei et al. (2017) reported that the color change in L. vannamei
could be retarded effectively with basil seed gum coating after
20 days of storage at 4 ∘ C and the higher concentration of coating
solution could brought a better color protection.70 Similar results
were also obtained in other coating studies including those with
chitosan coating,96 chitosan-carvacrol coating,60 chitosan coating
combined with pomegranate peel extract,59 and chitosan-gelatin
coating.62 The coating could be contributed to the maintenance of
shrimp color during the low-temperature storage was mainly due
to the isolation of oxygen and insulation of temperature.97,98
Texture
Texture is not only an important sensory property of shrimp
but also one of the important parameters for consumer
acceptability.48,54 It includes parameters of hardness, springi-
ness, cohesiveness, gumminess, and chewiness.48 Sometimes,
the chewiness is defined as the product of hardness, cohesive-
ness, and springiness, which indicates the tactile sensation of
labored mastication due to sustained elastic resistance from the
shrimp muscle.54 Zhang et al. (2015) reported that, even though
the springiness and chewiness values of shrimp muscle were
all decreased during storage, the slurry ice treatment showed
significant inhibition in texture loss compared with control or
flake-ice treatment.54 The changes of texture parameters could be
influenced by many factors, such as changes of pH, degradation of
myofibrillar proteins or the connective tissues.59,99,100 In agreement
with Zhang’s study, Annamalai et al. (2015) found a similar loss in
hardness, springiness, and chewiness of low-temperature stored
L. vannamei.48 It was notable that no differences were detected in
the adhesiveness and hardness of farm-raised L. vannamei during
icing storage.67 Moreover, deep water pink shrimp stored under
liquid icing condition showed similar minor changes in its texture
values (breaking force, hardness, and elasticity).101 Application of
chitosan-based coatings could also effectively retard the change
of texture parameters in shrimp during storage.59 Wang et al.
(2015) demonstrated that chitosan nanoparticle coating exhib-
ited the highest hardness and springiness of L. vannamei during
10 days of storage at 4 ∘ C.102 The retardation of texture loss was
also found in chitosan coating combined with pomegranate peel
extract59 and chitosan-carvacrol coating treatment.60 Interest-
ingly, Farajzadeh et al. (2016) declared that the chitosan-gelatin
coating showed an effect in improving the rheological properties
on shrimp compared with non-coating group after 14 days of
storage at refrigerated condition.62 The gelatin plays an important
role in the increasing of shrimp texture.
Water holding capacity (WHC)
Many physical properties such as color and texture are partially
dependent on the WHC of a given food material, especially in
aquatic foods.103 Driven by lipid oxidation and its derivatives, the
changes in protein are also somehow associated with WHC in
aquatic muscles.104 This was measured using the water retention
index (WRI): decreases in WRI reflect decreases in the release of
‘free water’, which mean an increase in WHC of meat flesh.16,105
Okpala et al. (2015) revealed that the WRI of L. vannamei decreased
in the first 4 days of icing storage and then there was no significant
change from days 6 to 12.67 They also found that the decrease in
the WRI of ozone-treated samples was apparent but to a lesser
extent when compared with icing storage.16 They suggested that
the ozone-treated shrimps could hold more moisture inside the
tissues, which could somehow reduce the decrease in WRI. Medina
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et al. (2012) hypothesized that proteins in muscle tissue could
oxidize before the lipids, thus allowing movement of water from
myofibrillar into extracellular spaces, which could enhance the
WHC.104
SENSORY EVALUATION
Sensory evaluation is a scientific discipline that applies the
principles of experimental design and statistical analysis to the
use of human senses (sight, smell, taste, touch, and hearing)
for the purposes of evaluating consumer products.2,106 In gen-
eral, the shrimps have a bright natural color, odor, and firmness
(excellent elastic and rigid characteristics), presenting very good
overall quality.48 Huang et al. (2016) reported that the sensory
evaluation of L. vannamei stored at 25 ∘ C for 12 h showed a
gray color with a pink tint, a semi-firm texture, and a light fishy
smell.45 The advanced decomposition was obtained after 18
h of storage. In this study, the shrimp was of acceptable qual-
ity for 6 h at 25 ∘ C and it increased to 3 days when stored at 4
∘ C.45 Icing-treated samples obtained higher scores for sensory
analysis, including color, firmness, and odor. Sensory properties
showed no significant change up to at least 6 days.48,54 Tem-
perature plays an important role in the rate of deterioration
of fish and fishery products. Within an increase of pH, TVB-N,
TMA, and TBARS values inside the shrimp muscle, the sensory
scores were decreasing significantly.21,107,108 Studies on coating
treatments combined with low-temperature storage revealed
that a reasonable coating process can effective resist the phys-
iological and biochemical reactions and metabolism of shrimp
tissue during storage, leading to a better sensory experience
and greater acceptance by the consumer.4,59–62,66,70 For instance,
Khazaei et al. (2017) described the effect of basil seed gum coating
on quality maintenance of L. vannamei during cold storage.70 It
was shown that the coating process could extend the shelf life
of shrimps for at least 2 days compared with non-coating group,
resulting in a total shelf life of 12 days.70 Similar observations
were reported for some other active edible coating treatments
combined with low-temperature storage, such as grape-seed
extract coating,4 pomegranate peel extract-chitosan coating,59
chitosan-carvacrol coating,60 chitosan-gelatin coating,62 and
green tea extract-chitosan coating.66 The combination of low
temperature with modified atmosphere packaging (MAP) could
also bring better sensory acceptance than separate use of them
over a given storage period.55,94
MICROBIOLOGICAL ANALYSIS
Many bacteria are attached to the organs of aquatic animals, such
as the skin, gills, and intestines. With the death of an animal,
the balance between bacteria and the defense system is bro-
ken, which can result in the growth of bacteria. Then the free
amino acids, glycogen, and organic acids will be degraded into
amines, alcohols, and aldehydes, assisted by the bacteria. Eventu-
ally, unpleasant flavors and a decrease in quality will be detected.42
About 30% of the world’s food production is lost due to micro-
bial spoilage.18 It is also responsible for the deterioration in the
quality of most fresh aquatic foods. Thus, the total number of
microorganisms has been used in mandatory seafood standards
in many countries, like China, Japan, and the USA.2,21 The aero-
bic mesophilic bacterial count of freshly harvested shrimp was
reported to range from 103 –106 in the tropical environment.109
Annamalai et al. (2015) investigated the effect of delayed icing
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on the quality of L. vannamei during chilled storage.48 They
demonstrated that the bacterial population did not exceed the
maximum limit for acceptability, indicating that the growth of
mesophilic bacteria was controlled by icing treatment.48 In addi-
tion to low temperature, edible coating processing can also retard
the growth of microorganisms.4,59–61,64–66,69,70 According to Khaz-
aei et al. (2017), edible coatings made from basil seed gum and
thymol had lower total bacterial count than uncoated samples
when stored at the same temperature (4 ∘ C).70 Nirmal and Ben-
jakul (2012) found that green tea extract in combination with
ascorbic acid could retard the increase in the psychrophilic bacte-
rial count during a 12-day storage period,69 and this is consistent
with the result from Banon et al. (2007), who found that ascor-
bate and green tea extract increased the shelf-life of beef pat-
ties by delaying microbial spoilage.110 Similar results were also
reported in the chitosan coating combined with pomegranate
peel extract,59 and chitosan-gelatin coating.62 Furthermore, Wang
et al. (2016) demonstrated that mesophilic and psychrotrophic
bacterial counts were inhibited by the presence of carbon diox-
ide combined with freezing-point storage at −0.8 ∘ C.55 A possible
explanation is that the solubilization of carbon dioxide in the water
present in samples tends to create weakly acidic conditions and
thus retards the growth of microorganisms.55,94
FUTURE PERSPECTIVES AND CONCLUSIONS
Aquatic products especially shrimps, are preferred by a great
number of consumers due to their high nutrition, low fat, and
easy digestion. Meanwhile, the increasing demand for fresh foods
is leading to an increase in the development of energy-saving,
quality-maintenance technology. Low-temperature preservation
technology has advantages in maintaining food freshness, pre-
serving nutritional quality, and extending shelf life.
Temperature is the most crucial factor that correlated with
the quality of aquatic foods. Different aquatic species and prod-
ucts should be stored at different temperatures. To maintain the
product quality, the precise control of time and temperature
distribution during the storage process is important. Any small
temperature fluctuation during preservation may cause the qual-
ity to decrease due to the melting and recrystallization of ice crys-
tals inside the aquatic products.2 Suitable packaging and precise
temperature control throughout the preservation will effectively
increase the shelf life of aquatic products.
Many works are currently focusing on how to understand
the changes in the microstructural and biochemical quality
of aquatic products. There is relatively little research on the
changes in the proteins in the muscles of aquatic products during
low-temperature preservation. Shrimps, like many other aquatic
foods, are famous for their high-protein contents. Proteolysis is
one of the main factors in the deterioration of aquatic products
during processing, transportation, and preservation. The protein
contained inside aquatic muscles is also one of the food-derived
allergens, like the sarcoplasmic protein that accounts for 20% of
shrimp muscle. Understanding the changes in muscle protein
during the preservation of aquatic products could improve the
shelf life of aquatic products and may also provide insight into
allergic reactions caused by proteins in aquatic flesh.
There is a strong demand for a fast, inexpensive, reliable, and
simple quality-evaluation method for aquatic food. Conventional
food analysis techniques are almost protein-based methods.111
However, they have some disadvantages: 1) some cross activ-
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ities with non-target proteins could be occurred due to the
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 www.soci.org
dependence of protein against target antibodies;112,113 2) for
highly processed aquatic products, protein-based assays may
yield false / negative results due to the irreversible denaturation
of proteins caused by high temperatures or other treatments;114
3) target protein may show various levels in different tissues,
which restricts the quantitative determination by protein-based
methods.115 Real-time polymerase chain reaction (RT-PCR) has
emerged as a novel choice for food analysis. It can be used for
a variety of samples, including food ingredients, modified food
organisms, food-borne bacteria or allergens, even treated at very
low concentrations.111 Furthermore, unlike conventional PCR
amplification, which shows the results on the agarose gel, the
RT-PCR allows direct online analysis of results, which could save
time and money. However, protein identification is required before
primer design, and these primers are non-universal because the
highly conserved regions among aquatic species are not the same.
Furthermore, samples for RT-PCR determination are relatively easy
to contaminate during pre-treatment.
In conclusion, quality changes in shrimp and its products during
storage cannot simply be determined by a few parameters; it is a
complex reaction correlated with many factors. Low-temperature
preservation with precise temperature control and energy saving
technology has not been applied widely in the shrimp industry and
consumer awareness of stored crustacean products is currently
very low especially in China. Extensive studies on the improvement
of preservation techniques and systematic analytical techniques
for commercial shrimp products during low-temperature process-
ing are needed in the future.
ACKNOWLEDGEMENTS
This work was partially supported by the Central Public-Interest
Scientific Institution Basal Research Fund, South China Sea
Fisheries Research Institute, CAFS (NO.2018YB02); National Key
Research and Development program (2018YFD0700901-2);
Project supported by the Research and Development
Projects in Key Areas of Guangdong Province, China (Grant
No.2019B020225001).
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