DOI: 10.1002/vms3.

979

REVIEW

A review on molecular detection techniques of white spot

syndrome virus: Perspectives of problems and solutions in
shrimp farming

Sk Injamamul Islam1 Moslema Jahan Mou2 Saloa Sanjida3 Sarower Mahfuj1

1
Department of Fisheries and Marine
Bioscience, Faculty of Biological Science, Abstract
Jashore University of Science and Technology,
This review aims to provide an update on the current scientific understanding of vari-
Jashore, Bangladesh
2
Department of Genetic Engineering and
ous aspects of White Spot Syndrome Virus (WSSV) formation, diagnostic procedures,
Biotechnology, Faculty of Life and Earth transmission, ecological effects, pathophysiology and management strategies. In terms
Science, University of Rajshahi, Rajshahi,
Bangladesh
of production and financial benefits, the WSSV has been the most virulent in shrimp
3
Department of Environmental Science and and several other crustacean sectors around the globe. It spreads vertically from dis-
Technology, Faculty of Applied Science and eased broodstock to post-larvae and horizontally by cannibalism, invertebrate vectors,
Technology, Jashore University of Science and
Technology, Jashore, Bangladesh freshwater and sediments. In the transfer of white spot disease (WSD) in newly stocked
ponds, the survivability of WSSV in sediment is the most important variable. In typical
Correspondence
cultural conditions, it is a highly infectious pathogen capable of inflicting total death
Department of Fisheries and Marine
Bioscience, Faculty of Biological Science, within 3–10 days after an outbreak. Some of the current biosecurity strategies used
Jashore University of Science and Technology,
to keep diseases out of shrimp ponds such as pond water disinfection, quarantine of
Jashore 7408, Bangladesh.Email:
6378506331@student.chula.ac.th new stocks before stocking and broader usage of specific pathogen-free shrimp. The
sequencing and characterisation of various WSSV strains have provided details about
pathogen biology, pathogenicity and disease. To develop successful control methods,
knowledge of these characteristics is essential. In several shrimp-producing countries
in Asia and the Americas, the infections produced by the WSSV have had disastrous
socio-economic consequences. As a result of international trade or migration of dis-
eased species, the World Animal Health Organization recognised several illnesses
as posing a substantial hazard to farmed shrimp. WSD is receiving much scientific
research due to the potential economic effects of the virus. Research is now being done
to understand better the molecular biology and pathophysiology of WSSV, as well as
how to treat and prevent the virus. However, further study should be conducted in
countries with more resilient host species to understand their role in mitigating disease
impacts since these revelations may aid in developing a WSD treatment.

KEYWORDS
biosecurity, diagnosis, economy, pathogenicity, transmission, WSSV

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided
the original work is properly cited.
© 2022 The Authors. Veterinary Medicine and Science published by John Wiley & Sons Ltd.

778 wileyonlinelibrary.com/journal/vms3 Vet Med Sci. 2023;9:778–801.

ISLAM ET AL . 779

1 INTRODUCTION out the disease’s fundamental causes and finding potential remedies
for disease control or mitigation (Clay, 1997; C. F. Lo, 2014; Nguyen
The pathogen that causes white spot disease (WSD) in cultured et al., 2009; Sánchez-Paz, 2010). Cultured shrimp have a maximum
penaeid shrimp is the White Spot Syndrome Virus (WSSV). This virus life span of 4–5 months. Shrimp must tolerate increasingly varied cir-
is a solitary individual from the Nimaviridae virus family, which was cumstances, including temperature and salinity fluctuations, during the
recently discovered (Pradeep et al., 2012). WSSV is a big double- culture stage (Broom, 2009; Joseph et al., 2013). Excessive salinities
stranded DNA baculovirus, and infection with the WSSV pathogen in are a significant environmental stressor that may increase shrimp sus-
shrimp farming can reach a 100% death rate in 3–10 days (Oakey et al., ceptibility to viral diseases (Abad-Rosales et al., 2018; Ramos-Carreño
2019). The white dots on the carapace of shrimp infected with WSSV et al., 2014). Additionally, genetic markers resulting from mutations
are immediately identifiable (Saravanan et al., 2017). On the other might be a possible cause of the molecular diagnostic error due to inter-
hand, the initial clinical signs arise after the animal has been infected ference of PCR primer binding, which could result in non-specific PCR
and is near death. As a result, diagnosis methods have evolved from products or incorrect negative PCR findings, limiting the utility of PCR-
morphological-based characterisation utilising electron microscopy based therapeutic agents (Mendoza-Cano and Sánchez-Paz, 2013a; X.
(EM) to exceedingly effective immunological and molecular methods Pan et al., 2016). Despite the hurdles posed by its unique genotypes and
that previously could only recognise the infection in asymptomatic car- infection process, which differs significantly from that of other viruses,
riers (Curry et al., 2006b). White spots, then again, may not appear in much progress has been made in WSSV research (L. Li et al., 2006;
all diseases or all species (Md Rahman, 2021). Witteveldt et al., 2004). Researchers revealed the complete genome
WSSV was initially discovered in farmed Penaeus japonicus in Japan; sequencing of the WSSV (2000) (van Hulten et al., 2001). The struc-
however, it is believed that it was brought with live diseased post- tural proteins of the virion, the genes that control their production
larvae (PL) from Fujian Province, Mainland China, in 1992 (Jiang et al., and their function in the host species have been identified (Chaivi-
2017; W.-B. Zhan et al., 1998). Subsequently, the virus soon spread suthangkura et al., 2010; Sánchez-Paz, 2010; Wan et al., 2008). Other
through most of Asia’s shrimp-producing countries, most likely due to research has looked into the virus’s transfer to the host body from
infected broodstock and PL of Penaeus monodon. The virus entered the one species to another, as well as the role of physicochemical qualities
Pacific Ocean in 1995, possibly due to the transfer of infected PL, as of water in infection (Selvam et al., 2012). WSD, like most other viral
it was discovered in Texas, North America and South Carolina, USA, infections, cannot be treated at this time; hence, it is recommended
1 year later (Rosenberg, 2000; Schalie, 1999). The virus did not make that the virus be kept out of shrimp farms (Menasveta, 2002). Several
it to the Latin American pacific shrimp farming countries until 1999, biosecurity strategies, antibiotics and vaccinations have been inves-
when it wreaked havoc in Ecuador, Peru and Mexico. Shrimp farming tigated to increase shrimp behaviour to WSSV infection (Dey et al.,
has been conducted on a small scale in coastal Southeast Asia for gen- 2019; Feng et al., 2017). Specific pathogen-free (SPF) shrimp brood-
erations when farmers grew wild shrimp as a side crop in tidal ponds stock domestication, insignificant water exchange and culture pond
(Kungvankij and Fi 1984; J. Lotz, 1997). dry-out procedure after each harvesting period are different biosecu-
Since the disease’s occurrence, the economic losses associated with rity methods that have been used to avoid the introduction of WSSV
the disease condition to the shrimp sector have been predicted to be (D. V. Lightner, 2005; Tendencia et al., 2011).
eight to fifteen billion dollars worldwide (D. Lightner et al., 2012; Shinn Most WSSV reviews have focused on advanced genetic analyses
et al., 2018). The financial losses had already been growing at a rate and biosecurity measures used in WSD maintenance (Bir et al., 2017;
of USD 1 billion every year (Dashtiannasab, 2020; Escobedo-Bonilla Dey et al., 2019; B. Dieu et al., 2021; Jana et al., 2018). On the other
et al., 2008; FAO, 2014). WSSV was blamed in China for 80% of farmed hand, shrimp farming practices are evolving, and recently, certain inno-
shrimp production losses (Briggs et al., 2004; Salehi, 2021) and the vative technologies have been applied to raise yields and better WSD
influence of WSSV on Ecuador’s farmed shrimp output was likewise monitoring. New experimental molecular detection methodologies, the
severe (Lucien-Brun, 2017). The transmission of WSSV to several other impact of WSSV in shrimp farming, development of SPF shrimp, the
shrimp-producing nations threatens the industry’s development. As a advantage of specific pathogen-resistant (SPR) shrimp, the function of
result, WSD jeopardises shrimp businesses such as farms, feed manu- WSSV viability owing to physio-chemical changes in pond sediment
facturing, processing factories and hatcheries, increasing employment. and their potentiality in dealing with the WSSV scourge in shrimp
Even though WSD has been causing havoc in shrimp farms around the production are all highlighted in this study.
world since 1991, surprisingly, the illness is still not within grip at the
farm level (Bondad-Reantaso and Subasinghe 2001; Flegel, 2009; D.
Lightner, 2011; D. V. Lightner and Redman, 2010; S Salim, 2012; Walker 2 WSSV GENOME
and Mohan, 2009).
WSSV infection and disease can affect a wide variety of crus- The genome of WSSV is a circular dsDNA molecule that is quite possi-
taceans. These hosts’ natural populations can also serve as a reservoir bly the most comprehensively sequenced viral genome (B. Dieu, 2010;
for the infectious agent. Given the severity of WSD in captive crus- Parrilla-Taylor et al., 2018; Vlak et al., 2004). Depending on the viral iso-
taceans, it is hardly unexpected that much effort has gone into figuring late, the genome size varies. Thailand, China and Taiwan had genome
780 ISLAM ET AL .

sizes of 292,967, 305,107 and 307,287 bp, respectively, according to

three total WSSV variants (accession numbers AF369029, AF332093
and AF440570) (Kang et al., 2009; Megahed et al., 2019; Sablok et al.,
2012). According to nucleotide sequence analyses, about 185 open
reading frames (ORFs) of 50 amino acids or more are encoded by
the WSSV genome. After successfully confirming the WSSV genomic
DNA sequence, researchers focus on quantitatively studying genetic
products, specifically the role of viral envelope proteins. A small num-
ber of functional and DNA metabolic genes have been found and
characterised (de-la-Re-Vega et al., 2011; Guevara-Hernandez et al.,
2012).

3 WSSV VIRION PROTEINS FIGURE 1 Structure of WSSV

A virion is a sophisticated assemblage of macromolecules adapted

to protecting and delivering viral DNA. Its structural proteins have
been highly significant, as they are the primary particles to inter- VP51A, VP51B, VP68, VP124, VP150, VP187, VP281, VP292 and a
face with the host, performing cell targeting functions and stimulat- collagen-like protein (L. Li et al., 2005), whereas the proteins VP35 (L.-
ing most defences (Tsai et al., 2004). Structural proteins and their L. Chen et al. 2002), VP466 (C. Huang et al., 2002), VP15, VP51 and
genetic sequence must be characterised to determine a virus’s taxo- VP76 (C. Wu and Yang, 2006) are present in the nucleocapsid (Yang
nomic categorisation. The structural function and collaboration of the et al., 2002). VP466 is one of these proteins that has been discovered
virion proteins of WSSV may clarify the virus’s unusual morphologi- to play a crucial function in viral penetration (W. Wu et al., 2005). P466
cal attributes. In response, diagnostic assays focused upon a few of is a glutathione S-transferase fusion protein denoted by ORF151 and
these cellular components may be created (Yang et al., 2002). Various is one of the latencies associated with the genes in WSSV (X. Xie et al.,
WSSV proteins have been recognised. Non-structural proteins have 2006). ORF112 encodes VP76, and it is implicated in WSSV infection. It
been distinguished as being engaged with transcriptional regulation involves the conserved domain of Class I cytokine receptors (Yang et al.,
(VP9) (Y. Liu et al., 2006), viral multiplication (WSV 021) as well as DNA 2002). ORF112 is a 2025-nt quality that codes for a 675-aa protein
replication regulation (WSV 477). The envelope contains 21 proteins, with a molecular mass of 76 kDa (R. Huang et al., 2005). ORF112 has
the nucleocapsid contains 10 proteins and the tegument contains five a signal peptide and multiple glycosylation sites but no transmembrane
proteins. The viral genome replication, virus particle production and domain, according to phylogenetic analyses. This suggests that WSSV
cell function inhibition depend on non-structural proteins found in the has designed techniques to circumvent the host’s defence mechanism
WSSV genome. As a result, these proteins are promising candidates (Table 1).
for medical research and vaccine development. Western blotting, mass
spectrometry and immunoelectron microscopy were used to identify
VP9, a complete WSSV protein encoded by ORF115 in diseased P. 4 MORPHOLOGY AND STRUCTURE OF WSSV
monodon shrimp gills and stomach as a novel, non-structural protein.
Even though the specific physiological role of VP9 is unknown, inves- WSSV has a diameter of 80–120nm and a length of 250–
tigations have shown that it is a common protein in host tissue which 380nm (Durand et al., 1996). The virions contain a distinct flagella-like
is WSSV (Y. Liu et al., 2006). VP9 has a DNA binding fold containing appendage at one point and are rod-shaped to elliptical (Walker and
Zinc ion active site that is distinct according to structural investigations Mohan, 2009). At least 45 structural proteins are found in virions,
using X-rays and NMR (Banci et al., 2002; Rosenzweig et al., 1999). which are organised into three morphologically different layers (Tsai
These findings suggest that VP9 may have a role in WSSV transcrip- et al., 2004). The virions proliferate without producing occlusion
tion. Because envelope proteins are commonly involved in viral entry, bodies inside the nuclei of infected cells. The WSSV virus attacks
assembly and budding, they are especially critical for enveloped viruses ectodermal and mesodermal tissues (Jesudhasan et al., 2000). The
(Chazal & Gerlier, 2003). A cell attachment motif is present in the enve- nucleocapsid proteins include a basic DNA binding protein (VP15)
lope proteins VP31, VP110 and VP281. This motif is involved in the and a big protein (VP664) that make up the stacking ring components
viral entrance Single-cell attachment motif can also be found in the (Verbruggen et al., 2016; Witteveldt et al., 2005). The viral envelope
tegument proteins VP36A and the nucleocapsid proteins VP664 (Leu does have a cellular structure that is 6–7 nm thick with 35 distinct pro-
et al., 2005;Tsai et al., 2004) and VP136 A (C. Huang et al., 2002; Tsai teins, the most numerous of which are VP28 and VP26, which account
et al., 2004; X. Xie et al., 2006). Other proteins discovered in the enve- for around 60% of the envelope (Hulten, 2001; Sánchez-Paz, 2010)
lope include VP28 (Yoganandhan et al., 2006), VP39B, VP31A, VP41B, (Figures 1–3).
ISLAM ET AL . 781

TA B L E 1 White spot syndrome virus (WSSV) proteins that have been identified so far

Amino acid Apparent Location in WSSV

Protein names residues size size (kDa) Putative function virion References
VP9 79 9 Transcriptional Non-structural Li et al. (2005)
VP11 433 11 – Not determined (Tsai et al., 2004)
VP12A (VP95) 95 11 Structural Tegument (X. Xie et al., 2006)
VP12B (VP68) 68 7 Structural Envelope (C. Wu and Yang 2006)
VP13A 100 13 Energy metabolism Not determined (Tsai et al., 2004)
VP13B (VP16) 117 13 Structural Envelope (X. Xie et al., 2006)
VP14 97 11 Structural Envelope (X. Xie et al., 2006)
VP15 80 15 DNA binding protein Nucleocapsid/core (Hulten and Vlak 2021)
VP19 121 19 Structural Envelope (Tsai et al., 2004)
WSV021 200 23 Regulation virus Non-structural1 (Zhu et al., 2007)
replication
VP22 (VP184) 891 100 – Not determined (Tsai et al., 2004)
VP24 (VP208) 208 24 Structural Nucleocapsid (M. C. W. Hulten et al., 2000)
VP26 204 26 Structural Tegument (Tsai et al., 2004)
VP28 204 28 Structural Envelope (M. C. W. Hulten et al., 2000)
VP31 261 31 Cell attachment Envelope (X. Xie et al., 2006)
VP32 278 32 Structural Envelope (L. Li et al., 2005)
VP33 (VP 281) 281 32 Cell attachment Envelope (L.-L. Chen et al., 2002)
VP35 228 26 Structural Nucleocapsid (X. Xie et al., 2006)
VP36A 297 36 Cell attachment Tegument (C. Huang et al., 2002)
VP38A 309 35 Structural Envelope (Tsai et al., 2004)
VP38B 321 38 Endonuclease Not determined (X. Xie et al., 2006)
VP39A 419 39 Structural Tegument (Tsai et al., 2004)
VP39B 283 32 Structural Envelope (X. Xie et al., 2006)
VP41A (VP292) 292 33 Structural Envelope (X. Xie et al., 2006)
VP41B (VP300) 300 34 Structural Envelope (X. Xie et al., 2006)
VP51A 486 51 Structural Envelope (Edgerton, 2004)
VP51B (VP384) 384 46 Structural Envelope (X. Xie et al., 2006)
VP51C (VP466) 466 50 Structural Nucleocapsid (Tsai et al., 2004)
VP53A (VP150) 1301 144 Structural Envelope (X. Xie et al., 2006)
VP53B 968 53 Signal transduction Not determined (Tsai et al., 2004; X. Xie et al., 2006)
pathway
VP53C 489 53 – Not determined (X. Xie et al., 2006)
VP55 (VP448) 448 55 – Not determined (Tsai et al., 2004)
VP60A (VP56) 465 60 Structural Envelope (Ramırez-Douriet et al., 2005)
VP60B (VP544) 544 60 Adenovirus fibre-like Nucleocapsid (Tsai et al., 2004)
protein
VP75 786 75 Structural Nucleocapsid (X. Xie et al., 2006)
VP76 (VP73) 675 76 Class 1 cytokine Nucleocapsid (X. Xie et al., 2006)
receptor
VP90 856 96 Structural Envelope (Tsai et al., 2004)
VP95 800 89 Structural Tegument (X. Xie and Yang 2006)
VP110 972 110 Cell attachment Envelope (Zhang et al., 2004)
VP124 1194 124 Structural Envelope (Tsai et al., 2004)

(Continues)
782 ISLAM ET AL .

TA B L E 1 (Continued)

Amino acid Apparent Location in WSSV

Protein names residues size size (kDa) Putative function virion References
VP136A 1219 136 Cell attachment Nucleocapsid (Tsai et al., 2004)
VP136B 1243 136 – Not determined (Tsai et al., 2004)
VP180 (VP16840) 1684 169 Collagen-like protein Envelope (Tsai et al., 2004)
VP187 1606 174 Structural Envelope (Tsai et al., 2004)
VP190 1565 174 Structural Nucleocapsid (X. Xie and Yang 2006)
WSV477 208 30 DNA replication Non-structural (Han and Zhang, 2006; Shi et al., 2008)
VP664 6077 664 Cell attachment Nucleocapsid (Tsai et al., 2004)
VP800 800 90 – Not determined (Tsai et al., 2004)

F I G U R E 2 The structure of a WSSV nucleocapsid has been drawn based on the information supplied in the morphological section. (a) The
WSSV nucleocapsid, which has 15 prominent vertical helices, was examined using electron microscopy. (b) A cross section of a single nucleocapsid
helices with a diameter of 70 nm (side view). (c) An empty nucleocapsid (cross section) having an 80 nm diameter (side view). The nucleocapsid’s
cross-hatched structure is a distinctive WSSV identifying characteristic (Verbruggen et al., 2016).

5 ORGANS OF TARGET AND MECHANISMS OF japonicus revealed that epithelium in the midgut trunk might serve as a
TRANSMISSION transitory site for WSSV replication, enabling the virus to circumvent
the surrounding basal lamina (Leonardo et al., 2005; S. Xie et al., 2015).
Ectodermal and mesodermal targets for WSSV transmission include WSSV challenge by immersion, on the other hand, revealed that at late
the skin, gill, antennal gland, hindgut, foregut, gonads, haematopoietic stages of infection, haemocytes moving to the gills and midgut were
cells, lymphoid organ and cells linked with the nervous system (A and WSSV negative (Arts et al., 2007). Cells in the antennal glands, as well
R, 2010; Kv et al., 1999). Endodermal organ epithelial cells, such as as epithelial cells in the foregut and cells in the gills, were revealed to be
those in the hepatopancreas, anterior and posterior midgut caeca and foci of WSSV replication when shrimp were per os challenged with viral
midgut trunk, are resistant to WSSV infection (Sahul Hameed et al., proteins (with a high infectious dosage) (Escobedo-Bonilla et al., 2008).
1998). The epithelia of the stomach, gills and integument may be exten-
sively injured in the late stages of illness. This could result in several
organ malfunctions and death in the worst-case scenario. The portals 6 DIAGNOSTIC METHODS OF WSSV
by which WSSV enters the shrimp have yet to be discovered. Varia-
tions in entrance sites have been discovered in studies. However, in situ Several diagnostic methods have been developed to identify WSSV.
hybridisation demonstrated that the most common locations of WSSV Gross observation, histopathological techniques (Reyes-López et al.,
reproduction in early juvenile P. monodon were subcuticular epithelial 2009), in situ hybridisation (D. Lightner et al., 2012), immunologi-
cells of the stomach and cells in the gills, integument and connective cal methods such as nitrocellose-enzyme immunoblot (Nadala et al.,
tissue of the hepatopancreas (upon os challenge using WSSV dam- 1998) and Western blot techniques (Durand et al., 1997) and poly-
aged tissue) (P. S. Chang et al., 1996). Further studies in Marsupenaeus merase chain reaction (PCR)-based approaches have lately become
ISLAM ET AL . 783

tions (mimiviruses, poxviruses and some iridoviruses), optical micro-

scope decision is insufficient to identify viruses; however, transmission
electron microscopy (TEM) resolution is a thousand times greater than
optical microscope resolution, enabling rapid identification of micro-
scopic viruses. TEM, on the other hand, can only identify the virus up
to the family or genus level; however, this is a rapid procedure (Curry
et al., 2006a). Even though the detection limit (DL) of viruses was origi-
nally 106 viruses per ml when using TEM alone, this limit has now been
reduced to 102 viruses per ml when employing the filtering approach
(Laue and Bannert, 2010). There are several benefits to utilising EM,
including remembering the capacity to see everything, for example
(Laue & Bannert 2010), the norm for the distinguishing proof of new
viral diseases (Biel and Gelderblom, 1999), identification of double
infections involving several viruses (C. Goldsmith and S. Miller, 2009;
Kennedy, 2005) and the capacity to rapidly recognise morphological
differences (C. Goldsmith and S. Miller, 2009; Kennedy, 2005). As a con-
sequence, EM can be coupled with multiple approaches and employed
as a cutting-edge method (Hazelton and Gelderblom, 2003).
F I G U R E 3 Some of the ORFs and the dominant envelope protein
coding gene VP28 are shown in the genome sequence of WSSV from
Thailand (WSSV-TH).
6.3 Immunological techniques

more accessible, sensitive and reliable (T. Kim et al., 1998). The sen- Employing western blot analysis, dot blotting and immunohistochem-
sitivity of these techniques varies depending on whether they are istry, specific monoclonal antibodies (MAbs) have been shown to
DNA-based. External appendages can be used to detect preliminary discriminate WSSV in shrimp while avoiding cross-reactions with other
WSSV. The inside surface of the carapace of severely infected shrimp viral or shrimp proteins (Vaniksampanna et al., 2016); Specific MAbs,
often has a loose cuticle with white spots ranging from 0.5 to 2.0mm in on the other hand, can bind to the WSSV target protein. For instance,
diameter (Hossain et al., 2015). The cuticular epidermis deposits cal- MAb targeting VP28 could only bind to WSSV VP28 and not to
cium salts abnormally, resulting in this white spot. The expansion of any other viral or shrimp antigens (Marappan et al., 2006). Com-
the cuticular chromatophores gave shrimp a pink to reddish-brown pared with a single-step PCR, these immunological approaches have
coloration in many cases. a higher diagnostic sensitivity (100%) (PCR). Dot blotting, a well-
known antibody-based protein detection approach, on the other hand,
is reported to require sufficiently large viral loads for detection (Sithig-
6.1 Histopathology orngul et al., 2011). Two MAb cocktails were 100 times more accurate
than one-step PCR and approximately equivalent to two-step PCR
Davidson’s fixative should be injected into live moribund prawns from detecting WSSV, focusing on its VP28 envelop protein. Additionally,
a probable WSSV epidemic. This extremely acidic fixative decalcifies when a MAb cocktail was used instead of a single MAb, the DL of
the tissue, making it easier to process and section. Traditional fixative WSSV was increased by two to four times (Inchara et al. 2018). In sev-
solutions, including 10% neutral buffered formalin, do not decalcify eral antibody-based tests, scientists recommended employing a mix of
or penetrate the shell, resulting in poor fixing and cutting difficulty. VP19 and VP28 (virion proteins) targeted MAbs to boost the speci-
Tissues can be repaired indefinitely without losing quality if the diag- ficity of WSSV diagnosis in shrimp (Chaivisuthangkura et al., 2010).
nosis is based on histology. However, after 24–48 h, specimens for An immunosensor has been designed to detect WSSV in shrimp pond
ISH should be changed from Davidson’s fixative to 70% ethyl alcohol, water quantitatively. That immunosensor was particularly sensitive for
or they will be inappropriate for ISH. The specimens are embedded testing WSSV in the linear range 1.6 × 10(1) – 1.6 × 10(6) copies/μl.
in paraffin wax and processed using standard procedures after fixing Moreover, the immunosensor was reusable up to 37 times and was
them. WSSV infection is indicated by the inclusion of bodies in target straightforward (Waiyapoka et al., 2015). Another study used gold
tissues(Reyes-López et al., 2009). nanoparticles coupled to a polyclonal anti-VP28 antibody, lateral flow
immunoassay (LFIA) was shown to be a rapid and convenient approach
for detecting WSSV in under 20 min (Underwood et al., 2013). Differ-
6.2 Electron microscopy ent shrimp viruses, such as infectious hypodermal and hematopoietic
necrosis virus (IHHNV), hepatopancreatic parvovirus (HPV) and mon-
Generally, EM seems to be the first line of defence regarding viral iden- odon baculovirus (MBV), showed no cross-reactivity with the LFIA (P.
tification (C. S. Goldsmith and S. E. Miller, 2009). With several excep- Kulabhusan et al., 2017). To date, the double antibody enzyme-linked
784 ISLAM ET AL .

immunosorbent assay (ELISA) has been used to detect specific mAbs evaluation of reaction products on a gel, this sort of assay needs more
and capture polyclonal antibodies that can identify various epitopes to hands-on time than the others.
quantify antigens at the protein level (Y. Chang et al., 2016; Mecham, It is an audit method that is the gold standard recommended for use
2006). These have been proven to be a reliable and sensitive approach in WSSV inspections in shrimp. After following the first-step PCR and
for detecting viral infections, and they might also be used to identify second step of the (nested) PCR, which relies on two pairs of primers,
WSSV in shrimps (X. Tang et al., 2018). In the Escherichia coli medium, the reading is performed by the electrophoresis gel method. The pos-
Murugan and Sankaran proposed a new ELISA approach for evalu- itive results in the first episode of the standard method criteria refer
ating recombinant bacterial lipid change in proteins (Murugan and to the infection of the severe white spot virus, while when the positive
Sankaran, 2018). In that study, the most frequently generated white results are obtained in the second period, it indicates that it is a latent
spot syndrome viral protein ICP11 was found as a new target in a novel or carrier infection (Nunan & Lightner, 1997). In addition to reading
ELISA method using a lipid-modified protein. The scientists discovered results under UV light or reading results with DNA strips, PCR out-
that this approach was a predictive factor of antigen in contaminated put can also be evaluated with loop-mediated isothermal amplification
shrimp tissues, with a DL of 250 pg for ICP11 protein and a dynamic (LAMP), providing easy and convenient readings, especially in cases
range of 15–240 ng. The benefits of this test are that it has a low assay of non-operational analysis, using a processing time of approximately
requirement, can be detected with the unaided eye and is the budget 60 min (Kono et al., 2004). After the PCR output has been acquired
(Murugan and Sankaran, 2018) yet, the researchers did not mention if from regular episodes, enter the LAMP technique, where the transla-
the detecting method is swift or takes a long time. Antibody-based indi- tion can be determined by the turbidity from the white sediment of
rect immunofluorescence is another approach commonly employed in magnesium pyrophosphate. If the gene is enlarged, white sediments
virological studies, though it is less sensitive than antibody-based indi- are found. In addition, it is considered fluorescent paint using fluores-
rect immunofluorescence. In a study, indirect immunofluorescence’s cent substances such as SYBR Green I, filled after the lamp technique
sensitivity and specificity were determined to be 29 and 96%, respec- will change the default colour (orange) to green when viewed under UV
tively, compared with PCR (Burgesser et al., 1999). As a result, the light (302 nm) (Marshall and Johnsen, 2017).
indirect immunofluorescence approach necessitates a huge density
of viral particles in the host tissue, making virus detection at an
early stage of infection impossible (Mott et al., 1997). Finally, despite 6.4.2 Quantitative real-time PCR
the many advantages of immunological WSSV detection techniques
outlined above (e.g., highly sensitive, specific and precise diagnosis) It is a technique to increase the amount of genetic material, which
(Poulos et al., 2001), these approaches are often expensive, time con- can measure the amount of genetic material based on fluorescence,
suming and need specialised equipment and expert workers making starting with mRNA and then turning it into cDNA using reverse tran-
them unsuitable for use in the field (Poulos et al., 2001). Recently, a scriptase enzymes. Increased DNA content Studies in shrimp often
LFIA was developed using gold nanoparticles and a polyclonal antibody used plasmids containing the genome part of the WSSV (U50923) for
targeting WSSV VP28’s main envelope protein. Through a simply spe- positive control and 10× diluted sensitivity tests to contain copies
cialised up-gradation of the approach for selective virus detection, this in the range of 2 × 105 copies in standard curves and often using
method was reported to concurrently detect WSSV, IHHNV (infectious TaqMan and FAM-BHQ1 probes (S. Durand and Lightner, 2002). In
hypodermal and haematopoietic necrosis virus), MBV, HPV (human real-time, PCR has a free fluorescence meter during the polymerase
papillomavirus) and others. Furthermore, the approach is relatively less DNA reaction, which digests fluorescent labels to fall out of the
expensive (USD 3 per sample), and the farmer can execute the test primer, becoming a free fluorescent colour. Independent measure-
without the assistance of an outside professional, reducing the farmers’ ments of variations with genetic material content in WSSV diagnosis
financial burden significantly. Consequently, the LFIA might provide have applied the qPCR technique (Mendoza-Cano and Sánchez-Paz,
practical assistance to farmers, fish farm operators and fisheries (P. K. 2013b; Y. L. Tsai et al., 2014). By relying on the reverse transcrip-
Kulabhusan et al., 2017). tase enzyme called recombinase polymerase amplification (RPA), this
reaction can be performed in one step, reducing the step for cDNA
preparation, temperature is also used by RPA (32-42◦ C), which is lower
6.4 Molecular techniques than that of conventional PCR. The number or synthesis of DNA rep-
resents a reaction that can happen even at normal temperature, so
6.4.1 Nested PCR this method is a quick way to diagnose a disease (Lobato & O’Sullivan
2018), the qPCR technique was also developed, can be done at the
To improve the efficacy and accuracy of one-step PCR, the findings of same time. The duplex qPCR technique connects the two. By using only
the first experiment can be amplified further in a nested second cycle 2–20 DNA concentrations in the plasmid copies/reaction and 2 × 105
of PCR (Peinado-Guevara and lopez-meyer, 2006). Since it includes copies/diagnosis of WSSV and IHHN, respectively, is considered high
two steps in the construction of the assay and typically requires the diagnostic sensitivity and specificity (K. F. Tang and Lightner, 2001).
ISLAM ET AL . 785

6.4.3 Fluorescent quantitative PCR 6.4.6 Insulated isothermal PCR assays

Another approach for quantifying DNA is quantitative fluorescence Each iiPCR test employs a unique PCR format that only necessitates
PCR (FQ-PCR), where the result is decided by the concentration of the usage of a simple source of heat to complete the PCR steps. It is
DNA templates having fluorescent sequences (Bustin 2000). RT-PCR is used in a portable nucleic acid analyser created especially for iiPCR
more expensive than this approach (Ririe et al., 1997); however, unlike and can detect products automatically, allowing for WSSV identifica-
dye-based qPCR, any type of double-strand DNA can produce fluo- tion at the pond’s edge. This is comparable to real-time qPCR in that it
rescence, making it less sensitive than TaqMan RT-PCR (Ririe et al., produces high-sensitivity data instantly with little or no post-reaction
1997), TaqMan probes, on the other hand, are employed to improve the processes (Wilkes et al., 2018).
RT-PCR sensitivity (Holland et al., 1991). The test becomes more efficient and consumer friendly as product
signals are automatically read, reducing human errors and data inter-
pretation conflicts. iiPCR experiments take about 1.5 h to complete.
6.4.4 Loop-mediated isothermal amplification Reagents and equipment for iiPCR assays are less expensive than those
for real-time qPCR tests. iiPCR tests, on the other hand, do not produce
The LAMP method is an innovative, sensitive and fast technology used quantitative results, and the machines can only execute eight reac-
in aquaculture to diagnose illness. The LAMP method developed a high tions per run. The OIE recently certified the I.Q. Plus WSSV Kit with
sensitivity and diagnostic specificity test for identifying the WSSV. The POCKIT System, the only iiPCR-based assay for WSSV, is appropriate
WSSV genome DNA was used to build a set of four primers, two outside for diagnosing WSSV.
and two interior primers. The reaction time and temperatures were
set at 1 h at 65◦ C. As contrasted to 10 fg using nested PCR, the DL
of the LAMP approach was up to 1 fg (PCR). As a response, a stan- 6.4.7 One-step PCR assay
dard LAMP technique was used to detect WSSV in the heart, stomach
and lymphoid organs of diseased shrimp (Notomi et al., 2000). The Traditional one-step PCR procedures only amplify target DNA at a
LAMP process is an auto-cycling strand displacement DNA synthesis basic level, resulting in lower specificity than other PCR techniques. Gel
that uses a high-strand displacement DNA polymerase and a set of spe- electrophoresis, which comprises the separation of products through
cialised primers to recognise six different patterns on target DNA. As a a gel of specific pore sizes in an electrical field (gel electrophoresis)
result, it is intended to amplify the target sequence selectively. The final and the staining and visual detection of product bands on the gel with
LAMP outputs are a mix of diverse stem-loop DNA lengths (Notomi the use of a fluorescent dye, is the most common method of product
et al., 2000). When the LAMP results are observed using agarose gel detection (Olive and Bean, 1999).
electrophoresis, numerous bands of various widths are visible up to the It usually takes 2–3 h to complete target amplification and product
loading well. detection. In addition to being labour intensive and time consuming,
these techniques carry the danger of cross-contaminating amplicons
in future assays, resulting in false-positive results. On the other hand,
6.4.5 Pre-amplification PCR experiments utilising this format are usually less expensive than other
PCR assays.
Researchers recently established a new pre-amplification PCR
approach, in which numerous short-strand sequences were employed
as adapters to ligate to the WSSV restriction pieces VP28, VP39 and 6.4.8 Duplex real-time PCR
VP108 via a link (X. Pan et al., 2017). Following that, using a universal
primer, the ligated product is used as a template for pre-amplification A duplex qPCR was standardised for WSSV and PstDV1 identification
PCR. The pre-amplification PCR considerably enriches the target in clinical specimens employing valid and reliable primers and probes.
DNA, and the pre-amplified product is then used as a template for Utilising the improved duplex qPCR settings, singleplex assays had
the particular PCR that completes the ultimate diagnosis. Such a specificities and sensitivities similar to those of multiplex tests, allow-
non-quantitative diagnosis methodology seems more accurate than ing the diagnosis of both singular and coinfections in infected white leg
nested PCR and can be used to diagnose WSSV at an early stage (X. shrimp. Because such coinfections in shrimp are widespread, the con-
Pan et al., 2017). The sensitivity of the PCR process, however, is greatly current identification of the WSSV and the PstDV1 is beneficial (Leal
influenced by the method of DNA extraction, template concentration et al., 2014).
and amplicon size. A positive PCR test that finds a DNA fragment The analytical sensitivity of the duplex qPCR assay for WSSV and
is also a poor predictor of viral opportunity (Cha and Thilly, 1993). PstDV1 was comparable to that of singleplex assays. According to the
Furthermore, in most situations, these tests necessitate long-distance researchers, this duplex qPCR technique discovered 10- and 627-fold
sample transfer from the area to the laboratories. And these tests lower quantities of PstDV1 DNA than traditional duplex and multi-
usually come at a high cost, take a long time and necessitate highly plex PCR procedures (Leal et al., 2014). Moreover, the duplex qPCR
skilled laboratory employees and devoted research facilities. test’s WSSV DL was onefold lesser than multiplex PCR techniques and
786 ISLAM ET AL .

10 copies lower than multiplex qPCR techniques. Several studies have

shown that the standardised duplex qPCR approach outperforms other
conventional and real-time PCR-based approaches. Furthermore, the
standardised duplex qPCR assay’s lower DL makes it a therapeutically
relevant tool for identifying animals with low virus loads, such as those
found in shrimp larvae and PL, minimising the risk of false-negative
results (Leal et al., 2014).

6.4.9 Optimised PCR assay

Based on the nested PCR procedure described by Chu Fang Lo et al.

(1996), a rapid PCR assay for the detection of WSSV was developed
and outlined as the recommended PCR diagnostic assay in the Man- FIGURE 4 Schematic diagram of WSSV detection method
ual of Diagnostic Tests for Aquatic Animals published by the Office of
International Epizootics. The second-step primers utilised in the nested
WSSV PCR were added to the optimised process. This modified assay food; (ii) cutaneous transmission via infected organisms or contami-
is significantly faster and as sensitive as the recommended OIE pro- nated food and (iii) cutaneous transmission via infected organisms or
cedure after lowering the annealing temperature and decreasing the contaminated food. Furthermore, the virus can be transmitted through
cycling periods. The two-step nested PCR methodology and a modified the consumption of affected tissues by parasitism and cannibalism, (ii)
nested process were directly compared to the modified PCR test. The through waterborne transmission, the virus enters the body through
sensitivity of the published assay was tested using template dilutions of the gills and external areas via direct contact with virus particles in
semi-purified WSSV virions that had been quantitated using real-time water and (ii) through airborne transmission via the lungs and exter-
PCR for WSSV detection. The modified technique was used to exam- nal areas by close contact with virus particles in water (Thuong, 2016)
ine various isolates to ensure that the assay could detect WSSV from and (iii) through vertical transmission (via oocytes) from an infected
various geographical regions. broodstock to offspring (De Gryse et al., 2020; Sánchez-Paz, 2010).
According to various findings, the transmission rates in both P. van-
namei and P. monodon species are similar. The authors believe, however,
6.4.10 Fluorescence in situ hybridisation that the major impact of direct and indirect transmission rates might
be different (Tuyen et al., 2014). Although pathogens have been found
It is a method of detecting DNA that contains nucleoli. The tidying is in oocytes in mature eggs they have not been found. This could signify
specific to cells that show signs or suspected infection. DNA probes that the virus-infected oocytes are unable to develop into ovarian fol-
of approximately 50–100 nucleotides and peraluminous substances licles (Y. G. Wang et al., 2000). The transportation of live and frozen
are attached. The probe will be able to capture the target nucleotide uncooked shrimp can help transmit WSSV between different geo-
acid that contains nucleotides that match the DNA probes within graphical locations (S. V. Durand et al., 2000), and broodstock imported
the cells to translate the results into consideration. From the appear- from different countries and regions (Stentiford et al., 2009). Environ-
ance of labelled colours on the probe, which requires the fluorescent mental factors such as temperature, fluctuation in salinity and pH have
microscope (Nunan & Lightner, 1997), with white colour with A WSSV been identified as major stressors that can influence WSSV infection
virus infection, appears dark silver sediments. In addition to diagnosing transmission and outbreaks, according to several studies (Van Thuong
viruses, fluorescence in situ hybridisation (FISH) is also used to study et al., 2016; Yu et al., 2006). Several studies in Thailand were under-
types of bacteria. The liver membranes, fillings and water are used to taken to assess the impact of weather conditions on WSD infection,
treat shrimp. FISH is a highly sensitive and cultured method. However, and the results showed a higher incident ratio of disease at the time of
it is not suitable for diagnosis or diagnosis. As such, a PCR method has mean ambient temperature fluctuating from 24.5 to 27.23◦ C, based on
been developed to analyse abnormalities at a specific genetic level and the data gathered (Gronski, 2000; Nakorn et al., 2017; Piamsomboon
is very sensitive (Bennour et al., 2016) (Figure 4 and Tables 2 and 3). et al., 2016).

7 TRANSMISSION 8 CLINICAL SYMPTOMS AND ESSENTIAL

NATURE OF WSSV
WSSV can spread horizontally and vertically (Leu et al., 2008; Prior
et al., 2003; Soowannayan and Phanthura, 2011). There are three WSSV takes a long period to manifest, and if it does, infected animals
ways for the virus to spread among shrimp and other decapod crus- die between 3 and 8 days, leading to a higher fatality rate (Corbel
taceans: (i) oral transmission via infected organisms or contaminated et al., 2001). WSSV-infected shrimp assemble at the pond’s side in the
ISLAM ET AL . 787

TA B L E 2 Summary of PCR methods for WSSV detection

PCR Nested PCR Real-time qPCR iiPCR

Specificity +++ +++ +++ +++
Sensitivity + +++ +++ +++
Quantitation Semi-quantitation possible Semi-quantitation possible Yes No
Product detection Visual Visual Automatic reading Automatic reading
Post-amplification Yes Yes No No
Cross contamination +++ +++ + +
Reaction time ++ +++ + +

+++, high; ++, moderate; +, low.

TA B L E 3 Primers used for detection of WSSV in shrimp

Product size
Methods Sequence 5′–3′ (bp) Reference
Nested PCR F1: ACTACTAACTTCAGCCTATCTAG 1441 (S. Durand and
R1: TAATGCGGGTGTAATGTTCTTACGA 941 Lightner, 2002)
F2: GTAACTGCCCCTTCCATCTCCA
R2: TACGGCAGCTGCTGGACCTTGT
Duplex qPCR WSSV-F: ′GCTGCCTTGCCGGAAATTA – (Leal et al., 2014)
WSSV-R; AGCCATGAAGAATGCCGTCTATCACACA
Prob-r: 6-carboxyfluorescein (FAM) at the 5′
Prob-q: BHQ1 at the 3′
IHHNV-F: TAC TCC GGA CAC CCA ACC A
IHHNV-R: GGC TCT GGC AGC AAA GGT AA
1
/Prob-q: N,N,N′,N′-tetramethyl-6-carboxyrhodamine (TAMRA)
1
/Prob-r: 6-carboxyfluorescein (FAM) at the 5′
qPCR F: ’GCTGCCTTGCCGGAAATTA - (S. Durand and
R; AGCCATGAAGAATGCCGTCTATCACACA Lightner, 2002)
Prob-r: 6-carboxyfluorescein (FAM) at the 5′
Prob-q: BHQ1 at the 3′
PCR F: AATGGTCCCGTCCTCATCTCA 71 (Y. L. Tsai et al., 2012)
R: GCTGCCTTGCCGGAAATT
RPA F: CATGGATGAAAACCTCCGCATTCCTGTGAC 124 (Xiaoming et al.,
R: CATCAGACTTTCCATTGCGGATCTTGATTTTG 2014)
Prob: TGCTGAGGTTGGATCAGGCTACTTCAAGA(BHQ1-dT)
G(THF)C(FAM-dT)GATGTGTCCTTTGAC (phosphate)

natural environment and display pathological signs 1 or 2 days process of white spot creation is unknown, it is conceivable that
before any death happens. Within 10 days of the commencement of the WSSV infection causes integument malfunction, leading to calcium salt
disease, cumulative death may exceed 100% (J. M. Lotz and Soto, 2002). accumulation inside the cuticle and the production of white spots (Y. G.
Juvenile shrimp of all ages and sizes are pathogenic in grow-out ponds, Wang et al., 2000). This disease also causes a red coloration of the skin
but significant mortality occurs either 1 or 2 months upon stocking and appendages due to the expansion of chromatophores (Alfaro et al.,
(Kasornchandra et al., 1998; Mishra and Shekhar, 2005). The most com- 2021; W. Zhan et al., 2000), fewer feed intake (Nadala et al., 1998),
mon sign of WSSV infection is the appearance of circular white spots or reduced preening and poor reaction to a stimulus (Durand et al., 1997),
patches of 0.5–3.0 mm in diameter, especially noticeable in the cuticle loose cuticle (Zhou, 1999), enlargement of branchiostegites due to
of the cephalothorax and tail section (Kim et al., 1998; Chu Fang Lo fluid buildup (W. Zhan et al., 2000) and thinning and delayed clotting of
et al., 1996). White to reddish-brown/reddish/pinkish/ to discoloration haemolymph (Q. Wang et al., 2001). Infected cells’ hypertrophied nuclei
are some of the characteristics of WSSV infection; over the head and have eosinophilic to increasingly basophilic inclusion masses (Afshar-
carapace, reduced appetite, congregation near the embarkation and nasab et al., 2014). Infected nuclei become increasingly basophilic and
other symptoms that are not unique to bacterial white spot syndrome expand over time (Takahashi et al., 2000; Y.-C. Wang et al., 1998). Dur-
(Corbel et al., 2001; Y. T. Wang et al., 2002). Although the specific ing the late stages of infection, karyorrhexis and cellular disintegration
788 ISLAM ET AL .

FIGURE 5 Infected with the white spot syndrome virus, symptoms of Penaeus monodon on the carapace (a) and last abdominal segment (b)

might occur, resulting in vacuolisation in necrotic regions (Karunasagar 10 WSSV VIABILITY in POND SEDIMENTS
et al., 1997; Kasornchandra et al., 1998) (Figure 5).
WSSV is found in large quantities in shrimp pond sediment. The most
significant metric in the transmission of WSD in newly stocked ponds is
9 WSSV IMPACTS ON SHRIMP FARMING WSSV viability in sediment. The physicochemical properties of the sed-
iment influence the viability and infectivity of WSSV. WSSV could last
WSSV is currently the biggest serious threat to Asia’s shrimp farming for up to 19 days after being exposed to the sun (De Gryse et al., 2020).
business and has been for some time. This is a very virulent disease that Under actual field conditions, WSSV could remain infectious in pond
can infect a broad host range. Moreover, profits were lost due to sec- sediments for up to 32 days after harvest. The significant difference
ondary losses in hatchery, feed and packing factory capabilities, among in WSSV viability in pond sediment could be related to the pond sed-
other things. iment’s early viral load, the physical and chemical features of the pond
Although there is little information on the existence and conse- sediment and the incidence of the disease epidemic during the pre-
quences of WSSV in wild shrimp communities in afflicted areas, it is vious crop, which contributed to this. Infectivity bioassays show that
predominantly located in Asia and Latin America (Walker & Mohan, acute to subacute WSSV disease in shrimp PL can be caused by fresh
2009). The cuticular epithelium, connective, neural, muscular, lymphoid and 7-day post-harvest sediment samples, with mortalities appearing
and haematopoietic tissues are among the ectodermal and mesoder- within 8–15 days. Sediment samples taken 14–32 days after harvest
mal tissues infected by WSSV. The stomach, gills, antennal gland, heart can cause chronic WSSV infection in shrimp PL, with mortality occur-
and eyes are all badly affected by the virus. In the final stages of ring between 18 and 25 days. An earlier study found that WSSV DNA
infection, these organs are damaged, and several cells are lysed. Hep- could be stored in the soil for up to 10 months under experimental
atopancreas of shrimp turns crimson and white spots (inclusions) with conditions (Natividad et al., 2008).
a diameter of 1–2mm appear on their carapace, limbs and interior Virus persistence in the environment is influenced by various phys-
body parts (Kv et al., 2005). They also exhibit sluggish behaviour, with ical, chemical and biological factors (C. P. Gerba, 2005). Temperature,
cumulative mortality reaching 100% between 2 and 7 days of illness. daylight, osmotic pressure, inorganic chemicals, microbial antagonism,
Although it has been increasingly evident since the late 1990s that the viral sorption state and virus type all influence virus survival in sedi-
occurrence of WSSV in an aquatic system does not invariably portend ment. The number and kind of organic matter, as well as pH, moisture
tragedy. Environmental changes, likely related to osmotic stress via salt content and electrical conductivity, are all factors to consider, among
concentrations or hardness, or abrupt temperature fluctuations, have other chemical components of sediment, which may influence the per-
generated outbreaks from latent P. monodon carriers in Thailand (de sistence of viruses in soil (Bosch et al., 2007). (Sobsey et al., 1986)
la Vega et al., 2007). Likewise, in Latin and North America, temper- compared hepatitis A virus inactivation in five different soil types,
ature fluctuations were shown to cause infected P. vannamei to die. including clay, clay loam, loamy sand, sandy and organic muck, the
Nevertheless, there have already been various claims about the ambi- soil clay had the best survival rate. Clay has a stronger affinity for
ent temperature that have been disclosed to limit WSSV death rate at viral particles due to adsorption and it protects the virus from viruci-
18 or 22◦ C and induce 100% fatalities at 32◦ C in the United States, but dal chemicals and thermal breakdown. The biological composition of
induce mortality at less than 30◦ C and prevent it at significantly larger a virus can have a significant impact on its survival and persistence
than 30◦ C in Ecuador (Verbruggen et al., 2016). Consequently, 3–4 in the environment. Heat, desiccation and acidic environments make
years of gene evaluation work on domestic P. vannamei stocks (selec- enclosed viruses more vulnerable than non-enveloped viruses, which
tion of shrimp surviving WSSV outbreaks) appears to have improved can tolerate these conditions. The enveloped virus, like WSSV, is likely
WSSV resistance in Ecuador. Since the outbreak of the WSSV in 1999, to be stabilised and protected in the aquatic environment by dissolved,
in Central and South America, the cultural sectors for P. vannamei have colloidal and solid organic matter. According to previous research, the
been steadily rebuilding. Ecuador, for example, exported 115,000 met- WSSV is stable at a pH of around 8, and it becomes utterly non-
ric tonnes in 1998, but only 38,000 metric tonnes in 2000 following infectious at pH levels as low as one and as high as 13 (Durand et al.
the arrival of WSSV in 1999. Ecuador has since recovered, with an 1997). By controlling the virus’s adsorption on soil particles in sedi-
estimated export of 50,000 metric tons in 2003 (Cuéllar-Anjel et al., ments, pH can indirectly influence virus viability (Yates et al., 1987).
2012). Nonetheless, it is well understood that the kind and amount of saline
ISLAM ET AL . 789

in the ecosystem substantially impact viral transmission at the bottom. (Mai et al., 2014;Yao et al., 2004). Shrimp are commonly exposed to
According to previous research, the WSSV remains stable at salinity heavy metals. The effects of heavy metal pollution on shrimp, as well
levels ranging from 0 to 10% (Durand et al., 1997). Viruses are des- as the mechanisms of heavy metal resistance, have mostly focused
orption from soil particles during rainfall due to a decrease in soil on metallothionein (MT). The sulfhydryl group of MT aggressively
salt concentration. In soil, humic acid and fulvic acid can inhibit virus chelates harmful metals and expels them from the body, making it a
pathogenicity, limit adsorption to organic matter by battling for adsor- metal-binding protein high in cysteine (Ecker et al., 1987). Environ-
bates and even function as an eluting agent, causing virus desorption mental stress has a well-documented impact on macroscopic markers
(C. P. Gerba, 2005). Most research on the survivability of several viruses of physiological status and immunological response. Strong or long-
in wet and dry soil, including poliovirus, enterovirus, coxsackievirus and term environmental stressors have a deleterious influence on shrimp’s
echovirus, discovered a positive link between virus survival and mois- innate immunity in general, and different environmental conditions
ture levels (C. Gerba, 1999). The viral nucleic acid is degraded in moist have somewhat different effects. The intestinal cell wall is the initial
soil before it is released from the virus capsid coat protein. The virus line of defence versus foreign matter invasion in L. vannamei (Vidal
loses its infectivity in soil under dry conditions due to degeneration of et al., 2001). The integrity of the cellular wall structure, the makeup
the capsid protein coat, but the nucleic acid can be restored undam- of intestinal bacteria and the immunological components in the intesti-
aged from the sediment (Mayer et al., 2015). Temperature, depth of nal mucus all play a role in the intestinal wall’s efficacy. However, in L.
the soil layer and distance from a nearby river basin are all said to vannamei, ammonia-nitrogen exposure damages the intestinal mucosa
impact soil moisture content. Numerous investigations on the influ- and disrupts the makeup of intestinal microorganisms, resulting in a
ence of temperature on poliovirus survival have discovered that as reduction in intestinal immunological function (Duan et al., 2018). In
the temperature increases, so does the inactivation rate (Cliver, 2009). Fenneropenaeus indicus, the pH of cultured water is directly connected
Denaturation of the viral capsid protein may cause viral inactivation to total hemocyte count (THC) and total hemocyte pPO (Sharma S R
at higher temperatures (Yan et al., 2004). The temperature indirectly et al., 2009). Plasma RB activity, malondialdehyde levels and hemocyte
affects virus survival by lowering the moisture content of the soil, par- DNA damage levels all rose significantly in L. vannamei exposed to cold
ticularly in the upper layer. The viral particles in the lower layers of the stress (12◦ C), although plasma THC, total protein content and osmotic
soil, on the other hand, remain viable due to the protection from sun- pressure were all reduced (Qiu et al., 2011). Changes in salinity within
light provided by the top layer of soil. The chemical disinfectant sodium a particular range (22–14) had little effect on L. vannamei THC, but
hypochlorite is commonly used in shrimp farms to disinfect water, significantly reduced pO activity, which resulted in enhanced WSSV
soil and equipment. It is an effective disinfectant for pond soil decon- replication (Yu et al., 2006). The immune system of shrimp has been
tamination. After 10 min of treatment with sodium hypochlorite at a demonstrated to be weakened by water pH, temperature and other
concentration of 100 ppm, WSSV becomes non-infectious (Oseko et al., physical and chemical variables. On the other hand, environmental
2006). According to another study, shrimp ponds rinsed with 100 ppm stress of the proper level has been demonstrated in certain experi-
bleaching powder were infected with WSSV-positive invertebrates 20 ments to generate immune-like effects in shrimp, offering a degree
days post disinfection (Burge, 1981). Disinfection of soil in real-world of protection. Heat stress (33◦ C) during the early stages of WSSV
farming conditions appears to be extremely difficult. The treatment’s infection was also found to boost WSSV resistance and reduce cumu-
efficacy would be determined by the amount of active chlorine in the lative shrimp mortality in L. vannamei (M. Rahman et al., 2007; Rahman
commercial preparatory work. et al., 2006). Indeed, both cold and heat shocks reduced the cumulative
mortality of WSSV-infected shrimp (Yuan et al., 2017). Furthermore,
injection with -glucan raised ROS and produced oxidative stress in
11 ENVIRONMENTAL IMPACT of WSSV WSSV-infected shrimp, but substantially decreased WSSV viral load;
yet, the cumulative death rate of the infected shrimp remained greater
Multicellular organisms’ typical physiological processes necessitate than the control category (Thitamadee et al., 2014). Some effects of
a reasonably steady internal redox status. External variables, for environmental conditions on viral illness in shrimp have been observed
instance, mycotoxin and heavy metal stress, on the other hand, might since the 1990s. Nitrogen content, nitrite nitrogen and saltiness of
cause an internal redox state imbalance, potentially leading to oxida- aquatic water all affected shrimp MBV infection in P. monodon, accord-
tive stress. Reactive oxygen and nitrogen species (ROS and RNS) ing to (G. Li et al., 2001). MBV was eventually superseded as the most
are produced as a result of oxidative stress, and they damage DNA dangerous virus to Shrimp by WSSV. Every year since the first breakout
and proteins, impairing normal cellular activity. As a result, organ- of WSS in 1992, it has resulted in significant losses of cultivated shrimp.
isms have devised several ways to counteract such harm. To eliminate Shrimp with WSSV is still frequent in shrimp farms today. Shrimp WSS
ROS and RNS, downstream enzymes such as superoxide dismutase has been proven to be directly related to acute factors in both pro-
(SOD) and glutathione reductase are often utilised, as well as upstream duction practice and laboratory study (Al-Bahry et al., 2011; Y. Chen
signalling recognition and signalling transduction molecules. Several et al., 2010; J. G. He et al., 2000) (in Chinese), and in recent years,
enzymes involved in redox reactions have been researched in shrimp, its mechanism has been increasingly elucidated. To better understand
with SOD being the most well researched. MnSOD and Cu/ZnSOD the processes of environmentally induced WSS in shrimp, researchers
have been identified in shrimp. FeSOD had yet to be discovered looked at the impacts of a variety of environmental stresses on WSSV
790 ISLAM ET AL .

infection. It has been demonstrated that heat stress (32◦ C) enhanced dependable demonstrative tests accessible (Powell et al., 2006). Tissue
HSP70 transcription, and that stressed shrimp used RNAi to lower lesions will be studied using DNA-based tools such as PCR and in situ
HSP70 expression. At the same temperature, the shrimp were more hybridisation (ISH). At least twice during the production cycle, sample
vulnerable to WSSV than the treatment group (Y.-R. Lin et al., 2011). hatcheries and keep samples for subsequent testing (3 weeks post ship-
Scientists previously demonstrated that increasing or decreasing the ping). Only purchase animals that have been PCR checked and stress
water temperature correctly improved L. vannamei resistance to WSSV tested (Sterchi, 2008). It’s critical to start with no or a very low viral
(Yuan et al., 2017). Shrimp are very poisonous to molecular ammo- load. It has been discovered that stressing PLs with formalin helps to
nia nitrogen. Shrimp exposed to ammonia nitrogen had much more screen out weaker animals, yet it is never a good idea to stock PLs that
WSSV copies in their hepatopancreas, hemolymph and swimming feet are known to be infected with the virus. It has been noticed that the
than control shrimp (Mohamed Fouzi et al., 2010). Shrimp’s capacity to virus in P. japonicus may not show pathogenicity until after PL6, necessi-
combat WSSV infections was decreased by osmotic stress. WSSV repli- tating the screening of later-stage animals. Reduce the shrimp’s stress
cation in shrimp is inhibited mostly by ROS produced under oxidative as much as possible. You can do it in a few different ways and cannot do
stress (H.-H. He et al., 2017; Su et al., 2014; Thitamadee et al., 2014). it in many of them. Some of the things you can do include:
Shrimps have also been found to be more sensitive to WSSV under
hypoxic settings, probably due to anoxic circumstances releasing fewer 1. Before stocking, increase acclimation times.
ROS. As a result, most ecological stresses impair shrimp susceptibility 2. Increase stress tolerance with non-specific immune stimulants
to WSSV (Lehmann et al., 2016) (NSIS) and supplemented mineral and vitamin diets.
3. Consider stocking during periods of the year when you know the
environment will not be subjected to extremes in temperature and
12 ECONOMIC CONSEQUENCES of the WSSV salinity. It has been claimed that these pressures can trigger an
VIRUS epizootic in a virus-infected population.
4. Maintain a high-quality diet and employ NSIS throughout your life
WSSV has a significant economic impact on aquaculture around the cycle.
world. After the first epidemic in 1992, production decreased by more 5. If conceivable, stock at lower densities.
than 70%, resulting in a 3-year production loss of more than US$2 bil- 6. Keep an eye out for WSSV-carrying vectors in the ponds and take
lion in China. Before the WSSV outbreak, Thailand’s yearly production steps to eliminate them.
rate was around 34,000 tonnes per year, but by 1994, it had dropped to 7. Before stocking, collect phytoplankton and zooplankton from the
265,000 tonnes, worth US$1.6 billion. Indonesia followed a similar pat- pond and test for WSSV using PCR. Positive ponds should be
tern, with production gradually increasing to 17,000 tonnes per year avoided at all costs.
from 1985 to 1991 until the WSSV outbreak. Due to the WSSV, their 8. Sample your ponds regularly. Ascertain that sick and dying animals
production fell in 1992, resulting in projected output losses of almost are routinely collected by PCR and histology, as well as any unex-
$1 billion over the next 10 years. In Ecuador, an outbreak occurred in pected patterns of death. If you can, harvest the shrimp at the first
1999, and production fell by over 60% in 2 years, resulting in losses of sign of a problem.
over $1 billion between 1998 and 2001 (Ghosh, 2014). A similar situ-
ation was recorded in Panama and Peru, where production decreased
by 90%, resulting in losses of over US$100 million and US$70 million 13.1 Biosecurity for shrimp farming
over 3 years, respectively. Since 1992, India’s shrimp aquaculture rev-
enue has decreased to a loss of INR 2700 million (Xu et al., 2007). In Biosecurity is more accessible to deploy in small, intensive and reg-
1994, the illness expanded to Bangladesh’s southwest area, damag- ulated farming systems than in large-scale outdoor activities. In the
ing roughly 90% of large shrimp farms and creating a 20% decline in shrimp industry, biosecurity measures are divided into two categories:
national shrimp production. As a result, Bangladesh’s shrimp exports excluding pathogens and eradicating infections when they are present.
fell from 25,742 tonnes in 1997–1998 to 18,630 tonnes (Debnath (D. Lightner, 2003) highlighted measures to keep diseases out of
et al., 2014). stock (i.e., PL and broodstock), including quarantine and SPF, certi-
fied stocks and limiting live and frozen shrimp imports. Removing
diseases from hatcheries and farms was advised by excluding vec-
13 MANAGEMENT TECHNIQUES FOR WSSV tors and external sources of pollution and protecting against internal
cross-contamination (Figures 6 and 7).
At first, avoid purchasing nauplii or PLs from sources that may or may
not be infected with the virus. When used on eggs, nauplii and PLs,
iodine and water washes will likely eradicate the virus. This procedure 13.2 Prevention
must be followed consistently. Incubation facilities should maintain
high biosecurity norms and inspect every animal batch. A hatchery The methods of prevention are similar to those used in the case of
should be constructed to prevent the infection from being presented in TSV. Before importing live or frozen shrimp from disease-infected to
the ocean. To diagnose and screen WSSV, utilise the most sensitive and disease-free zones, PCR testing needs to be performed on all live
ISLAM ET AL . 791

FIGURE 6 Farm biosecurity factors

FIGURE 7 Farm biosecurity – implementation

792 ISLAM ET AL .

and frozen shrimp (Dhar et al., 2004). Before breeding, PCR testing 15 ADVANTAGES OF SPF
of broodstock should be performed. Before stocking into ponds, PL
should also be PCR screened, resulting in an increased percentage SPF animals give some certainty to a nation presenting a species
of good harvests. Although PCR is not a perfect way of diagnosing interestingly that the imported animals will not spread the recorded
white spot virus, this is currently the most accurate diagnostic pro- diseases to local species. SPF stocks, on the other hand, may include
cedure. Vertical transmission of WSSV from infected broodstock to unidentified infections, which should be considered because they can
larval stages can also be prevented by washing and disinfecting eggs constitute a concern when the animals are stressed (Claydon et al.,
and nauplii. Fresh crab and other crustaceans should never be fed to 2010). Biosecurity techniques are used to combat the possibility of
broodstock (P. Liu et al., 2000). Polyculture strategies involving some- disease outbreaks in shrimp culture. The principle point of biosecurity
what carnivorous fish species (Tilapia) have also been demonstrated to frameworks is to keep illnesses from spreading and to aid in their elim-
reduce WSSV pathogenicity in the farm areas, so the fish can consume ination if they do. Because the specific pathogen can be eradicated and
infectious vectors before contacting live shrimp. Because this virus can contamination minimised, the particular pathogen-free stock is one
only survive 3–4 days in water, preventing transmission can be as sim- of the most important components in any biosecurity system (Aarat-
ple as disinfecting the water used for adjustments and using precise tuthodiyil and Wise, 2017). Because the specified interfering infections
filtration (H. Liu et al., 2019). Formalin doses of 70 ppm have been can be ruled out, SPF animals are incredibly beneficial for basic and
shown to prevent transmission while causing no damage to shrimp. applied scientific research, particularly immunological studies and vac-
Furthermore, all wastewater from farming or processing operations cination trials. SPF animals are also required for other bioassays; for
with the potential for WSSV infections should be treated before dis- example, a study of shrimp virus infections requires pathogen-free
charge (e.g., with formalin or chlorine). PCR and probes for dot-blot animals because the shrimp cell line is unavailable (Barman et al., 2012).
and ISH tests can detect WSSV. The presence of characteristic white
spots can likewise be utilised to analyse it visually (albeit these are
not generally present in infected animals) (McColl et al., 2004). The 16 LIMITATIONS OF SPF SHRIMP
presence of huge numbers of Cowdrey A-type nuclear inclusions and
hypertrophied nuclei in H&E-stained sectioned tissues, or simply fixing All microorganisms representing a critical danger should be identified
and staining gill tissue and studying it under a microscope, can be used and physically removed from the stocking area. It should be noted,
to confirm WSSV (especially for asymptomatic carriers) (Pazir et al., however, that the shrimp could in any case, be infected with a microbe,
2012). not on the list. SPF shrimp offspring are not SPF unless grown and kept
in a biosecure SPF facility (Washim et al., 2019; Wyban, 2010). They can
no longer be classified as SPF after they leave the institution. Managing
14 PRODUCTION OF SPF SHRIMP P. monodon must be more troublesome than working with L. vannamei
(V. Sedhuraman et al., 2014). Due to the difficulty of maturing P. mon-
The collection of apparently healthy wild stock from areas with low odon in captivity, a significantly longer holding period is needed before
disease frequency, followed by individual primary quarantine, where they arrive at a feasible size (D. V. Lightner, 2005).
the shrimp can be individually checked for specific infections and con-
taminated individuals eliminated, are significant components of an
SPF procedure(Barman et al., 2012). The shrimp are then moved to a 17 SPR SHRIMP
secondary quarantine facility, which is raised to broodstock size and
constantly checked. Healthy broodstock is subsequently sent to the SPR stands for SPR and is another frequently misunderstood acronym.
breeding centre, where many families from various sources are pro- It refers to a shrimp genetic feature that offers some resistance to a
duced (Barman et al., 2012). The larvae are then raised in biosecure particular virus. SPR shrimp are usually the outcome of a specialised
hatcheries from the selected families. Any diseased stock discovered breeding effort to increase viral resistance. SPF and SPR are two dis-
through continuous monitoring is discarded right away. The funda- tinct qualities. SPR shrimp may not all be SPF shrimp and vice versa.
mental steps for developing SPR lines of broodstock P. monodon are SPR shrimp strains, on the other hand, do not have to be SPF (Alday-
similar to those for the SPF program. A more rigorous genetic selec- Sanz, 2018). Because of their superior presentation in broodstock as
tion approach, including a more significant number of families, is well as in grow-out ponds, and frequent monitoring of disease out-
necessary to select for desired features. Whatever program is cho- breaks with the quarantine process for SPR, P. vannamei is currently
sen, developing SPF and/or SPR lines of P. monodon should be viewed almost exclusively used in Latin America. According to a recent FAO
as a long-term commitment (D. Lightner, 2011). It requires consis- survey, there were over a hundred maturation units delivering 15 bil-
tent authority over all components of culture, exceptionally gifted lion nauplii each 30 days and loading approximately 400 hatcheries,
logical individuals, the most significant levels of discipline and col- largely of SPR shrimp mostly in Ecuador and Mexico (Ghanaatian,
laboration, specialised staff training and progressing research centre 2015). TSV can create severe mortalities in P. vannamei-stocked farms
examination. and easily spread across ponds by insect or avian vectors. As a result,
ISLAM ET AL . 793

the introduction of strains against TSV, with the advancement of biose- tiny amount of water to the system. Filters that catch everything using
curity applications to prevent infections from other viruses, might plant fibres in combination with serial mesh filtration could be effec-
significantly boost the production of the P. vannamei in Asian countries. tive. Using mesh filters with a pore size of 250 microns or less, as well
The United States industry has embraced such a protocol, resulting in as small mesh bags, can be beneficial (Taslihan et al., 2021).
a 50% annual growth rate during the previous 3 years (Thongrak et al., Avoiding the exchange of water is another suggestion. Recent out-
1997). Recently, some work has been done on generating a P. chinen- breaks of WSSV in South Carolina, USA, appear to be linked to the
sis strain with SPR for WSSV. Ponds filled with PL produced by WSSV addition of water to ponds, therefore there appears to be some weight
survivors, pandemic saw an increase in survival rates from 0–0.8% to to this theory (Boyd et al., 2020). It is unclear if this stressed the shrimp
12–45%, whereas laboratory test examinations demonstrated 30–60% and caused an outbreak, or whether it transferred vectors and viruses
enhancements in overall survival for third and fourth-generation win- into the ponds. Aeration is used in intensive systems but not in semi-
ners. Both control and select animals exhibited a 60% infection rate intensive systems. In ponds where oxygen levels cannot be regulated
with WSSV, according to PCR testing, proving that this was related to without water exchange, the capacity to mechanically aerate the water
resistance (Kong et al., 2003). It is indeed possible that WSSV-resistant without resorting to water exchange could be quite valuable (Hopkins
P. vannamei strains will emerge. Even though WSSV is still the most et al., 1993). Animals that have been PCR positive for the infection are
severe disease risk in the Asian shrimp aquaculture industry, it could not bought. Screening supplied animals for the presence of the infec-
boost the sector itself well. The recent discovery of multiple molecu- tion has been accounted to be a successful methodology for following
lar markers (particularly microsatellites) linked to immune function and the disease’s status in the population in certain conditions. This can be
growth, as well as potential advancements in quantitative genetics to used to time harvests and prevent problems from spreading to adjacent
shrimp breeding, suggest that selecting fast-growing, disease-resistant ponds and/or neighbours.
variants could become much more productive shortly. This could also
provide insight into invertebrate antiviral immunity, which is currently
unknown. In P. stylirostris, such disease-related indicators have already 19 CONCLUSION AND SUGGESTIONS
been discovered (Hizer et al., 2002).
WSSV remains a crucial disease in shrimp production and a big dan-
ger to the global shrimp business. WSD is attracting a lot of scientific
18 RECOMMENDATION TO CONTROL WSSV AT attention these days, owing to the disease’s economic implications.
THE FARMING LEVEL Currently, research is being conducted to achieve a remarkable com-
prehension of the molecular and pathophysiology of WSSV, as well
The movement of infected PLs mostly determines the viruses and as the proper treatment and prevention of the virus. Nevertheless,
other diseases’ mobility. As a result, if this can be stopped, the virus’s additional research should be done in places with more resistant host
transmission will be slowed. Numerous nations have found a way to species to comprehend their function against disease effects, as these
accomplish this, however, the truth will surface eventually about how insights could be helpful in finding a cure for WSD. SPR shrimps are pre-
fruitful they will be. The virus appears to be here to stay once it takes ferred over SPF animals for cultivation in locations where the disease
a footing. Luckily, as with all other viral infections like BP, IHHN, TSV is endemic. As a result, more research should be done in the breeding
and MBV, the infection’s effect will blur over a long period (Mani et al., program to develop SPR animals which are resistant to WSSV. Notably,
2016). The Thais adapt to the infection with a range of devices, some of various studies in the domain of WSSV immunisation have been under-
which will be valuable for P. vannamei (C. Lin, 2020). However, it is criti- taken, although their field application has not yet succeeded. As a
cal to grasp the variations between the two types of shrimp cultivation, result, intervention and development of vaccines for commercial uses
as well as the difficulties that come with utilising the same methods to can help to minimise the risks. On the other hand, advanced tech-
try to limit the virus’s presence. Pesticides should be used to kill the niques such as shrimp farming in recirculating aquaculture systems and
vectors before replenishing the ponds, according to the advice given polyculture could be the potential strategies for WSD regulation.
to and received by Thai farmers (Szuster, 2006). Typically, a powerful
insecticide is added to the water to kill any crustaceans or other virus- AUTHOR CONTRIBUTIONS
carrying vectors that may be present in the pond. This is impossible to Conceptualisation, supervision, writing – original draft and writing – review
achieve on the scale required across most of America’s regions. The and editing: Sk Injamamul Islam; Conceptualisation and writing – review
expenses would be prohibitively exorbitant, and the massive volumes and editing: Moslema Jahan Mou. Conceptualisation and writing – review
of pesticides that would have to be poured into the environment would and editing: Saloa Sanjida.
be unwelcome. This should be avoided at all costs and only used as a
last option. One potential method for removing some vectors is to fil- ACKNOWLEDGEMENTS
ter incoming water entering the pond (Newman, 2019). Because of the The first author is sincerely grateful to the ASEAN and Non-ASEAN
high demand for water in the Americas, conventional filters might be scholarship authority at Chulalongkorn University, Thailand for giving
hazardous. This demand, however, can be controlled by adding only a financial support for pursuing master’s studies.
794 ISLAM ET AL .

CONFLICT OF INTEREST Bennour, A., Saad, A., & Sennana, H. (2016). Chronic myeloid leukemia:
The authors declare that they have no competing interests. Relevance of cytogenetic and molecular assays. Critical Reviews in
Oncology/Hematology, 97, 263–274. https://doi.org/10.1016/j.
critrevonc.2015.08.020
DATA AVAILABILITY STATEMENT Biel, S. S., & Gelderblom, H. R. (1999). Diagnostic electron microscopy is still
No. a timely and rewarding method. Journal of Clinical Virology, 13(1), 105–
119. https://doi.org/10.1016/S1386-6532(99)00027-X
Bir, J., Howlader, P., Ray, S., Sultana, S., Ibrahim Khalil, S. M., & Banu,
ETHICS STATEMENT G. (2017). A critical review on White Spot Syndrome Virus (WSSV):
Ethics statement is not applicable as no sample collection or question- A potential threat to shrimp farming in Bangladesh and some Asian
naires was conducted. countries. International Journal of Microbiology and Mycology, 6, 39–48.
Bondad-Reantaso, M., & Subasinghe, R. (2001). Aquaculture Development,
Health and Wealth.
ORCID Bosch, A., Pintó, R., & Abad Morejón de Girón, X. (2007). Survival and Trans-
Sk Injamamul Islam https://orcid.org/0000-0002-0888-6075 port of Enteric Viruses in the Environment. In Viruses in Foods. Springer:
Boston(pp. 151–187).
Boyd, C., D’Abramo, L., Glencross, B., Huyben, D., Juarez, L., Lockwood, G.,
PEER REVIEW
McNevin, A. A., Tacon, A. G., Teletchea, F., Tomasso Jr, J. R., Tucker, C.
The peer review history for this article is available at https://publons. S., & Valenti, W. (2020). Achieving sustainable aquaculture: Historical
com/publon/10.1002/vms3.979. and current perspectives and future needs and challenges. Journal of the
World Aquaculture Society, 51, 578–633. https://doi.org/10.1111/jwas.
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