Disease prevention strategies for penaeid shrimp culture

Abstract

Penaeid shrimp aquaculture expanded significantly over the past two decades. However, shrimp farmers have suffered significant economic losses because of viral diseases. Researchers from the U.S. Marine Shrimp Farming Program (USMSFP) have developed novel approaches to mitigate the devastating impact of shrimp viruses, including the use of specific pathogen free (SPF) and specific pathogen resistant (SPR) shrimp, as well as the establishment of biosecure production systems that rely on pathogen exclusion. These approaches have evolved over the past decade in response to changing disease problems faced by U.S. shrimp farmers. In the late 1980's and early 1990's, U.S. farmers observed Runt Deformity Syndrome (RDS), an economically significant and frequent disease problem of cultured Pacific white shrimp, Litopenaeus vannamei. RDS is characterized by reduced growth rates and cuticular deformities and is caused by Infectious hypodermal and hematopoietic necrosis virus (IHHNV). The increasing incidence of RDS on commercial farms catalyzed USMSFP researchers to develop SPF stocks of L. vannamei that were free of IHHNV. High health offspring from these SPF stocks were made available to U.S. shrimp farmers, resulting in a significant increase in U.S. farmed shrimp production from 1992 -1994. However, in mid-1995, Taura syndrome virus (TSV) was identified in south Texas, the major shrimp farming region in the U.S., and the resulting TSV epizootic contributed to a 164% decline in Texas shrimp production from 1994 to 1995. USMSFP researchers responded by initiating a selective breeding program to develop TSV-resistant L. vannamei. The use of these high-health SPR stocks, in conjunction with on-farm biosecurity practices, resulted in incremental increases in U.S. shrimp production from 1998 to the present. Although TSV-resistant shrimp improved production and profitability for those farmers who were experiencing crop losses from TSV, breeding shrimp for resistance to a single viral pathogen, using current selective breeding strategies, may not be the most effective course of action for the long-term viability of the shrimp farming industry. USMSFP researchers are now developing biosecure shrimp production systems which rely on pathogen exclusion, and research results indicate that it is possible to produce > 5 kg of market-sized shrimp (~ 20 g) per m 2 of raceway in about 12 weeks, using < 400 L of water per kg of shrimp. With advanced biosecure technologies available, the U.S. shrimp farming industry will be able to expand into areas away from the coast with greater control against the spread of disease and without adversely affecting the environment.

Full text

Disease Prevention Strategies for Penaeid Shrimp Culture
Shaun M. Moss
Steve M. Arce
The Oceanic Institute
41-202 Kalanianaole Highway
Waimanalo, Hawaii 96795 USA
The Oceanic Institute
41-202 Kalanianaole Highway
Waimanalo, Hawaii 96795 USA
smoss@oceanicinstitute.org
sarce@oceanicinstitute.org
Dustin R. Moss Clete A. Otoshi
The Oceanic Institute
41-202 Kalanianaole Highway
Waimanalo, Hawaii 96795 USA
The Oceanic Institute
41-202 Kalanianaole Highway
Waimanalo, Hawaii 96795 USA
dmoss@oceanicinstitute.org cotoshi@oceanicinstitute.org
Abstract
Penaeid shrimp aquaculture expanded significantly over the past two decades. However,
shrimp farmers have suffered significant economic losses because of viral diseases. Researchers
from the U.S. Marine Shrimp Farming Program (USMSFP) have developed novel approaches to
mitigate the devastating impact of shrimp viruses, including the use of specific pathogen free
(SPF) and specific pathogen resistant (SPR) shrimp, as well as the establishment of biosecure
production systems that rely on pathogen exclusion. These approaches have evolved over the past
decade in response to changing disease problems faced by U.S. shrimp farmers. In the late 1980’s
and early 1990’s, U.S. farmers observed Runt Deformity Syndrome (RDS), an economically
significant and frequent disease problem of cultured Pacific white shrimp, Litopenaeus vannamei.
RDS is characterized by reduced growth rates and cuticular deformities and is caused by
Infectious
hypodermal and hematopoietic necrosis virus (IHHNV). The increasing incidence of
RDS on commercial farms catalyzed USMSFP researchers to develop SPF stocks of L. vannamei
that were free of IHHNV. High health offspring from these SPF stocks were made available to
U.S. shrimp farmers, resulting in a significant increase in U.S. farmed shrimp production from
1992 - 1994. However, in mid-1995, Taura syndrome virus (TSV) was identified in south Texas,
the major shrimp farming region in the U.S., and the resulting TSV epizootic contributed to a
164% decline in Texas shrimp production from 1994 to 1995. USMSFP researchers responded by
initiating a selective breeding program to develop TSV-resistant L. vannamei. The use of these
high-health SPR stocks, in conjunction with on-farm biosecurity practices, resulted in incremental
increases in U.S. shrimp production from 1998 to the present. Although TSV-resistant shrimp
improved production and profitability for those farmers who were experiencing crop losses from
TSV, breeding shrimp for resistance to a single viral pathogen, using current selective breeding
strategies, may not be the most effective course of action for the long-term viability of the shrimp
farming industry. USMSFP researchers are now developing biosecure shrimp production systems
which rely on pathogen exclusion, and research results indicate that it is possible to produce > 5
kg of market-sized shrimp (~ 20 g) per m
2
of raceway in about 12 weeks, using < 400 L of water
per kg of shrimp. With advanced biosecure technologies available, the U.S. shrimp farming
industry will be able to expand into areas away from the coast with greater control against the
spread of disease and without adversely affecting the environment.
Introduction
Shrimp aquaculture expanded significantly during the 1980s and now represents a
multi-billion dollar a year industry. In 2002, the global shrimp farming industry
produced an estimated 1.6 million metric tons of shrimp, and production is projected to
increase at a rate of 12-15% per year over the next several years (Rosenberry 2003).
Although farmed shrimp now represent about 50% of the global penaeid shrimp supply,

farmers have suffered significant economic losses over the last decade, largely from viral
diseases that have plagued the industry (Table 1). In Asia, mortalities of cultured shrimp
due to White spot syndrome virus (WSSV) and Yellow head virus (YHV) have resulted
in significant economic losses (Flegel and Alday-Sanz 1998), and Taura syndrome virus
(TSV) is now spreading throughout this region. Similarly, in the Western Hemisphere,
both WSSV and TSV have caused catastrophic losses on shrimp farms (Lightner 2003).
In Ecuador alone, WSSV was responsible for an estimated 53% decline in shrimp
production from 1998 to 2000, resulting in a loss of export revenue in excess of $516
million (Rosenberry 2000).
Virus Year of Emergence to
2001
Estimated loss
WSSV - Asia 1992 $4 – 6 billion
WSSV - Americas 1999 > $1 billion
TSV 1991 – 1992 $1 – 2 billion
YHV 1991 $0.1 – 0.5 billion
IHHNV
a
1981 $0.5 – 1.0 billion
a
Includes Gulf of California fishery losses for 1989 – 1994.
Table 1. Estimated economic losses (in US$) since the emergence of certain viral
pathogens in penaeid shrimp aquaculture (modified from Lightner 2003).
In response to these viral pathogens, the global shrimp farming industry is
changing the way shrimp aquaculture is practiced. In the United States (U.S.),
researchers from the U.S. Marine Shrimp Farming Program (USMSFP) have developed
novel approaches to mitigate the impact of shrimp viruses on domestic farm production.
USMSFP member institutions involved in this effort include The Oceanic Institute (OI,
Waimanalo, Hawaii), University of Arizona (UAZ, Tucson, Arizona), University of
Southern Mississippi, Gulf Coast Research Laboratory (Ocean Springs, Mississippi),
Waddell Mariculture Research Center (Bluffton, South Carolina), Texas A&M
University (Port Aransas, Texas), and Tufts University (Boston, Massachusetts). Several
of the approaches developed by the USMSFP have been used successfully in other meat-
producing industries, and include the use of specific pathogen free (SPF) and specific
pathogen resistant (SPR) stocks, as well as the establishment of biosecure production
systems that rely on pathogen exclusion. Importantly, these approaches have evolved
over the past decade in response to changing disease problems faced by U.S. shrimp
farmers, and their evolution represents an interesting case study on the maturation of an
important industry.
Pre-SPF Era for the U.S. Shrimp Farming Industry (1980’s – 1991)
Although the first commercial shrimp farm in the U.S. was established in 1967, it
wasn’t until the late 1980’s and early 1990s when the industry began to expand
(Rosenberry 2003). During that time, the most commonly cultured species was the
Pacific white shrimp, Litopenaeus vannamei, because it was considered to be highly
resistant to Infectious hypodermal and hematopoietic necrosis virus (IHHNV), a member
of the Parvoviridae family (Bonami et al. 1990). IHHNV was first recognized in 1981
when it was associated with catastrophic losses of cultured blue shrimp, Litopenaeus
stylirostris, in Latin America (Lightner 1999). Despite the relative resistance of L.
1

vannamei to IHHNV, shrimp farmers in the Western Hemisphere observed reduced
growth rates and cuticular deformities in L. vannamei infected with IHHNV. This
condition is referred to as Runt Deformity Syndrome (RDS), and RDS represented an
economically significant and frequent disease problem of cultured L. vannamei
(Kalagayan et al. 1991). As much as 30% of cultured L. vannamei infected with IHHNV
exhibited RDS, and this reduced the market price of IHHNV-infected shrimp by 10-50%
relative to IHHNV-free shrimp.
Establishment and Benefits of SPF Shrimp (1991 – 1994)
The increasing incidence of RDS on commercial farms in the U.S. catalyzed
USMSFP researchers to develop SPF stocks of L. vannamei that were free of IHHNV.
Although the SPF concept was well established in other meat-producing industries
(Zavala 2001), it had not yet been applied to shrimp. Guidelines for establishing SPF
shrimp came from The International Council for the Exploration of the Sea’s (ICES)
“Code of Practice to Reduce the Risks of Adverse Effects Arising from the Introduction
of Non-indigenous Marine Species” (Sindermann 1990). Modifications of the ICES
guidelines were used to develop the first SPF stock of penaeid shrimp for the U.S. shrimp
farming industry from 1989-1991 (Wyban et al. 1993, Pruder 1994, Pruder et al. 1995).
The guidelines stipulate that only disease-causing organisms that can be reliably
diagnosed and physically excluded from a facility can be considered in an SPF program.
The working list of specific pathogens for SPF penaeid shrimp in the U.S. has changed
over time, as new pathogens have been identified and as more advanced disease
diagnostic tools have become available. The current SPF list for the U.S. includes eight
viruses, one prokaryote, and certain classes of parasitic protozoa (Table 2).
Pathogen Type Pathogen/Pathogen Group Pathogen Category
2
Viruses WSSV – nimavirus (new family) C-1
YHV, GAV, LOV – roniviruses (new family) C-1
TSV – picornavirus C-1
BP – occluded enteric baculovirus C-2
MBV –occluded enteric baculovirus C-2
BMN – nonoccluded enteric baculovirus C-2
IHHNV –systemic parvovirus C-2
HPV – enteric parvovirus C-2
Procaryote NHP – α-proteobacteria C-2
Protozoa Microsporidians C-2
Haplosporidians C-2
Gregarines C-3
Table 2. A working list of “specific” and excludable pathogens for penaeid shrimp.
1
1
Modified from D.V. Lightner, U.S. Marine Shrimp Farming Program FY03 Progress Report.
2
Pathogen category (modified from Lotz et al. 1995) with C-1 pathogens defined as excludable
pathogens that can potentially cause catastrophic losses in one or more penaeid species; C-2
pathogens are serious, potentially excludable; and C-3 pathogens have minimal effects, but may
be excluded from NBCs, multiplication facilities, and some types of farms.
2

To develop an SPF stock, shrimp are collected from the wild and transferred to a
primary quarantine facility where they are analyzed for specifically listed pathogens
using appropriate disease diagnostic tools (Fig. 1). If shrimp test positive for any of the
listed pathogens, they are destroyed in the primary quarantine facility. If shrimp test
negative for specifically listed pathogens after several successive screenings, they are
transferred to a secondary quarantine facility where they are matured and spawned to
produce an F
1
generation of captive shrimp. Because some viruses can be transmitted
from parent to offspring (vertical transmission), representative shrimp from the F
1
generation are analyzed for specifically listed pathogens. If shrimp from the F
1
generation test negative for specifically listed pathogens after several successive
screenings, they are transferred out of the secondary quarantine facility and can be
included as part of the germplasm in a nucleus breeding center (NBC). Shrimp that are
maintained in a well-established NBC (i.e. where there is a history of negative disease
status documented through a surveillance program) may be designated as SPF (Lotz
1997). However, once shrimp leave an SPF-NBC, they no longer are referred to as SPF
even though they may be free of specifically listed pathogens. If shrimp are transferred
from an SPF-NBC to a medium-biosecurity facility, their new designation is High Health
(HH), indicating that these shrimp are at greater risk of pathogen exposure and infection.
If shrimp are transferred to a low-biosecurity shrimp farm, they have entered the
Commodity Production (CP) stream, which is most vulnerable to pathogen outbreaks,
and the shrimp are neither SPF nor HH.
Figure 1. Procedures used to develop specific pathogen free (SPF) shrimp
collected from the wild.
3

Initial growout trials using HH L. vannamei indicated that these stocks out-
performed non-HH stocks when evaluated at commercial shrimp farms in the U.S.
(Wyban et al. 1993). In Texas, Jaenike et al. (1992) reported that HH shrimp obtained
from the USMSFP produced a greater yield, higher survival, a more uniform size
distribution, and a lower feed conversion ratio than non-HH shrimp. In Hawaii,
Carpenter and Brock (1992) reported that HH shrimp produced a greater yield and higher
survival than non-HH shrimp when cultured under semi-intensive and intensive culture
conditions. Importantly, the HH crop yielded a 62.5% higher return than non-HH crops,
and similar improvements were reported in South Carolina (Wyban et al. 1993). The
overall effect of using HH shrimp in the U.S. was a significant increase in production
from 1992-1994 (Fig. 2). This huge impact was most apparent in Texas where the
majority of domestic shrimp farming occurs. During this time, production increased from
1.66 million pounds in 1991 to 3.8 million pounds in 1992 and 4.2 million pounds in
1993 (Rosenberry 2003), and this represents a 153% increase in production over two
years.
Figure 2. U.S. farmed shrimp production from 1988 – 2002.
Emergence and Impact of TSV in the U.S. (1995 – 1998)
Despite these encouraging results, HH shrimp were not a panacea for the disease
problems plaguing the shrimp farming industry (Pruder 1994). In 1993, HH shrimp were
cultured with wild-caught seed at a commercial shrimp farm near Rio Guayas in Ecuador.
HH shrimp exhibited poor survival (7-43%) compared to wild seed (36-42%), and heavy
mortalities were attributed to TSV infection. From this experience, it was demonstrated
that HH shrimp cultured in environments with disease problems may not perform well.
In mid-1995, TSV was identified in south Texas and the presence of this virus resulted in
a significant decline in U.S. farmed shrimp production (Brock et al. 1997, Fig. 2). In
Texas alone, shrimp production went from 3.69 million pounds in 1994 to 1.4 million
pounds in 1995, a 164% decline in one year. Although shrimp production increased from
4

1995-1998, production levels never exceeded the pre-TSV years when SPF or HH shrimp
were available.
Breeding of SPR Shrimp (1998 – present)
In response to the devastating effects of TSV on cultured shrimp in the U.S.,
USMSFP researchers initiated a selective breeding program to develop a TSV-resistant
strain of L. vannamei. This approach seemed reasonable, especially in light of the
tremendous improvements made through selective breeding of commercially important
agriculture crops and animals (see Boyle 1999 for a review on the benefits of chicken
breeding). Based on research conducted at OI since 1995, there appears to be additive
genetic variation for resistance to TSV in L. vannamei, and significant improvements in
TSV resistance have been made. In a recent research trial, shrimp selected for TSV
resistance exhibited a mean family survival that was 18.4% higher than unselected
control shrimp after a TSV-challenge test (Argue et al. 2002). Similar challenge tests
conducted at UAZ from 1998-2000 revealed that mean survival of all TSV-challenged
families increased from 24% to 39% during this period (White et al. 2002). In addition,
mean survival of the best performing families increased from 65% in 1998 to 100% in
2000, and there are now commercial broodstock suppliers who claim to have families of
L. vannamei that exhibit >90% survival to TSV (e.g. Wyban 2000). The use of TSV-
resistant shrimp, in conjunction with on-farm biosecurity practices, contributed to a
significant increase in production from 1998-2002 (Fig. 2). Again, this impact was most
significant in Texas where production increased from 3.17 million pounds in 1998 to 8.27
million pounds in 2002 (Rosenberry 2003), representing a 161% increase in production
over four years.
Although there is no doubt that TSV-resistant shrimp can improve production and
profitability for those farmers who experience a TSV outbreak, there are compelling
reasons why breeding shrimp for resistance to a single viral pathogen, using current
selective breeding strategies, may not be the most prudent course of action for the long-
term viability of the shrimp farming industry (Moss et al. in press). Similar to other
organisms, there appears to be a trade-off between disease resistance and shrimp growth
(Chevassus and Dorson 1990, Henryon et al. 2002, CENIACUA and AKVAFORSK
2002). Also, there are concerns that results from laboratory disease-challenge tests may
not be predictive of survival in commercial ponds. Importantly, there are growing
concerns about viral mutations, whereby previously resistant shrimp strains may become
susceptible to evolving viruses. In fact, this situation occurred recently with TSV.
In 2001, significant mortalities of L. vannamei occurred at shrimp farms in Belize
resulting from TSV epizootics (Rosenberry 2001), and there were concerns that a new
TSV strain had emerged. Researchers from UAZ compared a TSV isolate from Belize
with the reference isolate from Hawaii to identify possible differences, using selected
OIE (Office of International de Epizooties) diagnostic methods and sequence analysis of
nucleotides and amino acids in the viral genome. These researchers concluded that the
two isolates exhibited different characteristics and thus represented different strains of the
virus (Erickson et al. 2004). Importantly, broodstock suppliers to Belize reported that
shrimp bred for resistance to the “old” Taura strain (Hawaii isolate) succumbed to the
“new” Belize strain. In response to these concerns, researchers from OI and UAZ
explored the possibility of developing selectively bred families of L. vannamei that
exhibited resistance both to the Hawaii and Belize TSV strains. In a recent trial, several
shrimp families from OI’s germplasm were identified as having high survival to the
Belize strain, and offspring from these families were produced and distributed to U.S.
5

broodstock suppliers and subsequently challenged with both TSV strains (Moss et al.
2003). Selectively bred shrimp exhibited 95% survival after exposure to the Hawaii
TSV. This was 75% higher than unselected control shrimp, which exhibited 20%
survival after exposure to the same virus. Importantly, selected shrimp exhibited 63%
survival after exposure to the Belize TSV, whereas all of the control shrimp died by day 4
of the challenge test. These results indicate that the Belize strain of TSV was more lethal
than the Hawaii TSV, although it is possible to develop lines of shrimp that exhibit some
resistance to both virus strains.
The Need for Biosecure Production Systems
In light of the limitations in breeding for disease resistance, selective breeding
should not be perceived as a panacea for the health problems plaguing the shrimp farming
industry. Rather, the industry needs to adopt strict biosecurity protocols to ensure its
future. On-farm biosecurity strategies were rapidly adopted by U.S. shrimp farmers in
the aftermath of the TSV epizootic in Texas in 1995, and included reduced water
exchange rates, filtering of pond influent, drying out of ponds over the winter, and
screening of postlarvae for diseases. Although these biosecurity measures, along with the
use of HH-SPR shrimp, contributed significantly to the increase in U.S. shrimp
production from 1998-2002, diseases continue to impact the domestic shrimp farming
industry. Recently, WSSV was reported in Hawaii and TSV has emerged again in south
Texas. In addition, necrotizing hepatopancreatitis (NHP) continues to be problematic in
Texas, for shrimp farmers, as its seasonal appearances have been ongoing since the mid-
1980s (Pantoja et al. 2003). Although no shrimp viruses were detected in Texas in 2002,
necrotizing hepatopancreatitis (NHP) continues to be problematic, as its seasonal
appearances have been ongoing since the mid-1980s (Pantoja et al. 2003). For shrimp
farmers to meet the growing demand for high-quality shrimp products, novel production
systems and management protocols must be designed to minimize the introduction and
spread of pathogenic agents, as well as to protect coastal resources. Biosecure shrimp
production systems represent an emerging alternative to traditional shrimp culture, and
provide a high degree of pathogen exclusion with minimal water exchange. In an effort
to develop biosecure technologies for the U.S. shrimp farming industry, several
researchers from the USMSFP are evaluating prototype systems that may have
commercial application (Browdy and Bratvold 1998, Moss et al. 1998, Ogle and Lotz
1998, Samocha and Lawrence 1998).
OI’s system consists of a concrete 58-m
2
raceway that is filled with seawater (34
ppt) from an underground aquifer to a depth of about 60 cm. A 2-HP, aspirator-type
aerator is used to provide aeration and to move water in a circular pattern around a central
baffle. Water flow produces a scouring velocity to keep solids in suspension. For
filtration, a 25-ft
3
propeller-washed bead filter is used for solids removal and biological
filtration (Malone et al. 1998). The bead filter allows a sufficient amount of microalgae
and other suspended particles to pass through so a “green water” environment was
maintained. This is important because shrimp reared in water with high concentrations of
microalgae and microbial-detrital aggregates grow better than shrimp reared in clean,
filtered seawater (Moss 2002). Clear, plastic sheeting (6 mil) is used to cover the
raceway as a biosecurity feature to reduce pathogen introduction by airborne vectors.
The cover also serves as an effective thermal insulator to maintain desirable water
temperatures.
Shrimp production data from this system are encouraging. In a recent trial,
juvenile shrimp were stocked at a density of 300/m
2
and were grown to a harvest weight
6

of 19.9 g in 12 weeks (Moss et al. 2002). During this trial, shrimp growth rate was 1.47
g/wk, survival was 86.3%, and production was 5.2 kg/m
2
(52,000 kg/ha/crop equivalent).
Importantly, the amount of water used to produce one kg of whole shrimp was about 352
L, and this is two to three orders of magnitude less than what is commonly used by the
existing shrimp farming industry. In 1992, Hopkins and Villalόn reported the volume of
water used by farmers to produce one kg of whole shrimp ranged from 39,000 to 199,000
L. Information used to determine these values came largely from semi-intensive shrimp
farms where liberal water exchange was a common management protocol. Over the past
decade, the global shrimp farming industry has made a concerted effort to reduce the
amount of water used during shrimp growout. The primary impetus for this change came
from efforts to mitigate the introduction of pathogens into the shrimp culture
environment, although the collateral benefits of reduced effluent discharge also were
recognized. Over the past several years, research institutions and commercial shrimp
farms have evaluated intensive shrimp production systems that rely on reduced or zero-
water exchange and results indicate that it is possible to reduce significantly the amount
of water used to culture shrimp (Table 3).
Table 3. Amount of water used to produce one kilogram of whole shrimp. Data are
from research institutions and commercial shrimp farms that culture shrimp under
intensive conditions and rely on reduced or zero-water exchange.
Species Water
Exchange
(%/Day)
Stocking
Density
(Shrimp/m
2
)
Water Use
(L/kg shrimp)
Reference
L. setiferus 25.0 40 64,000 Hopkins et al. (1993)
L. setiferus 2.5 40 9,000 Hopkins et al. (1993)
L. setiferus 0 20 6,000 Hopkins et al. (1993)
L.
vannamei
0 63-121 2,000 Fast & Menasveta
(2000)
L.
vannamei
< 10.0 35 1,500 Hamper (2000)
L.
vannamei
< 0.5 100 483 Moss et al. (2002)
L.
vannamei
< 0.5 200 370 Moss et al. (2002)
L.
vannamei
< 0.5 300 352 Moss et al. (2002)
Conclusions
Results from USMSFP research indicate that it is possible to produce > 5 kg/m
2
of market-sized shrimp (~ 20 g) in a biosecure production system in about 12 weeks,
using < 400 L of water per kg of shrimp. Although these results are encouraging, the
application of this capital-intensive technology only makes sense if shrimp can be
produced at a competitive price. Unfortunately, from June 2000 to June 2003, the Urner
Barry shell-on white shrimp index dropped from US$6.50 per pound to less than
US$3.50 per pound (Rosenberry 2003), thus making it very difficult for U.S. shrimp
farmers to make a profit. The good news is that per capita shrimp consumption in the
U.S. reached a record level in 2002 to 3.7 pounds. It is hoped that, with advanced
7

biosecure technologies available, the U.S. shrimp farming industry will be able to meet
these growing market needs by providing consumers with a high-quality product at a
competitive price. Such technologies will allow shrimp farmers to expand shrimp
production into areas away from the coast with greater control against the spread of
disease and without adversely affecting the environment.
Acknowledgments
We thank the Shrimp Program at the Oceanic Institute for their technical
assistance and commitment to excellence. USDA/CSREES Grant No. 99-38808-7431
awarded to the U.S. Marine Shrimp Farming Program supported this research.
Literature Cited
Argue, B.A., S.M. Arce, J.M. Lotz, and S.M. Moss. 2002. Selective breeding of
Pacific white shrimp (Litopenaeus vannamei) for growth and resistance to Taura
Syndrome Virus. Aquaculture 204:447-460.
Bonami, J.R., M. Brehelin, J. Mari, B. Trumper, and D.V. Lightner. 1990.
Purification and characterization of IHHN virus of penaeid shrimps. Journal of
General Virology 71:2657-2664.
Boyle, M.L. 1999. Chicken breeding and genetics. 1999. In: Controlled and Biosecure
Production Systems. Evolution and Integration of Shrimp and Chicken Models.
(R.A. Bullis and G.D. Pruder, eds.). The Oceanic Institute, Waimanalo, Hawaii,
pp. 23-28.
Brock, J.A., R.B. Gose, D.V. Lightner, and K. Hasson. 1997. Recent developments
and an overview of Taura Syndrome of farmed shrimp in the Americas. In:
Diseases in Asian Aquaculture III. (T.W. Flegel and I.H. MacRae eds.), Fish
Health Section, Asian Fisheries Society, Manila, the Philippines, pp. 275-284.
Browdy, C. L. and D. Bratvold. 1998. Preliminary development of a biosecure shrimp
production system. In: Proceedings of the US Marine Shrimp Farming Program
Biosecurity Workshop (S.M. Moss, ed.), The Oceanic Institute, Waimanalo,
Hawaii, USA, pp. 19-38.
Browdy, C. and S. Moss. (In press.) Shrimp culture in urban, superintensive closed
systems. In: Urban Aquaculture (B. Costa-Pierce, P. Edwards, D. Baker, and A.
Desbonnet, eds.), CABI.
Carpenter, N. and J.A. Brock. 1992. Growth and survival of virus-infected and SPF
Penaeus vannamei on a shrimp farm in Hawaii. In: Diseases of Cultured
Penaeid Shrimp in Asia and the United States. (W. Fulks and K.L. Main, eds.),
The Oceanic Institute, Waimanalo, Hawaii, pp. 285-294.
CENIACUA and AKVAFORSK. 2002. Selective breeding of Litopenaeus vannamei
in Colombia. Panorama Acuicola 7(2):30-31.
8

Chevassus, B. and M. Dorson, M. 1990. Genetics of resistance to disease in fishes.
Aquaculture 85:83-107.
Erickson, H.S., B.T. Poulos, D. Bradley-Dunlop, B. White-Noble, and D.V. Lightner.
2004. Diagnostic profile of Belize Taura Syndrome Virus. Global Aquaculture
Advocate 6(6):88-89.
Fast, A.W. and P. Menasveta. 2000. Some recent issues and innovations in marine
shrimp pond culture. Reviews in Fisheries Science 8:151-233.
Flegel, T.W. and V. Alday-Sanz. 1998. The crisis in Asian shrimp aquaculture: current
status and future needs. Journal of Applied Ichthyology 14:269-273.
Henryon, M., A. Jokumsen, P. Berg, I. Lund, P.B. Pedersen, N.J. Olesen, and W.J.
Slierendrecht. 2002. Genetic variation for growth rate, feed conversion
efficiency, and disease resistance exists within a famed population of rainbow
trout. Aquaculture 209:59-76.
Hopkins, J.S. and J. Villalón. 1992. Synopsis of industrial panel input on shrimp pond
management. In: Proceedings of the Special Session on Shrimp Farming. (J.
Wyban, ed.), World Aquaculture Society, Baton Rouge, Louisiana, pp. 138-143.
Hopkins, J.S., R.D. Hamilton, P.A. Sandifer, C.L. Browdy, and A.D. Stokes. 1993.
Effect of water exchange rate on the production, water quality, effluent
characteristics and nitrogen budgets in intensive shrimp ponds. Journal of the
World Aquaculture Society 24:304-320.
Jaenike, F., K. Gregg, and L. Hamper. 1992. Shrimp production in Texas using
specific pathogen-free stocks. In: Diseases of Cultured Penaeid Shrimp in Asia
and the United States. (W. Fulks and K.L. Main, eds.), The Oceanic Institute,
Waimanalo, Hawaii, pp. 295-302.
Kalagayan, H., D. Godin, R. Kanna, G. Hagino, J. Sweeney, J. Wyban, and J.
Brock. 1991. IHHN virus as an etiological factor in runt-deformity syndrome
(RDS) of juvenile Penaeus vannamei cultured in Hawaii. Journal of the Word
Aquaculture Society 22:235-243.
Lightner, D.V. 1999. The penaeid shrimp viruses TSV, IHHNV, WSSV, and YHV:
Current status in the Americas, available diagnostic methods, and management
strategies. Journal of Applied Aquaculture 9:27-52.
Lightner, D.V. 2003. Exclusion of specific pathogens for disease prevention in a
penaeid shrimp biosecurity program. In: Biosecurity in Aquaculture Production
Systems: Exclusion of Pathogens and Other Undesirables. (C-S Lee and P.J.
O’Bryen, eds.). The World Aquaculture Society, Baton Rouge, Louisiana, pp. 81-
116.
Lotz, J.M., C.L. Browdy, W.H. Carr, P.F. Frelier, and D.V. Lightner. 1995.
USMSFP suggested procedures and guidelines for assuring the specific pathogen
status of shrimp broodstock and seed. In: Swimming Through Troubled Water,
9

Proceedings of the Special Session on Shrimp Farming. (C.L. Browdy and J. S.
Hopkins, eds.), World Aquaculture Society, Baton Rouge, Louisiana, pp. 66-75.
Lotz, J.M. 1997. Disease control and pathogen status assurance in an SPF-based shrimp
aquaculture industry, with particular reference to the United States. In: Diseases
in Asian Aquaculture III. (T.W. Flegel and I.H. MacRae eds.), Fish Health
Section, Asian Fisheries Society, Manila, the Philippines, pp. 243-254.
Malone, R.F., L.E. Beecher, and A.A. DeLosReyes. 1998. Sizing and management of
floating bead bioclarifiers. In: Proceedings of the 2
nd
International Conference on
Recirculating Aquaculture. (G.S. Libey and M.B. Timmons, eds.), Virginia
Polytechnic Institute and State University, Roanoke, Virginia, pp. 319-341.
Moss, S.M., R.W. Doyle, and D.V. Lightner. (In press.) Breeding shrimp for disease
resistance: challenges and opportunities for improvement. In: Diseases in Asian
Aquaculture V. (P. Walker and R. Lester, eds.), Fish Health Section, Asian
Fisheries Society, the Philippines.
Moss, S.M., W.J. Reynolds, and L.E. Mahler. 1998. Design and economic analysis of
a prototype biosecure shrimp growout facility. In: Proceedings of the US Marine
Shrimp Farming Program Biosecurity Workshop (S.M. Moss, ed.), The Oceanic
Institute, Waimanalo, Hawaii, USA, pp. 5-18.
Moss, S.M. 2002. Dietary importance of microbes and detritus in penaeid shrimp
aquaculture. In: Microbial Approaches to Aquatic Nutrition within
Environmentally Sound Aquaculture Production Systems (C-S Lee and P.
O’Bryen, eds.), World Aquaculture Society, Baton Rouge, Louisiana, pp. 1-18.
Moss, S.M., C.A. Otoshi, A.D. Montgomery, and E.M. Matsuda. 2002. Recirculating
aquaculture systems for the production of market-sized shrimp. In: Proceedings
from the 4
th
International Conference on Recirculating Aquaculture (T.T
Rakestraw, L.S. Douglas, and G.F. Flick, eds.), Virginia Polytechnic Institute and
State University, pp. 245-254.
Moss, S.M., S.M. Arce, D.R. Moss, Y. Hwang, B. White-Noble, and D.V. Lighter.
2003. Selectively bred shrimp survive varied TSV exposure. Global Aquaculture
Advocate 6(4):16-17.
Moss, S.M., R.W. Doyle, and D.V. Lightner. in press. Breeding shrimp for disease
resistance: a panacea or pariah? In: Diseases in Asian Aquaculture V. (P.
Walker and R. Lester, eds.), Fish Health Section, Asian Fisheries Society, Manila,
the Philippines, pp. 243-254.
Ogle, J.T. and J.M. Lotz. 1998. Preliminary design of a closed, biosecure shrimp
growout system. In: Proceedings of the US Marine Shrimp Farming Program
Biosecurity Workshop (S.M. Moss, ed.), The Oceanic Institute, Waimanalo,
Hawaii, USA, pp. 39-48.
10

Pantoja, C.R. and D.V. Lighter. 2003. Necrotizing Hepatopancreatitis: diagnosis,
distribution in shrimp. Global Aquaculture Advocate, 6(4):18.
Pruder, G.D. 1994. High health shrimp stocks: an advance, an opportunity – but not a
panacea. World Aquaculture, 25(3):26-28.
Pruder, G.D., C.L. Brown, J.N. Sweeney, and W.H. Carr. 1995. High health shrimp
systems: seed supply – theory and practice. In: Swimming Through Troubled
Water, Proceedings of the Special Session on Shrimp Farming. (C.L. Browdy
and J. S. Hopkins, eds.), World Aquaculture Society, Baton Rouge, Louisiana, pp.
40-52.
Rosenberry, B. World Shrimp Farming 2000. 2000. San Diego, California, Shrimp
News International.
Rosenberry, B. World Shrimp Farming 2001. 2001. San Diego, California, Shrimp
News International.
Rosenberry, B. World Shrimp Farming 2003. 2003. San Diego, California, Shrimp
News International.
Samocha, T.M. and A.L. Lawrence, 1998. (Browdy and Moss in press). Preliminary
design and operating specifications for a biosecure shrimp growout facility in
Texas. In: Proceedings of the US Marine Shrimp Farming Program Biosecurity
Workshop (S.M. Moss, ed.), The Oceanic Institute, Waimanalo, Hawaii, USA, pp.
49-58.
Sindermann, C.J. 1990. Principle diseases of marine fish and shellfish. Volume 2.
Academic Press, New York, 516 pp.
White, B.L. , P.J. Schofield, B.T. Poulos, and D.V. Lightner. 2002. A laboratory
challenge method for estimating Taura Syndrome Virus resistance in selected
lines of Pacific white shrimp Litopenaeus vannamei. Journal of the World
Aquaculture Society 33:341- 348.
Wyban, J.A. , J.S. Swingle, J.N. Sweeney, and G.D. Pruder. 1993. Specific pathogen
free Penaeus vannamei. World Aquaculture 24:39-45.
Zavala, G. 1999. Biosecurity in the poultry industry. In: Controlled and Biosecure
Production Systems. Evolution and Integration of Shrimp and Chicken Models.
(R.A. Bullis and G.D. Pruder, eds.). The Oceanic Institute, Waimanalo, Hawaii,
pp. 75-78.
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