Current opinion

Parasitic diseases in marine cage culture An example

of experimental evolution of parasites?
Barbara F. Nowak
*
School of Aquaculture, Tasmanian Aquaculture and Fisheries Institute, University of Tasmania, Locked Bag 1370, Launceston 7250 Tasmania, Australia
Received 4 October 2006; received in revised form 9 January 2007; accepted 9 January 2007
Abstract
Rapid development of sh culture in marine cages has been associated with an emergence of parasitic diseases. There is a general trend
to an increase in infections with ectoparasites with direct life cycles and a reduced diversity of parasites in aquaculture. Some mariculture
creates conditions that are similar to serial passage experiments, which are used to study adaptation during experimental evolution of
pathogens. In particular, increased density of sh, repeated introduction of naive hosts, homogenous host populations, fast growth
and a potential decrease in genetic diversity are attributes of both aquaculture and serial passage experiments. Some free-living organ-
isms, for example Neoparamoeba spp. and Uronema spp. parasitise sh in culture, but have not been reported from wild populations.
Farming sh in marine cages can increase the risk of outbreaks of parasitic diseases, including those caused by opportunistic parasites.
However, aquaculture has the potential to control parasitic diseases through selective breeding, vaccination and general sh health
management.
2007 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.
Keywords: Parasites; Aquaculture; Experimental evolution; Neoparamoeba; Uronema; Amoebic Gill Disease; Swimmer syndrome
1. Introduction
As wild sheries continue to decline (Hutchings, 2000;
Worm et al., 2006), aquaculture has been expanding world-
wide (Eng and Tech, 2002; FAO, 2004). For many sh spe-
cies one of the grow-out options is the use of marine cages
(Table 1). While sh mariculture is mostly based on hatch-
ery-supplied sh, in some cases wild juveniles are captured
and grown in marine cages. For example, some tuna spe-
cies are captured then fattened in cages for a number of
months (Deveney et al., 2005; Aiken et al., 2006). Cage
design, farm size and intensity of the operation vary
depending on sh species and the location of the farm
(Eng and Tech, 2002). For most sh species breeding tech-
niques have been well established and manufactured feeds
are available. Stocking density depends on sh species
and carrying capacity. Almost all intensive cage maricul-
ture focuses on a single species and therefore can be treated
as monoculture. Marine cages have the advantage of lower
running costs than land-based facilities of equivalent
production capacity. However, there is less potential for
control over water quality and ow as well as the presence
of other marine life in the pens, including planktonic or net
fouling organisms.
As in other monocultures, disease outbreaks can become
an issue for mariculture industry. Increased host popula-
tion densities in aquaculture have been reported to result
in an increased virulence of microorganisms (Murray and
Peeler, 2005). For example, on the basis of phylogenetic
analysis of nucleotide sequences of the haemagglutinin
gene from 70 isolates, it has been suggested that the Infec-
tious Salmon Anaemia (ISA) virus has evolved from a wild
avirulent ancestor (Nylund et al., 2003). ISA outbreaks
could occur as a result of mutation of benign isolates in
farmed salmon following transmission from wild salmon
(Nylund et al., 2003). Aquaculture can therefore provide
0020-7519/$30.00 2007 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.ijpara.2007.01.003
*
Tel.: +61 3 6324 3814; fax: +61 3 6324 3804.
E-mail address: b.nowak@utas.edu.au
www.elsevier.com/locate/ijpara
International Journal for Parasitology 37 (2007) 581588
a suitable environment for the emergence, establishment
and transmission of new pathogens (Murray and Peeler,
2005).
2. Emergence of parasitic diseases in cage culture
The development of cage culture has been associated
with emergence of parasitic diseases (Kent, 2000). As anti-
parasitic treatment of sh in cages is often impractical or
expensive; parasitic diseases can have a serious economic
impact on marine pen culture. In some cases the parasitic
disease may not cause a direct loss due to sh mortality,
but increase production costs through treatment (for exam-
ple freshwater bathing against Amoebic Gill Disease) or
reduction in the product quality (for example due to Kudoa
spp. infections). Many parasites aecting sea cage maricul-
ture are ectoparasites (Table 2). Cultured sh are usually
fed manufactured feeds and as a result trophic transmission
of parasites is limited. However, even sh fed on manufac-
tured diets may sometimes consume marine invertebrates
(planktonic or from net biofouling) or wild sh. Trophic
transmission can therefore occur in aquaculture but it is
not as common as in the wild. As a result, cultured sh
have signicantly lower loads of parasitic species requiring
trophic transmission than wild sh. An overall reduction in
parasite diversity and a shift from endoparasites to ecto-
parasites is at least partly due to the use of manufactured
feed. While wild marine sh are hosts to a wide range of
parasites, sometimes the dominant parasite in culture
is either rare or absent in the same species in the wild
(Table 2). In cases where the dominant parasite species in
aquaculture is present in wild sh populations, adverse
eects are more obvious in farmed sh. For example, dis-
ease and mortality due to monogenean infections can be
common in aquaculture but are rare in wild populations
(Thoney and Hargis, 1991).
Some free-living organisms have parasitised cultured
sh, however they have never been reported from wild sh
(Table 2). This emergence of new parasites in mariculture
suggests evolution from free living organisms into opportu-
nistic parasites. For example, Neoparamoeba spp. are pres-
ent in marine and estuarine sediments in Tasmania
(Crosbie et al., 2003) as well as on cage netting (Tan
et al., 2002; Dykova et al., 2005) and have been found in
marine invertebrates such as sea urchins and crabs
(Dykova et al., in press). Neoparamoeba pemaquidensis
and Neoparamoeba branchiphila have been isolated from
Atlantic salmon with Amoebic Gill Disease and it is
unclear if both species or only one of them are causing
the disease (Dykova et al., 2005). Amoebic Gill Disease
aects farmed salmonids and farmed turbot, Psetta maxi-
ma, in a number of countries, including Australia, USA,
France, Ireland, Spain, Chile and New Zealand (Munday
et al., 2001), however the disease or the parasite have never
been reported from wild sh surveys, even when these sur-
veys included salmonids (Kent et al., 1998b) or other wild
sh species co-habiting with infected salmon in the same
cage (Douglas- Helders et al., 2002; Nowak et al., 2004).
It is currently unclear if the free living paramoebae are as
virulent as those colonising sh gills. So far, molecular
techniques have failed to dierentiate between isolates of
dierent origin (Dykova et al., 2005; Dykova et al., in
press). As Neoparamoeba spp. loose their virulence in cul-
ture (Morrison et al., 2005) it is currently impossible to test
infectivity of dierent isolates.
Similarly, Uronema spp. are free living scuticociliates
which were considered to be environmental scavengers
(Munday et al., 1997). There are 25 known species of
Table 1
Major mariculture sh species produced in cages ranked by production value
Fish species Scientic name Global aquaculture production (t) Global aquaculture
value (US$ 000)
Maximum stocking density
production (kg/m
3
)
Atlantic salmon
c
Salmo salar 1244637 4085052 25
Yellowtails and amberjacks Seriola spp. 150113 1275893 528
Milksh
a,b,d
Chanos chanos 573732 707836 4
Gilthead seabream Sparus aurata 90995 482922 1015
Coho salmon
c
Oncorhynchus kisutch 100967 344455 10
European seabass Dicentrarchus labrax 49103 313891 15
Japanese seabass Lateolabrax japonicus 219341 238962 30
Tunas
b
Thunnus spp. 11508 158150 4
Chinook salmon
c
Oncorhynchus tshawytscha 21517 90302 8
Barramundi
a
Lates calcarifer 29899 77733 1525
Turbot
a
Psetta maxima 6138 49060 30
e
Cobia Rachycentron canadum 20461 36206 14
Aquaculture production and value for 2004 based on FAO Fisheries Global Information System. Stocking densities: Atlantic salmon Crawford 2003,
yellowtails and amberjacks PIRSA, tunas PIRSA, other stocking densities sources listed in Acknowledgements.
a
Cages only one of the forms of grow-out, production data provided for all types of culture.
b
Wild seed used.
c
Freshwater hatchery.
d
Maximum stocking density calculated on the basis of information provided in Eng and Tech, 2002.
e
As turbot is a bottom dwelling sh stocking densities are provided in kg/m
2
.
582 B.F. Nowak / International Journal for Parasitology 37 (2007) 581588
Uronema-like ciliates. All these species are free living, how-
ever, some of them have been reported to parasitise cul-
tured sh, including southern bluen tuna Thunnus
maccoyii, olive ounder Paralichthys olivaceus, sea bass
Dicentrarchus labrax, seahorse Hippocampus abdominalis
and turbot (Yoshinaga and Nakazoe, 1993; Munday
et al., 1997; Sterud et al., 2000; Jee et al., 2001). Uronema
nigricans causes swimmer syndrome, resulting in mortali-
ties of southern bluen tuna in growout cages (Munday
et al., 1997). This parasite is invasive and causes encepha-
litis in farmed tuna and general tissue damage in other
farmed sh. While no specic surveys have been undertak-
en for Uronema in wild sh, so far surveys of wild sh have
failed to detect this species and clinical signs of this disease
have not been reported from wild tuna.
Neoparamoeba spp. and Uronema spp. are known to
only infect farmed sh. Both are free living protistans
with direct life cycles and no resting (cyst) stage. While
Neoparamoeba spp. is an ectoparasite that colonises gills,
Uronema spp. is an endoparasite infecting nervous tissues
of tuna. In both infections temperature appears to be
one of the risk factors. Neoparamoeba spp. gill infections
in Atlantic salmon cause more severe eects in summer
when temperatures are higher and salmon are close
to their upper temperature limit (Munday et al., 2001).
Uronema nigricans infects the brain of tuna in winter
when the temperatures are below 18 C, with most cases
occurring at temperatures below 15 C (Munday et al.,
1997). While Neoparamoeba spp. is a persistent problem
in salmon mariculture, Uronema spp. infections are much
more sporadic. The reasons why these parasites are
reported only from farmed sh are unknown. One possi-
ble explanation could be that parasites (particularly
protistans) can easily be overlooked in wild populations
as the health of these populations is monitored much less
frequently than farmed sh and diseased wild sh cannot
be tracked (Bakke and Harris, 1998). However, targeted
sampling failed to nd Amoebic Gill Disease in wild sh
despite 310 individual sh from 12 dierent species being
examined, including sh caught within the cages holding
infected salmon (Douglas-Helders et al., 2002; Nowak
et al., 2004). Therefore, explanations other than low sam-
pling frequency and diculties of detection of diseases in
wild populations should be considered.
3. Serial passage experiments as a form of experimental
evolution
Parasite performance and its local adaptation can be
measured either as infectivity (infection success) or
Table 2
Examples of parasites having a signicant eect on commercial marine cage culture of sh
Parasite Type of parasite Host Reported from wild
populations?
Reference
Neoparamoeba spp. Ectoparasite
Sarcomastigophora
Atlantic salmon, turbot No Kent et al. (1988a), Dykova et al. (2005)
and Douglas-Helders et al. (2002)
Uronema nigricans Endoparasite
Ciliate
Southern bluen tuna No Munday et al. (1997) and Munday et al. (2003)
Loma salmonae Endoparasite
Microsporidian
Chinook salmon Yes Kent et al. (1998b)
Kudoa spp. Endoparasite
Myxozoan
Salmonids, tuna, yellowtail Yes Munday et al. (1998)
Benedenia seriolae Ectoparasite
Monogenean
Yellowtail Yes Egusa (1983), Whittington et al. (2001),
Sharp et al. (2003) and Chambers and Ernst (2005)
Zeuxapta seriolae Ectoparasite
Monogenean
Yellowtail, amberjack Yes Egusa (1983), Sharp et al. (2003)
and Mansell et al. (2005)
Heteraxine heterocerca Ectoparasite
Monogenean
Amberjack Yes Egusa (1983)
Neobenedenia melleni Ectoparasite
Monogenean
Barramundi, cobia,
amberjack
Yes Mueller et al. (1994), Deveney et al. (2001)
and Liao et al. (2004)
Lepeophtheirus salmonis Ectoparasite
Copepod
Atlantic salmon Yes Beamish et al. (2006)
Caligus spp. Ectoparasite
Copepod
Salmonids, tunas,
yellowtail, barramundi,
milksh
Yes Sepu lveda et al. (2004)
Ceratothoa spp. Ectoparasite
Isopod
Salmonids, yellowtail,
European seabass,
sea bream
Yes Papapanagiotou and Trilles (2001)
and Mladineo (2003)
Yellowtail is Seriola spp.
B.F. Nowak / International Journal for Parasitology 37 (2007) 581588 583
virulence (damage to host), both of which are important
in infection processes (Dybdahl and Storfer, 2003). Serial
passage experiments are a form of experimental evolution
that is useful for studying adaptations. These experiments
are based on transmission in dense host cultures which
may present fewer constraints on the infection processes
found in nature (Ebert, 1998). These host cultures usually
have low genetic diversity and may be clonal or inbred,
so parasites may achieve higher virulence through
adaptation to specic genotypes (Ebert, 1998). Virulence
increases in more homogenous host populations. Further-
more, repeated introduction of naive hosts results in host
mortality having no cost to the parasite and this contrib-
utes to an increase in virulence in serial passage experi-
ments. A similar escalation of virulence could be
responsible for outbreaks of Amoebic Gill Disease and
swimmer syndrome in farmed sh. Are parasitic diseases
in aquaculture an example of experimental evolution of
parasites?
3.1. Low genetic diversity of host
While hatchery management usually ensures that genetic
variation is maintained, reduction in genetic diversity has
been reported in some farmed stocks of Atlantic salmon
(Norris et al., 1999), chinook salmon Oncorhynchus
tshawytcha (see Kim et al., 2004), turbot (Coughlan
et al., 1998) and sea bass (Bahri-Sfar et al., 2005). As par-
asites adapt to specic host genotypes, they can develop
greater virulence in monoculture where the host popula-
tions usually have lower genetic diversity (Ebert, 1998).
Greater parasite loads were observed in clonal sh than
in sexual populations in natural infections of Poecilopsis
sp. with trematode larvae from Uvulifer sp. (see Lively
et al., 1990). The only exception was when the sexual sub-
population was highly inbred. This suggests that parasites
evolve to disproportionately infect genetically uniform
strains as they become common (Lively et al., 1990). Het-
erogeneity within livestock populations is an important
factor stopping continued outbreaks of parasitic diseases
(Galvani, 2003). This also applies to cultured sh
populations.
3.2. Repeated introduction of naive host
In mariculture, na ve sh are introduced from hatcheries
to sea cages, in some cases not just once during the year,
but all year round. For example, in Atlantic salmon cul-
ture, the availability of spring smolts, out of season smolts
and triploids, collectively provide for a continuous whole
year supply of market sh, which relies on continuous
introduction of naive sh to the sea cages. This mimics
the situation in serial passage experiments (continued
introduction of a na ve host) and can increase the frequen-
cy of infections. In serial passage experiments, any factor
that increases the frequency of multiple infections will lead
to an increase in virulence (Ebert, 1998).
3.3. Dense host populations
Dense host populations are another aspect of serial
passage experiments and experimental evolution of para-
sites. While there are species-specic dierences in opti-
mum stocking density (and sometimes too low a
stocking density can be stressful to the sh) intensively
cultured sh are usually kept at greater densities than
occur in the wild (Conte, 2004, Table 1). Some species,
which do not school in the wild, form schools in cages.
This increases host proximity and therefore improves
the ability of a parasite to locate a host (Kabata, 1981).
For example, density of juvenile coral reef sh, Haemulon
avolineatum, had a positive eect on the success of estab-
lishment of a monogenean parasite, Haliotrema sp., in the
host population (Sasal, 2003). This increased availability
of a host due to increased population density, reduces
the negative eect of host mortality on parasite transmis-
sion. Cultured sh are also conned to a particular space
and have little opportunity to avoid infective stages of
parasites in the water column. Wild sh may be able to
avoid habitats associated with infection risk or avoid
schools infected with parasites (Barber et al., 2000). For
example, under laboratory conditions, uninfected stickle-
backs Gasterosteus aculeatus preferred to associate with
an uninfected shoal over one infected with the cestode
Schistocephalus solidus (Barber et al., 1998). Few avoid-
ance responses are possible for farmed sh. Leaping
behaviour in salmon in sea cages was signicantly corre-
lated with the number of sea lice on the sh, suggesting
that leaping behaviour is a response to the presence of
ectoparasites (Furevik et al., 1993).
3.4. Eect of host mortality
Most parasitic diseases in aquaculture are horizontally
transmitted. Unlike vertical transmissions, horizontal
transmissions do not rely on host survival or reproduc-
tion (Galvani, 2003). However, parasite-induced host
mortality has adverse eects on parasite transmission in
the wild. On the other hand, parasite transmission may
not be signicantly aected by host mortality in high
host densities or where naive hosts are repeatedly intro-
duced to the system. Some parasites can be transmitted
from dead hosts or even colonise dead hosts. For exam-
ple, Neoparamoeba spp., a causative agent of Amoebic
Gill Disease, was shown to colonise dead Atlantic salm-
on (Douglas-Helders et al., 2000). The freshwater mono-
genean Gyrodactylus salaris survived on a dead host and
the attached ukes maintained their infectivity for a
longer period than detached ukes (Olstad et al., 2006).
This shows that dead sh, particularly under culture con-
ditions, can be a signicant reservoir of infection even if
they were infected post-mortem. Under these circum-
stances an increase in virulence can be benecial for
the parasite, as in serial passage experiments (Ebert,
1998).
584 B.F. Nowak / International Journal for Parasitology 37 (2007) 581588
3.5. Host growth rates
The common aim of aquaculture is to grow sh quickly.
Cultured sh are usually fed more and grow faster than
wild sh. Gene transcription proles suggested a reduction
of basal metabolic rate and an increased metabolic ecien-
cy in juvenile farmed Atlantic salmon (Roberge et al.,
2006), which would result in allocation of resources
towards growth and fat deposition. Increased growth rate
could have an inuence on parasitic infections. For exam-
ple, in a laboratory infection the fastest growing stickle-
backs had the largest cestodes S. solidus, indicating that
at least for helminth parasites,the growth rates of the par-
asites and their hosts are related (Barber, 2005). Further-
more, host size seemed to determine parasite size for
some monogeneans, particularly polyopisthocotylean spe-
cies with more than four clamp pairs (Cone and Burt,
1985; Thoney, 1986, 1988; Hayward and Rohde, 1999).
For example, larger individuals of spot Leiostomus xanthurus
had larger monogeneans Heteraxinoides xanthophilis on
their gills (Thoney, 1988). Additionally, a positive regres-
sion between temperature-corrected maximum parasite
biomass and vertebrate host mass showed that a larger host
can support larger biomass of parasites. For sh, the same
relationship was also true for average parasite biomass and
host biomass (Poulin and George-Nascimento, in press).
4. Parasites biodiversity in aquaculture
A shift from endoparasites to ectoparasites and a reduc-
tion of parasite diversity can be observed in aquaculture.
For example, 17 ectoparasite species and 41 endoparasite
species have been recorded in wild Seriola spp. in Australia
(Hutson, K., personal communication), however only two
species of monogeneans (ectoparasites) are a serious prob-
lem in aquaculture of yellowtail kingsh Seriola lalandi
(Whittington et al., 2001; Chambers and Ernst, 2005; Man-
sell et al., 2005; Tubbs et al., 2005). In general, most para-
sites which have a potential economic eect on marine
aquaculture are ectoparasites with direct life cycles and
short generation time (Table 2). This is most likely due to
the use of manufactured feed and almost complete elimina-
tion of trophic interactions required by some metazoan
parasites for their transmission.
This shift to ectoparasites is also consistent with the
eects of eutrophication, which often also increases
parasitism (Laerty, 1997). In disturbed ecosystems, for
example by pollution, there is usually an increase in ecto-
parasitic infections and a decrease in endoparasitic infec-
tions (MacKenzie, 1999). Eutrophication promotes
parasites with simple life cycles and reduces biodiversity
of parasites (Kesting and Zander, 2000). However, there
was no environmental impact from sea bream Sparus
aurata and sea bass D. labrax farms in semi-exposed
locations in the Mediterranean (Maldonado et al., 2005).
Eutrophication may not even be an issue in cage culture
due to increased environmental monitoring (Crawford,
2003), fallowing and lower stocking densities (Maldonado
et al., 2005).
5. Parasite interactions between farmed and wild sh
populations
Parasite interactions between farmed and wild popula-
tions could be one potential outcome of increased viru-
lence. There are numerous known cases of transmission
of pathogens from domesticated animals to wildlife
(Daszak et al., 2000). It has been suggested that some par-
asites can be transmitted from cultured sh to wild popula-
tions (Butler, 2002; Morton et al., 2004, 2005; Krkosek
et al., 2005, 2006). Similarly, cases of transmission of
parasites from wild sh to cultured populations have been
postulated (Ho and Nagasawa, 2001; Rae, 2002). For
example, migrating wild chum salmon, Oncorhynchus keta,
was suggested to be a source of sea lice, Lepeophtheirius
salmonis, infection for cultured rainbow trout, Oncorhyn-
chus mykiss and coho salmon, Oncorhynchus kisutch, in
Onmae Bay, Japan (Ho and Nagasawa, 2001). One of the
most contentious issues is the eect of salmon farming on
wild salmon in British Columbia. In particular, it has been
suggested that farmed salmon is the source of sea lice which
threaten the existence of wild pink salmon, Oncorhynchus
gorbuscha, populations (Morton et al., 2004, 2005; Krko-
sek et al., 2005, 2006). However, survival of wild pink salm-
on was exceptionally good in 2003, indicating that this
species was not aected by sea lice from salmon farms in
that year (Beamish et al., 2006). Furthermore, experimental
infections of pink salmon with L. salmonis resulted in low
parasite abundance and no sh mortalities, suggesting little
obvious adverse impact of sea lice on pink salmon during
laboratory exposure of up to 28 days (Jones et al., 2006).
Interactions with wild populations and their parasites
may dier depending on whether the cultured sh species
is endemic or exotic. Exotic host species usually show a
reduced diversity of parasites, as few parasite species are
introduced with them and only some local parasites colo-
nise them (Torchin et al., 2003). In a survey of 26 host spe-
cies, including sh, native populations had twice as many
parasite species as introduced populations (Torchin et al.,
2003). If the cultured sh species was exotic, interactions
between the cultured and wild sh may be limited due to
the host-specicity of some parasites. However, the reduc-
tion in numbers of parasite species and the shift to ectopar-
asites may be still observed. For example, Atlantic salmon
Salmo salar has been introduced to Pacic Canada, Chile
and Australia. A survey in Chile showed that seven wild
sh species were hosts to 30 parasite species (including nine
ectoparasites and 21 endoparasites) and Atlantic salmon
had two main parasite species (including one endoparasite
and one ectoparasite) (Sepu lveda et al., 2004). A survey of
gill parasites in Atlantic salmon and wild sh in Australia
found two ectoparasites in farmed Atlantic salmon: an
amoeba, Neoparamoeba spp., and an isopod, Cerathotoa
sp.; however, these parasites could not be found on wild
B.F. Nowak / International Journal for Parasitology 37 (2007) 581588 585
sh collected on farms including from cages with infected
salmon (Douglas-Helders et al., 2002; Nowak et al.,
2004). These wild sh harboured seven species of parasites
on their gills, which were all absent from farmed Atlantic
salmon (Nowak et al., 2004). While this would at least partly
reect host-specicity of the parasites, it may not always be
the case. For example, Neoparamoeba spp. is known to infect
other sh species, unrelated to Atlantic salmon (Dykova
et al., 1995, 1998) but was absent from wild sh examined
in the survey in Australia (Douglas-Helders et al., 2002;
Nowak et al., 2004).
The degree of virulence of a horizontally transmitted
parasite is determined by selection to maximise parasite
transmission (Ebert, 1994). Local parasites show greater
infectiveness, reproductive success and virulence than
introduced parasites (Ebert, 1994). It has been suggested
that the high virulence of some introduced pathogens is
an exception and not a rule. This conrms the unpredict-
ability of interactions between novel combinations of hosts
and parasites (Ebert, 1994).
6. Conclusions
Most parasites are associated with a specic ecological
niche and diet (Marcogliese, 2002). Aquaculture alters both.
Therefore, parasites aecting aquaculture may be dierent
from those common in the same species in the wild, making
it dicult to assess risks of parasitic infections in new cul-
ture species. This is also true for agricultural pests due to
monoculture and management of culture systems which
can lead to localised environmental changes (Jones, 2001).
Aquaculture oers the potential for parasite control
through selective breeding, vaccination and general sh
health management, therefore limiting the experimental
evolution of parasites. Selective breeding of farmed sh
resistant to parasitic infections and the search for molecu-
lar markers has shown some promising results (Jones et al.,
2002). Although heritability markers to most parasites
show a moderate genetic component for this trait, there
is a potential for improving resistance to parasitic diseases.
The heritability value for resistance to Amoebic Gill Dis-
ease was moderate in Atlantic salmon families (Wynne
et al., in press). The heritability value for resistance in
Atlantic salmon families to sea lice L. salmonis varied from
0.06 in a natural infection (Kolstad et al., 2005) and 0.074
in another natural infection (Glover et al., 2005) to 0.26 in
a challenge (Kolstad et al., 2005). It was 0.22 for Caligus
elongatus (Mustafa and MacKinnon, 1999). Heritability
values larger than 0.20 suggest that selective breeding could
result in an improved resistance to those parasites
(Gjedrem, 2000). While vaccination against ectoparasites
still requires more research, there are a number of
approaches being tested to develop a sea lice vaccine
(Raynard et al., 2002). General improvements in farm
management, in particular cage positioning and setting
above the seabed together with improved lease location,
signicantly reduced the impact of U. nigricans on southern
bluen tuna farming (Munday et al., 2003; Nowak, 2004;
Nowak et al., 2006). These examples illustrate dierent
strategies aquaculture industry uses to control outbreaks
of parasitic diseases. These strategies would reduce the
potential for accelerated evolution of parasites.
However, aquaculture can contribute to the emergence
of new parasitic diseases, particularly those caused by
opportunistic parasites. Aggregation of sh in cages in
the marine environment can facilitate the spread of para-
sites that are rare or unreported in wild populations. Ecto-
parasites with direct life cycles are often promoted under
farming conditions. In comparison with wild sh of the
same species, there is usually a reduction in diversity of
parasites in cultured animals. As interactions between par-
asites of wild and cultured sh are not easy to predict, a
priori risk analysis for culture in marine cages, based only
on parasites of wild populations of the same species, may
be inadequate.
Acknowledgements
I would like to thank Dr. Rob Adlard and Dr. Tom
Cribb for inviting me to give a symposium presentation
on diversity of parasites in aquaculture at the Parasites in
Aquaculture and Marine Biodiversity Symposium at the
Australian Society for Parasitology and ARC/NHMRC
Research Network for Parasitology conference at the Gold
Coast, July 2006. This, together with the invitation from
IJP to submit a manuscript encouraged me to write this pa-
per. I would like to thank Dr. Craig Hayward for inspiring
discussions and providing references. I am grateful to Dr.
Brian Jones for sharing with me his ideas on tuna parasites
and to Dr. Rob Gurney for his comments. I thank Kate
Hutson for providing unpublished information on para-
sites of Seriola spp. in Australian waters, Dr. Francesc
Padros for information on stocking densities in European
seabass, Dr. Sonja Saksida for information on stocking
densities for salmonids farmed in Canada, Dr. Erik Vis
for information on stocking densities for Seriola spp. and
Japanese seabass in Japan, Dr. Carlos Zarza for informa-
tion on turbot culture in Spain. I am grateful to Dr. Craig
Hayward and two anonymous reviewers for their helpful
comments on this manuscript.
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