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Multitrophic Integration for Sustainable Marine Aquaculture

Chapter · December 2008

DOI: 10.1016/B978-008045405-4.00065-3

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T Chopin, S M C Robinson, M Troell, A Neori, A H Buschmann, and J Fang. Multitrophic

Integration for Sustainable Marine Aquaculture. In Sven Erik Jørgensen and Brian D. Fath (Editor-
in-Chief), Ecological Engineering. Vol. [3] of Encyclopedia of Ecology, 5 vols. pp. [2463-2475]
Oxford: Elsevier.
 Author's personal copy
Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture 2463

Multitrophic Integration for Sustainable Marine Aquaculture

T Chopin, University of New Brunswick, Saint John, NB, Canada
S M C Robinson, Department of Fisheries and Oceans, St. Andrews, NB, Canada
M Troell, The Royal Swedish Academy of Sciences, Stockholm, Sweden
A Neori, Israel Oceanographic and Limnological Research Ltd., Eilat, Israel
A H Buschmann, Universidad de Los Lagos, Puerto Montt, Chile
J Fang, Chinese Academy of Fishery Science, Qingdao, China
ª 2008 Elsevier B.V. All rights reserved.

Introduction The Development of Innovative Practices, such as

Geographical Expansion IMTA, Will Require Defining the Appropriate
Intensification of the Existing Sites Regulatory and Policy Frameworks, and Financial
Diversification Incentive Tools
Changes in Attitudes Are Needed Innovative Practices Conclusions
Need to be Developed Examples of IMTA Further Reading
Development in Several Countries

Introduction the industry’s large-scale dependency and impact on

The seafood industry is at a crossroads: while capture
fisheries are stagnating in volume and decreasing in prof-
itability, they are also falling short of world demand, as the
annual consumption of seafood has been rising, dou-bling in
three decades. This trend is expected to persist in the
decades to come. As was the case on land with the
acquisition of food by hunter–gatherer societies evolving
into food production with the development of agriculture, we
can no longer depend solely on fishing. Considering the
depletion of natural stocks, people developed the idea that
aquaculture could be increasingly important as the solution
for providing the difference between the demand for
seafood and the biomass available. Aquaculture pro-
duction, which already supplies 40% of the seafood
consumed worldwide, has increased by nearly 10% per year
over the last few decades, making it the fastest growing
global food production sector: from 40 million tonnes in
2004 to an anticipated 70 million tonnes by 2015. By
comparison, terrestrial farmed meat production grew by only
2.8% annually. The majority of aquaculture production still
originates from extensive and semi-intensive systems;
however, the rapid development of intensive marine fed
aquaculture (e.g., carnivorous finfish and shrimp)
throughout the world is associated with con-cerns about the
environmental, economic and social impacts these often
monospecific practices can have, especially where activities
are highly geographically concentrated or located in
suboptimal sites whose assim-ilative capacity is poorly
understood and, consequently, prone to being exceeded.
Without a clear recognition of
natu-ral ecosystems and traditional societies, the
aquaculture industry is unlikely to either develop to its
full potential or continue to supplement ocean fisheries.
For many aquaculture operations, monoculture is, spa-
tially and managerially, often the norm. Species are
cultivated independently in different bays or regions.
Consequently, the two different types of aquaculture (fed vs.
extractive) are often geographically separate, rarely
balancing each other out at the local or regional scale, and,
thus, any potential synergy between the two is lost. As the
volume of production of these monoculture opera-tions
increases to higher levels, the sector generally moves into a
commodity market where prices to the producer and
consumer generally drop because of the volumes involved.
This drop in profit margins to the producer encourages
practices that decrease the cost of production such as
implementation of more efficient automated technologies,
labor-saving strategies, less-expensive inputs, and subsidy-
based systems, whether they be natural (e.g., irrigation on
land or capture of wild species) or social (e.g., government
financial support). Other responses of the industry may be
to investigate the possibilities of securing a larger market
share via company consolidation, new markets, or new
products. If the industry has a high capital equipment and
operational overhead, shareholders will further encourage
this response. One problem with this approach is that the
solutions are not ‘open-ended’ and are ultimately limited in
their effectiveness (e.g., technology can only get you so far).
Another problem is that in an aquaculture environment with
fixed geographic limits (e.g., lease boundaries), this
increased production generally comes at the expense of the
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2464 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
natural environment, as the farmer tends to squeeze more
and more production from a fixed area. Once the natural
system is destabilized, the chances for the entire operation
to collapse increase (e.g., the shrimp industry in several
Asian countries). The underlying, erroneous assumption of
the above business model is that there is capacity within the
ecosystem for almost continuous expansion; otherwise, the
commodity-based system will ultimately fail at regio-nal
scales as someone will always be able to produce for less
cost than you. It is baffling to understand why we promote
these unsustainable practices as most people realize that it
goes against the concepts of balanced eco-systems,
conservation of energy, and carrying capacity of coastal
ecosystems. In the natural world, to avoid pro-nounced
shifts in coastal processes, the solution to nutrification by
fed aquaculture is not dilution, but con-version of the excess
nutrients and energy into other commercial crops produced
by extractive aquaculture (e.g., seaweeds and shellfish).
To continue to grow, while developing better manage-ment
practices, the aquaculture sector needs to develop more innovative,
responsible, sustainable, and profitable practices that optimize its
efficiency, create diversification, and ensure the mitigation of the
consequences of its activ-ities to maintain the health of coastal
waters. Maintaining sustainability, not only from an environmental,
but also from economic, social, and technical perspectives, has
become a key issue, increased by the enhanced awareness of more
and more demanding consumers regarding quality, traceability, and
production conditions. What, then, are the options for the
aquaculture sector to face these challenges, grow, and meet the
environmental, economic, and social concerns?
Geographical Expansion
Geographical expansion of monocultures is still possible in
some areas of the world, but for how long? In other areas,
site access and availability are already limited and public
resistance is growing against further expansion of the
current aquaculture model. Moving from sheltered
nearshore sites to exposed nearshore sites and offshore
sites is being contemplated, but technical and economic
challenges remain, especially in regions where the coastal
zone is already highly used by many stakeholders with
different and competing needs. Real offshore develop-ment,
proposed by some as the next frontier for development in
aquaculture, is not necessarily the appro-priate solution for
all regions. Moreover, present designs for offshore farms
are almost entirely for the development of fed monoculture
of ‘high-valued’ fish and rarely consider the association with
extractive aquaculture operations and their specific
requirements. For example, seaweed aquaculture needs
infrastructures near the sur-face to capture solar energy
needed for photosynthesis
and, consequently, nutrients available near the
surface (i.e., if the fed aquaculture infrastructure is
submerged, then upwelling water circulation is
needed to bring the nutrients near the surface). It is,
therefore, obvious that, sooner or later, the scope for
geographic expansion will be limited for the existing
monoculture technologies and practices.
Intensification of the Existing Sites
If the possibility for expansion of the existing finfish aqua-
culture sector is limited in spatial extent by biological,
economic, and social factors, the only other solution to the
expansion issue is to increase the production from existing
sites. This is analogous to the issue that faces human
populations in urban areas. The terrestrial solution has been
to increase the tridimensional surface area by using
buildings with multiple levels. When one considers the
seawater volume available at a lease site and the volume of
the water column actually occupied by a series of salmon
cages, it is obvious that a cultivation unit is not optimized.
The area of a lease site also has to accommodate room for
the anchoring systems, vessel access, water flow, etc.
Advanced technology will thus be a prerequisite for
intensification. As with concentrated housing for humans,
there will have to be a high degree of surface area for
organisms, and efficient systems for food delivery, waste
treatment, and energy supply. Consequently, intensification
will require: (1) innovative and environmen-tally friendly
technologies, (2) new and better management practices and
codes, and (3) recognizing aquaculture within a broader
integrated coastal management framework.
Diversification
It is amazing to realize how very little the aquaculture
sector is diversified in some countries or significant pro-
ducing regions. For example, the salmon aquaculture in
Canada represents 68.2% of the tonnage of the aquacul-
ture industry and 87.2% of its farmgate value. In Norway,
Scotland, and Chile, the salmon aquaculture represents
88.8%, 93.3%, and 81.9% of the tonnage of the aquacul-
ture industry, and 87.3%, 90.9%, and 95.5% of its
farmgate value, respectively. Conversely, while Spain
(Galicia) produces only 8% of salmon in tonnage (16% in
farmgate value), it produces 81% of its tonnage in
mussels (28% in farmgate value). Why do we think that
the common old saying ‘‘Do not put all your eggs in one
basket’’, which applies to agriculture and many other
businesses, would not also apply to aquaculture? Having
too much of your production in a single species leaves a
business vulnerable to issues of sustainability because
of low prices due to oversupply, and the possibility of
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Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
catastrophic destruction of your only crop (diseases,
damaging weather conditions). Consequently,
diversifica-tion of the aquaculture industry is
imperative to reducing the economic risk and
maintaining its sustainability and competitiveness.
The traditional view of diversification often means
producing another product along the same lines of the
first, that would fit into the existing production and mar-
keting systems. In finfish aquaculture, this has usually
meant salmon, cod, haddock, or halibut. However, from
an ecological point of view, these are all ‘shades of the
same colour’. No synergies are created; rather, these
situations compound the impacts on the system. True
ecological diversification means changes at more than
one trophic level, that is, switching from another species
of finfish to another group of organisms of lower trophic
level (e.g., shellfish, seaweeds, worms, bacteria, etc.)
more resembling a natural ecosystem. Staying at the
same eco-logical trophic level will not address some of
the environmental issues because the system will
remain unbalanced due to the nonstable distribution of
energy and nondiversified resource needs.
Economic diversification should also mean looking at
seafood from a different angle. Aquaculture products on the
market today are very similar to those obtained from the
traditional fishery resources, and are, thus, often in direct
competition. While this may be part of the market forces at
work, the opportunity exists to diversify from the fish fillets,
or mussels and oysters on a plate in a restaurant, to a large
untapped array of bioactive compounds of mar-ine origin
(e.g., pharmaceuticals, nutraceuticals, functional foods,
cosmeceuticals, botanicals, pigments, agrichemicals and
biostimulants, and industry-relevant molecules). Research
and development on alternative species should no longer be
considered as R&D on alternative finfish species, but rather
on alternative marine products.
Moreover, diversification should be viewed as
an invest-ment portfolio, with short-term, long-term,
high-risk, and low-risk components, and with long-
term growth and stability as the main objectives.
Changes in Attitudes Are Needed
There is a paradoxical situation when looking at current
worldwide food production. In agriculture, 80% of the
production is made up of plants and 20% of animal
products (meat, milk, eggs, etc.), while in aquaculture,
80% of the production is animal biomass and 20% is
plant biomass. Considering only mariculture, the world-
wide production is made up of 45.9% seaweeds, 43%
mollusks, 8.9% finfish, 1.8% crustaceans, and 0.4% of
varied other animals. Consequently, in many parts of the
world, aquaculture is not synonymous to finfish aquacul-
ture, as so many people in affluent western countries
2465
believe. If one is reasonably assessing, based on the need
for balancing the cultured species functions within the
surrounding ecosystem functions, marine herbivores, car-
nivores, and omnivores cannot be cultivated while
neglecting marine plants – as efficient biofilters, a crop on
their own, or a food component for other organisms – a fact
apparently missed by a certain number of the ‘Blue
Revolution’ proponents. Several species of seaweeds cul-
tivated under the right conditions, especially near sources of
high levels of nitrogen as in proximity to finfish farms, can
be excellent sources of proteins, important amino acids, and
unsaturated oils. We need to be aware of the other food
production systems in the rest of the world if we want to
understand our present prevailing system and correctly
position it in perspective with other systems. Seaweeds and
microalgivores represent 59% of the world aquaculture
production, followed by the production of 30% of omnivores
and detritivores. In tonnage, the three leading aquacultured
species are the seaweed Laminaria japonica, and two
microalgivores, the Pacific cupped oyster, Crassostrea
gigas, and the silver carp, Hypophthalmichthys molitrix.
Vocal public opposition to aquaculture has been generated
by ‘high-value’ salmonids and other carnivorous marine fish
and shrimp, which, in fact, represent only 11% of the world
aquaculture production.
From the above numbers for mariculture, one may be
inclined to think that at the world level, the two types of
aquaculture, fed and extractive, are relatively balanced.
However, because of the predominantly monoculture
approach, these different types of aquaculture production
are often geographically separate, and, consequently, rarely
balance each other out on the local or regional scale. For
example, in Eastern Canada, fed salmon aqua-culture is
presently located in the Bay of Fundy in southern New
Brunswick, while extractive mussel and oyster aquaculture
is located in the Northumberland Strait and the Lower Gulf
of St. Lawrence, along the coastlines of Prince Edward
Island and northeastern New Brunswick. In Japan,
aquaculture is mostly carried out with various bays
dedicated to either shellfish, sea-weed, or finfish
aquaculture. An interesting situation has emerged in
southern Chile with the recent development of mussel
(Mytilus chilensis) cultivation. Mussel long lines can be
found between salmon cages in channels and fjords;
however, the decisions regarding siting were not based on
scientific data for prevailing currents, suspended matter and
nutrient circulation, oxygen availability, etc., and the IMTA
concept was not explicitly considered. There are, however,
examples in China of bays managed according to the IMTA
approach.
It is also important to consider that while fish do
generally have a higher price per unit value (but not
necessarily at a higher return on investment), sustainable
ecosystems are not based on the price/value of the return
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2466 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
to humans, but on a balance of biomass between organ-
isms having different complementary functions and a
balance of energy flows. One can wonder if the needed
evolution (not revolution, which often takes you back to
the starting point as its etymology indicates!) in aquacul-
ture practices is more a conceptual question of
rethinking how food production systems should work
more than a technological question. In other words, how
to make the ‘Blue Revolution’ greener . . .?! It would also
be appropri-ate to ask the question ‘‘Blue Revolution, but
for whom?’’ Depending on the answer, different species
and different systems should be focused on.
Innovative Practices Need to be
Developed
The challenge, then, is how to increase the production
capacity of an existing site (increased production per area
without exceeding the ecosystem assimilative capacity)
when the other options have shown their limitations. One of
the possible answers is to increase the level of technol-ogy
involved in the production of seafood so that food and waste
handling systems are all actively considered in the system
design and operation protocols from the start, and are
modeled after natural ecosystems (or, at least, that resource
utilization is maximized by adding functionally different
species to the cultivation system).
One of the innovative solutions being proposed for
environmental sustainability, economic diversifica-tion, and
social acceptability, is integrated multitrophic aquaculture
(IMTA). This practice combines, in the appropriate
proportions, the cultivation of fed aquaculture species (e.g.,
finfish) with organic extractive aquaculture species (e.g.,
shellfish) and inorganic extrac-tive aquaculture species
(e.g., seaweed) for a balanced ecosystem management
approach that takes into consid-eration site specificity,
operational limits, and food safety guidelines and
regulations (Figure 1). The aim is to increase long-term
sustainability and profitability per cultivation unit (not per
species in isolation as is done in monoculture), as the
wastes of one crop (fed animals) are converted into
fertilizer, food, and energy for the other crops (extractive
plants and animals), which can in turn be sold in the market.
Feed is one of the core operational costs of finfish
aquaculture operations. Through IMTA, some of the food,
nutrients, and energy considered lost in finfish monoculture
are recaptured and converted into crops of commercial
value, while biomitigation takes place. In this way all the
cultivation components have an economic value, as well as
a key role in services and recycling processes of the
system, the harvesting of the three types of crops
participating in the export of nutri-ents outside of the coastal
ecosystem (Figure 2). It is important to consider that these
systems do not work in
Integrated multitrophic aquaculture
(IMTA)
Fed aquaculture Extractive aquaculture
(Finfish) + Organic Inorganic
(Shellfish) (Seaweed)
DIN
POM
Effluent plume
Figure 1 Conceptual diagram of an integrated multitrophic
aquaculture (IMTA) operation including the combination of fed
aquaculture (e.g,. finfish) with organic extractive aquaculture
(e.g., shellfish), taking advantage of the enrichment in
particulate organic matter (POM), and inorganic extractive
aquaculture (e.g., seaweeds), taking advantage of the
enrichment in dissolved inorganic nutrients (DIN).
isolation and that solar energy and atmospheric and
ter-restrial inputs must also be factored in. Moreover,
the biomass and functions of the fed and extractive
species naturally present in the ecosystem in which
aquaculture farms are operating must also be
accounted for or this will lead to the development of
erroneous carrying capacity models.
The IMTA concept is extremely flexible. It can be applied
to open-water and land-based systems, and mar-ine and
freshwater systems (sometimes then called ‘aquaponics’ or
‘partitioned aquaculture’). What is impor-tant is that the
appropriate organisms are chosen based on the functions
they have in the ecosystem and, moreover for their
economic value or potential. What is quite remarkable, in
fact, is that IMTA is doing nothing other than recreating a
simplified, cultivated ecosystem in bal-ance with its
surroundings instead of introducing a biomass of a certain
type one thinks can be cultivated in isolation of everything
else. By using extractive species for the biomitigation of fed
species activities, the envir-onmental costs of a fed
monoculture are internalized, hence increasing the overall
profitability of the IMTA farm, especially when the costs and
benefits to nature and society of aquaculture wastes and
their mitigation will be quantified and associated with
discharge regulations.
The paradox is that IMTA is not a new concept. Asian
countries, which provide more than two-thirds of the
world’s aquaculture production, have been practicing
IMTA, through trial and error and experimentation, for
centuries. Even if the cultured species are different
(Figure 3), why, then, is this common-sense solution not
more widely implemented, especially in the western
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Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture 2467

Ecosystem-based
integrated multitrophic aquaculture
management concept

Atmospheric
inputs
Terrestrial
inputs Feed

Macroalgivores Macrophytes Nutrients Fed organisms

Abalone Seaweeds Fish
Dissolved
Sea urchin Aquatic plants Shrimp
Fish inorganic
Crop 2: Crop 1:
extractive plants
Planktivores fed animals
and Plankton
omnivores Microalgae
Shellfish Bacteria
Fish Protozoa
Sea cucumber Particulate
Worms Zooplankton organic
Crop 3:
extractive animals

Figure 2 Ecosystem-based integrated multitrophic aquaculture (IMTA) management concept. The wastes of one crop (fed animals) are
recaptured and converted into fertilizer, food, and energy for the other two crops of commercial value (extractive plants and animals), while
biomitigation takes place and the harvesting of the three types of crops participates in the export of nutrients outside of the coastal ecosystem.
Solar energy and atmospheric and terrestrial inputs must also be factored in. The biomass and functions of the fed and extractive species
naturally present in the ecosystem in which aquaculture farms are operating must also be accounted for.

% modern mariculture systems was initiated in the 1970s.

100
80
60
40
20
0
World Asian Western
countries countries
Figure 3 World aquaculture biomass production of the four
major farmed groups (fish, mollusks, crustaceans, and marine
plants) and differences in their distribution (%) between Asian
and Western world countries.
world? The reasons for this generally center around social
customs and practices that we are already familiar with,
even if common sense tells us that we should modify them.
Human society does not change quickly unless there are
compelling reasons to. The conservative nature of our
marine food production industries is a good exam-ple of the
relative slowness with which changes are adopted,
especially when dealing with a complex aquatic
environment, which we mostly see only the surface of, and
have difficulty understanding the processes taking place
beneath it over considerable distances and volumes.
Western countries are regularly reinventing the wheel.
Research on integrated methods for treating wastes from
Fish After that period, the scientific interest in IMTA stagnated,
Mollusks
and it was not until the late 1980s and early 1990s that a
renewed interest emerged, based on the common-sense
Crustaceans
approach that the solution to nutrification is not dilution but
Marine conversion within an ecosystem-based management
plants
perspective. In recognition of this growing interest, the
Aquaculture Europe 2003 Conference in Trondheim,
Norway, whose theme was ‘Beyond Monoculture’, was the
first large international meeting (389 participants from 41
countries) with IMTA as the main topic. In 2006, at the joint
European Aquaculture Society and World Aquaculture
Society Conference in Florence, Italy, IMTA was recognized
as a serious research priority and option to consider for the
future development of aquaculture prac-tices. The
determination to develop IMTA systems will, however, only
come about if there are some visionary changes in political,
social, and economic reasoning. This will be accomplished
by seeking sustainability, long-term profitability, and
responsible management of coastal waters. It will also
necessitate a change in the attitude of consumers toward
eating products cultured in the marine environment in the
same way that they accept eating products from recycling
and organic production systems on land, for which they are
willing to pay a higher price. At the present time, several
organizations are trying to modify seafood consumption
tendencies by incorporating such concepts as food safety,
and environmental and social sustainability.
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2468 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
Examples of IMTA Development in Several
Countries
Canada
An interdisciplinary team of scientists from the University of
New Brunswick in Saint John and from the Department of
Fisheries and Oceans in St. Andrews has been working
together on an Atlantic salmon (Salmo salar)/kelp
(Saccharina latissima and Alaria esculenta)/blue mussel
(Mytilus edulis) IMTA project in the Bay of Fundy since 2001
(Figures 4 and 5). The project, initially supported by
AquaNet, the Canadian Network of Centres of Excellence
for Aquaculture, is now moving from the R&D phase
towards the C (commercialization) phase with the support of
the Atlantic Innovation Fund from the Atlantic Canada
Opportunities Agency, and industrial and government
Figure 4 Harvesting of kelps (Saccharina latissima) cultivated in
proximity to Atlantic salmon (Salmo salar) cages at an IMTA site
in the Bay of Fundy, Canada. Photo by M. Sawhney.
Figure 5 Blue mussel (Mytilus edulis) raft and a mussel sock
grown next to Atlantic salmon (Salmo salar) cages at an IMTA
site in the Bay of Fundy, Canada. Photo by S. Robinson.
partners (Cooke Aquaculture Inc., Acadian Seaplants
Limited, the Canadian Food Inspection Agency, and the
New Brunswick Innovation Foundation). Because AquaNet
was funded by three Canadian federal granting agencies –
the Natural Sciences and Engineering Research Council of
Canada (NSERC), the Canadian Institutes of Health
Research (CIHR), and the Social Sciences and Humanities
Research Council of Canada (SSHRC) – and Industry
Canada, it was the ideal programme for addres-sing the key
interdisciplinary aspects of the IMTA concept: environmental
sustainability, economically viable diversi-fication, food
safety/security, and social acceptability (Figure 6). This
project, which has five modules (seaweeds, mussels, food
safety/health indicators, environment, and socioeconomics),
is on the verge of demonstrating the biological and
economic validity of the IMTA concept.
The first phase of the project (2001–04) (Figure 7) was a
period of testing the IMTA concept and of trying various
aquaculture sites having different oceanographic regimes in
order to understand how three species (Salmo salar,
Saccharina latissima, and Mytilus edulis), with different bio-
logical and physiological requirements, can be cultivated in
proximity to each other and what range of site
Exogenous Endogenous
(e.g., therapeutants) (e.g., harmful algae)
Dollar-
Regulation Safety I specific
changes
S n species
o d
c u
i s
a t
l r
Acceptability Biological y Cost-
effective
practices
Organic Inorganic
loading loading
Figure 6 The different interdisciplinary aspects covered by the
IMTA project in the Bay of Fundy, Canada: environmental
sustainability (through nutrient organic and inorganic loading),
economically viable diversification (through the choice of
commercial second crops and cost-effective practices), food
safety security (through the study of exogenous and
endogenous sources), and social aspects (adapting regulations
and policies for the development of acceptable practices).
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Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
EI-17
Production Filter Safety Seaweeds Production
feeders
Feeding
Physical
Social
Economics Acceptability O2 Modeling
- Benthos
- Salmon
- Disease Carrying capacity/
transfer industrial scale-up
Figure 7 Development of the IMTA project in the Bay of Fundy,
Canada, in three phases (see text for explanation of the
different phases).
characteristics could be applicable to a multitrophic opera-
tion (intermediate exposed sites worked the best in terms of
biomitigation potential and growth of the additional crops).
Kelp culture techniques have been improved. The
laboratory phase of the growth of the microscopic stages of
kelps has been reduced from 113 to between 30 and 40
days. Shortening and refining the required laboratory phase
is key for the development of commercial cultivation
(substantial operational savings, earlier transfer to sea for
higher biomass production, and potential production of
multiple crops for increased nutrient removal capacity of the
IMTA system). The kelp biomass production at the grow-out
sites has been increased from 8.01 to 20.7 kg m 1 of rope.
The onsite cultivation system has evolved from a system of
monolines at different depths deployed between the
compensator buoys of the salmon cages to parallel double-
and triple-line systems, and presently a system of raft units,
which can be placed near, but independently of the salmon
grid structure, according to the nutrient plume and
hydrodynamic conditions identified for each site. The
juvenile mussels required for growing purposes in IMTA
operations were found to be readily available from wild
mussels fouling the salmon nets. These were simply col-
lected during net changes and put into growing socks (mesh
tubes). The cultured mussels opportunistically took
advantage of the increased levels of particulate organic
matter at salmon sites during periods of feeding, using
particles in the 2–100 mm range, particularly in the fall-early
winter season when natural food levels decreased in the
natural environment. Due to the increased energy available
to the mussels, their growth rates and reproduc-tive rates
increased and some of their biochemical constituents
(glycogen, omega-3 fatty acids) were signifi-cantly elevated,
resulting in a very high-quality aquaculture product.
Increased growth rates of kelps
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Phase 1: (46%) and mussels (50%) cultured in proximity to fish farms,
Testing
concepts
compared to reference sites 1250 and 200 m away,
respectively, reflect the increase in food availability and
energy. In fact, in the hydrodynamically complex system of
the Bay of Fundy, it appears that to take full advantage of
the additional organic particulate matter available mussel
Phase 2: rafts need to be very close to the salmon cages (within 50
Linkages m), whereas seaweed rafts do not need to be in immedi-ate
proximity to salmon cages as long as they are within the
inorganic nutrient plumes (within a few hundred meters).
Phase 2 of the project, which is investigating the lin-
kages between the different species, is underway. Nutrient,
biomass, and oxygen levels are being monitored to model
Phase 3 the biomitigation potential of an IMTA site. Over 6 years,
none of the therapeutants used in salmon aquaculture have
been detected in kelps or mussels collected from the IMTA
sites; levels of heavy metals, arsenic, PCBs, and pesticides
have always been below the regulatory limits prescribed by
the Canadian Food Inspection Agency, the USA Food and
Drug Administration, and the European Community
Directives. A taste test of market-size mussels between
salmon-farm versus reference-site samples showed no dis-
cernable difference. Monitoring of phycotoxins (domoic acid
(DA) produced by Pseudo-nitzschia pseudodelicatissima
and paralytic shellfish poisoning (PSP) toxins produced by
Alexandrium fundyense) over 6 years has revealed the
absence of DA in mussel and kelp tissues. Levels of PSP
toxins in mussel tissues follow the seasonal patterns of A.
fundyense abundance with peaks lagging by 3–8 days.
Mytilus edulis also has the capacity to readily depurate PSP
toxins over time. Levels of PSP toxins in mussels can
exceed regulatory limits in late spring and summer, but with
appropriate management and monitoring regimes similar to
those in other shellfish production regions in other parts of
the world, the harvesting of mussels for human consumption
could take place safely during the other times of the year.
These encouraging results regard-ing the food safety of
organisms cultivated under IMTA conditions are presently
being used to help define the appropriate regulatory and
policy framework for the com-mercial development of IMTA
in Canada.
Phase 2 was also a period for testing the social accept-
ability of the IMTA concept and initiating its economic
modeling. A survey of aquaculture attitudes found that the
general public is more negative toward current mono-culture
practices and feels positive that IMTA would be a better
path to follow for marine food production. A focus group
social study revealed that most participants felt that IMTA
has the potential to reduce the environmental impacts of
salmon farming (65%), benefit community economies and
employment opportunities (93%), and improve the industry
competitiveness (95%) and sustain-ability (73%). All felt that
seafood produced in IMTA systems would be safe to eat
and 50% were willing to pay 10% more for these products if
labeled as such, which
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2470 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture

opens the door to developing markets for premium pro-

ducts, either environmentally labeled or organically certified.
Preliminary data of a bioeconomic model, in which net
present value calculations are conducted over 10 years to
portray long-term variability, show that the addition of
seaweeds and mussels to existing salmon farm-ing is more
profitable and helps reduce economic risks through
diversification. A preliminary economic scenario for the Bay
of Fundy shows that farming these three types of organisms
in an IMTA setting would provide CDN$44.6 millions in extra
revenues and 207 new jobs in a sector presently worth
CDN$223 millions and already employing 1683 people
directly and 1322 indir-ectly. Including the value of the
bioremediative benefits would make the case even more
convincing. While the revenues generated from IMTA
operations mentioned above are gross revenues, it is
important to note that the additional expenses associated
with these operations are incremental. Much of the capital
infrastructure (boats, barges, storage facilities,
transportation, etc.) is already in place with the existing
aquaculture operations as is some of the labor. Therefore,
the cost to produce an additional crop is often much less
than that required to produce it in a monoculture operation.

Site selection for the best compromise between site

characteristics, species selection, and market demands will
be key to optimizing IMTA systems. In phase 3, further
scaling-up of cultivation systems (rafts), species diversifica-
tion, economic analysis, and development of niche markets
will be implemented. Scaling-up to commercial level will
allow investigation into the impacts of IMTA on the carry-ing
capacity of the coastal environment, water and benthos
quality, potential for disease transfer, and animal and plant
health. The present IMTA system has three components
(salmon, kelps, and mussels), but this is admittedly an initial
simplified system; there is room for a more advanced sys-
tem with several other components by growing sea
cucumbers, polychaetes, sea urchins, etc. These organisms
can have different functions, or similar functions but in a
different size bracket of particles, for example.
South Africa
Abalone (Haliotis midae) aquaculture in South Africa is an
example in which IMTA is practiced at a large scale. These
systems have evolved from both the ‘trial and error’
approach and small-scale scientific experiments. It is still in
its early stage but the potential is high as South Africa is the
largest producer of cultured abalone outside Asia. Abalone
is cultured in land-based flow-through tank systems and fed
kelps (Ecklonia maxima) or pellet feeds. Over 6000 tonnes
of kelps are harvested annually on the South African west
coast for abalone feed, and some kelp beds are now
reaching sustainable limits of exploitation (Figure 8). A
growing body of evidence suggests that a
Figure 8 Seaweed farming integrated with abalone operations
will lessen the pressure on harvesting wild seaweed (Ecklonia
maxima) beds, close to their sustainable exploitation limits, in
South Africa. Photo by M. Troell.
mixed diet of kelps and other seaweeds can induce
growth rates at least as good as with artificial feed, can
improve abalone quality, and reduce parasite loads. Ulva
and Gracilaria are cultivated in the wastewater from the
aba-lone and one farm on the southeast coast currently
grows most of its feed (4 tonnes per working day of
Ulva) in shallow raceways (Figure 9). Seaweeds grown
in abalone wastewater have an increased nitrogen
content, resulting in value-added seaweeds with over
40% protein dry weight content and, hence, of excellent
quality to feed abalone.
The general benefit from IMTA, that is, reduction of
nutrient release to the environment, is also true for inte-
grated seaweed–abalone culture. Furthermore, as
seaweeds remove ammonium from the seawater and add
oxygen, the abalone wastewater passing through seaweed
ponds can be partially re-circulated back to the abalone
tanks, thus potentially reducing pumping costs. The abil-ity
to operate in re-circulation mode is important as red tides
occasionally occur along the South African coast. Moreover,
some coastal areas experience heavy traffic of tanker boats,
which represent potential risks for oil spills. It has been
shown that a farm can operate successfully at
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Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
Figure 9 Shallow seaweed (Ulva lactuca, U. rigida, and U.
fasciata) raceways receiving the effluents of the abalone
(Haliotis midae) covered tanks (left) as source water at the
Haga Haga farm, 70 km from East London, on the southeast
coast of South Africa. Photo by R.J. Anderson.
50% re-circulation, and even higher recirculation (up to
100%) can be sustained for shorter periods. This can, of
course, be optimized, depending on what the main
objec-tive is with re-circulation. The re-circulation
through seaweed tanks/ponds also has the potential to
raise water temperature, which can stimulate abalone
growth in areas of cold coastal waters. Compared to
many other aquaculture operations, there is currently no
real envi-ronmental pressure from abalone wastewater
release in South Africa. Wastes from abalone operations
are differ-ent from those of fish, with significantly lower
concentrations of both nitrogen and phosphorus. This
implies that the seawater in the seaweed tanks needs to
be fertilized to sustain seaweed growth. This additional
input of nutrients would not be needed if seawater from
fish tanks were to be used (this has been tested with
success). The development of IMTA in South Africa has,
in fact, been driven by other incentives, such as future
limitation of wild kelp harvesting and the proven eco-
nomic benefits from improved abalone growth and
quality with seaweed diets.
Israel
The development of first a two-stage, and then a three-
stage, on-land IMTA farm started as an R&D project at the
Israeli National Center for Mariculture in Eilat in the late
1980s to mid-1990s. Once the integration of the cultivation
of the fish, seaweed, and abalone was demon-strated, and
the basic design criteria experimentally defined, the
commercial farm, SeaOr Marine Enterprises Ltd., was
designed and constructed by private investors (with financial
support from the Israeli government and the European
Commission), about 0.5 km from the
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Figure 10 The SeaOr Marine Enterprises Ltd. IMTA farm in
Mikhmoret, Israel. The abalones (Haliotis discus hannai) are
grown in the white building in the background, the green-
covered fishponds (Sparus aurata) are in the middle, and the
elongated seaweed ponds (Ulva sp. and Gracilaria sp.) are in
front. Photo by M. Shpigel and B. Scharfstein.
Mediterranean Sea coast, at Mikhmoret, 35 km north of
Tel Aviv (Figure 10). The first phase of the farm began in
early 2001. It has been growing steadily and improving
its operation protocols to become a modern versatile
inten-sive IMTA farm; the technology is generic and
modular, adaptable to several fish/shrimp, shellfish, and
seaweed culture combinations at any level of
intensification. The farm recently changed ownership
and was renamed Sakura, Products from Nature.
Seawater is pumped from a deep well, next to the sea,
into a header tank; it then drains into the abalone (Haliotis
discus hannai) tanks and from there into the fishponds
(gilthead seabream, Sparus aurata), sedimentation ponds,
seaweed ponds (Ulva sp. and Gracilaria sp.) and, finally,
into the sump. From there, the seawater can be recycled or
released into a nearby estuary, as it meets all environ-
mental regulations for point-source discharge. The algal
biofilters efficiently extract from the fishpond effluents
dissolved nutrients, CO2, and BOD, and recharge the
effluents with dissolved oxygen. The nutritious algae
produced are marketed, or fed to the algivores, abalone in
this particular case. The farm has sold its products in the
local market; however, its main market will be Japan.
Chile
In Chile, the IMTA approach started in the late 1980s.
The first attempt considered the development of land-
based intensive marine systems using pumped seawater
to intensively culture trouts (Oncorhynchus mykiss) at a
stock-ing biomass of 35 kg m 3. The fish effluents were
then used for the cultivation first of oysters (Crassostrea
gigas) and second of the agar-producing alga Gracilaria.
The results showed that tank cultivation of trouts was
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2472 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
Figure 11 A typical modern Atlantic salmon (Salmo salar) farm,
with an automated feeding barge, in the Los Lagos Region of
southern Chile. Photo by A. Buschmann.
technically feasible and economically viable if a high
density of fish could be reached and the size of the
farm was sufficiently large to pay for the investment.
In addi-tion, the oysters and the algae were able to
reduce nitrogen and phosphorus significantly.
The cage culture approach for salmon species started
in the early 1990s and developed at an explosive rate, to
the point that Chile is now the second largest salmon
producer in the world, after Norway (Figure 11). This
explosive growth prevented the adoption of alternative
farming strategies, like IMTA, as the industry had no
immediate incentive to modify a very successful financial
story. Following the tendencies in Northern Hemispheric
countries, concerns about the unsustainable nature of
fed aquaculture in Chile started to be raised in the
second half of the 1990s. This encouraged the
development of studies integrating the cultivation of
oysters and seaweeds (Gracilaria and Macrocystis). The
first trials were successful and demonstrated that an
IMTA approach was an addi-tional way for developing a
more sustainable aquaculture approach for the country.
It has, however, not been easy to adopt an IMTA
approach in Chile. Like oriental countries, Chile has a long
tradition of shellfish and seaweed consumption; however,
the price for these goods is very low, therefore, they cannot
be suggested as an interesting business for investors. For
this reason, the Research Centre for Coastal Resources
and Environment (i-mar) of the Universidad de Los Lagos,
in Puerto Montt, has a plan of action parallel to the IMTA
research programme, which involves publishing and
obtaining patents related to novel uses of seaweeds to
encourage the farming of these organisms. The
development of abalone cultivation is presently emerging in
Chile, adding an extra pressure on natural resources of
seaweeds as a source of feed. A
Figure 12 The kelp (Macrocystis pyrifera) cultivated on long
lines near Atlantic salmon (Salmo salar) cages in southern
Chile. Photo by A. Buschmann.
pilot scale farm (4–5 ha) is already producing the brown
alga Macrocystis (Figure 12) and has demonstrated its
technical and economic feasibility. The implementation
of these new cultivation approaches will probably take
place in the coming years and it is important that
research keeps up with development in order to promote
better understanding of the results.
Basic and applied research must come together to
create the conditions necessary for the establishment of a
sustainable aquaculture activity in Chile and other parts of
the world. Besides the development of demonstration units
and applied research to clearly show farmers and regulators
the benefits of IMTA, basic research on, for example,
nutrient uptake and assimilation, remains neces-sary. The
environmental requirements for the growth of seaweeds and
shellfish may not be the same as for maxi-mizing their
biofiltering capacity. Their nutrient recycling efficiency can
be optimized by manipulating the culture conditions (depths,
relative position with respect to the salmon cages in relation
to the prevailing currents, distance from the cages, and
culture density), allowing for both a high biofiltering
efficiency and increased biomass production capacity to
make the IMTA approach commercially attractive.
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Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
China
Two variations of the IMTA approach have been
devel-oped in China: suspended multispecies
aquaculture, generally in shallow nearshore
waters, and multispecies large-scale sea ranching
in more offshore and deeper waters.
An example of suspended multispecies aquaculture is
what is being developed in Sungo Bay, in the East of the
Shandong Peninsula. Scallops (Chlamys farreri, 2100
tonnes fresh weight in 2005) and oysters (Crassostrea
gigas, 110 000 tonnes fresh weight) are cultivated, on the
same long line system, with the kelp, Laminaria japonica (80
000 tonnes fresh weight) (Figures 13 and 14). The cultivation
zone extends to 8 km offshore with a water depth of around
20–30 m. The co-cultivation of abalone (Haliotis discus
hannai, 1000 tonnes fresh weight) with L.
Figure 13 Large-scale IMTA of scallops (Chlamys farreri),
oysters (Crassostrea gigas), and kelps (Laminaria japonica) in
Sungo Bay, Shangong Province, China. Photo by M. Troell.
Figure 14 Harvesting of kelps (Laminaria japonica) in Sungo
Bay, Shangong Province, China. Photo by M. Troell.
2473
japonica is also being developed, with abalones kept in
lantern nets hanging vertically from the long lines, while
kelps are grown on ropes maintained horizontally
between long lines so that the abalones can feed on the
kelps by manual feeding. Once the kelps have been har-
vested, the abalones are fed with dried kelps.
An example of multispecies large-scale sea ranching is
taking place near Zhangzidao Island, 40 miles offshore in
the northern Yellow Sea (water depth from 10 to 40 m). Sea
ranching is usually practiced for the enhancement of natural
stocks, but the scale and intensity at which it is practiced in
some Chinese waters means, in fact, that one is really
talking about aquaculture on natural substrates. The
Zhangzidao Fishery Group Co. Ltd. is authorized to farm up
to approximately 40 000 ha, and presently culti-vates 26
500 ha of scallop (Patinopecten yessoensis), 10 000 ha of
arkshell (Scapharca broughtonii), 660 ha of sea cucumber
(Apostichopus japonicus), and 100 ha of abalone (Haliotis
dis-cus hannai). The company has been in existence for
more than 10 years. The total harvest in 2005 reached 28
000 tonnes, valued at more than US$60 million (US$18 mil-
lion in net profit). To improve ecological conditions and the
sustainability of the operation, the company is now thinking
of developing seaweed cultivation and the con-struction of
artificial reefs in more offshore environments. To date, about
13 300 ha have been optimized in this way.
The Development of Innovative Practices,
such as IMTA, Will Require Defining
the Appropriate Regulatory and Policy
Frameworks, and Financial Incentive
Tools
For IMTA to develop to a commercial scale, appropriate
regulatory and policy frameworks need to be put in place.
Present aquaculture regulations and policies are often
inherited from previous fishery frameworks and reason-ings,
which have shown their limitations. To develop the
aquaculture of tomorrow, the present aquaculture regula-
tions and policies need to be revisited. Adaptive regulations
need to be developed by regulators with flex-ible and
innovative minds, who are not afraid of putting in place
mechanisms that allow the testing of innovative practices at
the R&D level, and, if deemed promising, mechanisms that
will take these practices all the way to C
(commercialization). As the IMTA concept continues to
evolve, it is important that all sectors of the industry be
aware of the implications of the changes involved so that
they can adapt in a timely and organized manner. To move
research from the ‘pilot’ scale to the ‘scale–up’ stage, some
current regulations and policies may need to be changed or
they will be seen as impediments by industrial partners who
will see no incentive in develop-ing IMTA. For example, an
earlier version of the
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2474 Ecological Engineering | Multitrophic Integration for Sustainable Marine Aquaculture
Canadian Shellfish Sanitation Program (CSSP) prevented
the development of IMTA because of a clause that spe-
cified that shellfish cannot be grown closer than 125 m of
finfish netpens. This paragraph was never written with IMTA
in mind, but impinges seriously on its develop-ment. It is
currently being reviewed and amended so that IMTA
practices can develop to commercial scale legally, based on
recent, reliable, and relevant data and informa-tion provided
by the IMTA project in the Bay of Fundy and similar projects
in other parts of the world.
It is also important to note that present aquaculture
business models do not consider and recognize the
economic value of the biomitigation services provided by
biofilters, as there is no cost associated with aqua-
culture discharges/effluents in open seawater-based
systems. Regulatory and financial incentives may there-
fore be required to clearly recognize the benefits of the
extractive components of IMTA systems (shellfish and
seaweed). A better estimate of the overall cost/benefits
to nature and society of aquaculture waste and its
mitigation would create powerful financial and regula-
tory incentives to governments and the industry to jointly
invest in the IMTA approach. For example, Denmark,
after the initial development of finfish aqua-culture in the
1970–1980s, is now reconsidering more finfish
aquaculture development, but the condition for that to
occur is proper planning for biomitigation and the
recommended use of biofilters, such as seaweeds and
shellfish, is being considered. This means that extractive
species would now become one’s license to operate, or
not, in Denmark, and that the nutrient reduction services
provided by these organisms would finally be recognized
and valued for their ecosystem functions. These services
need to be quantified; for example, in Denmark, the cost
of remediating 1 kg of nitrogen is estimated at E33. If
laws or regulations were implemented to have
aquaculture operations responsibly internalize their
environmental costs, a sig-nificant reduction of their
profitability would occur. A study in Chile showed that by
integrating the culture of the algal nutrient biofilter
Gracilaria, environmental costs of waste discharges are
significantly reduced and profitability is significantly
increased. The introduction of a nutrient tax, or its
exemption through the imple-mentation of biomitigative
practices (nutrient credits), would make the economic
demonstration of the validity of the IMTA approach even
more obvious. Moreover, by implementing better
management practices, the aquaculture industry should
increase its social accept-ability, a variable to which it is
very difficult to give a monetary value, but an imperative
condition for the development of its full potential.
Reducing environ-mental and economic risk in the long
term should also make financing easier to obtain.
Conclusions
Several projects in different parts of the world, like those
presented above, have now accumulated enough data to
support the biological demonstration of the IMTA con-
cept. The next step is the scaling-up of the experimental
systems to make the biological demonstration at a com-
mercial scale, and to document the economic and social
advantages of the concept, which will be key to convin-
cing practitioners of monospecific aquaculture to move
toward IMTA practices. Underlying this demonstration
will be the development of a better understanding of the
major ecological interactions involved with IMTA sys-
tems. Working on appropriate food safety regulatory and
policy frameworks in the respective countries will be
essential for enabling the development of commercial
scale IMTA operations in a more universal fashion.
IMTA farms should be engineered as complete sys-tems,
rather than as clusters of different crops, to maximize the
benefits of the complementing ecological functions of the
different species toward the profitability of the entire
operations. Economic analyses need to be inserted in the
overall modeling of IMTA systems as they get closer to
commercial scale and their economic impacts on coastal
communities are better understood. It will, then, be possible
to add profitability and economic impacts to the comparison
of the environmental impacts between IMTA and
monoculture settings. These models will need to be
sensitized for the most volatile parameters and explicit
assumptions so as to develop models for IMTA systems
with built-in flexibility to be tailored to the environmental,
economic, and social particulars of the regions where they
will be installed. They could be modi-fied to estimate the
impact of organic and other eco-labelings, the value of
biomitigation services, the savings due to multitrophic
conversion of feed and energy which would otherwise be
lost, and the reduction of risks by crop diversification and
increased social acceptability.
There is still a large amount of education required to
bring society into the mindset of incorporating IMTA into
their suite of social values. Some of the social surveys
conducted in Canada indicate that the general public is in
favor of practices based on the ‘recycling concept’. Whether
this will translate into a greater appreciation of the
sustainable ecological value of the concept, a willing-ness to
support it tangibly with their shopping money, and demands
to their elected representatives will be the ulti-mate test. The
degree to which researchers and extension people become
creatively involved with this educational component will be
vital to the success of IMTA practices.
The ecological, engineering, economic, and social chal-
lenges remaining to be solved are for some maybe
daunting. However, the goal is to develop modern IMTA
systems, which are bound to play a major role worldwide in
 Author's personal copy
sustainable expansions of the aquaculture operations of
tomorrow, within their balanced ecosystem, to respond to a
worldwide increasing seafood demand with a new paradigm
in the design of the most efficient food production systems.
See also: Carrying Capacity; Energy Balance;
Mariculture Waste Management; Monocultures Versus
Polycultures; Technology for Sustainability.
Further Reading
Buschmann AH, Troell M, and Kautsky N (2001) Integrated algal
farming: A review. Cahiers de Biologie Marine 42: 83–90.
Chopin T and Bastarache S (2004) Mariculture in Canada: Finfish,
shellfish and seaweed. World Aquaculture 35: 37–41.
Chopin T and Robinson SMC (2004) Proceedings of the Integrated Multi-
Trophic Aquaculture Workshop. Saint John, NB, 25–26 March 2004.
Bulletin of the Aquaculture Association of Canada 104(3): 1–84.
Chopin T, Buschmann AH, Halling C, et al. (2001) Integrating
seaweeds into marine aquaculture systems: A key towards
sustainability. Journal of Phycology 37: 975–986.
Costa-Pierce BA (2002) Ecological Aquaculture: The Evolution of the
Blue Revolution, 382pp. Oxford: Blackwell Science.
Fang J, Oivind S, Liang X, and Zhang J (2001) Carrying capacity
and optimizing measures for mariculture in Sungo Bay. Marine
Fisheries Research 22(4): 57–63.
FAO (2002) The state of world fisheries and aquaculture 2002. Food
and Agriculture Organization of the United Nations, Rome, Italy;
http://www.fao.org/docrep/005/y7300e/y7300e00.htm.
Mutagenesis
C W Theodorakis, Southern Illinois University Edwardsville, Edwardsville, IL, USA
ª 2008 Elsevier B.V. All rights reserved.
Introduction
DNA Damage
Mutations
Introduction
Mutagenesis is the formation of mutations in DNA mole-
cules. There are a variety of mutations that can occur in
DNA, such as changes in the DNA sequence or rearran-
gement of the chromosomes. Such mutations may occur
spontaneously, as a result of ‘mistakes’ that occur during
DNA replication or mitosis. Spontaneous mutations are
essential to produce genetic variation necessary for
natu-ral selection. Mutations may also occur as a result
of environmental exposure to genotoxins (chemicals that
alter the structure of DNA). Mutagenesis is of concern
because it may lead to irreversible effects that can affect
fitness of organisms, which in turn may affect population-
level processes.
View publication stats
Ecotoxicology | Mutagenesis 2475
FAO (2004) The state of world fisheries and aquaculture 2004.
Food and Agriculture Organization of the United Nations, Rome,
Italy; ftp://ftp.fao.org/docrep/fao/007/y5600e/y5600e01.pdf
FAO (2006) State of world aquaculture 2006. Food and Agriculture
Organization of the United Nations, FAO Fisheries Technical
Paper 500, Rome, Italy.
Folke C, Kautsky N, Berg H, et al. (1998) The ecological footprint
concept for sustainable seafood production: A review.
Ecological Application 8(supplement): S63–S71.
McVey JP, Stickney RR, Yarish C, and Chopin T (2002) Aquatic
polyculture and balanced ecosystem management: New
paradigms for seafood production. In: Stickney RR and McVey
JP (eds.) Responsible Marine Aquaculture, pp. 91–104. Oxon:
CABI Publishing.
Naylor RL, Goldburg RJ, Primavera JH, et al. (2000) Effects of
aquaculture on world fish supplies. Nature 405: 1017–1024.
Neori A, Chopin T, Troell M, et al. (2004) Integrated aquaculture:
Rationale, evolution and state of the art emphasizing seaweed
biofiltration in modern mariculture. Aquaculture 231: 361–391.
Ryther JH, Goldman CE, Gifford JE, et al. (1975) Physical models of
integrated waste recycling-marine polyculture systems.
Aquaculture 5: 163–177.
Tacon AGJ and Forster IP (2003) Aquafeeds and the environment:
Policy implications. Aquaculture 226: 181–189.
Troell M, Ro¨ nnba¨ ck P, Halling C, et al. (1999) Ecological engineering
in aquaculture: Use of seaweeds for removing nutrients from
intensive mariculture. Journal of Applied Phycology 11: 89–97.
Troell M, Halling C, Neori A, et al. (2003) Integrated mariculture: Asking
the right questions. Aquaculture 226: 69–90.
Troell M, Neori A, Chopin T, and Buschmann AH (2005)
Biological wastewater treatment in aquaculture – more than
just bacteria. World Aquaculture 36: 27–29.
Effects on Fitness and Ecological Parameters
Further Reading
There are potentially thousands of mutagenic and
genotoxic agents to which organisms are exposed.
Examples of the classes of mutagenic compounds,
the DNA damage they elicit, and their sources in the
envi-ronment are listed in Table 1. Each genotoxin
may elicit many different types of DNA damage.
DNA Damage
Types of DNA Damage
Because most environmentally induced mutations origi-
nate as DNA damage, any discussion on mutagenesis
must begin with a discourse on this subject. For the sake
of clarity, the structure of DNA bases is given in Figure 1.