Received: 23 December 2022 Revised: 1 February 2023 Accepted: 1 February 2023

DOI: 10.1111/jwas.12948

REVIEW ARTICLE

Harvesting the benefits of nutritional research to

address global challenges in the 21st century

Brett Glencross 1,2 | Débora Machado Fracalossi 3 |

4 5
Katheline Hua | Marisol Izquierdo | Kangsen Ma 6 |
Margareth Øverland 7 | David Robb 8 | Rodrigo Roubach 9 |
10 11 12
Johan Schrama | Brian Small | Albert Tacon |
13,14 15
Luisa M. P. Valente | Maria-Teresa Viana | Shouqi Xie 16 |
Amararatne Yakupityage 17

1
Institute of Aquaculture, University of
Stirling, Stirling, UK Abstract
2
IFFO – The Marine Ingredients Organisation, Over the past 20 years, substantial progress has been
London, UK made in improving feeds and feeding technologies for
3
Departamento de Aquicultura, Universidade
most aquaculture species. Notable improvements in feed

polis, Brazil
Federal de Santa Catarina, Floriano
4
conversion efficiency (through a better understanding of
Tropical Futures Institute, James Cook
University, Singapore City, Singapore requirements and improved feed management) and ingre-
5
GIA, ECOAQUA Institute, Universidad de Las dient sustainability (through increased capability to use a
Palmas de Gran Canaria, Telde, Spain wider range of ingredients) have been achieved. While
6
Department of Aquaculture, Fisheries
advances have been made in understanding the require-
College, Ocean University of China, Qingdao,
China
ments of many of the main aquaculture species, there is
7
Faculty of Biosciences, Norwegian University still much to be done in defining requirements, especially
of Life Sciences, Aas, Norway for many of the species being farmed in the developing
8
Cargill Animal Nutrition and Health, world. Gains in the efficiency of feeds are slowing for
London, UK
9
developed species, but potential gains are still apprecia-
Food and Agriculture Organisation, Rome,
Italy
ble for less developed species. There is a growing need to
10
Aquaculture and Fisheries Group, more precisely prescribe the required levels of essential
Wageningen University and Research, nutrients and various additives in the diet based on age,
Wageningen, The Netherlands
genotype, environment, and immune status to deliver a
11
Aquaculture Research Institute, University
“precision nutrition” approach to farming aquaculture
of Idaho, Hagerman, Idaho, USA
12
Aquahana LLC, Kailua, Hawaii, USA
species. There is still further need to diversify our ingre-
dient options to provide greater resilience, as the

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and
reproduction in any medium, provided the original work is properly cited.
© 2023 The Authors. Journal of the World Aquaculture Society published by Wiley Periodicals LLC on behalf of World Aquaculture
Society.

J World Aquac Soc. 2023;1–21. wileyonlinelibrary.com/journal/jwas 1

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2 GLENCROSS ET AL.

13
CIIMAR/CIIMAR-LA, Centro Interdisciplinar
de Investigação Marinha e Ambiental, sustainability of different feed ingredient sources, includ-
Universidade do Porto, Matosinhos, Portugal ing possible climate change impacts, is becoming a grow-
14
ICBAS, Instituto de Ciências Biomédicas de ing issue. There is a growing demand for biocircularity in
Abel Salazar, Universidade do Porto, Porto,
our feed ingredient supply chains. Ultimately, what is
Portugal
15
Institute of Oceanology, Autonomous
needed to sustain future feed ingredient needs are sus-
University of Baja California, Ensenada, tainable sources of cost-effective protein, some essential
Mexico
amino acid additives, some omega-3 fatty acid resources,
16
Institute of Hydrobiology, Chinese Academy
and various minerals and vitamin additives. The increas-
of Sciences, Wuhan, China
17
ing use of new and varied resources will ensure that food
Asian Institute of Technology, Bangkok,
Thailand safety remains an important issue throughout the world.
Feed manufacturing has evolved from a simplistic exer-
Correspondence
Brett Glencross, Institute of Aquaculture, cise to a highly complex science with state-of-the-art
University of Stirling, Stirling, UK.
engineering, but its application is not consistent across all
Email: [email protected]
sectors, as there is still widespread use of pelleting, mash,
and trash fish feeding in the developing world. Similarly,
feed management has also dichotomized between the
developed and developing world, with a high reliance on
manual skilled labor in the developing world, whereas
more advanced aquaculture systems are becoming
increasingly reliant on automated computer-controlled
feeding systems.

KEYWORDS
aquafeed, feed conversion efficiency, nutrition, sustainability

1 | I N T RO DU CT I O N

With any science-based prediction of the future and of which path to follow (essentially a navigation exer-
cise), we need to understand where we are and where we have been. It is only once we understand this
context of our situation that we can then begin to make some rational prediction about where we are head-
ing. As such, in this article, we examine the three key themes that underpin the feeds and feeding process
(requirements/ingredients/management) and have framed our review of this journey in terms of the past–
present–future. Additionally, it needs to be noted that the top seven cultured species groups (carps, tilapia,
catfish, shrimp, other freshwater fish spp., salmonids, and other marine fish spp.) collectively comprise close
to 90% of all aquaculture production and as such form the focus of the article. While increasingly, each of
these species' groups is being produced intensively and are reliant on compounded feed for their sole nutri-
tional inputs, this is not equally the case among all seven groups. Substantial production, particularly of
carps, which comprise almost 50% of global aquaculture production, are still produced in small-scale exten-
sive, semi-intensive, and integrated polyculture systems with little to no manufactured feed inputs. Notably,
feed specifications, ingredient use, and feeding practices also vary regionally across the world. As such, in
this article, we have attempted to generalize on those issues that affect the three key themes that underpin
the feeds and feeding process across the world.
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GLENCROSS ET AL. 3

2 | REQUIREMENTS

2.1 | Past (prior to 2010)

The terms nutrient “requirement” and “specification” have frequently been used interchangeably. While “specifica-
tion” is the detailing of the design instructions for something (e.g., a diet containing 40% protein), the actual nutrient
requirement of an animal is more based around the need for a particular nutrient required at a certain metabolic
body weight and for a defined duration (e.g., mg protein/kg0.7/d), and for defined physiological functions such as
maintenance, maximal growth, or optimal health. The process of establishing these nutritional requirements (opti-
mum dietary levels) for aquaculture species has been a big challenge not only because of difficulties associated with
feeding an aquatic animal but also because of considering the increasing number of species of commercial impor-
tance (Boyd et al., 2020; Hua et al., 2019). This contrasts with what has happened with terrestrial domestic species,
such as poultry and pigs, where the definition of nutritional requirements is generally focused on a single species,
and has varying recommendations based on production stage and/or objective (e.g., broilers or layers in poultry)
(NRC, 1981). Additionally, finfish and shellfish requirements can vary not only among species but also on their devel-
opmental stage and in some cases different environmental constraints (NRC, 2011). Moreover, requirements may
further depend on the farming system used and regionality. In many cases, production of key aquaculture species
was and still is undertaken in extensive, semi-intensive, and integrated polyculture systems, meaning that a holistic
systems approach to nutrition had to be used rather than a monoculture approach. Aquaculture was notable in being
much more varied in the nutritional requirements of its species than those in livestock production.
Traditionally the definition of requirements for essential nutrients (energy, protein, lipid, amino acids, fatty acids,
vitamins, and minerals) for most species was undertaken on a gross nutrient basis only (Boonyaratpalin, 1997;
Cowey, 1992; NRC, 2011). Requirements were typically defined based on empirical experiments and limited to
young animals and laboratory environments. This served well in terms of defining the distinct differences among
many of the species and between aquatic and terrestrial animals (NRC, 2011).
Initially, the definition of nutrient requirements for many species led to big gains in feed efficiency, with
improvements in FCR values from 1.8–3 to 1.2–1.8 being achieved for many species (Hua et al., 2019; Boyd
et al., 2020; Tacon, 2020; Naylor et al., 2021; Hardy et al., 2021). However, as time has passed, there have been
diminishing returns on any improvements in the definition of nutrient requirements across most species' groups
(Figure 1).
The use of various modeling systems in aquaculture to define requirements for energy and some macronu-
trients and explore various physiological processes was established prior to 2010 (Dumas et al., 2010;
Shearer, 1995). These models were largely derived from similar approaches used for terrestrial species but had
to include various empirically derived features to account for the differences between ectotherms and endo-
therms (Cho & Bureau, 1998; Lupatsch et al., 2001). Different iterations of either nutrient flow or nutrient
demand models were developed, which were largely based around factorial bioenergetic processes (Cho &
Bureau, 1998; Glencross, 2008; Han et al., 2011).

2.2 | Current status (2010–2020)

Since 2010, smaller improvements in feed efficiency have been made, with typical changes from FCRs of 1.5–2 to
1–1.5 for some species (Figure 1). Such gains were mainly because of the increasing knowledge about nutritional
requirements but also because of better management practices following the intensification of growing systems
(NRC, 2011). An increasing level of replacement of fish meal with plant proteins as the main ingredients in commer-
cial feeds demanded a better knowledge of amino acid requirements. Although fish meal has an amino acid balance
close to that of many fish requirements most plant proteins do not, thus meeting fish demands for growth with plant
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4 GLENCROSS ET AL.

FIGURE 1 Changes in economic feed conversion ratios (eFCR) of major aquaculture species groups over the past
45 years.

proteins required a deeper knowledge of amino acid requirements in terms of amounts and balance (Kaushik &
Seiliez, 2010).
This was especially important for carnivorous species, which have higher demand for amino acids, but it was also
important for omnivores such as tilapia and shrimp, where the high cost of fish meal limited its use in commercial
feeds. A deeper knowledge about the requirement of other nutrients such as taurine, n-3 fatty acids, and various
vitamins and minerals also became important with increasing levels of fish meal replacement, as these nutrients are
less available or even absent in plant sources (Domínguez, Montero, et al., 2020; Domínguez, Sehnine, et al., 2020;
Gatlin III et al., 2007).
During the 2010–2020 decade, after the key NRC publication (NRC, 2011), there was an increasing level
of definition of requirements for animals of different sizes and in some cases different environmental constraints
(e.g., hypoxia, temperature, salinity, etc.). Both age and environmental conditions may markedly affect fish
physiology and modulate the expression of key genes, affecting nutrient utilization and nutritional requirements
(Fernández-Palacios et al., 2011; Hamre et al., 2013; Izquierdo & Koven, 2011; Lund et al., 2019; Torno et al., 2019).
However, such information is still scattered or absent for some species (e.g., Seriola, Sole) (Sicuro & Luzzana, 2016;
Valente et al., 2019). Industrial breeding programs are rapidly progressing for a few species, based on recently devel-
oped customized genomic tools for marker-assisted selection, and are producing fast-growing animals with improved
FCR that may have different requirements for specific nutrients (Glencross et al., 2013; De Verdal et al., 2018; Lee
et al., 2020; Ferosekhan et al., 2021). Moreover, nutritional history (protonutrition) and nutritional programming
affect the utilization of dietary nutrients and therefore could affect fish requirements (Turkmen et al., 2019). The
main drivers for such development were the high cost of aquafeed, which typically represents 50% of the variable
production cost of intensive aquaculture, together with the increasing pressure for producing more sustainable diets
able to address the Sustainable Development Goals established by UN. This implies knowing more about the require-
ments of the various species to sustain optimal growth and produce high quality and healthy animals. In many of
those species already produced at large scale, requirements for macronutrients and most micronutrients are now
well established for early/larval, juvenile, grower, and brood stock stages of the production cycle. But most non-
mainstream species still need clearer definition of many of the basic nutrient requirements and their response to
environmental constraints remains largely unknown. Modern aquaculture uses diets that rely on a variety of alterna-
tive raw materials that need to be balanced with additives including certain amino acids, minerals, and vitamins to
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GLENCROSS ET AL. 5

ensure the optimal use of the feed. However, to face various growing environmental and production challenges, the
currently established requirements of many amino acids and other micronutrients may need to be revisited to assure
fish growth and robustness into the future (Aas et al., 2019; Berntssen et al., 2018; Hamre et al., 2016; Kousoulaki
et al., 2021; Prabhu et al., 2014).
The last comprehensive compilation of nutritional requirements of fish and shrimp was published in 2011
(NRC, 2011), although a recent revision of the “Fish Nutrition” book by Hardy et al. (2021) has been timely. This
publication included the requirements of various freshwater and marine fish as well as shrimp of commercial impor-
tance. Despite organizing and facilitating access to nutritional requirement information for various important aqua-
culture species, it also made clear several unknowns not only on micronutrient (vitamins and minerals) but also on
amino acid requirements. Notably, the compiled requirement specifications were largely defined from animals in
juvenile stages and based on using highly digestible diets and animals maintained in optimal growing conditions. In
most cases, requirements were presented as a single recommendation for each species and did not consider changes
in requirements throughout the various production stages or production systems. Therefore, despite the increasing
use of farming system environments to undertake research and provide industrially relevant information for some
species such as salmon, this is not so for many other species (Ng & Romano, 2013).
Most feeds in the world are still formulated on a crude composition basis, although there is increasing formula-
tion of feeds on a digestible nutrient basis in the developed world for some species such as Atlantic salmon. The
quality of the formulation depends on various knowledge sets, including the accurate understanding of the nutrient
requirements, the nutritional composition of the ingredient being used and their nutrient digestibility and/or bioavail-
ability, and the processes being used to form the feed into a physical pellet. While the feed of salmon is formulated
to be highly efficient, the feed efficiency of most other cultured fishes or shrimp lags that of salmon.
This is mainly because of the rapid development of aquaculture with high diversity and complexity of cultured
species and feed ingredients used in developing countries, and the lack of comprehensive knowledge of essential
nutrient requirements of so many cultured animals and of the digestibility or the bioavailability of nontraditional
ingredients or their nutrients. For example, nutritional studies published in China have been conducted on more than
200 species, representing a wide geographic distribution, different feeding habits (herbivorous, carnivorous, omnivo-
rous, and filter feeders), and different culture modes. It is evident from the information provided in the NRC (, 2011)
that the tables listing the nutritional requirements of salmonids are mostly complete, while those for many other
important marine or freshwater species are either incomplete or mostly blank (Hardy et al., 2021). Despite the almost
700 species produced in aquaculture, increasing pressure in the last decade for further species diversification has
brought attention to several either low trophic species or very fast-growing or endangered species (Boyd
et al., 2020; Cottrell et al., 2021; Naylor et al., 2021; Newton et al., 2021).
Over the past decade, there has been a growing use of factorial and nutrient-flow (or demand) modeling systems
to define requirements for energy and macronutrients for an increasing number of species. Such models are now
being widely used to iteratively define nutritional requirements for various species across their production size
ranges and within varying environmental conditions (Chowdhury et al., 2013; Glencross & Bermudes, 2012). There
have been increasing levels of adoption of modeling by the commercial aquaculture sector as a means of refining
feed design and improving feed management. Furthermore, the development of mechanistic models has further
advanced our understanding of the interrelationships between different metabolic pathways and nutrient use and
how this influences growth and body composition (Bar et al., 2007; Bar & Radde, 2009).
A growing understanding of the role of the microbiome in nutrient supply/utilization has emerged. Brought
about by low-cost methods to evaluate microbial diversity using 16 S ribosomal RNA sequencing technologies, this
has enabled an increased understanding of the physiological interaction between the microbiome and its host. How-
ever, its potential to mediate how organisms respond to multiple environmental factors remains poorly understood.
For example, most aquafeeds contain carbohydrate-based binders (e.g., starches), which do not exist in the aquatic
environment. These complex carbohydrates are influential in the development of particular microbiomes. Differences
in microbiome structures have been observed between fish fed either extruded or nonextruded pellets containing
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6 GLENCROSS ET AL.

starch (Barreto-Curiel et al., 2018). So, it is apparent that the gelatinization process is influential to the structure of
the different microbial communities. Understanding the impacts of such dietary changes on the resulting microbial
communities will be helpful to identify those strategies that are beneficial or not, so long as the feed is included as a
control element (Fuentes-Quesada et al., 2018; Karlsen et al., 2022). Another important aspect is the differences in
microbiome structures in organisms subjected to stress, whether because of effects of system intensity, environmen-
tal contamination, temperature, and/or nutritional deficiencies. Studies have shown that stress, particularly chronic
stress, and microbial manipulation (probiotics or prebiotics) influence the microbiome structure of the fish gut
(Anker-Ladefoged et al., 2021; Rimoldi et al., 2020; Serradell et al., 2020). However, there is still insufficient informa-
tion on the influence of nutritional, environmental, or genetic factors on the microbiome on fish growth and health
to assess the economic benefits of adding probiotics in the feed or water.
The growing need for immune-enhancing diets comes back to issues of poor definition of micronutrient supply
under prevailing conditions linked to large-scale marine ingredient replacement without tacking all the required
issues comprehensively. Consequently, functional feeds are being developed to enhance animal health. Such func-
tional feeds must have optimized nutrient composition and are supplemented with some functional additives, all-
owing the farmer to not only meet the needs for better growth but also improve the immunity, stress resistance, and
health status of aquaculture animals, while ensuring the quality and safety of the products.
The most common functional additives used in aquafeeds include preservatives (antioxidant), nutritional supple-
ments (amino acids, vitamins, and trace elements), enzymes, pigments (carotenoids), palatants, and immunomodula-
tors (probiotics, prebiotics, plant extracts, and nucleotides). The mechanistic action of such functional additives is
multidimensional. They can act directly and/or indirectly on the animal, such as antioxidation, modification of gut
microbe profiles, improvement of digestive tract morphology, elevation of digestive enzyme activities and nutrient
absorption, and enhancement of immunity and disease resistance (Aguirre-Guzmán et al., 2012; Hayatgheib
et al., 2020; Hoseinifar et al., 2017; Tacchi et al., 2011; Torrecillas et al., 2014). In the future, further studies are
needed on the functional mechanisms of individual additives and their interactions, especially their synergistic
effects. Usage of many such additives is however under strict regulations that vary widely among countries, although
there are levels of regulation exerted on their use through global efforts such as Codex Alimentarius (http://www.
fao.org/fao-who-codexalimentarius/home/it/).

2.3 | Future (2020 and beyond)

Analysis of the changes in the FCR values for most mainstream species demonstrates that improvements have been
plateauing for some time and only smaller gains are now being made, leading to a point of diminishing returns for
many species (Figure 1) (Naylor et al., 2009; Tacon, 2020; Ytrestøyl et al., 2015). Any future work on defining
requirements needs to focus more precisely on refining estimates with further consideration of nutrient interactions
and the use of new and more stable delivery forms, which help reduce nutrient wastage and increase efficiency. In
the coming decades, aquaculture sustainability will remain a major concern. Water quality remains vital for aquacul-
ture production and inadequate nutrition and feeding practices are known to deteriorate water quality. Thus, design-
ing more efficient feeds through better knowledge of nutrient requirements, their appropriate balance, and effective
delivery forms will improve nutrient stability and absorption and minimize nutrient waste. Likewise, refining feeding
management practices to avoid excess nutrient load in the water will limit (or reduce) the increased oxygen demand
following eutrophication, which can compromise fish health, welfare, and growth.
There is a growing need to prescribe the required levels of essential nutrients in the diet more precisely based
on age, genotype, nutritional history, environment, farming system, and immune status, and the fish quality to deliver
a “precision nutrition” approach to farming targets. This will necessitate the use of various modeling (using big data),
internet-of-things (IoT), and artificial intelligence (AI) approaches to support aquaculture production and feed specifi-
cation management. Integration between traditional models with more mechanistic approaches will lead to better
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GLENCROSS ET AL. 7

prescription of nutrient and energy requirements based on an increasing number of input parameters. Advances in
such modeling systems are now already accommodating demands for amino acids and net energy demands for vari-
ous species (Hua & Bureau, 2019; Glencross, 2021).
The improved definition of requirements for key nutrients (energy, amino acids, and fatty acids) on a digestible
and net basis is still required for most species and especially those in the developing world. The future of aquaculture
nutrition will increasingly be based on precision and smart farming that will require a clear definition of nutrient
requirements on a digestible and a net basis. This is particularly important in the context of encouraging the utiliza-
tion of locally sourced ingredients. The shift toward increasing use of alternative ingredients in recent years has
made it even more important to formulate diets on a digestible nutrient basis, not only because of the presence of
antinutritional factors that can interfere with nutrient utilization but also because of the variability in the bioavailabil-
ity of the nutrients required to formulate balanced diets (Boyd et al., 2020).
Some of these new ingredients have complex matrices that limit digestion and/or absorption of nutrients
(e.g., algae) (Valente et al., 2021), highlighting the relevance of formulating on a digestible basis as is already per-
formed in many other species.
Tailoring fish diets to produce value-added products able to respond to consumer's expectation will also gain
importance in the future. This will include nutrient fortification of fish via their diet for children, pregnant women,
and elderly people, by enrichments in omega-3 (especially EPA and DHA), selenium, vitamin D, and iodine. The
capacity of farmed fish to not only increasingly supply protein for the world, but to also do so in a way that enhances
the nutrition of the young and at risk will become increasingly important (Tacon & Metian, 2013; Thilsted, 2012).
This food fortification/enhancement combined with a growing focus on food safety will enhance the role of nutrition
as a critical control point in aquaculture.
Functional diets for challenging production periods and conditions will gain increasing prominence. Climate
change and growing levels of intensification and the use of recirculating aquaculture systems (RAS) will
increase the susceptibility of fish to alterations in environmental conditions that will increase stress, which will
need to be mitigated. Functional diets that contain ingredients or additives that affect animal robustness and
health will become increasingly important. Because production system environmental changes are expected
with the growing intensification of aquaculture, it will be necessary to further investigate what positive effects
functional diets will have, by identifying the most adequate ingredients and/or additives that result in improved
animal performance and health. The formulation of diets for challenging times in the fish production process
will be increasingly required. The use of various feed additives to help or mitigate the effects of stress through
the year or seasonally will become increasingly common. Although the potential for formulating marine
ingredient-free diets already exists for a vast majority of species, it is usually not cost-effective in most cases
for carnivorous species. Such diets are also often associated with chronic health issues and poor animal robust-
ness, so formulating based on nutrients (amino acids and fatty acids) will further require the search for additives
that can be used to improve the general health status and robustness of different aquaculture species in a
future with constraints on marine ingredient use.
Over the past 40 years, the rapid development and increasing intensification of aquaculture across a broad range
of species and geographies have led to problems arising in some cases. Issues such as environmental deterioration
and stress, sometimes leading to disease outbreaks and then subsequent abuses of antimicrobials and other chemical
drugs, have been reported in various sectors. These issues threaten the food safety and sustainability of aquaculture.
Misuse of antimicrobials not only leads to drug residues, affecting food safety, but also potentially leads to drug-
resistant pathogens, which can then lead to public health issues. Many countries have legislated to prohibit the use
of antimicrobials as growth promotors in animal feed (since 2006 in European Union), and many are gradually
restricting its use for medical purposes to tackle the emergence of bacteria and other microbes resistant to antimi-
crobials. In addition, because of the constraints on further growth in the production of fish meal, fish oil, and other
marine-derived ingredients, many nontraditional ingredients have been used to replace fish meal and fish oil in
aquafeeds.
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8 GLENCROSS ET AL.

In many cases, this has resulted in growth retardation and impaired health of fish and shrimp as the changes
were often not supported by science or backed up with required alternatives to bring a positive outcome. Hence, in
the future, we must look for new technologies or nutritional approaches (e.g., functional feeds) to mitigate the nega-
tive aspects of substitutions of marine-derived ingredients and the reduction in the use and overall stewardship of
antibiotics to ensure the good growth, health, product safety of aquaculture animals, and to enable the sustainable
development of aquaculture. In parallel, there is also an opportunity to tailor flesh quality to increasing consumer's
demand for healthier products. The fortification of aquafeeds with omega-3 (especially EPA and DHA), vitamin D,
and minerals (e.g., selenium and iodine) can help provide adequate nutrition for vulnerable population groups
(Kwasek et al., 2020; Thilsted et al., 2014).
Having defined the nutritional requirements of the various aquaculture species, there is additionally a need to
find the appropriate raw materials (ingredients) from which to provide these nutrients and energy. This supply of raw
materials also needs to develop sustainable supply chains to enable the aquaculture sector to continue to grow
among the various sustainability challenges that the production of aquaculture feeds faces (MacLeod et al., 2020;
Naylor et al., 2021).

3 | INGREDIENTS

There have been marked shifts in ingredient use for aquaculture feeds over the past several decades. In this section,
we review the practices of the past and the present to predict future trends and needs. While not all aquaculture
uses industrially produced compounded feeds, this is the fastest growing feed sector in the world and indicators sug-
gest that more and more aquaculture production is becoming increasingly intensified and reliant on the provision of
external feed inputs.

3.1 | Past (before 2010)

From the late 20th century to 2010, global aquafeed production grew from <5Mtonnes in 1990 to 30Mtonnes in
2010 (Naylor et al., 2021; Tacon, 2020). Fish meal and fish oil were considered critical ingredients in feeds for many
carnivorous fish and crustacean species, and their global production averaged 6Mtonnes and 1MTonnes per annum
respectively over this period. Fish meal use in aquaculture feeds peaked during 2005–2010 at 3.6 Mtonnes per
annum (Figure 2). Fish oil use peaked a bit before this at around 0.9Mtonnes per annum. Recognition that such
marine ingredients were limited in supply led to a global effort to move toward more use of alternative ingredients in
aquafeeds. Early fish meal alternatives were primarily of animal origin because of availability, cost, and protein con-
tent. Since then, the perceived risks associated with inclusion of rendered animal by-products in aquafeeds have
seen their restriction in several geographic regions (primarily within European countries) because of concerns of
pathogens and zoonotic disease transmission (Glencross, Baily, et al., 2020). Plant proteins were initially only a minor
contributor to aquaculture feed protein and lipids for most carnivorous species, but by 2010, inclusion levels reached
about 28% of the total feed inputs for salmon feeds globally, up from 12% in 2000. At the same time fish meal
reduced from up to 45% in 2000 to 23% of the diet in 2010 (Ytrestøyl et al., 2015). This switch was initiated by sev-
eral El Nino events, which reduced the availability of fish meal and forced the prices up. The feed production sector
responded primarily by increasing the use of soy and some (wheat and corn) gluten meals. While much of the use of
fish meal and fish oil prior to 2010 was driven by the economy of nutrients, plant proteins were seen to have signifi-
cant limitations, often containing lower protein content, antinutritional factors, and imbalanced nutrient profiles
(Gatlin III et al., 2007).
Although these constraints still existed for plant ingredients, reduced fish meal supplies and increased costs
drove the demand for alternative protein sources. Terrestrial plant ingredients were generally considered as being
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GLENCROSS ET AL. 9

F I G U R E 2 Estimated feed production for all fed aquaculture species sector from 1975 to 2020, with concurrent
marine ingredient use across the aquaculture sector. Shown in the inset is a magnified set of the marine ingredient
(fish meals + fish oils) use data across the same time. Data derived from FishStat 2020 and IFFO, 2021.

more sustainable and gained a more significant position in aquafeeds with new processing technologies leading to
the creation of a variety of plant protein concentrates that could compete with the higher price and protein content
of fish meal. Although still not widely used in feeds for carnivorous species, plant proteins and oils were already by
this time the major contributor to aquaculture feed protein and lipids for omnivorous and herbivorous species. Soy-
bean meal was already a widely used ingredient in feeds for omnivorous and herbivorous species, with other grains
being used included a range of cereal, oilseed, pulse, and various grain legume seed products. Typical inclusion levels
of soybean meal in feeds for such omnivorous and herbivorous species ranged from 15% to 45% (Tacon et al., 2011).

3.2 | Current status (2010–2020)

Presently there is 55Mtonnes of aquafeed production globally, of which fish meal is now only 4MT, fish oil is
1MT, and the rest (55MT) is mostly plant-derived resources (IFFO, 2021). Use of marine ingredients (proteins
and oils) in aquafeeds has stabilized since 2010 and they are now seen more as strategic ingredients, with a growing
use of by-product fish meal now contributing to >30% of all marine ingredient use in manufactured feeds (Hamilton
et al., 2020; IFFO, 2021).
Plant proteins continue to be the main contributor to aquaculture feed protein and lipids for most aquaculture
species, but their use is also seen as contentious in some circumstances (e.g., soybean use and associated deforesta-
tion concerns). Notably, feeds for carnivorous species are now predominantly composed of plant proteins and oils
(Ytrestøyl et al., 2015). The increasing use of processing to produce protein concentrates from soy, peas, and other
legumes as well as cereal glutens has occurred. This processing maximizes the protein concentration, quality, and
suitability for use in feeds, while minimizing the non-nutritive factors (Drew et al., 2007). To balance the amino acids,
the use of plant proteins and the concurrent reduction in fish meal has led to a much more complicated formulation,
now involving a greater number of protein sources with addition of some crystalline essential amino acids to create a
balanced diet to support high growth performance and healthy fish. This broader raw material array brings some
advantages to the feed sector, in that it is not reliant on any one ingredient—so there is more resilience to supply
chain threats (such as poor harvests). But plant proteins also bring some sustainability challenges to address
(Malcorps et al., 2019).
 17497345, 0, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/jwas.12948 by UNIVERSIDADE FEDERAL DE GOIAS, Wiley Online Library on [29/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
10 GLENCROSS ET AL.

The focus on soy and deforestation is probably foremost, which has led the feed industry to work with their soy
suppliers in Brazil to set deforestation and conversion-free cut-off dates in 2020 for their entire supply chain. Plant
proteins also typically have a much higher carbon footprint than fish meal, which has pushed the overall footprint of
the feed up over the last 20 years (MacLeod et al., 2020). Carbon footprint is increasingly of interest, as the carbon
impact of food systems has been publicly highlighted. Here aquaculture is in a very good position, being a typically
low greenhouse gas (GHG) emitter compared with other animal protein production systems, but it also has a great
opportunity to improve (MacLeod et al., 2020). It has been suggested that replacing 20%–30% of fish meal in shrimp
feeds with plant protein ingredients could result in a 63% increased demand for freshwater, 81% increase in land
requirements, and a need for 83% more phosphorus (Malcorps et al., 2019).
Across a broader range of life cycle assessment impact factors, it has been argued that there is insufficient land
for the expansion of animal feed crop production (Popp et al., 2017), as it currently occupies roughly one third of
global croplands (Robinson et al., 2011). Furthermore, there are additional pressures on land use because of popula-
tion growth, climate change, and demands for food and biofuels (Godfray et al., 2010; Spiertz & Ewert, 2009). In their
discussion, Malcorps et al. (2019) remind us that the Sustainable Development Goals of the United Nations include
food security, hunger reduction, and protection of life on land and in the sea. They emphasize that minor price
changes to crops resulting from increased pressure on land-based food production systems could have dire conse-
quences in developing countries, where 50% of the household income is spent on food (Spiertz & Ewert, 2009).
The current focus is not only on the reduced reliance on fish meal and fish oil derived from capture fisheries but
also increasingly on the overall sustainable supply of terrestrial and plant feed ingredients. Some of the raw materials
used in aquaculture feeds can be consumed by humans, resulting in food-feed competition. There is also a consider-
ation of whether raw materials that could be consumed directly by humans should be used for animal feeds.
The shift towards plant-based ingredients in aquaculture feed also faces competition from livestock and agricul-
ture sectors, as well as biofuel production. Marine and terrestrial ingredient sources are both vulnerable to climate
change, which may cause disruption and decrease in supply and higher costs for aquaculture feeds (FAO, 2018).
There is thus a trend toward increased use of by-products as feedstuffs in aquaculture and livestock feeds with a
growing focus on biocircularity of resources. However, consumer concerns have been raised in some markets around
the use of by-products, in particular animal by-products that could bring a good nutritional input but are blocked
from use in some supply chains because of perceived health risks and social objections (Glencross, Baily, et al.,
2020). In parallel, circular proteins such as insects and microbial biomass are being increasingly advocated as sustain-
able alternatives, as they can convert various waste streams such as food waste, household waste, plant by-products,
and sludge into high-quality nutrients. High production and processing costs still limit their large-scale production,
and a range of regulations still limit their application in certain markets.
Both microalgal and genetically modified crops have been commercialized that produce long-chain omega-3 in
industrial volumes (>1000 tonnes). Because of an urgent need for fish oil alternatives, a range of omega-3-rich ingre-
dients are emerging, for example, microbial ingredients (microalgae), oil seeds with high level of LC-PUFAs (rapeseed
and camelina), and increased use of fish by-products, and lower trophic marine species (e.g., krill, mesopelagic fish)
(Hamilton et al., 2020). Up to eight relatively new sources of EPA and DHA with industrial potential for aquafeeds
have been recently described (Tocher et al., 2019). These products included five microalgal sources, two genetically
modified seed crop oils, and one yeast biomass. Identification of cost-effective, alternative lipids high in the LC-
PUFAs EPA and DHA remains a substantial hurdle for the future of aquaculture. Both microalgal and genetically
modified crops will undoubtedly play an increasingly important role in aquafeeds for those species requiring LC-
PUFAs. Perhaps equally important is the need to fortify fish feeds toward the improvement of human health.
Food safety is a considerably overlooked aspect of much of fish nutrition research, but it is increasingly
highlighted socially and politically (Glencross, Baily, et al., 2020). As an example, perceptions over the use of food
waste for livestock and fish feeding have limited efforts in this area, particularly in developed countries, and yet there
is an estimated 1.3 billion tons of human food lost and wasted each year (Gustavsson et al., 2011). Concerns over
genetically modified feed ingredients also limit options in certain parts of the world, but others are readily adopting
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GLENCROSS ET AL. 11

this technology. Food safety risks are associated with the possible chemical contaminants and biologically hazardous
materials present in ingredients and feeds, which might be passed to humans who consume aquaculture products.
Concerns have been raised about antimicrobial residues, persistent organic pollutants, heavy metals, mycotoxins,
and industrial contaminants (Tacon & Metian, 2008; Glencross, Baily, et al., 2020). Additional concerns over micro-
plastics in fish meal have been raised recently (Gündo
gdu et al., 2021; Hanachi et al., 2019; Thiele et al., 2021).

3.3 | Future (after 2020)

It has been suggested that by the year 2050 aquaculture production is going to double and intensify (FAO, 2020).
Fundamentally what will be needed to feed that production is sustainably sourced, economic, good quality protein,
some essential amino acid (EAA) additives, some omega-3 options, various mineral and vitamin additives, and cost-
effective energy sources. To achieve this will mean we need another 50MT of resources that we currently do not
have or that are presently being used in other sectors. Although gains in crop productivity have kept pace with
demand through most of the 20th century, they are not keeping up with demands more recently (Grassini
et al., 2013; Schauberger et al., 2018). Notably, most currently used plant protein resources are also used in pig and
poultry feeds (so a competition issue exists).
There is additionally the concern that many of these plant protein resources can be used directly to feed humans
rather than animals. Add into the equation that there is a declining availability of freshwater, phosphates, and arable
land, and it is obvious that terrestrial crop products are not the only solution going forward. Further competition for
ingredients of plant origin is also occurring for human food and nonfood products such as biofuels. While many
plant-source ingredients are considered economically sustainable, questions have been raised about their social and
environmental sustainability, notably products from soy and palm oil production (Hospes, 2014; Okereke &
Stacewicz, 2018). These constraints provide a strong argument for the development of nontraditional protein and oil
sources.
Fish meal and fish oil are still considered among the most nutritious ingredients as they are rich sources of
essential nutrients and have a high level of palatability in virtually all aquaculture species. Increasingly fish meal and
oil production will come from by-product resources from fish caught for direct human consumption (e.g., Alaskan
pollock) or aquaculture by-product. By 2030, more than one third of all fish meal and oil will come from by-product
sources (IFFO, 2021). The continued reduction in the reliance on fish meal and fish oil, combined with the concurrent
increased use of alternative raw materials in aquaculture feeds, highlights the need for an approach based on com-
plementarity of ingredients (Turchini et al., 2019). The increasing use of alternatives brings about the need for a
range of feed additives to supplement specific essential nutrients including essential amino acids, essential fatty
acids, and trace elements. We may also see greater usage of bioactive compounds, prebiotics, probiotics, and other
immunostimulants (Boyd et al., 2020). It is critical to find additional, cost-effective ingredient sources to meet the
growing nutrient demand. This burgeoning demand, growing ingredient competition, and heightened sustainability
awareness provide a strong argument for the development of nontraditional protein and oil sources, especially those
part of the circular bioeconomy like insects, microalgae, microbial biomass, and food waste.
Technologies that produce protein, amino acids, and omega-3 using noncompetitive processes based on non-
food grade resources (e.g., bacteria, yeasts, and algae; single-cell ingredients [SCI]) perhaps offer the most potential
to generate the additional resources needed (Glencross, Huyben, & Schrama, 2020). As biotechnology advances, a
broader range of substrates from various waste streams will be used in the fermentation process to reduce costs and
increase profitability of SCI production. This will include the downstream processing to enhance the nutritional value
of the microbial ingredients. As the competition for natural resources increases and technology advances, production
of microbial ingredients will shift from being dependent on photosynthesis and products from this process as sub-
strates toward use of cheaper input factors (e.g., organic acids, CH4, H2, and CO2 gas) from industrial waste or other
renewable sources.
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12 GLENCROSS ET AL.

Additionally, there will be increasing pressure to source ingredients locally for feed production and reduce
dependence upon imported sources. This will demand more effort be given to use of local ingredients, the adoption
of nutrient recycling, and use of innovative raw material processing techniques. This will probably be coupled with
the use of renewable energies as the type of fuel used for processing has a remarkable life cycle impact on such
ingredients (Campos et al., 2020). In the future, there is likely to be more competition for natural resources; driven
by factors such as population growth, development of the bioeconomy, and climate change. Aquaculture production
will play an increasing role in meeting the global protein supply and the need for feed to sustain this production will
clearly increase.
Sustainability of different feed ingredient sources, including possible climate change impacts, is becoming a
growing issue. For the global aquaculture sector to grow sustainably, it must have a sustainable supply of nutrients
to make the feeds. The key sustainability issues vary from ingredient to ingredient and the historical approach was to
focus just on one feed ingredient at a time, such as fishmeal or soy. Sustainability of different feed ingredient combi-
nations will become a key defining characteristic of their utility, with increasing use of independent certification sys-
tems to verify claims. Going forwards, we will see an increasing focus on the sustainability of the feed ingredient
supply chains. The discussion should move beyond whether an individual ingredient is sustainable or not, to one of
whether that supply chain is. For example, fishmeal can be supplied sustainably, from sustainably managed fisheries.
Soy can be farmed sustainably, on land that was not recently converted from native vegetation. Independent certifi-
cation schemes have been established to provide verification of claims. There are a broad range of schemes with dif-
ferent levels of value and credibility, but all add cost to the overall supply chain. However, the expectation is that the
use of certification to verify sustainability claims will continue to grow. As the topics covered by sustainability
increase, the number of schemes is likely to continue to grow, for example, through moving beyond fisheries and
deforestation issues to addressing issues of human rights, carbon footprint, land, and water use. The level of com-
plexity to be managed will increase, but the information to do this has to keep pace, so good decisions are made on
the latest reliable data. This can only be supported by a full value chain commitment to change, led from the con-
sumer and retail end to support the upstream supply side's transformation. Without commitment from the market,
changes will be harder to implement and value. Assessment of the sustainability of feed ingredients varies depending
on the type of ingredients as their social, economic, and environmental impacts all differ considerably. There is a
need for harmonization of various aquaculture-related sustainability certification standards to ensure consistency
(Kok et al., 2020). The adoption of certification schemes will continue to reduce or eliminate ingredients from
unsustainable sources.
Food safety will also become an increasingly important issue in the future. Driven by consumer perception, poli-
tics, and some level of science, consumers will gravitate toward those products that align with their attitudes, prefer-
ences, and expectations. Food safety will continue to be important for feed manufacturers and fish producers to
meet consumer demand for disclosure of credence attributes. In addition to safety, these include origin, sustainabil-
ity, and nutritional content. As such, both mandatory and voluntary labeling will continue to be strong drivers of the
seafood market.

4 | FEED MANAGEMENT AND MANUFACTURING

4.1 | Past (before 2010)

Feed rationing systems and management of how much and when feed was delivered to the animal were largely
based on prior experience or demand-based, relying on the animal providing feedback to the person feeding them.
Most feeding was manual, using hand, blower, or other simple delivery systems. As such, feeding was a labor-
intensive process (Ibrahim & Sultana, 2006). Traditional feeding regimes were based on the experience of the opera-
tors, considering factors such as weather, watercolor, season, and animal behavior among other things. The diet was
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GLENCROSS ET AL. 13

selected based on price and/or feed manufacturer recommendations. The focus was on fish growth. Traditional end
points for manual feeding were judged by farmers making decisions based on their experience and skills. This primar-
ily entailed deciding what feed, when, and how much was delivered to the animal (Jobling et al., 1995; Paspatis &
Boujard, 1996). Assessments of animal behaviors and interactions with feeds and the feeding process were some-
what subjective, although for certain species some simple tools, like feed trays, were used to assist the process
(Tacon, 2002). For technologically advanced sectors, sensor-based feeding systems were emerging (Kadri
et al., 1998).
Historically, many feeds were based on the use of trash fish and/or made on farm by the farmers. These may
have initially been fed as intact trash fish or made into a moist mash with the use of other materials and binders or
made into dried pellets using simple pelleting manufacturing systems (Hasan & New, 2013). With the growth and
increasing technical demands on the sector, specialist feed compounders emerged and brought advancements in the
various processes used in feed manufacturing. Additionally, the feed manufacturing sector played an important role
in helping farmers, not only in the provision of feed inputs but more importantly in the management of the feed on
the farm and in the use of appropriate environmentally sound husbandry practices. In many countries, the feed man-
ufacturers were the closest contact in the value chain between the farmer and the government legislature (Tacon
et al., 1995). Notable among the manufacturing introductions during this period was that of expansion extrusion,
which was introduced into some sectors from 1980s onwards and by the 2000s became widely used across the
developed world for most fish species, and increasingly was being applied in the developing world. Extrusion brought
with it a range of technical advantages, including the ability to control buoyancy of feeds, dramatically increase feed
fat levels, improve water stability, and reduce wastage because of dust (Barrows et al., 2007; Obaldo et al., 2000).

4.2 | Present (2010–2020)

In the present day, feed management systems in technologically advanced sectors are becoming increasingly com-
puterized in modern developed-world aquaculture. Automated centralized pneumatic feeding systems using in-cage
cameras, acoustics, and computer-based decision-making tools are widespread, resulting in a reduced level of labor
(Aas et al., 2011; Reis et al., 2020; Waagbø et al., 2013). However, in the developing world where the majority of
aquaculture still occurs, feeding is still largely a manually controlled and managed process with important labor
demands and continues to be a significant rural employer (El-Sayed, 2013; Hung & Quy, 2013; Ng & Romano, 2013).
Feeding end points are still largely judged by farmer decisions in combination with computerized algorithms. The
interpretation of the fish responses is experience-based and depends on the experience and skills of the individual
farmer.
The use of growth and energetics models is increasingly being applied for the evaluation and management of
feeding regimes for various species (Cho & Bureau, 1998; Glencross & Bermudes, 2012; Liu et al., 2018; Zhou
et al., 2005). The application of modeling helps reduce both the feed cost and the waste being discharged (Bueno
et al., 2017). Historically, modeling focused only on evaluating the feeding rate, but increasingly, diet formulation,
environmental factors, feeding frequency, feeding rhythm, and even animal behavior are all being taken into
consideration.
Manufacturing of feeds in the developed world is now mostly based on modern extrusion technologies (Barrows
et al., 2007). There has been considerable development of engineering technological capabilities in this area, allowing
considerable control over the forming, cooking, and densification attributes of the feeds. Such engineering systems
are highly complex but offer a substantial level of control over the pellet-forming process, enabling the user to have
significant control over pellet density, durability, and oil infusion capacities (Sørensen, 2012). Shrimp feeds, despite
still being predominantly pelleted, are beginning to emerge as extruded products through some companies (Obaldo
et al., 2000; Soares et al., 2021). In the developing world, there is still widespread use of pelleting, mash, and trash
fish feeding. Feed manufacturing in these regions, particularly in small-scale aquaculture operations, has continued
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14 GLENCROSS ET AL.

to rely on an on-farm mash feed manufacturing approach, although extrusion is increasingly being used by many
commercial feed suppliers in these parts of the world (Edwards et al., 2004; Merican, 2021; Tacon & Metian, 2009;
Xu et al., 2007).

4.3 | Future (after 2020)

To increase sustainability of fish production and optimize fish product quality and animal welfare, it is becoming even
more important to monitor and control the animal production process (Antonucci & Costa, 2020). By reducing the
feed conversion ratio, farmers will have less waste and loss of nutrients to the water they farm in, and they will tend
to reduce their overall carbon footprint. To achieve this, we need a better understanding of feed intake and its physi-
ological points of regulation and how this is affected by production system (e.g., RAS, pond, or cage) and a range of
other abiotic factors. In the future, we will see increasing use of mechanized and automated feeding in the develop-
ing world, matching that already being undertaken in much of the developed world. This use of mechanized and
automated feeding will aid in improving production efficiencies (through less wastage) and animal growth rates
(through higher intakes), and commercial quality (achieving the market requirement) by allowing for the better align-
ment of feeding with the demand and needs of the animals being farmed, but this will correspondingly reduce labor
demands and employment in the sector. Already there are signs of increasing use of autonomous feeding systems in
the developing world, occurring through the development of a variety of low-cost automatic feeding systems
(e.g., https://www.efishery.com/).
The use of AI in feeding systems is emerging. AI is going to impact businesses of all shapes and sizes across all
industries (Marr, 2020). The increasing use of centralized computer-controlled feeding systems and in-cage/in-pond
sensors and cameras will increasingly make an internet-of-things system of control more feasible for feeding aqua-
culture species (Måløy et al., 2019; Martos-Sitcha et al., 2019; Mustapha et al., 2021). The development of complex
algorithms and AI that monitors feeding behavior will be used to help make decisions about feeding management
(Jones et al., 2012; Zhou et al., 2018). Coupled with more precisely defined energy and nutrient demands over vari-
ous size classes, environments, genotypes, and in situ bio-loggers, the inclusion of a precision nutrition approach will
complement completely automated feeding (Hvas et al., 2020; Zhang et al., 2020).
The use of automation and AI will lead to precision and smart farming increasingly becoming a focus. The use of
big data applications may play a major role in improving the efficiency of the entire supply chain, with focus on food
security, safety, and sustainability (Gilpin, 2015). These will provide an increased focus on the application of princi-
ples of precision fish farming (PFF) to shift feeding management from comprising largely experience-driven processes
to become a more knowledge-driven procedure (Føre et al., 2018). PFF feeding regimes based on animal demands
and farm targets, including specialized diet formulations, considerations of environmental factors, animal health,
and/or quality and/or water quality will be included in the AI system to provide precise decisions for management of
computer-controlled systems. Biological and system informatics will become combined to increase the precision con-
trol of fish farming. The use of intelligent sensors and monitoring will support the application of predictive models
and simulation systems, leading to improved decision management and integration of operations that will enhance
the overall precision control of fish farming operation (Føre et al., 2018). The use of models to predict environmental
fluctuations and adjust feed distribution accordingly and autonomously to minimize feed waste and improve effi-
ciency should be the primary goal for improved production efficiency in aquaculture. The optimal solution should be
the use of real-time techniques that can determine the actual feeding behavior based on the appetite status of
farmed fish (Adegboye et al., 2020).
There will continue to be increasing use of extrusion feed production, with increasing uptake in the developing
world and development of novel extrusion applications for nontraditional species based on improved feed technical
qualities and flexibility of the manufacturing systems used. This increasing shift to manufactured feeds across the
developing world will result in an overall improvement in feed quality and resource utilization by the sector, further
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GLENCROSS ET AL. 15

aiding a move away from the use of trash fish and farm-made feeds. However, further support to promote this in
developing regions of the world is required. Additionally, because of the increased use of automatic feeders, there
will be increased demand for the use of high physical quality nonclogging extruded feeds in these feeding systems.
The increasing demand for functional feeds with specialist additives will see the emergence of nanotechnology to
precision deliver certain high-valued compounds to the fish, by preventing them to be digested or metabolized at
certain stages of the absorption process, allowing for targeted nutrient and nutraceutical delivery. Such novel encap-
sulation methods will enable the protection of certain labile functional ingredients from the damaging forces involved
in extrusion.

5 | C O N CL U S I O N

Many of the advances in nutritional understanding of aquaculture species over the past 20 years have come from an
intense focus on a few species. However, it should be clearly acknowledged that part of the challenge of aquaculture
nutrition is the diversity of species involved and that their requirements can be just as diverse. There is an urgent
need to bring our understanding of many of the nonfocus species into the same level of understanding. Even with
the well-researched species, there is a growing need to adopt a “precision nutrition” approach to the supply of
essential nutrients and various additives in the diet based on age, genotype, environment, and immune status. Addi-
tionally, although marine-derived ingredients are still widely used in feeds, even salmon farming has now become a
net producer of seafood. The widespread use of plant proteins and oils has underpinned the growth of aquaculture
over the past 20 years, although there is a need to further diversify this resource base. The capacity for aquaculture
to achieve this has been the result of decades of nutrition research (Naylor et al., 2021). The future will require fur-
ther work in developing the potential of a range of additional sustainable feed protein and oil options, particularly
those from the circular bioeconomy. Progress in feed manufacturing remains important, even though it has evolved
from a simplistic exercise to a highly complex science with state-of-the-art engineering. Further application of
advanced feed manufacturing is needed, as there is still widespread use of pelleting, mash, and trash fish feeding in
the developing world. As ever, feed management remains critical to ensure the best use of the feeds produced.
While aspects of this may have dichotomized between the developed and developing world, with a high reliance on
manual skilled labor in the developing world, the use of advanced automated computer-controlled feeding systems
in the developed world is a growing reality.

ACKNOWLEDGMENTS
We acknowledge the editorial contributions of Ron Hardy, Sadasivam Kaushik, Hasan Mohamed, Michael B. New,
Giovanni Turchini, and Johan Veretth.

ORCID
Brett Glencross https://orcid.org/0000-0003-1167-8530

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How to cite this article: Glencross, B., Fracalossi, D. M., Hua, K., Izquierdo, M., Ma, K., Øverland, M., Robb,
D., Roubach, R., Schrama, J., Small, B., Tacon, A., Valente, L. M. P., Viana, M.-T., Xie, S., & Yakupityage, A.
(2023). Harvesting the benefits of nutritional research to address global challenges in the 21st century.
Journal of the World Aquaculture Society, 1–21. https://doi.org/10.1111/jwas.12948