Reviews in Aquaculture (2017) 9, 342–368 doi: 10.1111/raq.

12141

The utilization and mode of action of organic acids in the

feeds of cultured aquatic animals
Wing-Keong Ng and Chik-Boon Koh
Fish Nutrition Laboratory, School of Biological Sciences, Universiti Sains Malaysia, Penang, Malaysia

Correspondence
Wing-Keong Ng, Fish Nutrition Laboratory,
School of Biological Sciences, Universiti Sains
Malaysia, Penang 11800, Malaysia.
Email: wkng@usm.my
Received 15 December 2015; accepted 22
December 2015.
Abstract
The use of antibiotics in the feeds of cultured aquatic animals to mitigate infec-
tious diseases or to boost growth performance is commonly practised. In recent
years, the prophylactic use of antibiotics has been criticized which eventually led
to their ban in animal production by many countries. In view of the current
restrictions on antibiotic use, there is a critical need to evaluate alternatives. One
potential substitute for antibiotic growth promoters in aquafeeds is organic acids
and/or their respective salts. The use of dietary organic acids in the culture of
aquatic animals has been the focus of much research and commercial interest
recently. This review provides a comprehensive compilation of the current knowl-
edge on the use of organic acids in aquafeeds, with emphasis on its impacts on
growth, nutrient utilization, mineral availability, gut microbiota and disease resis-
tance. Many studies have reported that some organic acids can significantly
enhance the growth performance and health status of fish. Contradictory results
have also been reported, which seems to depend on the aquatic animal species,
type and concentrations of organic acids and the culture conditions used. Recent
research indicating the beneficial impact of dietary organic acids on shrimp
growth and health is highlighted. Possible mechanisms of action of these func-
tional feed additives are discussed. Strategies for enhancing the effectiveness of
dietary organic acids are given. This review aims to contribute to the sustainability
of global aquaculture production by providing a viable alternative to harmful
antibiotics, thereby meeting the growing consumer demand for antibiotic-free
seafood products.
Key words: antibiotics, antimicrobials, aquaculture, disease, feed additive, organic acids.

infectious diseases caused by bacterial pathogens, following

Introduction
The aquaculture industry is one of the fastest growing
food-producing sectors in the world and is expected to
continue growing rapidly in the foreseeable future. The
rapid growth in modern aquaculture is fuelled by a variety
of factors, which include the increasing use of formulated
aquafeeds and intensification of culture systems. As a result
of increasing intensification of farming practices, one of the
most limiting problems in the aquaculture sector is disease
outbreaks (Bondad-Reantaso et al. 2005). The sustainabil-
ity and scalability of the global aquaculture industry are sig-
nificantly constrained by the spread of infectious diseases
caused by various pathogens and cause major economical
losses to farmers. Large amounts of antibiotics are often
used in the aquaculture industry to prevent and/or control
the discovery of the growth-promoting and disease-fighting
capabilities of antibiotics (Hern
andez-Serrano 2005). The
extensive use of a wide variety of antibiotics in the aquacul-
ture industry, both as therapeutic and growth-promoting
agents, has increased the potential harmful effects on
human and animal health as well as the aquatic environ-
ment (Cabello 2006). The emergence of antibiotic resis-
tance in various bacterial pathogens associated with fish
disease had been documented (De Paola et al. 1988; Sch-
midt et al. 2000; Miranda & Zemelman 2002; Agersø et al.
2007). The extensive use of antibiotics in aquaculture has
the potential to threaten public health due to antibiotic
residue bio-accumulation in consumer-ready aquaculture
products (Nawaz et al. 2001). In a recent review, Cabello
et al. (2013) indicated that several antibiotic-resistant

342 © 2016 Wiley Publishing Asia Pty Ltd


genetic elements had been reported to be shared between
aquatic bacteria, fish pathogens and human pathogens, and
might have originated in aquatic bacteria. They went on to
caution against the excessive use of antibiotics in aquacul-
ture farms for short-term economic gains. Moreover,
research in farmed shrimp has shown that the long-term
use of antibiotics can eventually reduce subsequent antibi-
otic efficacy due to the emergence of resistant bacterial
strains (Reboucßas et al. 2011), while excessive levels can
even negatively affect shrimp growth and overall condition
(Bray et al. 2006).
Public awareness regarding the prophylactic use of
antibiotics in animal feeds, which may lead to the transfer
of immunity of bacterial species pathogenic to humans, has
led to their ban in animal feed formulations. A worldwide
effort to minimize and eventually eliminate the use of
antibiotics for growth-promoting purposes in the aquacul-
ture and livestock industry started with the ban on the use
of subtherapeutic antibiotics in the European Union on
January 2006 (European Parliament and Council Regula-
tion (EC) No 1831/2003). The development of effective
nonantibiotic compounds as an alternative to the prophy-
lactic use of antibiotics to control infectious diseases and
enhance growth performance is therefore paramount for
the continued expansion of the global aquaculture indus-
try.
Short-chain organic acids (C1–C7) and their salts or
mixtures, commonly known as acidifiers, are promising
alternatives for antibiotic growth promotants (AGP) and
have been receiving growing attention from aquaculture
researchers (L€uckst€adt 2008a; Ng & Koh 2011). Organic
acids, such as benzoic, formic, lactic and propionic acids,
have traditionally been used as storage preservatives in food
and feed ingredients for preventing product deterioration
caused by fungi and microbes (Ricke 2003; van Dam 2006).
Some organic acids have been shown to have strong
antibacterial effects against important food borne patho-
Figure 1 Number of published papers in
mainstream peer-reviewed scientific journals
regarding the use of organic acids in aquacul-
ture between 1981 and 2014.
Reviews in Aquaculture (2017) 9, 342–368
© 2016 Wiley Publishing Asia Pty Ltd
Dietary organic acids in aquaculture
gens such as Escherichia coli and Salmonella spp. (Cherring-
ton et al. 1991b; Van Immerseel et al. 2003; Skrivanova
et al. 2006) and are currently employed in terrestrial live-
stock feeds to control bacterial pathogens (Van Immerseel
et al. 2002, 2003). Terrestrial livestock such as poultry and
swine fed organic acid-supplemented diets have consis-
tently been reported to show improved feed intake, growth,
feed utilization efficiency and health (Alp et al. 1999; Parta-
nen & Mroz 1999; Kluge et al. 2006). Although the use of
dietary organic acids and their salts has been extensively
studied in various terrestrial animals, the research on aqua-
tic animals has only intensified in the last 10 years (Fig. 1).
Research and commercial interest in the use of organic
acids in aquafeeds is expected to significantly increase in
the coming years due to changing global regulatory con-
trols on the use of antibiotics. We also observe that many
companies traditionally dealing with organic acid additives
targeting the terrestrial livestock feeds are currently market-
ing their products for the aquafeed sector, often times with-
out scientific proof that it works equally well for aquatic
animals. This is based on our personal observations at vari-
ous international trade exhibitions on animal feeds. A com-
prehensive review on what is currently known about the
use and efficacy of organic acids in aquaculture is much
needed to address some of these concerns.
Reviews on the use of organic acids in aquaculture have
previously been published as proceedings or book chapters
(L€uckst€adt 2008a; Ng & Koh 2011). These reviews were rel-
atively brief, narrowly focused and deal only with finfish.
The present review aims to update the current knowledge
on the use of organic acids and includes recent research
with shrimp feeds. A comprehensive review on the effects
of organic acids as a functional feed additive on feed palata-
bility, gut pH, digestive enzyme activity, gut microbiota
and morphology, growth, nutrient utilization, hepatopan-
creas pathohistology and disease resistance will be pre-
sented. Even though the antimicrobial effects of organic
343
W.-K. Ng and C.-B. Koh
acids are well understood, the explanation for the growth-
promoting effects of these compounds remains to be eluci-
dated (de Wet 2005). Possible modes of action of dietary
organic acids will be discussed. Practical strategies for
increasing the effectiveness of dietary organic acids will be
given. This single comprehensive review will likely benefit
researchers, feed manufacturers, feed additives companies,
farmers, seafood processors and policymakers related to the
aquaculture industry. It includes peer-reviewed research
published in mainstream journals (Fig. 1) as well as
selected research from conference proceedings, magazine
articles and other grey literature. This review seeks to pro-
vide scientific evidence for a viable alternative to the use of
antibiotics, thereby contributing to the sustainability of
aquaculture and safety of the resultant seafood products.
Organic acids
The global market value of acidifiers in the animal feeds
industry in 2015 was estimated to be about €1.0 billion
(FEFANA 2014). The most commonly used organic acids
in animal feeds are propionic acid followed by fumaric, for-
mic and lactic acids and/or their salts. With its emphasis on
food safety and animal health, Europe represents the largest
market (41%) followed by other developed countries such
as the United States (24%) and Japan (12%). For example,
about 67% of worldwide demand for formic acid/salt in
animal nutrition is from the EU countries (FEFANA 2014).
With the ban on the use of AGP in animal feeds in Europe
and with increasingly stringent regulations on the use of
antibiotics as prophylactic agents in the rest of the world,
the use of alternatives such as organic acids is expected to
increase.
Organic acids are organic compounds with one or more
carboxyl groups. These include saturated straight-chain
monocarboxylic acids (C1–C18) and their respective
derivatives, such as unsaturated (cinnamic, sorbic), hydrox-
ylic (citric, lactic), phenolic (benzoic, cinnamic, salicylic)
and multicarboxylic (azelaic, citric, succinic) acids (Cher-
rington et al. 1991a), with a general molecular structure of
R-COOH, where R represents the monovalent functional
group. These acids are commonly referred to as short-chain
fatty acids, volatile fatty acids or weak carboxylic acids.
Organic acids are produced through the microbial fer-
mentation of carbohydrates by various bacterial species
under different metabolic pathways and conditions. Some
lower molecular weight organic acids, for example acetic,
propionic and butyric acids, are also formed within the
large intestine of humans and animals at high concentra-
tions by anaerobic microbial communities (Cummings
et al. 1987; Macfarlane & Macfarlane 2003). Many of these,
particularly the short-chain organic acids (C1-C7), are nat-
urally present as normal constituents of plants or animal
344
tissues. However, most organic acids commercially used in
the food and feed industry are produced synthetically.
Organic acids may also form into single or double salts of
their acid through combining with potassium (K), sodium
(Na), calcium (Ca), etc.
Weak lipophilic organic acids and their salts are consid-
ered as ‘generally regarded as safe’ (GRAS) substances and
have been used for centuries as preservatives in foods and
beverages (Russell & Gould 2003). They are listed in the EU
regulations as permissible feed additives in food animal
production. Organic acids, their salts or combinations
thereof, have been successfully used in livestock feeds as
alternatives to antibiotics (Alp et al. 1999; Partanen &
Mroz 1999). Commercial mixtures of organic acids are
widely used to control pathogenic bacteria, such as Sal-
monella species, in feed ingredients and livestock feeds
(Van Immerseel et al. 2002). Table 1 presents a list of
organic acids that are currently being used singly or in
combination with commercial feeds for terrestrial livestock.
The strength/acidity of an acid in solution is represented
by the logarithmic constant (pKa) value, which is equal to -
log10 (Ka = [H+][A
]/[HA] = acid dissociation constant).
The smaller the value of pKa, the stronger is the acid. In
general, since most organic acids are considered weak acids
and only partially dissociate into their ions in aqueous solu-
tions, the majority of organic acids will remain in their
undissociated form (i.e. free organic acid). In an aqueous
solution, weak organic acids dissociate and form a pH-
dependent dynamic equilibrium between undissociated
acid molecules and dissociated anions as in the chemical
equilibrium below:
HA þ H2 O $ A
þ H3 Oþ
where HA represents a weak acid, A
is a conjugate base
(acid anion), and H3O+ is a hydroxonium ion. Organic
acids change from undissociated forms to dissociated
forms depending on the environmental pH and its pKa
value.
The pH at which 50% of the acid is dissociated is called
its pKa value and is unique for each individual acid
(Table 1). At a pH below their individual pKa values,
organic acids will mostly be present in an undissociated
form, thus increasing the proportion of free acids that can
readily enter the bacterial cells by simple diffusion. Lower
molecular weight organic acids, for example formic, acetic
and lactic acids, are miscible in water, whereas higher
molecular weight organic acids such as benzoic acid are
insoluble in water due to their hydrophobicity. The physi-
cal and chemical characteristics of some organic acids that
are commonly used in feeds for monogastric animals are
listed in Table 1. These physicochemical attributes are
important in our understanding of the antimicrobial
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 Dietary organic acids in aquaculture

Table 1 Common organic acids and their physicochemical properties (based on Mroz 2005)

Organic acid Molecular formula pKa value Molecular mass (g mol
1) Physical form Odour CR*

Formic CH2O2 3.75 46.03 Liquid Pungent +++

Acetic C2H4O2 4.6 60.05 Liquid Pungent +++
Propionic C3H6O2 4.88 74.08 Oily liquid Pungent ++
Butyric C4H8O2 4.81 88.12 Oily liquid Rancid +
Lactic C3H6O3 3.86 90.08 Liquid Sour milk +
Sorbic C6H8O2 4.76 112.1 Solid Mildly acrid +
Fumaric C4H4O4 3.02, 4.76 116.1 Solid Odourless 0 to +
Malic C4H6O5 3.40, 5.1 134.1 Solid Apple +
Citric C6H8O7 3.13, 4.76, 6.49 192.1 Solid Odourless 0 to ++

*CR = corrosiveness rate: high (+++), medium (++), low (+), negligible (0).

efficacy of organic acids which will be discussed in a later
section of this review.
Organic acids in cultured aquatic animal feeds
Despite the known beneficial effects of dietary organic acids
in improving the performance and health of terrestrial live-
stock, limited comprehensive research has been conducted
to evaluate the efficacy of organic acids or their salts in
aquatic animals, until recently. As far as we know, only a
few studies had been published on the use of organic acids
in aquafeeds prior to the ban of AGP use in livestock pro-
duction (Ringø 1991; Gislason et al. 1994, 1996; Vielma &
Lall 1997; Vielma et al. 1999). More recently, many studies
(Fig. 1) have been conducted to determine the effects of
dietary organic acids and their salts on growth perfor-
mance, nutrient utilization and disease resistance in several
commercially important farmed fish species, such as rain-
bow trout, salmon, carp and tilapia. In the last 5 years, this
research has been extended to shellfish. A general overview
of the major organic acids and their salts tested in aqua-
feeds to date is presented in Tables 2–7. The major impacts
of these dietary organic acids in fish and shrimp are sum-
marized in their respective Tables and described in this sec-
tion. A more detailed critical analysis and discussion of
some of these effects is given in a later section of this review
to better understand the mode of action of these dietary
organic acids.
Citric acid and its salt
Citric acid or its salts are by far the most investigated
organic acid in aquaculture (Table 2). Numerous studies
have reported that citric acid can improve growth, feed
utilization and mineral availability, particularly phosphorus
(P) in various fish species (Sarker et al. 2005, 2007; Baruah
et al. 2005, 2007a,b; Pandey & Satoh 2008; Castillo et al.
2014), while some reported contradictory findings (Fau-
conneau 1988; Vielma et al. 1999). The earliest study
regarding the use of citric acid was conducted on rainbow
trout by Fauconneau (1988). In this study, citric acid at
120 g kg
1 was supplemented in a fishmeal-based diet to
partially replace the protein content in trout diets. While
acidification of the diets appeared to lower the voluntary
feed intake, this did not affect the efficiencies of protein
and energy utilization. Since then, several studies with more
promising results have been reported in rainbow trout,
red sea bream (Pagrus major, Temminck & Schlegel, 1843),
rohu (Labeo rohita, Hamilton, 1822), beluga sturgeon
(Huso huso, Linnaeus, 1758) and yellowtail (Seriola qui-
queradiata, Temminck et Schlegel, 1845) (Table 2).
Sugiura et al. (1998) investigated the effects of dietary
acidification on the mineral availability from fishmeal in
rainbow trout diets. The 12-day preliminary study found
that citric acid at a 50 g kg
1 inclusion level significantly
increased the availability of P, Ca, Mg, Fe, Mn and Sr in
fishmeal-based diets. Also, the availability of Mn in the diet
was increased by the inclusion of 50 g kg
1 Na-citrate. In
their follow-up study, the ability of citric acid (20 or
50 g kg
1) in improving mineral utilization was further
confirmed in trout. However, using the same diets, no
effect on mineral availability was reported for the agastric
goldfish (Carassius auratas, Linnaeus, 1758). The faecal P
content of trout fed a diet supplemented with 50 g kg
1
citric acid was about half that of fish fed the nonacidified
diet indicating increased P absorption for those provided
with citric acid-supplemented diets. At this inclusion level,
citric acid had no effect on feed intake in trout during the
five-week trial, but a marked reduction was observed in
goldfish. Sugiura et al. (1998) speculated that the lack of
effect on mineral availability in the goldfish trial might be
due to the insufficient acclimation period for the fish to this
acidifier. In another study, Sugiura et al. (2001) similarly
reported that the addition of 50 g kg
1 citric acid to a high
ash moist diet increased the availabilities of ash, Ca, total P,
Sr and Zn for rainbow trout.
In contrast with the findings of Sugiura et al. (1998) with
goldfish, positive effects of citric acid on mineral utilization

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© 2016 Wiley Publishing Asia Pty Ltd 345
 346
Table 2 Use of citric acids or their salts in aquafeeds

Organic acids/salts Dose (g kg
1) Species and weight (g) Impacts* Reference

Citric acid 0, 7.5 and 15 Red drum, Tended to lower the pH of diets and stomach contents Castillo et al. (2014)
W.-K. Ng and C.-B. Koh

Sciaenops ocellatus (1.26  0.01) Improved weight gain, feed efficiency and PER
Enhanced the digestive enzyme activities
0 and 10 Rainbow trout, Improved growth, SGR, FCR, phosphorus absorption/retention andez et al. (2012)
Hern

Oncorhynchus mykiss (13.2  2.4) Reduced P load to the environment
0, 10, 20 and 30 Beluga sturgeon, Huso huso (25.1–25.7) Improved weight gain, SGR, PER and FCR Khajepour and
Improved apparent protein, Ca and P digestibility Hosseini (2012a,b)
0 and 5 Yellowtail, Increased retention of P Sarker et al. (2012a,b)
Seriola quinqueradiata (129.7–135.8) Tended to increase whole-body ash content
Increased the retention/availability of minerals (Ca, Mg, Na, K, Zn and Mn)
10 Tilapia, Oreochromis niloticus 9 Increased the activities of protease and amylase in digestive tract Li et al. (2009)
Oreochromis aureus (158.9  18.8) Did not affect lipase activities
0 and 10 Rainbow trout, O. mykiss (2.67  0.01) Improved growth and P retention with low fishmeal-based diets Pandey and Satoh (2008)
0 and 30 Rohu, Labeo rohita (~13.16) Increased Zn, Na, P, K, Mn, Mg, Fe, Cu and Ca absorption Baruah et al. (2007a)
Activated microbial phytase
0 and 30 Rohu, L. rohita (12.6–13.7) Improved weight gain, SGR, FCR and P availability Baruah et al. (2007b)
No effect on PER and apparent NPU
0 and 50 Rainbow trout, O. mykiss (~152) Increased availability of ash, Ca, total P, Sr and Zn Sugiura et al. (2001)
No negative impact on feed intake
0 and 10 Red sea bream, Pagrus major (5.46  0.01) Improved P retention and tended to improve weight gain Hossain et al. (2007)
0, 10 and 20 Red sea bream, P. major (12.6  0.4) Improved weight gain, SGR, FCR and P absorption Sarker et al. (2007)
Lowered N and P loading
0 and 30 Rohu, L. rohita (12.6–13.7) Improved mineral utilization Baruah et al. (2005)
Decreased intestinal digesta pH
0 and 30 Red sea bream, P. major (7.14  0.01) Improved fish growth, FCR, nutrient retention and reduced N and P loading Sarker et al. (2005)
0, 4, 8 and 16 Rainbow trout, O. mykiss (3.0  0.1) No effect on weight gain Vielma et al. (1999)
Improved P utilization and whole-body Fe deposition linearly
Citric acid and Na-citrate 10 and 50 Rainbow trout, O. mykiss (232.0  37.1) Improved availability of Ca, P, Mg, Fe, Mn and Sr Sugiura et al. (1998)
Citric acid 0, 20 and 50 Rainbow trout, O. mykiss (256.4) Improved apparent availabilities of Ca and P Sugiura et al. (1998)
0, 20 and 50 Goldfish, Carassius auratus (149.7) Negatively influenced feed intake at 50 g kg
1 inclusion Sugiura et al. (1998)
0 and 120 Rainbow trout, O. mykiss Lowered voluntary food intake, but did not affect the Fauconneau (1988)
efficiencies of protein and energy utilization
0, 1, 2, 3, 4 and 5 White shrimp, Significant growth improvement at 2 g kg
1 Su et al. (2014)
Litopenaeus vannamei (5.57  0.21) Significantly higher survival at 1–4 g kg
1
Higher gut protease at 2 g kg
1 inclusion
Enhanced immune response and disease resistance

*FCR, feed conversion ratio; PER, protein efficiency ratio; SGR, specific growth rate; NPU, net protein utilization.

© 2016 Wiley Publishing Asia Pty Ltd

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 Table 3 Use of formic acids or their salts in aquafeeds

Organic acids/salts Dose (g kg
1) Species and weight (g) Impacts* Reference

Formic acid 0, 4 and 10 mL kg
1 Rainbow trout, Oncorhynchus mykiss (~520) Improve availability of P, Mg and Ca Vielma and
Increased the intestinal pH Lall (1997)

© 2016 Wiley Publishing Asia Pty Ltd

0, 4 and 10 Yellowtail, Seriola Slightly increased the whole-body ash content Sarker et al.
quinqueradiata (129.7–135.8) Increased the retention/availability of minerals (2012a,b)

Reviews in Aquaculture (2017) 9, 342–368

(P, Ca, Mg, Na, K, Zn and Mn)
0, 3 and 6 Pacific white shrimp, Litopenaeus No difference in growth performance Chuchird
vannamei (postlarvae 12) Higher survival after Vibrio parahaemolyticus challenge et al. (2015)
Lowered gut bacterial counts
Potassium 0, 1, 2 and 3 Nile tilapia, Oreochromis niloticus (53.5  6.2) Improved growth performance, FCR, PER and apparent protein digestibility Abu Elala and
diformate (KDF) Decreased stomach and upper intestinal tract pH Ragaa (2015)
Stimulated immune response
Enhanced resistance against Aeromonas hydrophila
0, 7.5 and 15 Red drum, Sciaenops ocellatus (1.26  0.01) Tended to lower the pH of diets and stomach contents Castillo et al. (2014)
Better growth, feed and PER
Enhanced the digestive enzyme activities
12 Atlantic salmon, Salmo salar (1500  200) No effects on digestibility of nutrients or amino acids Morken et al. (2012)
Reduced the physical quality of soya bean meal-based diets
0, 2, 3 and 5 Hybrid tilapia, (~16.7) Improved feed intake, weight gain, FCR and resistance Ramli et al. (2005)
against Vibrio anguillarum infection
0 and 13.5 Atlantic salmon, S. salar (650) Improved body weight, SGR, digestibility of protein, dry matter and gross energy Lu adt (2008b)
€ckst€
No effect on the GI tract pH
0, 3, 6, 9 and 12 Hybrid tilapia, Oreochromis niloticus 9 No effects on growth performance, FCR or survival Zhou et al. (2009)
Oreohromis aureus (2.67  0.00) Alter the intestinal bacterial community
0, 2.5, 5.0, 7.5, 10.0, Nile tilapia, O. niloticus (7.05  0.14) Tended to improve weight gain and feed efficiency Lim et al. (2010)
12.5 and 15.0 with up to 10 g kg
1 inclusion
No effects on haematological parameters, innate immune responses
and resistance against Streptococcus iniae
Sodium 0 and 10.6 Rainbow trout, O. mykiss (520) Improved digestibility of major nutrients, amino acids and physical Morken et al. (2011)
diformate (NaDF) quality of barley protein concentrate-based diets
KDF and NaDF 0, 3 and 5 Nile tilapia O. niloticus, (33.9  0.1) Numerically improved growth and feed efficiency Liebert et al. (2010)
Improved feed efficiency, PER and protein
retention efficiency with 3 g kg
1 inclusion

*FCR, feed conversion ratio; PER, protein efficiency ratio; SGR, specific growth rate.
Dietary organic acids in aquaculture

347
W.-K. Ng and C.-B. Koh

Table 4 Use of lactic acids or their salts in aquafeeds

Organic Dose Species and weight (g) Impacts Reference

acids/salts (g kg
1)

Lactic acid 0 and 10 Rainbow trout, Increased bone Zn Pandey and Satoh (2008)
Oncorhynchus
mykiss (2.67  0.01)
0 and 10 Red sea bream, Improved absorption of P but not in weight gain Hossain et al. (2007)
Pagrus major (5.46  0.01) and feed utilization
Ca-lactate 0, 15 Red drum, Sciaenops Lower the pH of diets and stomach contents Castillo et al. (2014)
and 30 ocellatus (8.01  0.29) Depressed growth performance at 30 g kg
1 diet
No effects on body indices and composition
Na-lactate 0 and 15 Arctic charr, Salvelinus alpinus; Improved growth of Arctic char, but not in salmon Gislason et al. (1996)
and Atlantic salmon, Salmo salar Doubled the feed retention time in Arctic
char digestive tract
0 and 15 Atlantic salmon, S. salar (~8) No effects on fish growth, mortality or gross Gislason et al. (1994)
chemical composition of the faeces
0 and 10 Arctic charr, S. alpinus (~100) Improve weight gain and feed efficiency Ringø et al. (1994)
No effects on lipid/fatty acid composition
of muscle tissue and body proximate composition
0 and 10 Arctic charr, S. alpinus (~310) Improved growth and nutrient digestibility Ringø (1991)

Table 5 Use of butyric acids or their salts in aquafeeds

Organic acids/salts Dose (g kg
1) Species and weight (g) Impacts* Reference

Microencapsulated 0 and 0.3 Common carp, Tended to improve weight gain and reduced FCR Liu et al. (2014)
Na-butyrate Cyprinus carpio (6.22  0.0) No effects on organ indices and intestinal
mucosal morphology
No effects on adherent bacterial population
Improved immune response
Partially protected 0 and 3 Sea bream, Sparus aurata (15) Enhanced growth, but no effects on SGR, Robles et al. (2013)
Na-butyrate feed intake and feed utilization
Altered the metabolic pattern at the intestinal level
Na-butyrate 0, 5, 10 Pacific white shrimp, Improved growth performance and feed efficiency Silva et al. (2016)
and 20 Litopenaeus No effect on P retention
vannamei (2.53  0.03) Altered intestinal microbiota
Increased serum agglutination titre
0 and 20 Pacific white shrimp, Increased the diet attractiveness and feed intake Silva et al. (2013)
L. vannamei (7.4–11.7) No effect on P digestibility
0, 2 and 20 Catfish, Clarias No effects on weight gain, SGR, PER and FCR Owen et al. (2006)
gariepinus (64  1.4) Tended to alter the bacterial community
in the hindgut of catfish

*FCR, feed conversion ratio; PER, protein efficiency ratio; SGR, specific growth rate.

and growth in another agastric fish, rohu, have been well
documented. Baruah et al. (2005) evaluated the effect of
dietary citric acid (30 g kg
1) on gut digesta pH, bone ash
and bone mineralization of rohu juveniles. At this inclusion
level, citric acid resulted in a significant decrease in the pH
of gut digesta from 6.53 to 5.70 and was suggested as a con-
tributing factor for significantly improved growth as well as
mineral utilization in this agastric carp species. Similar
reductions to the pH of diet and faeces with subsequent
improvements in the utilization of some minerals have also
been reported for rainbow trout (Sugiura et al. 1998;
Vielma et al. 1999; Pandey & Satoh 2008). Meanwhile, Bar-
uah et al. (2007b) reported significant improvements to the
growth performance, feed conversion ratio (FCR) and P
availability in rohu fed a soya bean meal-based diet supple-
mented with 30 g kg
1 citric acid. However, it did not
affect the protein efficiency ratio (PER) or apparent net
protein utilization (NPU) of fish. Baruah et al. (2007b)
attributed the growth improvement to the beneficial effect
of citric acid in releasing minerals from the phytic acid
complex of the soya bean meal-based diet.
In another study evaluating the effect of citric acid at var-
ious dietary concentrations (0, 4, 8 or 16 g kg
1) on the
availability of minerals to rainbow trout, Vielma et al.

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348 © 2016 Wiley Publishing Asia Pty Ltd
 Dietary organic acids in aquaculture

Table 6 Use of other organic acids or their salts in aquafeeds

Organic acids/salts Dose Species and weight (g) Impacts* Reference

(g kg
1)

Acetic acid 0 and 50 Rainbow trout, Enhanced P digestibility Sugiura et al. (2006)
Oncorhynchus No effects on endogenous acid secretion
mykiss (234.4  24.2) Tended to decrease stomach pH,
but no effect on intestinal pH
Malic acid 0 and 10 Red sea bream, Improved absorption of P but not in weight gain Hossain et al. (2007)
Pagrus major (5.46  0.01) and feed utilization
Na-propionate 0, 5, 10 Pacific white shrimp, Improved growth performance and yield Silva et al. (2016)
and 20 Litopenaeus No effect on P retention
vannamei (2.53  0.03) Altered intestinal microbiota
Increased serum agglutination titre
Na-propionate† 0 and 20 Pacific white shrimp, Increased diet attractiveness and feed intake Silva et al. (2013)
L. vannamei. (7.4–11.7) Lowered the Vibrio species counts in the gut
Improved apparent digestibility of energy and P
0 and 10 Arctic charr, Depressed growth of fish fed 10 g kg
1 Ringø (1991)
Salvelinus alpinus (~310) Na-propionate-added diet
Na- fumarate‡ 73 mM Pacific white shrimp, Increased in vitro protein digestibility Silva et al. (2015)
L. vannamei (0.49  0.02) Increase growth by 53% and improved FCR by
23% compared to control
Succinic acid 0 and 120 Rainbow trout, O. mykiss Lowered voluntary food intake Fauconneau (1988)
No effects on protein efficiencies and energy utilization

*FCR, feed conversion ratio; PER, protein efficiency ratio; SGR, specific growth rate.
†Best performing organic acid salt compared to Na-acetate, Na-butyrate, Na-citrate, Na-formate or Na-lactate.
‡Best performing organic acid salt in terms of growth performance compared to Na-formate, Na-acetate, Na-lactate, Na-propionate, Na-butyrate,
Na-succinate or Na-citrate.

(1999) reported a significant increase in whole-body Fe
deposition in fish fed citric acid-acidified low-P fishmeal-
based diets. Citric acid at all tested levels had no effect on
weight gain but tended to increase the body P content. In
contrast, Pandey and Satoh (2008) reported that rainbow
trout fed with a low fishmeal-based diet supplemented with
citric acid showed significant growth improvements. Con-
sistent with other trout studies, bone ash, P, Ca and Zn
contents, and P retention were significantly increased,
which indicated a better mineral utilization in fish fed the
acidified diet. Pandey and Satoh (2008) concluded that
adding a P supplement (in the form of monocalcium or
dicalcium phosphate) to low fishmeal-based diets for rain-
bow trout might not be necessary if citric acid was added.
Positive effects on the growth performance and mineral
utilization in fish fed citric acid-supplemented diets
have also been reported in red sea bream (Sarker et al.
2005; Hossain et al. 2007). Hossain et al. (2007) reported
that even without additional 5 g kg
1 P supplementation,
citric acid at 10 g kg
1 inclusion level could significantly
increased the absorption and retention of P while
improving the weight gain and feed utilization when
compared to red sea bream fingerlings fed the nonacidified
diet. Similarly, Sarker et al. (2005) found that citric acid at
a 30 g kg
1 inclusion in fishmeal-based diet without addi-
tional P significantly increased fish growth, FCR and nutri-
ent retention, besides lowering N and P loading that would
otherwise been excreted into the aquatic environment. In a
subsequent study, even at a lower dietary inclusion of
10 g kg
1 and without inorganic P supplementation, citric
acid significantly improved fish growth, FCR and nutrient
retention, while also lowering the N and P loading when
used in a plant protein-based diet for red sea bream (Sarker
et al. 2007). Several recent studies with juvenile beluga
sturgeon (Khajepour & Hosseini 2012a,b) and yellowtail
(Sarker et al. 2012a,b) have also confirmed that dietary
supplementation of citric acid to plant protein-based diets
is very effective in improving the growth performance and
retention/availability of minerals, particularly P. Organic
acid-supplemented feeds will therefore contribute to the
formulation of more eco-friendly aquafeeds.
In a dose–response growth study using graded dietary
levels of citric acid, Su et al. (2014) reported that an inclu-
sion level of 1.9 g kg
1 diet would elicit maximum weight
gain of the Pacific white shrimp (Litopenaeus vannamei,
Boone, 1931). Based on other parameters such as
survival, digestive enzyme activities, immune response and
resistance to vibriosis, a supplementation level of citric acid
at 2–3 g kg
1 was recommended for white shrimp
(Table 2). In contrast, Silva et al. (2015) reported that when
sodium citrate was added to a commercial shrimp feed at
73 mM kg
1, no growth improvement was observed.

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© 2016 Wiley Publishing Asia Pty Ltd 349
W.-K. Ng and C.-B. Koh

Table 7 Use of commercial and prototype organic acids blend in aquafeeds

Organic acids/salts Dose Species and weight (g) Impacts* Reference

(g kg
1)

Organic acids 0, 5 and 10 Red hybrid tilapia, Lowered gut content pH Koh et al. (2016)
blend (OAB) Oreochromis sp. Reduced total cultivable bacterial in faeces and gut
(9.45  0.01) Improved apparent nutrient and mineral digestibility
Enhanced resistance against Streptococcus agalactiae
OAB 0, 2 and 4 Yellow catfish, No effects on growth performance, Zhu et al. (2014)
Pelteobagrus fulvidraco P utilization and mineral retention
(1.3  0.0) No effects on body composition
Enhanced antioxidant capacity of fish
OAB 0 and 20 South African abalone, Enhanced growth and SGR under ideal conditions Goosen
Haliotis midae (2.1  0.8) No effects on FCR, feed intake and gut microbiota et al. (2011)
OAB 0 and 20 Tiger shrimp, Penaeus No effects on growth performance Ng et al. (2015)
monodon (postlarvae 12) Enhanced nutrient and P digestibility
Enhanced resistance to Vibrio harveyi
Less hepatopancreatic damage
OAB 0, 10, 20 Pacific white shrimp, Improved growth at 20 g kg
1 diet Romano
and 40 Litopenaeus vannamei Lower hepatopancreas Vibrio counts et al. (2015)
(postlarvae 12) Improved nutrient and P digestibility
Hepatopancreatic protective properties
Higher survival during Vibrio harveyi challenge
Formic acid + 0 and 4 Olive flounder, No effects on growth performance Park et al. (2011)
propionic acid Paralichthys olivaceus Reduced total gut bacterial counts and tended
(3.5  0.05) to lower the Vibrio counts
Increased resistance against Edwardsiella tarda
Na-formate + Na-butyrate 0 and 10 g Rainbow trout, No effects on growth rate or feed utilization Gao et al. (2011)
(ratio 2:1 on acid moiety acid Oncorhymchus mykiss Decreased nutrient digestibility in fish
weight basis) moiety/kg (230 and 950) fed plant protein-based diets
No adverse effects on gut health
OAB 0, 1, 2 and 3 Hybrid tilapia, Tended to improve growth, feed Ng et al. (2009)
Oreochromis sp. utilization and nutrient digestibility
(6.90  0.03) Lowered diet pH and total faecal bacterial counts
Increased resistance against S. agalactiae
OAB 0, 5, 10 Nile tilapia, No effects on weight gain, total length and FCR Petkam
and 15 Oreochromis niloticus (~7) Numerically improved body weight up to et al. (2008)
11% with 15 g kg
1 OAB diet
OAB 0, 5, 10 Rainbow trout, Tended to improve growth de Wet (2005)
and 15 O. mykiss (~40) Improved SGR at 10 and 15 g kg
1 inclusion levels
Na-acetate, 0, 5 Atlantic salmon, No effects on SGR, macronutrient digestibility, total lipid Bjerkeng
Na-propionate and 20 S. salar (L.) (725  7) content, fatty acid compositions of fillet and gut tissues et al. (1999)
and Na-butyrate
(5:5:2 w/w/w)

*SGR, specific growth rate; FCR, feed conversion ratio.

of red hybrid tilapia, Oreochromis sp. In contrast, Zhou

Formic acid and its salt
Several studies presented at conferences reported that
graded levels of dietary salts of formic acid such as K-difor-
mate (KDF) (Ramli et al. 2005; Lim et al. 2010) and
Na-diformate (NaDF; Liebert et al. 2010) up to 10 g kg
1
enhanced the growth, feed utilization efficiency and protein
retention efficiency of tilapia (Table 3). Ng et al. (2009)
demonstrated that soya bean meal-based diets supple-
mented with 2 g kg
1 KDF tended to positively impact
growth, feed utilization efficiency and nutrient digestibility
et al. (2009) reported no significant effect on growth
performance of hybrid tilapia (Oreochromis niloticus,
Linnaeus, 1758 9 Oreochromis aureus, Steindachner, 1864)
fed the same acidifier, at various dietary levels. These
contradictory results may be due in part to the variation in
culture and rearing conditions. More recently, Abu Elala
and Ragaa (2015) reported that the inclusion of 2 or
3 g kg
1 KDF in the diets of Nile tilapia (O. niloticus) sig-
nificantly enhanced the growth performance, feed utiliza-
tion efficiency and apparent crude protein digestibility,

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350 © 2016 Wiley Publishing Asia Pty Ltd

which may be attributable to the significant decrease in the
pH of stomach and upper intestinal tract.
Vielma and Lall (1997) reported that dietary formic acid
had a significant effect on gut pH. The changes to
hydrochloric acid and bile accretion in fish fed the formic
acid-supplemented diet led to higher gut pH. In spite of
this, improved absorption of P, Mg and Ca in rainbow
trout was still observed. L€ uckst€adt (2008b) noted that at
13.5 g kg
1, dietary KDF did not influence the gut pH of
Atlantic salmon (Salmo salar, Linnaeus, 1758), but it did
significantly elevate the digestibility of protein, dry matter
and gross energy of fish. Morken et al. (2011) demon-
strated that the inclusion of 10.6 g kg
1 NADF in the diet
of rainbow trout together with high extrusion temperature
(141°C) improved the digestibility of major nutrients (pro-
tein, lipid and ash) and individual amino acids as well as
the physical quality (expansion ratio, durability and water
stability) of barley protein concentrate-based diets. In con-
trast, Morken et al. (2012) noted no significant interactions
between KDF and extrusion temperature with respect to
both nutrient digestibility and physical quality of extruded
feeds when 12 g kg
1 KDF was supplemented into diets
containing soya bean meal for Atlantic salmon. They attrib-
uted the inconsistent findings to differences in the concen-
trations of organic acids, animal species, diet composition
as well as the culture conditions.
In a study with crustaceans, despite no significant impact
on growth, Chuchird et al. (2015) provided evidence that
Pacific white shrimp fed formic acid-supplemented diets
showed enhanced resistance to Vibrio parahaemolyticus
challenge. The positive role of various dietary organic acids
in mitigating fish and shrimp bacterial diseases will be fur-
ther discussed in a later section of this review. Similarly,
Silva et al. (2015) did not observe any improvement in
growth response, feed utilization or survival of white
shrimp when fed a commercial shrimp feed with added
Na-formate.
Lactic acid and its salt
In a study with Arctic charr (Salvelinus alpinus, Linnaeus,
1758), Ringø (1991) showed that inclusion of Na-lactate in
a commercial diet resulted in a significant improvement
(21%) of fish body weight compared to fish fed an unsup-
plemented diet. Moreover, fish fed a diet supplemented
with 10 g kg
1 Na-lactate had significantly lower amounts
of water, energy, lipid, protein and free amino acids in their
gut contents (Ringø 1991). The authors suggested this
finding to be indicative of improved nutrient utilization. In
their follow-up study, dietary Na-lactate was found to
improve growth of Arctic charr without affecting the
chemical composition of the fish (Ringø et al. 1994). To
validate these findings, Gislason et al. (1994) supplemented
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Dietary organic acids in aquaculture
15 g kg
1 Na-lactate into diets of another salmonid spe-
cies, Atlantic salmon. Contrary to Ringø (1991), no effects
on growth, mortality or gross faecal chemical composition
were observed in salmon juveniles fed the lactate-supple-
mented diet. Gislason et al. (1994) speculated that the
effect of Na-lactate on growth might be species dependent.
In a more comprehensive study using larger Atlantic
salmon and Arctic charr, Gislason et al. (1996) found a
significant improvement in the growth of Arctic charr fed a
diet supplemented with 15 g kg
1 Na-lactate, while no
effect was observed in Atlantic salmon fed the same diet.
They attributed the growth differences between these two
species to the doubled retention time of feed in the digestive
tract of Arctic charr. In contrast, no significant differences
in growth performance were observed in rainbow trout
(Pandey & Satoh 2008) and red sea bream (Hossain et al.
2007) fed diets supplemented with 10 g kg
1 lactic acid.
However, the absorption of P was significantly increased in
sea bream fed diets acidified with lactic acid (Table 4).
More recently, Castillo et al. (2014) reported that the addi-
tion of Ca-lactate to diets tended to lower the pH of experi-
mental diets and stomach contents as well as improve the
growth, feed efficiency and protein efficiency ratio of red
drum (Sciaenops ocellatus, Linnaeus, 1766). However,
growth performance of fish was reduced when fed diet sup-
plemented with 30 g kg
1 of Ca-lactate. Moreover, dietary
Ca-lactate did not alter the pH of intestinal contents, despite
the gastric pH has been numerically lowered.
Similar to studies reported for citric and formic acids or
their salts, these studies with lactic acid/salt once again
showed the importance of using the correct type and diet-
ary concentration of organic acid for different fish species.
No beneficial effects or even detrimental effects on fish
growth and/or physiology can occur if the wrong type and/
or dosage of organic acids are used. In a recent study, when
supplemented at graded levels of dietary lactic acid from 0
to 16 g kg
1, we did not observe any further improvement
in growth performance above 2 g kg
1 in the feeds of the
freshwater prawn, Macrobrachium rosenbergii (unpublished
data). Considering the multitude of farmed aquatic ani-
mals, much more research is required to further elucidate
this basic tenet of organic acid supplementation in
aquafeeds.
Butyric acid and its salt
A study on African catfish (Clarias gariepinus, Burchell,
1822) (Owen et al. 2006) reported that dietary Na-butyrate
did not significantly increase the weight gain, specific
growth rate (SGR), PER and FCR of fish (Table 5). Never-
theless, this study showed that Na-butyrate diets altered the
intestinal microbiota of catfish. Robles et al. (2013)
demonstrated that inclusion of 3 g kg
1 butyrate into the
351
W.-K. Ng and C.-B. Koh
diets of sea bream (Sparus aurata, Linnaeus, 1758) signifi-
cantly increased weight gain and led to some potentially
beneficial changes in intestinal metabolites. The glycolysis
intermediates, 2-phosphoglycerate and 3-phosphoglycerate,
were found to be accumulated in fish fed the butyrate-sup-
plemented diet through inhibition on glycolysis, as the
butyrate was a preferential energy substrate over glucose.
Dietary butyrate was shown to spare the oxidation of some
amino acids and increased their bioavailability into arterial
circulation, resulting in enhanced absorption of certain
essential amino acids in the fish gut. Dietary butyrate also
significantly increased the concentration of some nucleo-
tide derivatives in the fish gut and Robles et al. (2013) sug-
gested that butyrate may have a potent effect on the
absorption of nucleotides. The detailed knowledge pro-
vided by this study on the metabolic consequences of buty-
rate supplementation in sea bream diets allows us to better
understand the many reported effects of dietary organic
acids altering the microbiota population in fish gut, often-
times with potential beneficial effects.
Na-butyrate supplementation has been reported to posi-
tively affect the growth performance of Pacific white
shrimp (Silva et al. 2016). The study reported that inclu-
sion of 20 g kg
1 butyrate in the diets significantly
improved the feed efficiency, N retention, protein efficiency
rate, survival and yield of shrimp as compared to the con-
trol diet. Apart from these, dietary butyrate at 5, 10 or
20 g kg
1, significantly lowered the Vibrio sp. counts in the
gut of shrimp and increased their serum agglutination titre.
Silva et al. (2016) suggested that butyrate supplementation
can alter the gut microbiota of shrimp which contributes to
better growth performance. Similarly, when Na-butyrate
was added to a commercial shrimp feed at 73 mM kg
1,
significant improvements in growth and feed intake were
observed (Silva et al. 2015).
According to Liu et al. (2014), dietary Na-butyrate at a
level of 0.3 g kg
1 increased the weight gain and improved
the FCR of juvenile common carp (Cyprinus carpio, Lin-
naeus, 1758) prefed with or without oxidized oil. Simulta-
neously, immune responses and intestinal condition of the
common carp were positively affected by the dietary buty-
rate with a more pronounced effect in the repairing of
intestinal mucosal damage in fish fed the oxidized oil.
Other organic acids and their salts
Several other organic acids and their salts have also been
investigated (Table 6). Hossain et al. (2007) investigated
the effect of dietary malic acid on growth and P utilization
in red sea bream. Although there was no significant
improvement in weight gain and feed utilization, fish fed a
10 g kg
1 malic acid-supplemented diet were found to
have significantly better P absorption.
352
Succinic acid supplementation in rainbow trout diets
had been reported to negatively affect the voluntary feed
intake of fish, although there was no apparent effect on
protein efficiency and energy utilization (Fauconneau
1988). In another study with rainbow trout, Sugiura et al.
(2006) reported that dietary acetic acid at 50 g kg
1 inclu-
sion level enhanced P digestibility and tended to decrease
stomach pH but not gut pH. Additionally, dietary Na-pro-
pionate at a level of 10 g kg
1 depressed the growth of Arc-
tic charr, which might be due, in part, to their inferior
utilization of lipid and protein which led to higher water
and gross energy contents in the gut contents (Ringø 1991).
In contrast, Silva et al. (2013) demonstrated that among
various organic acid salts (acetate, formate, lactate, citrate,
propionate and butyrate), diets supplemented with
20 g kg
1 Na-propionate led to significantly improved feed
intake, apparent gross energy and phosphorous availabili-
ties of Pacific white shrimp and were recommended as the
organic acid salt with the greatest potential for use in mar-
ine shrimp feeds. In their subsequent study, Silva et al.
(2016) showed that dietary Na-propionate enhanced
growth performance and feed utilization efficiency of
shrimp irrespective of the inclusion levels (5, 10 or
20 g kg
1). However, dietary acidification did not improve
P availability for shrimp. Nevertheless, when Na-propio-
nate was later compared with Na fumarate, shrimp fed diets
with Na fumarate significantly outperformed shrimp fed
with propionate (Silva et al. 2015). This is in spite of the
observed increased in enzyme activities in shrimp fed the
Na-propionate added diet. Needless to say, much more
research is needed to fully understand and to evaluate the
most suitable organic acid to be used for each species of
farmed aquatic animal as well as its mode of action.
Organic acids blend
Each organic acid has its own spectrum of antimicrobial
activity due to their specific physical and chemical prop-
erties (Dibner & Buttin 2002). Therefore, the advantage
of using organic acids blend (OAB) in animal feeds is that
the OAB can have a broader spectrum of antimicrobial
activity against a wider range of disease-causing bacteria
and with potential synergistic effects on growth perfor-
mance and nutrient utilization. Moreover, OAB may
allow further reduction of the dose used in animal feeds,
thereby reducing costs. OAB constitutes a potential strat-
egy to overcome the inconsistency of findings on the use
of single organic acids in the feeds of various aquatic ani-
mals (Tables 2–6). Commercial and prototype OAB is
usually proprietary formulations of the suppliers and
researchers, respectively.
An 8-week feeding trial comparing the growth perfor-
mance of all male Nile tilapia fingerlings fed diets
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© 2016 Wiley Publishing Asia Pty Ltd

supplemented with 5 g kg
1 oxytetracycline (OTC) or an
organic acid/salts blend containing citric acid and Ca salts
of formic, propionic, lactic and phosphoric acid at various
dietary levels (0, 5, 10 or 15 g kg
1) was conducted by Pet-
kam et al. (2008). Tilapia fed a diet supplemented with
15 g kg
1 organic acid/salts blend tended to have higher
body weight but no statistical differences in weight gain,
total length and FCR among the treatments were detected
(Table 7). Petkam et al. (2008) attributed the lack of signif-
icant effects to the shorter experimental period and optimal
culture conditions.
Similar improvements in the body weight of red hybrid
tilapia fed diets supplemented with a commercial OAB at 1
or 2 g kg
1 have been reported by Ng et al. 2009. The
addition of OAB tended to improve feed utilization, nutri-
ent digestibility and reduce total cultivable bacterial counts
in the faeces and gut of tilapia in a dose-dependent manner.
Total cultivable bacterial count in the faeces, mainly
Aeromonas hydrophila, was significantly reduced with OAB
inclusion. Furthermore, cumulative mortality of 16 days
postchallenge with Streptococcus agalactiae was lower in
fish fed OAB-supplemented diets. When raised from finger-
lings to near market size, we observed that hybrid tilapia
fed diets supplemented with a prototype OAB at 5 or
10 g kg
1 tended to have improved growth and feeding
efficiency while those fed the 10 g kg
1 OAB diet had sig-
nificantly enhanced P, dry matter and ash utilization over
tilapia fed the control diet (Koh et al. 2016). No significant
growth differences were detected between tilapia fed the
OTC- or OAB-added diets. Moreover, the prophylactic
effects of the 5 g kg
1 OAB or 5 g kg
1 OTC diets in tila-
pia subsequently challenged with S. agalactiae were similar
and led to significantly enhanced disease protection than
fish fed the control diet. Due to these encouraging labora-
tory-based results, we recently conducted a field trial of the
prototype OAB in a commercial Nile tilapia farm and
observed improved nutrient digestibility, reduced bacterial
counts in faecal matter, enhanced immune response, less
skin bacterial infections and significantly reduced antibiotic
use by the tilapia farmer (unpublished data). Similarly,
Park et al. (2011) also observed significantly improved
Edwardsiella tarda resistance in the olive flounder (Par-
alichthys olivaceus, Temminck & Schlegel, 1846) when fed
diets with OAB supplementations, although the implica-
tions to growth were not reported.
In contrast, Gao et al. (2011) reported no significant
improvement in growth or feed utilization of rainbow trout
fed fishmeal or plant protein-based diets added with 10 g
acid moiety/kg organic acid salt blends at two different feed
production stages (before and after extrusion). Their
24-day digestibility trial showed that trout (~950 g) fed a
plant protein-based diet with the acid salt blend added
before extrusion had significantly lower nutrient digestibility
Reviews in Aquaculture (2017) 9, 342–368
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Dietary organic acids in aquaculture
(dry matter, organic matter, crude lipid and amino acids).
Similarly, Bjerkeng et al. (1999) reported that Atlantic
salmon (725  7 g) fed diets supplemented with 0, 5 or
20 g kg
1 of an OAB containing Na salts of acetic, propi-
onic and butyric acid (5:5:2 w/w/w) showed no significant
effects on growth, or apparent digestibility coefficients of the
macronutrients. More encouraging results on salmonids
were reported by de Wet (2005) who investigated the use of
a commercial OAB at 0, 5, 10 or 15 g kg
1 on the growth of
rainbow trout fingerlings. After four months, a trend of
improved growth was observed with graded levels of dietary
acidification. The SGR of trout fed diets supplemented with
10 or 15 g kg
1 acid blends was significantly improved
when compared to the unsupplemented diet. The final aver-
age body weight of trout fed these diets was nearly similar to
that of fish fed diet supplemented with an AGP.
Goosen et al. (2011) investigated the effects of different
types of OAB (10 g kg
1 acetic acid + 10 g kg
1 formic
acid, 10 g kg
1 benzoic acid + 10 g kg
1 sorbic acid and
10 g kg
1 Na-benzoate + 10 g kg
1 K-sorbate) supple-
mentations at 20 g kg
1 on growth performance and feed-
ing efficiency of South African abalone (Haliotis midae,
Linnaeus, 1758). Growth performance was significantly
enhanced, but no notable effect on FCR, feed intake and
gut microbiota community was observed when cultured
under favourable growth conditions (16.5°C). However, no
significant growth improvement was recorded under sub-
optimal growth conditions (20.5°C). They attributed the
lack of growth stimulating effect to the disruption of physi-
ological function in abalone when exposed to temperature
outside their optimal range of 12–20°C.
In a study with juvenile yellow catfish (Pelteobagrus ful-
vidraco, Richardson, 1846), a commercially available OAB
product significantly reduced the serum reactive oxygen
species (ROS) level, serum superoxide dismutase (SOD)
activity and catalase (CAT) activity of fish, indicating that
organic acids could protect the fish body from cell oxida-
tive damage when fed mainly with plant protein-based diets
(Zhu et al. 2014). The mechanism by which organic acid
reduces the risk of oxidative damage in fish remains to be
elucidated.
Recent studies with marine shrimp indicated that under
laboratory conditions, the correct dietary OAB concentra-
tion can significantly improve growth performance
(Romano et al. 2015). However, when the same dietary
concentration was field tested in a commercial shrimp
farm, the beneficial impact on growth was not evident,
probably due to the multiple factors that masked the
growth beneficial effects under pond culture conditions
(Ng et al. 2015). Nevertheless, in both laboratory- and
field-based trials, marine shrimp fed an OAB-supplemented
feed showed better nutrient utilization efficiency, lower
hepatopancreas bacterial counts, enhanced immune
353
W.-K. Ng and C.-B. Koh
response and higher survival under Vibrio bacterial infec-
tions.
A review of current literature on aquatic animals would
indicate that in many instances, researchers have reported
that some organic acids, their salts and/or mixtures thereof
could improve growth, feed utilization, mineral availability
and disease resistance. Nevertheless, despite the generally
consistent improvement in nutrient availability of organic
acid-supplemented diets, contradictory results have been
reported for the growth-promoting effects of dietary
organic acids, which seem to depend on the aquatic animal
species, physiology, age and/or the type and dosage of
organic acid used. By further examining the potential mode
of action of these organic acids, we hope that this review
will help stimulate more focused and comprehensive
research on this subject matter for the benefit of the aqua-
culture industry.
Mode of action and effects of dietary organic acids
Antimicrobial properties of organic acids
The antimicrobial activity of lipophilic weak acids was tra-
ditionally explained by the perturbation of membrane func-
tion, which blocks the transport of substrate molecules
(amino acids, organic acids, phosphate, etc.) into cells
(Freese et al. 1973). Other mechanisms believed to be
responsible for the ability of organic acids to limit microbial
growth have been reviewed by Cherrington et al. (1991a)
and Booth and Stratford (2003). The most obvious mode
of action of these lipid-soluble weak acids is via direct acidi-
fication of the extracellular pH through its ability to disso-
ciate into ions and release hydrogen ions (protons) to the
surrounding medium. However, the general consensus is
that the predominant mode of action of these acidifiers is
mainly based on their ability to lower the cytoplasmic pH
once they traverse across the cell membranes of microbes.
The majority of bacterial species has specific pH require-
ments for optimal growth and is unable to grow under
extreme acidic conditions (pH < 4.5). Organic acids can
exert their antimicrobial activity on microbes by directly
lowering the pH of the environment via releasing hydrogen
ions and thus preventing and/or impeding the growth and
proliferation of acid-sensitive bacteria. Weak organic acids,
such as acetic, citric, benzoic, sorbic and lactic acids, are
commonly employed to lower the pH of foods or beverages
in order to limit microbial growth (Stratford & Eklund
2003).
It is now generally accepted that the antimicrobial effi-
cacy, both bacteriostatic and bactericidal effects, of organic
acids is mainly due to their ability to traverse across the
semi-permeable membrane of bacteria and to dissociate in
the pH neutral cytoplasm (Cherrington et al. 1991a; Booth
& Stratford 2003). Despite their various molecular struc-
354
tures (Table 1), all organic acids seem to have a similar
mode of action against microorganisms. Organic acids are
believed to be more effective at low pH when they are
mainly in the undissociated form, and are thus the most
effective form in killing microorganisms (Brul & Coote
1999; Lambert & Stratford 1999). This is because the undis-
sociated form of an organic acid is lipophilic and can pas-
sively diffuse through a bacterium’s cell membrane. Once
inside the nearly pH neutral cytoplasm, it dissociates,
releasing charged acid anions and protons that are imper-
meable back across the cell’s membrane (Cherrington et al.
1991a; Lambert & Stratford 1999).
Accumulation of excess protons within the cell will lower
the cytoplasmic pH, thereby causing an inhibition of bacte-
rial cell metabolism through the suppression of cell
enzymes (Warth 1991), particularly the pyruvate decar-
boxylase enzyme which contributes to energy metabolism
(Sava 2011). Cell death may occur when the cytoplasmic
pH drops below the physiological optimal range for growth
(Smigic et al. 2009). Lowering the cytoplasmic pH might
also neutralize the electrochemical gradient, for example
the pH gradient (DpH) across the plasma membrane,
which is required for the active transport of nutrients. Bac-
teria that do not tolerate changes in trans-membranous pH
gradients such as E. coli, Salmonella and Campylobacter
(Dibner & Buttin 2002) will undergo cellular stress and
eventually die (Jensen 2001). To restore the intracellular
pH within a physiological optimal range for growth and to
sustain functional macromolecules, the bacterial cell is
forced to pump out the excess protons, released by the
acids, via the membrane-bound H+-ATPase. Proton pump-
ing by the H+-ATPase requires substantial metabolic energy
in the form of adenosine triphosphate (ATP) (Holyoak
et al. 1996) and could therefore lead to depletion of cellular
ATP, and the cells will eventually die of energy depletion
(Warth 1991; Ricke 2003).
Depletion of ATP is not the sole mechanism for inhibit-
ing cellular growth or inducing death since proton removal
also leads to an accumulation of weak acid anions within
the cytoplasm, which is also responsible for the growth
inhibition mechanisms of organic acids at low pH (Russell
1992; Brul & Coote 1999; Lambert & Stratford 1999). Accu-
mulation of organic acid anions can inhibit the synthesis of
macromolecules, for example nucleic acids (Cherrington
et al. 1990), proteins, lipids and carbohydrates (Jensen
2001) as well as enzyme activity (decarboxylases and cata-
lases) and nutrient transport systems within the cytoplasm
(Russell & Diez-Gonzalez 1997; Roth & Kirchgessner 1998;
Partanen & Mroz 1999), and eventually results in cell death.
A summary of the inhibitory mechanisms of some organic
acid anions is shown in Table 8.
The substantial accumulation of acid anions in the cyto-
plasm may also cause hyperosmotic stress on the bacterial
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Table 8 Inhibitory mechanisms of some organic acid anions (based on
van Dam 2006)
Acid anion Mode of action
Formate Inhibits enzyme activity, specially decarboxylase and
catalase
Acetate Inhibits enzyme activity
Increases heat sensitivity
Propionate Influences membrane transport
Inhibition on synthesis of some amino acids
Lactate Inhibits enzyme activity
Sorbate Inhibits a series of enzymes, amino acid uptake and
synthesis of RNA/DNA
Cell membrane damage
Benzoate Inhibits a series of enzymes and amino acid uptake
Cell membrane damage
Changes membrane fluidity
cell, which may contribute to growth inhibitory effects
(Roe et al. 1998). The high anion concentration within the
cytoplasm has the potential to increase the turgor pressure
on the cell via increasing the osmotic pressure (Kroll &
Booth 1983). To maintain the turgor pressure at a constant
level, the cell appears to reduce other anion pools, for
example intracellular glutamates to compensate for this
accumulation (Roe et al. 1998). Thus, perturbation of
anion balance may contribute to inhibitory effects on
growth and eventually results in cell death (Roe et al.
1998).
Effects of organic acids on gastrointestinal microbiota of
aquatic animals
Several studies have demonstrated that the inclusion of
organic acids, their salts and/or mixtures thereof to aqua-
feeds can influence the bacterial community of the gas-
trointestinal tract in aquatic animals. The first evidence of
appreciable reduction in the cultivable microbiota in the
expelled faeces and gut surfaces of fish fed organic acid-
supplemented diets was reported by Ng et al. (2009). Later,
the antimicrobial efficacy of dietary organic acids on gut
microbial community was reported in olive flounder (Park
et al. 2011), Pacific white shrimp (Silva et al. 2013, 2016)
and red hybrid tilapia (Koh et al. 2016). The inclusion of
organic acids in fish and shrimp diets has been shown to
reduce the population of potentially deleterious bacteria
such as Vibrio spp. in the intestinal tract (Park et al. 2011;
Silva et al. 2013, 2016; Chuchird et al. 2015). Total viable
bacterial and Vibrio counts in the hepatopancreas of marine
shrimp were found to be lower when fed diets with
20 g kg
1 OAB compared to the control treatment without
any organic acids (Ng et al. 2015; Romano et al. 2015).
Furthermore, it has been reported that the inclusion of
3 g kg
1 KDF to plant protein-based diets stimulated the
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Dietary organic acids in aquaculture
growth of beneficial lactic acid bacteria in the gut of tilapia
(Abu Elala & Ragaa 2015).
Other than a reduction in cultivable bacterial population,
changes in the population composition of the gastrointesti-
nal microbiota have also been associated with organic acids
supplementation. Liu et al. (2014) observed slight alter-
ations in the adherent bacterial communities within the gut
segments of common carp fed 0.3 g kg
1 Na-butyrate
diets. Zhou et al. (2009) observed that dietary supplemen-
tations of KDF stimulated the colonization of some benefi-
cial gut bacteria and inhibited the growth of others, but
this did not lead to significant improvement in growth per-
formance of hybrid tilapia. Alterations in the gut micro-
biota have also been reported in Pacific white shrimp when
fed diets supplemented with Na-propionate or Na-buty-
rate, which might have contributed to the observed higher
growth compared to shrimp fed a control diet (Silva et al.
2016). In contrast, Owen et al. (2006) reported that
Na-butyrate supplementation did not significantly affect
the gut bacterial community of African catfish. In hybrid
tilapia, the addition of 1–3 g kg
1 OAB showed no dietary
effect on the gut microbiota composition, while 2 g kg
1
KDF tended to lower the relative population of potentially
harmful A. hydrophila (Ng et al. 2009). Similarly, Goosen
et al. (2011) reported that supplementation of various
OAB, at 20 g kg
1 inclusion level, had no effect on the gut
microbiota of the South Africa abalone.
Changes in the population and composition of gut
microbiota can have significant impact on the cultured
aquatic animal growth performance, nutrient utilization,
immune response and resistance against pathogens. Further
focused research is needed to elucidate the effects of dietary
organic acids as mediated through changes in gut micro-
biota. The microbial community of the gastrointestinal
tract needs to be characterized with greater precision using
more sophisticated methodology in order to provide deeper
insight into the underlying mechanisms.
Organic acids enhance nutrient availability
Addition of organic acids in aquafeeds had generally
resulted in improved nutrient digestibility (Tables 2–7).
Various hypotheses have been proposed on the mode of
action of organic acids in enhancing nutrient utilization in
terrestrial livestock which includes, among others, (i) low-
ering gastric pH leading to increase pepsin activation, (ii)
lowering diet and intestinal pH, which may increase solubi-
lization of minerals, (iii) acting as chelating agents which
binds various cations within the intestine resulting in
increased mineral absorption or (iv) inhibiting the colo-
nization of harmful microbes in the intestine which may
otherwise utilize nutrients meant for the host animal
(Sch€oner 2001; de Wet 2005). In animal nutrition, organic
355
W.-K. Ng and C.-B. Koh
acids and their salts act as a growth promotant primarily
through their impact in the feed and gastrointestinal tract
of the animal.
Organic acids are commonly used as preservatives to
help prevent feeds and feed ingredients from deterioration
caused by bacteria, moulds and yeasts (Ricke 2003; Skri-
vanova et al. 2006; van Dam 2006). Inclusion of organic
acids reduces the pH value of the feed and therefore pre-
vents the growth of undesired harmful microbes during
storage and/or release of toxic metabolites (especially
mycotoxins) produced by fungi. It is generally known that
the ingestion of even small amounts of mycotoxins from
contaminated feedstuffs can cause serious nutritional and
health problems during animal production, especially in
aquatic animals. Therefore, acidification enhances the
hygienic quality of the feed and reduces the loss of its nutri-
tional value during storage. In addition to improving feed
hygiene, organic acids reduce the buffering capacity of the
dietary feed ingredients. This is important as it ensures
optimal intestinal pH, which results in better feed diges-
tion, especially in young animals (Metzler & Mosenthin
2007).
In the gastrointestinal tract, organic acids exert their effects
on performance via two main mechanisms. Firstly, it reduces
the pH within the stomach and possibly small intestine of
animals (Sch€ oner 2001) and, secondly, through their ability
to inhibit and kill harmful bacteria, as described earlier. In
terrestrial animals, the addition of organic acids reduces the
gastric pH which favours proteolytic enzyme activity, thereby
increasing protein digestibility and animal performance
(Roth & Kirchgessner 1998; Dibner & Buttin 2002). More-
over, organic acids added in feeds may slow down the empty-
ing rate of the stomach, therefore allowing more efficient
hydrolysis of proteins and absorption of nutrients in the
small intestine. These effects are generally more pronounced
in younger animals, where the pancreatic enzyme secretion
and the hydrochloric acid production are inadequate com-
pared to adults (Freitag 2007).
Another beneficial effect of lowering gastric acidity is an
improvement of phosphorus (P) availability from phytate-
P in plant feed ingredients (Dibner & Buttin 2002). The
majority of organic P in plant ingredients exists in the form
of phytic acid or phytates which are not digestible to
monogastric animals due to lack of phytase activity in the
digestive tract. The addition of organic acids can increase
the phytate-P utilization by inducing microbial phytase
activity, which is more efficient in lower pH environment
(Dibner & Buttin 2002; de Wet 2005). By lowering the
intestinal pH, organic acids may also increase mineral
solubility leading to improved absorption of minerals.
Furthermore, the anions of weak acids can act as chelating
agents by binding up various cations to form mineral
and acid complexes along the intestine, which results in
356
increased mineral absorption by the intestinal cells
(Ravindran & Kornegay 1993).
Another possible mode of action by organic acids in
enhancing nutrient availability is through a direct stimula-
tion of mucosa proliferation activity in the gastrointestinal
tract. In rats, organic acids (acetate, propionate and buty-
rate) stimulated the proliferation of gastrointestinal muco-
sal cells by inducing the expression of plasma glucagon-like
peptide (GLP-2), ileal proglucagon mRNA, glucose trans-
porter (GLUT2) and c-myc, c-jun and c-fos, which can
potentially mediate mucosal proliferation (Tappenden &
McBurney 1998). It has been suggested that the positive
effects of organic acids, particularly butyrate, in stimulating
the growth of intestinal epithelium cell can lead to
increased nutrient absorptive capacity (Topping & Clifton
2001). Adil et al. (2010) reported that the dietary supple-
mentation of organic acids resulted in increases in villus
height in the duodenum and jejunum of broiler chicken.
The thickness of muscularis on the intestinal mucosa was
also decreased in all the segments of the small intestine.
These alterations in gastrointestinal structure may facilitate
nutrient absorption in the small intestine, thereby improv-
ing growth performance (Adil et al. 2010; Samanta et al.
2010).
Finally, improved nutrient utilization may also be attrib-
uted to the strong antimicrobial activity of organic acids
that can inhibit the colonization of harmful microbes
within the digestive tract (Kluge et al. 2006). A positive
outcome of reducing harmful microbial counts is a health-
ier gut, and the nutrients, which may otherwise be utilized
by the microbes, are now spared for the host animal.
Reduced competition for nutrients between microbes and
the host animal is one of the mechanisms responsible for
improved nutrient utilization (Partanen & Mroz 1999; Dib-
ner & Buttin 2002; Adil et al. 2010).
Effects of organic acids in aquatic feeds
Organic acids have been reported to affect feed palatability
in aquatic animals. Citric and malic acids were shown to be
very effective in stimulating the feeding responses in red-
belly tilapia, Tilapia zillii (Adams et al. 1988). Xie et al.
(2003) also observed a positive stimulating effect of dietary
citric (10
2–10
6 M), propionic (10
4–10
6 M) and lactic
acids (10
2–10
5 M) on the feeding responses of juvenile
Nile tilapia, while dietary 10
3 M acetic and propionic acids
had repulsive effects. A study with Pacific white shrimp
showed that among various Na salts of organic acids (for-
mate, acetate, propionate, lactate, butyrate and citrate), the
supplementation of propionate and butyrate salts to a
commercial shrimp diet at 2 g kg
1 acted as feed attrac-
tants and significantly enhanced feed intake (Silva et al.
2013). However, it has also been reported that high
inclusion levels (≥ 50 g kg
1) of certain organic acids
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 Dietary organic acids in aquaculture

(citric acid and succinic acids) tended to negatively impact
the palatability of the feed and reduced feed intake
(Fauconneau 1988; Sugiura et al. 1998). Improved or
reduced feed palatability associated with dietary organic
acid supplementation might have contributed to some of
the variations in growth performance reported in aquatic
animals (Tables 2–7). Therefore, more research is needed
to determine the inclusion level of organic acids to formu-
late organoleptically pleasing diets that will positively affect
feed intake and growth performance of aquatic animals.
Effects of organic acids on gut pH
A summary of the impact of dietary organic acids or their
salts on diet and stomach/gut pH in fish is given in Table 9.
Organic acids consistently reduced the pH value of experi-
mental diets, but its impact on stomach and/or gut pH was
not always significant. Sugiura et al. (2006) reported that
dietary supplementation of 50 g kg
1 acetic acid in trout
diets tended to decrease gastric and caecal pH, but did not
profoundly reduce the pH of gastrointestinal tract contents,
indicating that trout are efficient in regulating endogenous
gastric acid secretion to maintain their normal gut pH. It
was shown that dietary organic acid did not alter the
steady-state mRNA expression of H+/K+-ATPase as well as
the Na+/bicarbonate cotransporter and had no effect on
gastrin-like mRNA and somatostatin mRNA abundance
which are involved in the secretion and regulation of gastric
acid. Nevertheless, it was observed that the addition of
50 g kg
1 acetic acid to trout diets significantly improved
P availability. Based on these results, the researchers specu-
lated that the hydroxyapatite-P in acidified diets might
have been converted into more absorbable forms such as
di- or mono-calcium phosphates as the observed gut pH
changes were very minor. Comprehensive research focusing
on the actual contribution of a lower pH in the stomach
and gut of aquatic animals to improving nutrient and min-
eral availability (Tables 2–7) in much needed. The role of
feed ingredient types in buffering the pH lowering effects of
various organic acids at different dosages remains to be elu-
cidated. A comparative study between monogastric and
agastric fish might also shed new information on this sub-
ject matter.
Effects of organic acids on digestive enzyme activity
To date, very few studies have been conducted to investi-
gate the effect of dietary organic acids on the digestive
enzyme activity of aquatic animals, but it is a growing
research area. Li et al. (2009) reported that dietary citric
acid (10 g kg
1) increased the protease activity in tilapia
stomach. Moreover, activities of amylase in hepatopancreas

Table 9 Summary of published data on the impacts of dietary organic acid or their salts on pH of experimental diets and gastrointestinal tract
content

Organic acid Level (g kg
1) Species Δ pH diet Δ pH gastrointestinal tract Reference

Stomach Proximal Middle Distal

Ca-lactate 15 Red drum
0.16
0.01 ND Castillo et al. (2014)

30
0.25
0.25 ND
Ca-lactate 15
0.13
0.32 +0.66†
KDF 7.5
0.18
0.21 +0.21†
KDF 15
0.36
0.18 +0.09†
Citric acid 7.5
0.52
0.18 +0.64†
Citric acid 15
0.79
0.29 +0.81†
KDF 1 Nile tilapia ND
0.14
0.07
0.00
0.01 Abu Elala and Ragaa (2015)
KDF 2 ND
0.38*
0.23*
0.03
0.11
KDF 3 ND
0.47*
0.45*
0.05
0.22
OAB 1 Red hybrid tilapia
0.06
0.11
0.08† Ng et al. (2009)
OAB 2
0.10
0.12
0.32†
OAB 3
0.15
0.22
0.53†*
KDF 2
0.14
0.20
0.33†
OAB 5 Red hybrid tilapia
0.24
0.03
0.16†* Koh et al. (2016)
OAB 10
0.48
0.01
0.23†*
Citric acid 30 Rohu
1.01
0.00
0.83†* Baruah et al. (2005)
Acetic acid 50 Rainbow trout
1.00
0.50
0.00
0.00
0.10 Sugiura et al. (2006)
Formic acid 4 mL kg
1 Rainbow trout
0.50 ND +0.16 +0.15* Vielma and Lall (1997)
10 mL kg
1
1.00 ND +0.31 +0.41*
Citric acid 20 Rainbow trout
1.75 ND
0.37‡ Sugiura et al. (1998)
Citric acid 50
2.43 ND
0.88‡

Δ, unit change in the pH value relative to the basal diet; *, significantly different from the control group;
, decrease in pH; +, increase in pH;
†, pH value of entire gut content; ‡, pH value of faecal matter; ND, not determined.

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W.-K. Ng and C.-B. Koh
and gut were found to be elevated due to changes in the gut
pH induced by dietary citric acid. This subsequently
affected the release of cholecystokinin and exocrine secre-
tions of the pancreas. Similarly, Su et al. (2014) reported a
significant increase in intestinal protease activity of white
shrimp fed 2 g kg
1 citric acid compared to the control
group. However, no significant differences were observed
for the amylase activity among the shrimp fed graded levels
of citric acid. Castillo et al. (2016) showed that supplemen-
tation of organic acids at 15 g kg
1 diet, especially citric
acid, improved the growth performance in juvenile red
drum, which they attributed in part to the enhanced activ-
ity of digestive enzymes. In this study, pepsinogen activity
was found to be greater in the stomach of fish fed the acidi-
fied diets. Nevertheless, the researchers suggested that pH
of the diets was not the only factor affecting the synthesis of
pepsin, as it might also be influenced by other factors. Diet-
ary citric acid and KDF also resulted in higher activities of
trypsin, lipase and amylase in fish. It was hypothesized that
gut acidification by organic acids may increase secretin
concentration which ultimately stimulate the secretion of
pancreatic enzymes. With respect to intestinal digestive
enzymes, leucine-aminopeptidase and phosphatases were
found to be higher in fish fed with organic acid-added
diets. Given that hydrolysis of P is influenced by the both
acid and alkaline phosphatases, the enhanced enzyme activ-
ities might have been a contributing factor for the
improved mineral digestibility observed.
Silva et al. (2015) reported that white shrimp showed
increased trypsin and chymotrypsin activity when fed diets
with added sodium acetate or propionate, but the activity
of these enzymes was decreased in shrimp fed diets with
sodium lactate or citrate. Sodium formate, butyrate, fuma-
rate and succinate did not alter the activities of these diges-
tive enzymes in shrimp. Despite the lack of impact on
digestive enzyme activities by fumarate and succinate, the
highest in vitro protein digestibility results occurred in
diets supplemented with these organic acid salts and the
lowest digestibility in diets supplemented with acetate or
propionate. Factors contributing to these discrepancies in
shrimp remain to be fully elucidated.
Further studies are needed to better understand the rela-
tionship between gut pH, digestive enzymes and nutrient
availability in aquatic animals.
Effects of organic acids on gastrointestinal morphology
It had been proposed that organic acids, particularly buty-
rate, can stimulate the growth of intestinal epithelial cells
(by acting as an energy source) which ultimately leads to
increased nutrient absorptive capacity (Topping & Clifton
2001). Apart from being an energy source for colonocytes,
butyrate also influences a variety of cellular functions asso-
ciated with intestinal health such as mitigating mucosal
358
inflammation and oxidative stress, and improving the
intestinal epithelial barrier function (Hamer et al. 2008).
Empirical data on the effects of dietary organic acids on gut
morphology in aquatic animals are very scarce. Gao et al.
(2011) observed that despite the lack of impact on growth
or feed utilization, the 10 g acid moiety/kg mixture of
Na-formate and Na-butyrate (ratio 2:1 on acid moiety
weight basis) added before extrusion into fishmeal- or plant
protein-based diets significantly increased the intestine
weight of rainbow trout, indicating that organic acids may
have stimulated the intestinal growth. The morphology of
intestinal mucosal folds was, however, not affected by the
dietary inclusion of the acid salt blend. In common carp, it
had similarly been reported that the intestinal mucosal
morphology was not significantly altered when fish were
fed diets supplemented with Na-butyrate although some
differences were observed in microvillus height (Liu et al.
2014). Apart from the slight improvement in weight gain,
FCR and immune responses, it was reported that dietary
Na-butyrate seemed to be able to prevent or repair the
intestinal mucosal damage and increase the microvillus
density in carp prefed with oxidized soya bean oil.
Organic acids as functional feed additives in
aquafeeds
Organic acids and their salts are considered functional feed
additives since they have additional functions such as
health-promoting or disease-preventing properties beyond
their nutritional value. Their use in terrestrial animal feeds
to control bacterial pathogens and to promote health of
livestock is well documented (Partanen & Mroz 1999; Van
Immerseel et al. 2002, 2003). However, despite their known
antimicrobial properties, the role of organic acids in disease
prevention in farmed aquatic animals has only been investi-
gated recently. Nevertheless, the information regarding
their potential impact against pathogenic bacteria in aqua-
tic animals has been very encouraging.
Ramli et al. (2005) reported that the addition of KDF in
the diet of hybrid tilapia significantly improved their resis-
tance to Vibrio anguillarum and that the effects were dose
dependent. The authors concluded that KDF at 20 g kg
1
inclusion might act as an effective tool to control V. anguil-
larum infection in tilapia culture. In contrast, Lim et al.
(2010) reported no significant improvements in the
immunological responses of Nile tilapia or their survival to
Streptococcus iniae challenge when fed diets supplemented
with different levels (0–15 g kg
1) of KDF for 12 weeks. It
should be noted that the full description of the experimental
methodology including the disease challenge tests is lacking
since the results of both these studies were presented at vari-
ous conferences. Nevertheless, Ng et al. (2009) found that
even at relatively low doses of 1–3 g kg
1 OAB or 2 g kg
1
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© 2016 Wiley Publishing Asia Pty Ltd

KDF in the diets of red hybrid tilapia, it was sufficient to
increase their resistance to S. agalactiae challenge and was
suggested to be linked with a significant reduction in the total
gut bacterial counts. The prophylactic ability of organic acids
in enhancing disease resistance in tilapia was further con-
firmed by another study with hybrid tilapia using a propri-
etary OAB (Koh et al. 2016). It was observed that the
cumulative mortality of hybrid tilapia was significantly
reduced in fish fed diets supplemented with 5 g kg
1 OAB
or OTC when compared to fish fed the control diet at
21 days after challenge with S. agalactiae. We concluded that
the inclusion of 5 g kg
1 OAB was as effective as 5 g kg
1
OTC in protecting tilapia against Streptococcal infections.
Field trials in commercial tilapia farms indicated that part of
this increased resistance to Streptococcus infections was due to
enhanced immune response of the farmed tilapia fed OAB-
supplemented feeds resulting in a significant reduction in the
amount of antibiotics used by the farmer during the grow-
out period (unpublished data). Similarly, Park et al. (2011)
reported that OAB was comparable to OTC in providing
protection against Edwardsiella tarda infection in olive floun-
der. Abu Elala and Ragaa (2015) showed that inclusion of
KDF (1–3 g kg
1) in the diets increased the numbers of ben-
eficial gut lactic acid bacteria and enhanced the cellular and
humeral nonspecific immune responses of Nile tilapia, as well
as reduced the cumulative mortality after 15 days of chal-
lenge with A. hydrophila. Based on their results, the research-
ers attributed the better resistance to the eubiotic effect of
KDF, which play an important role in the activation of the
host immune response.
Several organic acids, particularly formic, acetic, propi-
onic and butyric acids, are the most commonly used and
have been shown to inhibit the growth of various marine
pathogenic Vibrionaceae, including Vibrio campbellii, Vib-
rio harveyi and Vibrio parahaemolyticus in vitro, but their
efficacy is often dependent on the type, pH as well as the
dose used (Defoirdt et al. 2006; Immanuel et al. 2011,
2012; Mine & Boopathy 2011; Adams & Boopathy 2013;
Silva et al. 2013). Short-chain organic acids (formic, acetic,
propionic, butyric or valeric acid), when added to the cul-
ture water of Artemia nauplii at a concentration of 20 mM,
significantly enhanced their resistance to the pathogenic
V. campbellii (Defoirdt et al. 2006) and V. parahaemolyti-
cus (Immanuel et al. 2012). Similarly, Immanuel et al.
(2011) found that medium chain organic acid (caprylic
acid) protected Artemia nauplii from virulent V. harveyi
and V. parahaemolyticus.
More recent research demonstrated that dietary OAB at
20 g kg
1 inclusion level resulted in significant improved
cumulative survival of Pacific white shrimp and tiger
shrimp (Penaeus monodon, Fabricius, 1798) after being
intramuscularly challenged with V. harveyi (Ng et al. 2015;
Romano et al. 2015). Moreover, in the Vibrio challenged
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Dietary organic acids in aquaculture
shrimp, phenoloxidase activity was significantly higher for
shrimp fed the OAB-supplemented diets as well as having
less hepatopancreatic damage. These were the first studies
to show the ability of dietary organic acids to protect and
reduce the structural damage to the hepatopancreatic cells
during a bacterial infection. This is important considering
the fact that the hepatopancreas is the main digestive organ
of crustaceans. We also observed an increase in lipid storing
R cells indicating enhanced nutritional health of the hep-
atopancreas in shrimp fed organic acid-added diets.
Su et al. (2014) reported that the activities of serum phe-
noloxidase, superoxide dismutase and lysozyme in white
shrimp fed 2 or 3 g kg
1 citric acid were significantly
higher compared to the control group. The enhanced
shrimp immune response most likely contributed to the
higher survival of citric acid-fed shrimp during a challenge
with Vibrio alginolyticus. In contrast, with the exception of
serum agglutination titre, Silva et al. (2016) did not find
any significant differences in all immune response parame-
ters measured in white shrimp fed Na-butyrate or Na-pro-
pionate at graded levels up to 20 g kg
1. It was also
interesting to note that the serum antimicrobial titre
against V. alginolyticus did not show any significant differ-
ences between treatments.
These findings show substantial promise, as vibriosis can
contribute to mass mortalities during the grow-out of
shrimp (Lavilla Pitogo et al. 1998; Soto-Rodriguez et al.
2010a,b) and recently had been implicated as a causative
agent of acute hepatopancreatic necrosis syndrome
(AHPNS) or more commonly known as early mortality
syndrome (EMS) that is decimating shrimp production
globally (Tran et al. 2013; Ng 2015; Kondo et al. 2015).
Chuchird et al. (2015) reported that white shrimp cultured
for 30 days in water containing V. parahaemolyticus col-
lected from an EMS-infected farm in Thailand showed
lower gut Vibrio spp. and total bacteria counts in shrimp
fed 3 or 6 g kg
1 formic acid diets. Survival was signifi-
cantly higher in shrimp fed organic acid-added diets com-
pared to the control group. However, dietary formic acid
did not influence growth performance or immune response
of the white shrimp. We are currently conducting more in-
depth research on the efficacy of organic acids in the pre-
vention of AHPNS/EMS in marine shrimp.
Research findings augment well for the use of organic
acids as a viable alternative to harmful antibiotics in com-
mercial aquaculture. Dietary organic acids can be used as a
component of a sustainable fish and shrimp health manage-
ment programme in the aquaculture industry. Organic
acids, as functional feed additives, can impart disease
protective properties to farmed aquatic animals via several
modes of action. The pH modulating and antimicrobial
properties of organic acids can inhibit proliferation of
harmful bacteria while encouraging the growth of beneficial
359
W.-K. Ng and C.-B. Koh
bacterial in the gastrointestinal tract of aquatic animals
which in turn can affect the overall health of the animal.
Dietary organic acids and/or their salts have been shown to
have immunostimulation effects on several species of fish
and shrimp which ultimately lends itself to imparting
greater disease resistance and recovery from infections. Cer-
tain organic acids have been reported to stimulate feed
intake and improve nutrient utilization giving rise to well-
fed fish and shrimp which are then less prone to stress and
pathogen infections. The hepatopancreatic protective prop-
erties of dietary organic acids have also been demonstrated
in shrimp during bacterial infections. These factors, both
singly or in combinations thereof, will contribute to the
reduction and ultimately elimination of the current abuse
and misuse of antibiotics in aquaculture.
Strategies to enhance efficacy of organic acids in
aquafeeds
The use of organic acids in aquafeeds has three major chal-
lenges. Most free organic acids are highly corrosive
(Table 1), some are easily soluble in water, and organic
acids become less effective in an alkaline environment (e.g.
in distal gut). The corrosiveness and pungent smell of most
free organic acids requires special handling, storage and
transportation of these compounds. Other than workers’
safety, free organic acids can also cause corrosive damage to
the aquafeed mill machineries. Leaching of organic acids
from the feed pellets into the surrounding culture water
will be a major issue for aquatic animals. This is especially
challenging for crustaceans since they do not swallow the
feed pellets whole but break up and masticate their feed
before ingesting. In experimental shrimp pellets, Silva et al.
(2013) reported that leaching observed with dietary organic
acid salts was high with Na-acetate exhibiting 100% leach-
ing within 30 min. As previously mentioned, organic acids
are most effective in their undissociated form which enables
them to passively diffuse into bacterial cells. In the stom-
ach, when the pH value is usually lower than the pKa value
of most organic acids (Table 1), organic acids remain
undissociated. However, in the intestinal tract, under more
alkaline conditions, some of the organic acids dissociate
and therefore lower its antibacterial efficacy.
A schematic illustration summarizing the reported bene-
fits, effects and mode of action of dietary organic acids in
aquatic animals as discussed in the previous sections of this
review is given in Fig. 2. Among other factors, a full under-
standing of the antibacterial mode of action of organic
acids is pivotal in designing strategies to enhance the
efficacy of organic acids in aquafeeds. Strategies to reduce
the corrosiveness, leaching and dissociation of organic acids
should be considered when these feed additives are used in
aquafeeds.
360
The corrosiveness of free organic acids can be reduced or
avoided by converting them into their natural salts such as
salts of potassium, sodium and calcium. The efficacy of an
organic acid can be maintained even when part of the free
acid is in its salt form. For example, the commercial potas-
sium diformate (KDF, Aquaformâ) is a double-salt acidi-
fier of formic acid and formate bound to potassium and
had been tested successfully in the feeds of several aquatic
animals (Table 3). This product exists as 35.4% formic
acid, 34.6% formate and 30% potassium in granulated
form (Fig. 3). The organic acid salts will also reduce the
pungent smell and loss from vaporization during pelleting
and storage of the product compared to its corresponding
free acid form.
To reduce corrosiveness and increase ease of mixing with
other dietary ingredients, free organic acids can be
adsorbed onto a solid inert carrier. By processing liquid
organic acids into powder form will also allow them to be
packed, stored and transported more easily. Carrier materi-
als are selected based on their ability to ensure high acid
load adsorbed onto the carrier particles. Special carrier
materials are sometimes used to ensure slow release of free
organic acids throughout the gut for optimal antimicrobial
efficacy. The scanning electron micrograph (SEM) of a
commercial organic acids product (BiotronicTM, BIOMIN,
Getzersdorf, Austria) shows the stacked, latticed, intercon-
necting pores of the special carrier used to adsorb organic
acids (Fig. 4). This product is being marketed as an organic
acid product with slow-releasing capabilities. To increase
the efficacy of organic acids, some manufacturers will add
inorganic acids such as phosphoric acid to the mix so as to
reduce the pH of the product and allow the organic acids
to be in their effective undissociated form further down the
digestive tract. This will add costs to the product as inor-
ganic acids are not inexpensive.
To enhance the efficacy of dietary organic acids, a new
range of organic acids called coated or microencapsulated
organic acids has been developed. Other than increasing
ease of use and eliminating corrosiveness, encapsulation
will eliminate leaching of the soluble organic acids. This is
important for aquatic animals, especially for crustaceans
such as shrimp as they are slow feeders and masticate feed
pellets before ingestion. The SEM of a prototype microen-
capsulated organic acids product (OrgacidsTM-AQUA, Sun-
zen Biotech, Shah Alam, Selangor, Malaysia) is shown in
Fig. 5. Protection of the organic acids may also prevent
their interaction with other ingredients in the feed mix dur-
ing processing. Furthermore, in studies with swine, it had
been reported that lipid microencapsulation enables the
weak acids to be released gradually along the intestinal tract
and reach far into the hindgut where the highest abundance
of bacterial populations is located (Piva et al. 2007). We
have also shown that a similar situation may exist when
Reviews in Aquaculture (2017) 9, 342–368
© 2016 Wiley Publishing Asia Pty Ltd
 Dietary organic acids in aquaculture

Stomach with pH < 3.8 O

DIET + ORGANIC ACIDS: O
C
↓ pH and buffering capacity C
O R O H
↓ harmful bacteria intake R O H
↑ feed hygiene C
R O H

Passive diffusion

O
Dissociates Cytoplasm with pH ≈ 7
C O
R O- H+ H+
STOMACH: O
C
R O-
↓ harmful bacteria H+ ↓↓↓ pH
C Bacteria
↑ enzyme activity R O-
↑ mineral solubility
Interrupts enzymes activity and DNA replication
ATP

H+-ATPase
pump

H+ H+

INTESTINE:
ANTI-BACTERIAL MECHANISM OF ACTION
↑ epithelial cell proliferation
OF ORGANIC ACIDS IN ANIMAL CELLS
↑ nutrient absorption/ digestibility
↑ mineral availability
↑ beneficial microbiota
FECES: ↓ colonization of harmful bacteria
↓ phosphorus load ↑ gut health
↓ microbiota load
POSSIBLE MECHANISM OF ACTION OF
↓ risk of cross infection
DIETARY ORGANIC ACIDS IN AQUATIC ANIMALS
↑ water quality

Figure 2 Mode of action of organic acids against microorganisms. The undissociated organic acids traverse across the cell wall of bacteria via pas-
sive diffusion and dissociate inside the cytoplasm, causing the cytoplasmic pH to decrease. Eventually, the cell enzymes and nutrient transport systems
are suppressed resulting in irreversible damage to the microbial cell, often causing death.

used in the feeds for tilapia (Teoh et al. 2013) and the pro-
totype microencapsulated OAB was successfully field tested
in a large commercial tilapia farm (unpublished data). So
far, the effect of coated or microencapsulated organic acids
has been reported in several aquatic animals, including sea
bream (Robles et al. 2013), common carp (Liu et al. 2014)
and penaeid shrimps (Ng et al. 2015; Romano et al. 2015).
Further research is needed to better understand the benefi-
cial effects of coated/protected organic acids versus
uncoated versions in terms of transporting undissociated
dietary organic acids further along the digestive tract of
various aquatic animals. The encapsulation process is usu-
ally a closely guarded secret by the manufacturers as it
entails very specific technical knowledge in ensuring the
fine particle size of the microcapsules (Fig. 5) and that the
coating material used can be slowly degraded by the aquatic
animal species to allow slow release of the active organic acids
to their target site in the gastrointestinal tract.
Other than blending several different organic acids/salts
together to increase efficacy as previously described,
another trend in acidifier strategies is to combine organic
acids with other feed additives for potential synergistic
effects. In a study with rohu, additions of 30 g kg
1 citric
acid and microbial phytase (500 FTU kg
1) in low protein
diets were found to have synergistic effect on growth per-
formance, nutrient utilization (Baruah et al. 2007b), bone
mineralization (Baruah et al. 2005) and mineral bioavail-
ability (Baruah et al. 2007a). Zhu et al. (2014) reported
that the addition of organic acid (2 or 4 g kg
1) to plant
protein-based diets tended to improved growth perfor-
mance and significantly enhanced antioxidant capacity of
juvenile yellow catfish. However, when organic acid was
used in combination with phytase (1000 FTU kg
1), no
synergistic effect on growth performance was observed and
the addition of 4 g kg
1 organic acid resulted in a signifi-
cant increase in oxidative stress parameters on the juvenile

Reviews in Aquaculture (2017) 9, 342–368

© 2016 Wiley Publishing Asia Pty Ltd 361
W.-K. Ng and C.-B. Koh
Figure 3 Scanning electron micrograph of potassium diformate gran-
ules of a commercial organic acid salt product (Aquaformâ, ADDCON,
Bonn, Germany). Bars = 100 lm.
yellow catfish. Sugiura et al. (2001) noted that the addition
of 50 g kg
1 citric acid to high ash diets (12.3%) contain-
ing both herring fish meal and soya bean meal as main pro-
tein source decreased the activity of phytase, whereas
organic acid markedly increased the effect of the enzyme
with respect to both phytate-P and mineral availability
when added in low ash diets (6.7%) containing soya bean
meal as the sole protein source. It was reported that white
shrimp fed diets with a combination of formic acid and
astaxanthin was no better than using singly or in combina-
tion in terms of preventing V. parahemolyticus infections
(Chuchird et al. 2015). Despite the contradictory reports to
date on the synergistic effects of organic acids and other
Figure 4 Scanning electron micrograph showing the stacked, latticed,
interconnecting pores of the special carrier used to adsorb organic acids
used in the production of an organic acids blend (BiotronicTM, BIOMIN).
This product is marketed as an organic acid product with slow-releasing
capabilities. Bars = 30 lm.
362
Figure 5 Scanning electron micrograph showing the fine spherical
microcapsules of a prototype microencapsulated organic acids blend
(OrgacidsTM-AQUA, Sunzen Biotech). This product is targeting the
shrimp feeds market as it prevents leaching of organic acids.
Bars = 3 lm.
functional feed additives, recent trends indicate that
organic acids manufacturers are combining organic acids
with additives such as essential oils, enzymes and plant/her-
bal extracts for potentially stronger antimicrobial and
health-promoting effects on aquatic animals. More research
is needed to provide a stronger scientific foundation for
such strategies in enhancing the effectiveness of dietary
organic acids in the feeds of aquatic animals.
Conclusion and perspectives
There is currently great interest in the commercial use of
organic acids in aquafeeds, to enhance both growth perfor-
mance and control disease. As evident from research
reviewed, many studies have reported that organic acids,
their salts or mixtures thereof can improve growth, feed uti-
lization, gut health and disease resistance in aquatic ani-
mals. Nevertheless, despite the reported improvement in
nutrient availability of organic acid-supplemented diets in
most studies, contradictory results have been reported for
the growth-promoting effects, which seem to depend on the
aquatic animal species and/or type and dosage of organic
acids tested. The reduction in P and N excretion due to
improved mineral utilization as a result of diet acidification
will greatly promote the formulation of more environmen-
tally friendly aquafeeds. The reduction in the microbial load
of excreted faecal matter from farmed fish fed organic acid-
supplemented feeds will benefit the farming of fish in water
reservoirs and closed recirculating water culture systems.
Increasingly, scientific evidence is being accumulated as to
the positive effects of dietary organic acids on the health of
farmed fish and shrimp, imparting to them increased resis-
Reviews in Aquaculture (2017) 9, 342–368
© 2016 Wiley Publishing Asia Pty Ltd

tance to pathogenic diseases that is commonly encountered
in most modern aquaculture nowadays. However, unlike
terrestrial livestock where a limited number of improved
strains are currently being farmed, aquaculture constitutes a
multitude of combinations of farmed species, feeding prac-
tices and culture systems. This means that research results
on the success or failure in the use of organic acids in one
study might not be applicable to other species of aquatic
animals cultured under different conditions. Based on
research conducted so far, organic acids appear to be a
promising candidate to replace AGP in aquaculture. More
research is needed to fully understand the mechanism of
action of dietary organic acids on the growth- and health-
promoting benefits to cultured aquatic animals for the con-
tinued and sustained growth of the global aquaculture
industry. It is hoped that this comprehensive review will
help researchers conduct more focussed studies in address-
ing the current gaps in knowledge as highlighted through-
out the various sections of this review. It is also hoped that
this review will provide some guidance to aquafeed manu-
facturers in formulating effective functional feeds by using
the right type and dosage of organic acids for each farmed
aquatic animal species.
It is anticipated that the use of dietary organic acids as
functional feed additives in aquafeeds will increase mark-
edly in the foreseeable future. The continuous expansion of
the global aquaculture industry constitutes a sizable market
potential. The intensification of aquaculture systems cou-
pled with global warming is likely to increase the incidence
of disease outbreaks. For example, the recent outbreak of
AHPNS/EMS that has been traced to Vibrio bacteria as the
causative agent has decimated many shrimp farms around
the world with farmers suffering huge economic losses. The
increasingly compelling evidence linking the development
of antibiotic resistance genes in bacteria of aquatic origins
to animal and human pathogens (Miranda & Zemelman
2002; Cabello 2006; Agersø et al. 2007; Reboucßas et al.
2011; Cabello et al. 2013) has already prompted many
governmental and nongovernmental agencies to issue bans
and/or restrictions on the use of antibiotics as growth pro-
motants in many countries. The issue now is one of educa-
tion and enforcement, especially in many developing
nations where a significant amount of aquaculture produc-
tion still comes from small-scale farmers who may be una-
ware of the dangers posed by the excessive use and misuse
of antibiotics. For larger scale producers, other than effi-
cacy, the use of antibiotic alternatives such as organic acids
will also come down to costing. The cost of dietary organic
acids depends on the type of product and the country
where they are sold (due to transport and taxes). In Asia,
the average price of an organic acid product is about
USD2.50 to 5.00 per kg (Encarnacao, pers. comm.). Some
shrimp feed manufacturers find it difficult to consider any
Reviews in Aquaculture (2017) 9, 342–368
© 2016 Wiley Publishing Asia Pty Ltd
Dietary organic acids in aquaculture
feed additive that will cost more than USD20.00 per metric
ton feed unless the shrimp farmer insists and willing to pay
a premium price for the feed (Guerin, pers. comm.). Cur-
rently, organic acids are recommended to be added at 1.5–
5.0 kg per metric ton of aquafeed by various feed additives
companies. The economic window for the incorporation of
organic acids in aquafeeds is therefore narrow. Convincing
farmers and aquafeed manufacturers as to the actual bene-
fits of organic acid supplementation is crucial and this can
only be done through scientifically proven laboratory- and
field-based research data.
Through decades of research and development, the use
of dietary organic acids in the feeds and drinking water of
terrestrial livestock such as swine and poultry is now con-
sidered standard animal husbandry management practice.
The efficacy and cost-effectiveness of dietary organic acids
to livestock farmers are well established. Due to the sheer
number of aquatic animal species being farmed and under
various culture conditions, the path to establishing dietary
organic acids as part of a standard disease management
programme in any aquaculture farm is anticipated to be a
long one but crucial. It is hoped that this review will con-
tribute towards this endeavour.
Acknowledgements
The research grants awarded by various Malaysian funding
institutions that allowed our foray into organic acids
research in aquaculture are acknowledged. These include
the Technofund Project (TF310M172) from the Ministry of
Science, Technology and Innovation, the Knowledge Trans-
fer Program by the Ministry of Higher Education and a
Research University grant (1001/PBIOLOGI/815031) from
Universiti Sains Malaysia. The postdoctoral fellowship pro-
vided by USAINS Holding Ltd (Universiti Sains Malaysia)
through a research grant from Sunzen Feedtech Pte Ltd to
the second author is gratefully acknowledged. We thank
Drs. C. Luckstadt (ADDCON Group GmbH), P. Encar-
nacao (formerly BIOMIN Singapore Pte Ltd) and PC Kok
(Sunzen Biotech Ltd) for the potassium diformate, Biotro-
nicTM and OrgacidsTM-AQUA samples, respectively.
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