Available online at www.sciencedirect.
com
ScienceDirect
Recent advances in single cell protein use as a feed
ingredient in aquaculture
Shawn W Jones1, Alon Karpol2, Sivan Friedman2,
Biniam T Maru1 and Bryan P Tracy1
The global demand for high-quality, protein-rich foods will
continue to increase as the global population grows, along with
income levels. Aquaculture is poised to help fulfill some of this
demand, and is thus the fastest growing animal protein
industry. A key challenge for it, though, is sourcing a
sustainable, renewable protein ingredient. Single cell protein
(SCP) products, protein meals based on microbial or algal
biomass, have the potential to fulfill this need. Here, we review
potential sources of SCP strains and their respective
production processes, highlight recent advances on
identification of new SCP strains and feedstocks, and, finally,
review new feeding trial data on important aquaculture species,
specifically Atlantic salmon, rainbow trout, and whiteleg
shrimp.
Addresses
1
White Dog Labs, Inc., 239 Lisa Dr., New Castle, DE 19720, USA
2
White Dog Labs, Israel Prof. A.D. Bergman St. 2b, Rehovot, 7670504,
Israel
Corresponding author: Tracy, Bryan P (btracy@whitedoglabs.com)
Current Opinion in Biotechnology 2020, 61:189–197
This review comes from a themed issue on Food biotechnology
Edited by Mark Blenner and Jan-Peter van Pijkeren
For a complete overview see the Issue and the Editorial
Available online 25th January 2020
https://doi.org/10.1016/j.copbio.2019.12.026
0958-1669/ã 2020 The Author(s). Published by Elsevier Ltd. This is an
open access article under the CC BY license (http://creativecommons.
org/licenses/by/4.0/).
Introduction
Seafood, wild-catch and aquaculture, is the largest animal
protein industry in the world (189 million tons in 2013 -
Figure 1a) [1]. Wild-catch tonnage has been stable since
1990 at 90 million tons; thus, aquaculture accounts for
all growth. In fact, aquaculture has grown faster than any
animal protein sector at 7% compound annual growth
rate over past two decades compared to 4% for poultry
[1]. Feed is a major cost in livestock production, and
protein ingredients particularly dominate aquaculture
feed cost. Thus, efficient feed and particularly protein
conversion is essential to manage production costs and
improve the sustainability of aquaculture. Fortunately,
aquaculture species exhibit low feed conversion ratios
(FCRs) of 1.1–1.6 kg of feed per kg of edible seafood
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versus 1.4–1.8 for poultry, 2.6–4.4 for pork, and 3.5–9 for
beef. Consequently, aquaculture contributes to a more
sustainable animal protein industry, and as we review,
single cell protein (SCP) is poised to play a major role in
its future success.
Aquaculture diets generally have higher crude protein
content, ranging from 35 60 wt%, compared to terres-
trial livestock at 12 26 wt%. Moreover, many aquacul-
ture markets are carnivorous species, whereas most
terrestrial livestock are herbivores. Thus, plant-based
protein ingredients are better suited to terrestrial live-
stock than many aquaculture species. Historically, fish-
meal (i.e. ground, dried forage fish and/or fish trimmings
and waste) has been the preferred protein ingredient in
aquaculture because of high crude protein, high essen-
tial amino acid (EAA) and ‘completeness’ of the ingre-
dient. However, aquaculture competes with fishmeal
use in swine, poultry, and other animal diets, and fish-
meal production cannot scale with growth of all these
industries without jeopardizing forage fish fisheries.
Even with diminishing inclusion of fishmeal in aqua-
culture feeds (Figure 1b), an estimated shortage ranging
from 0.4 to 1.32 million metric tons of fishmeal could
occur by 2050, significantly impairing aquaculture
industry growth. Plant-based ingredients can be refined
to improve compatibility with aquaculture diets, for
example, removing anti-nutritionals like phytic acid,
but this increases cost and most ‘early’ wins have been
accomplished for species with greatest nutritional
knowledge, for example, salmonids. Thus, there is
demand for more suitable protein ingredients that main-
tain feed performance, benefit aquaculture health and
stabilize supply and economics during industry expan-
sion. SCP has the potential to deliver multiple solutions
through a myriad of products and production
approaches, but considerable research, development
and particularly scale-up is still required. Here, we
review recent developments in SCP production modali-
ties and feed trials, in addition to a brief introduction on
organisms and feedstocks.
SCP organisms
SCP products can be prepared from different microbial
sources, including microalgae, yeast and other fungi, and
bacteria (Table 1). All are actively being investigated and
commercialized and exhibit unique advantages and
challenges.
Current Opinion in Biotechnology 2020, 61:189–197
190 Food biotechnology

Figure 1

(a)
500

Total
400 Seafood
Million Metric Tons

300

200

100

0
1990 1995 2000 2005 2010 2015 2020 2025
Beef Pork Poultry Wild-catch Seafood Aquaculture Seafood

(b)
3.5
6.0%

Fishmeal Inclusion (wt%)

3.0
5.0%
Fishmeal (MMT)

2.5
4.0%
2.0

1.5 3.0%

1.0 2.0%

0.5 1.0%

0.0 0.0%
2000 2005 2010 2015 2020 2025 2030 2035 2040 2045 2050

Fishmeal Consumption in Aquaculture Fishmeal Inclusion Level

Fishmeal Inclusion Level (Scenario 1) Fishmeal Shortage (Scenario 1)
Fishmeal Inclusion Level (Scenario 2) Fishmeal Shortage (Scenario 2)
Current Opinion in Biotechnology

Growth of aquaculture industry and potential shortage of fishmeal. (a) Total animal protein production in a million metric tons from 1990 to
2025 [62]. (b) Fishmeal consumption for aquaculture applications through 2015 and projected consumption through 2050 [62] is presented (blue
line). Average fishmeal inclusion (green line) from known data through 2015 and projected growth of aquaculture through 2050 [62] with the
assumed flat supply of fishmeal. Inclusion is calculated by dividing the fishmeal consumption (blue line) by total feed, which is calculated by
multiplying aquaculture output (Figure 1a) by an average feed conversion ratio (FCR) of 1.2. Two alternative scenarios, in which fishmeal inclusion
levels do not achieve targeted reductions, are considered: Scenario 1 (solid red) and Scenario 2 (dashed red). Scenario 1 assumes only 85% of
the reduction target is met, and Scenario 2 assume only 50% of the reduction target is met.

Microalgae acid (EPA) and docosahexaenoic acid (DHA)) [3] and

As recently reviewed by Tibbetts [2], microalgae can be carotenoids, namely astaxanthin — an antioxidant, immu-
an SCP source with relatively high crude protein content. nostimulant [4,5] and pink pigment for salmon and
However, interest seems more motivated by potential shrimp [6] for example by the strain Aurantiochytrium
production of omega-3 fatty acids (i.e. eicosapentaenoic Sp. [7].

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 Single cell protein advances for aquaculture use Jones et al. 191

Table 1

Summary of SCP sources and current examples

SCP Protein content Special characteristics Example of specific Challenges Current commercial
sources range organisms activities
Microalgae 60–70% [2,10]  Phototrophic growth  Chlorella vulgaris  Economical scale-up  Cellana
 Production of  Desmodesmus sp.  Cell disruption to  Pond Technologies
omega-3 fatty acids release nutrients  BioProcess Algae

Yeasts 30–50% [10]  Use of a  Saccharomyces  Improve protein  ADM

variety of feedstocks cerevisiae and EAA content  Alltech
 Production of  Candida utilis  Flint Hills
vitamins and Resources
micronutrients  ICC Brazil
 Pacific Ethanol

Bacteria 50–80% [10]  High protein content  Methylococcus  Palatability issues  Calysta
 Growth on C1 capsulatus  Kiverdi
substrates  Cupravidus nectar  KnipBio
 NovoNutrients
 White Dog Labs

Protists 10–20% [63,64]  Production of  Schizochytrium  Improve protein  Veramaris

omega-3 fatty limacinum content
acids

Yeast and other fungi Production processes

Yeasts and fungi have long histories as animal feed
ingredients, particularly for terrestrial livestock and direct
human consumption, like Quorn. The most widely
known species are Saccharomyces cerevisiae, various Asper-
gillus and Fusarium venenatum, but other strains are attract-
ing interest for protein replacement such as Candida utilis
and Kluyveromyces marxianus [8]. In most cases though, the
aim is to deliver additional benefits such as Rhodotorula
mucilaginosa [9] biomass exhibiting immunomodulation
and antioxidant benefits, and Yarrowia lipolytica produc-
ing EPA, which was previously commercialized by
DuPont and AquaChile as Verlasso salmon [3].
Bacteria
Bacterial SCP strains can produce very high crude protein
content (>80 wt%) and EAA values, along with vitamins,
phospholipids, and other functional compounds and can
also utilize a wide-range of feedstocks [10,11,12]. As we
detail later, feedstock diversity has motivated deeper
exploration of bacteria, largely those that utilize methane,
methanol, syngas, CO2 and H2, and second generation
sugars.
Protists
Previously classified as microalgae or fungi, Labyrinthulo-
mycetes are actually heterotrophic marine protists [13] and
constitute three different categories: thraustochytrids, apla-
nochytrids, and labyrinthulids. Thraustochytrids, like
Schizochytrium limacinum (Aurantiochytrium limacinum), are
of particular interest because of their ability to produce
omega-3 fatty acids [14,15].
The goal of SCP production is to maximize cellular
growth and co-product yields in economically viable
approaches, and the feedstock has a tremendous, if not
the greatest, influence on economics. A sampling of
recent research into various feedstock use is summa-
rized in Table 2, and includes general advantages and
disadvantages. We recommend Ritala et al. [10] for a
more comprehensive review on various feedstock
usage. Our updated survey of the literature, since
2017, confirms strong interest to utilize ‘wastes,’ resi-
dues and by-products, as they are presumed to support
a more circular economy and lower cost. However,
practical experience urges deeper consideration on
capital and processing costs to control and upgrade
such feedstocks, along with the potential regulatory
challenges, and constrained supply both in quantity
and geography. Worth noting, recent trends continue
to promote methane, syngas and off-gas feedstocks
such as the classic methane to SCP approach [18] or
knallgas bacteria [61]. Also, there is an emerging
opportunity to better valorize low-value co-product
streams from dry-grind corn ethanol plants, like the
evaporated solubles [27].
Such feedstock diversity requires diverse modalities
of growth including autotrophs, photoautotrophs,
chemoautotrophs, methylotrophs, heterotrophs and
mixotrophs. Consequently, many different production
processes are being explored (Table 3), and we review
some of the recent advances and current areas of
research focus.

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192 Food biotechnology

Table 2

Potential feedstocks for SCP production

Feedstock sources Examples Advantages Disadvantages

Wastewaters  Municipal waste [28]
 Industrial wastewater [65]
 Potential for tipping fees
 Most circular economy appeal
 Limited control or significant
processing
 Supply constrained and
defined in scale
 Major regulatory challenges

Industrial & Agricultural  Off-gases (e.g. steel mill)
Residues  Biogas [66]
 Agricultural wastes
[22,26,29,67]
 Cellulosic biomass
 Lowest cost  Less control or significant processing
 Enhanced valorization  Supply constrained and defined in scale
 Bio-potential  Significant regulatory challenges
 Circular economy appeal

Bioindustry  Brewery residues [24]  Lower cost  Limited supply

By-Products  Dry-grind ethanol  Enhanced valorization  Processing often required
co-products  Large quantity point sources  Variable composition
 Starch processing waters  Established processing and  Output of other processes that
 Biogas infrastructure could change
 CO2 [68]  Regulatory challenges

Fit for Purpose  Dextrose  Low cost  Higher comparable cost

 Sucrose  Best defined  More limited to locations with
 Corn starch  Most controlled developed supply logistics
 Molasses  Abundant and expandable supply  Less circular economy appeal
 Soybean meal  More regulatory certainty
 Methane  More conversion options
 Methanol  Developed infrastructure
 Ethanol
 Syngas
 H2 + CO2

Photoautotrophs known as the Wood-Ljungdahl pathway), methanogens

In this modality, growth utilizes inorganic carbon sources
and light as the energy source. The major feedstocks are
CO2, water, and light, making this a desirable platform
from a renewable, sustainable point-of-view, but several
hurdles must be overcome for broader commercialization.
First is maintaining a high surface area to allow for light
penetration into the bioreactor, second is achieving high
gas flow rates into the bioreactors and third is promoting
higher cell densities to reduce water volumes. Yen et al.
[16] present comprehensive updates, beyond the scope
of this review, on where the technology stands, how it is
being commercially deployed and where the challenges
persist. Microalgae are the primary focus of this modality,
but some bacteria, particularly purple photosynthetic
bacteria (PPB) [17], are also capable of photoautotrophic
growth.
Chemoautotrophs
In this modality, an inorganic carbon source is required
along with an inorganic energy source. Bacteria mainly
constitute this modality, though there is great diversity on
how growth is achieved. For SCP production, there are
three main categories: acetogens which utilize CO2/CO/
H2 through the reductive acetyl-CoA pathway (also
which convert CO2/H2 to methane, and hydrogen-oxidiz-
ing bacteria (also known as knallgas bacteria), which
oxidize H2 with O2 as the final electron acceptor and
fix CO2 through different pathways, such as the Calvin-
Benson-Bassham cycle or the reverse Krebs cycle. Similar
to photoautotrophs, chemoautotrophs are attractive
because of the relatively inexpensive feedstock and
ability to fix CO2. There are several significant challenges
though including: 1) solubilizing the gases, particularly
H2, 2) achieving sufficient growth rates to realize ade-
quate volumetric productivity, and 3) obtaining high cell
mass yields. Acetogens and methanogens must produce
other carbon metabolites as the final electron acceptors,
such as acetate or butyrate for acetogens and methane
for methanogens, so these co-products must also be
valorized. Some commercial progress has been made
on hydrogen-oxidizing bacteria, for example, Cupravidus
nectar and related organisms are being pursued by Novo-
Nutrients (Sunnyvale, CA), Solar Foods (Helsinki,
Finland), and Kiverdi (Hayward, CA).
Methylotrophs
Methane utilizers and some methanol utilizers do not
easily fit into either the traditional autotroph or

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 Single cell protein advances for aquaculture use Jones et al. 193

Table 3

SCP production processes

Cultivation Operation Growth modality Capital & operational considerations Emerging commecial examples
Aerobic Bioreactor  Heterotrophs  High cell mass yield  Methanol, glycerol or ethanol –
 Mixotrophs  High capital costs KnipBio
 High energy consumption  Glucose – Veramis
 Sterile operation  Cellulose – Arbiom & Menon
 Significant installed industrial capacity

Anaerobic Bioreactor  Heterotrophs  Low cell mass yield  Glucose or glycerol – White Dog
 Mixotrophs  Low capital costs Labs
 Low energy consumption  Yeast sepatiation – Fluid Quip
 Requires metabolite production and valorization Technologies & ICM
 Non-sterile operation
 Most installed industrial capacity

Gas Bioreactor  Methylotrophs  Variable cell mass yield  Methane – Calysta, Unibio, &
 Chemoautotrophs  High capital costs String Bio
 Mixotrophs  High energy consumption  CO2, H2 & O2 – Kiverdi, NovoNu-
 Could require metabolite production and trients & Solar Foods
valorization  CO2 & H2 – LanzaTech
 Sterile & non-sterile operation  Glucose & syngas – White Dog
 Limited installed industrial capacity Labs

Photosynthetic Bioreactor  Photoautotrophs
 Mixotrophs
 High cell mass yield  CO2 & light – BioProcess Algae &
 High capital costs Pond Technologies
 High energy consumption
 Sterile operation
 No known installed industrial capacity

Open Cultivation Systems  Photoautotrophs  Variable cell mass yield  Brewing by-products – iCell Sus-
 Heterotrophs  Low capital costs tainable Nutrition
 Mixotrophs  Low energy consumption  Open photosynthetic system –
 Non-sterile operation Cellana
 Limited installed industrial capacity

heterotroph category, as they consume an organic C1
feedstock both for carbon and energy. Carbon fixation
can occur through a number of different pathways, such as
the ribulose biphosphate cycle, the ribulose monopho-
sphate cycle or the serine cycle, but the key intermediate
is formaldehyde, made by oxidation of the methane or
methanol. Similar to chemoautotrophs, a major technical
hurdle is dissolving the feedstocks methane and oxygen
into an aqueous broth. Another significant challenge is
heat removal from the fermentation as this approach is
very exothermic.
Methylotrophs have an established commercial history.
The first bacterial SCP product was Pruteen, made from
Methylophilus methylotrophus by Imperial Chemical Indus-
tries (ICI) from a methanol feedstock from the late 1970s
through the mid-1980s [18,19]. This process was revived
in the 1990s by Norferm, a Statoil subsidiary, using
Methylococcus capsulatus and a methane feedstock to pro-
duce Pronin. This product was targeted for aquaculture
applications, and a joint venture with DuPont produced a
strain that also produced carotenoids and specifically
targeted astaxanthin. However, economics again caused
this process to be canceled. Recently, this process has
been revived by several companies: Unibio (Lyngby,
Denmark) making UniProtein1 from methane, Calysta
(Menlo Park, CA) making FeedKind1 from methane,
String Bio (Bangalore, India), and KnipBio (Lowell, MA)
making KnipBio Meal from methanol. KnipBio engineers
their microbes to also make carotenoids [20] and taurine
[21], and announced plans to scale-up with ICM (Col-
wich, KS). It is important to note that KnipBio Meal, to
the best of our knowledge, is the first genetically engi-
neered SCP product to obtain GRAS (generally regarded
as safe) recognition by the US Food and Drug
Administration.
Heterotrophs
Heterotrophs must consume organic carbon as a feed-
stock and encompass all possible SCP sources. Arguably
the most advanced process scheme, a wide range of
feedstocks can be used to achieve SCP production, giving
this modality a great deal of flexibility. Recent research
interest has largely focused on waste or cellulosic

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194 Food biotechnology
feedstock utilization that is well discussed elsewhere
[22–26,27,28–31], while current commercial products
have largely focused on valorizing biomass from other
biological processes, most notably corn ethanol processes.
Several S. cerevisiae SCP products are on the market today
including LysCell1 by ICC Brazil, Aventine YeastTM by
Pacific Ethanol, NuPro1 by Alltech, and ProPlex1 DY
by ADM, though most are targeted for high-value appli-
cations in terrestrial livestock. Recently, an enhanced
Dried Distillers’ Grain (DDG) product, marketed as
NexPro1 by Flint Hills Resources, has been generated
using Fluid Quip Technologies’ Maximized Stillage Co-
Products (MSCTM) technology. MSCTM mechanically
separates a protein fraction from the solubles (i.e. partially
concentrated thin-stillage) from dry-grind ethanol opera-
tions, which is effectively enriched for the ethanol yeast
cell mass. Marketing literature by Flint Hills Resources
shows NexPro1 can be used on several aquaculture
species such as tilapia, Pacific white shrimp, and rainbow
trout. Similar to these products, ProPlex1 T is a bacterial
SCP by-product of ADM’s threonine fermentation pro-
cess. Recently, a dedicated sugar to SCP process was
announced by White Dog Labs (New Castle, DE) to
produce a SCP product called ProTytonTM [32]. This
process largely uses the dry-grind ethanol process to
produce a SCP product along with a co-product called
MiruTytonTM.
Mixotrophs
Many of the autotrophic modalities can also be operated
in a mixotrophic mode where both autotrophic and het-
erotrophic modalities are utilized by the same microbe in
the same process. This is most common for microalgae,
where in addition to CO2 and light, sugars can also be
added to further boost growth and biomass production
rates and yields. For example, the cell mass yields for
Spirulina and Chlorella sp. in a mixotrophic production
mode are higher than in pure photoautotroph mode [33].
Mixotrophy can also be applied to chemoautotrophs [34],
though this has been mainly in the context to improve
metabolite yields rather than cell.
SCP feeding trial results
Extensive feeding trials are necessary to validate new
products, and aquaculture consists of a wide variety of
species and growth conditions. For this review, we
primarily focus on species of greater commercial and
nutritional knowledge — whiteleg shrimp (Litopenaeus
vannamei), Atlantic salmon (Salmo salar), and rainbow
trout (Oncorhynchus mykiss). In most trials, the goal was
to evaluate growth and feed conversion when fishmeal is
replaced by the SCP product, but trials also investigate
broader nutritional effects.
Microalgae
In regards to commercial interests, Cellana tested both a
defatted Desmodesmus sp. SCP product and an algal meal
Current Opinion in Biotechnology 2020, 61:189–197
derived from Nanofrustulum and Tetraselmis on salmon. In
the first study, defatted Desmodesmus sp. SCP could be
included up to 20% of the feed without any adverse effect
on salmon growth or the final composition [35], and in the
second, the algal meal could be included at 10% without
affecting the performance [36]. For rainbow trout, a
Spirulina algal meal could be incorporated at 10% of
the diet without an effect on performance [37]. In another
trout study, a biomass of Scenedesmus almeriensis could
replace up to 40% of fishmeal in the diet, though growth
was on the lower end of acceptable performance [38]. A
trial by Gamboa-Delgado et al. on shrimp, testing differ-
ent ratios of Spirulina (Arthrospira platensis), Nannochlor-
opsis ocultata, and fishmeal, showed that Nannochloropsis
ocultata was a poor fishmeal replacement ingredient [39].
Nutrient accessibility (i.e. digestibility) for algal meals
can be an issue because of the rigidity of the cell wall [40
]. To improve the digestibility of these meals [41], the cell
wall can be disrupted by a variety of methods [40,42]:
enzymatic (such as cellulases), chemical (such as organic
solvents or acids), and physical and mechanical (such as
bead milling, high-pressure homogenization or microflui-
dics). Physical and mechanical methods are generally
preferred [42,43], as enzymatic and chemical methods
can impact intracellular nutrients. In a recent study, a
microfluidizer was used on a Chlorella vulgaris meal and
then fed to salmon [40]. The whole-cell meal and the
cell-ruptured meal had the same nutrient and protein
quality, but the cell-ruptured meal had improved digest-
ibility over the whole-cell meal for EAAs, fats, and
carbohydrates. Interestingly, starch digestibility was
actually higher than the control diet.
Yeast and other fungi
Salmon and shrimp have been the major focus of recent
yeast feeding trials. For salmon, several different yeast
meals have been tested: S. cerevisiae, Candida utilis, and
K. marxianus [8,44]. S. cerevisiae was found to be a poor
protein meal, while C. utilis and K. marxianus could
replace up to 40% of the fishmeal in diets without
effecting growth performance or nutrient retention [8].
In a follow-up study, the researchers investigated whether
C. utilis could overcome soybean meal induced enteritis, a
common side effect in carnivorous aquaculture species,
but when fed a diet consisting of 40% soybean meal and
up to 20% C. utilis meal (replacing wheat gluten and
starch), the fish still displayed signs of enteritis [44].
For shrimp, several S. cerevisiae products were successful
in replacing fishmeal (from 15 to 24%, depending on the
product) or soybean meal (up to 24%) in shrimp diets with
no effect on growth performance [45–47]. NexPro1
product was also successful in replacing up to 20% of
corn protein concentrate [46]. Finally, shrimp fed a diet of
about 50% C. utilis had no adverse side effects, and
actually displayed higher growth rates compared to a
complete fishmeal diet [48]. So far, these studies indicate
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 Single cell protein advances for aquaculture use Jones et al. 195
C. utilis is a better SCP source than S. cerevisiae for salmon
and shrimp diets. Another fungi tested as SCP source (as
well as unsaturated fatty acids, beta-glucan, and mannane
polymers) is Yarrowia lipolytica. The strain was tested on
shrimp [49] and salmon [50] and found to support increase
fish weight as the control.
Bacteria
Several different methanotroph-based SCP products have
been tested successfully on Atlantic salmon. In one study,
salmon fed a diet containing up to 36% bacterial protein
meal (BPM) displayed a higher growth rate and feed
efficiency ratio than the control diet, though nutrient
digestibility was reduced [51]. In another study, the
authors found that M. capsulatus SCP can make up to
52% of the dietary protein in a salmon diet and 38% in a
trout diet with no adverse growth effects [52]. Interest-
ingly, the inclusion of M. capsulatus SCP in a diet with
soybean meal prevented the development of soybean
meal-induced enteritis in salmon, suggesting further ben-
efits of SCP products [53]. Finally, KnipBio Meal (Methy-
lobacterium extorquens) could replace 55% of fishmeal in
salmon diets with no adverse side effects upon growth
[54] and could replace up to 10% of soybean meal in
trout diets [55]. Shrimp have also been a key target for
bacterial SCP. Inclusion of a novel mixture of two purple
non-sulfur bacteria at 1% of a diet gave modest growth
improvements over the control [56], a Corynebacterium
ammoniagenes SCP could replace 10–20 % [57], and Knip-
Bio Meal was found to be able to completely replace
fishmeal in shrimp diets [54]. A biofloc meal, prepared
from biofloc grown on shrimp farm tanks, could replace up
to 30% of fishmeal in diets [58]. A microbial biomass
mixture of bacteria and microalgae (marketed at
NovacqTM, CSIRO Canberra, Australia) has been exten-
sively tested on black tiger shrimp (Penaeus monodon).
One study suggests it can overcome the growth disad-
vantages when fishmeal and fish oil are removed from the
prawns’ diet [59], and another shows improved growth
rates when Novacq is included at 10% of the diet [60].
These studies are highly encouraging of SCP’s role in the
growing aquaculture industry. Not only do they offer
another high-quality protein ingredient applicable across
a wide range of species, but several studies suggest
further health benefits to the fish and shellfish.
Conclusions
The demand for animal protein continues to increase
along with the world’s population, and aquaculture is
poised to play an ever-larger role in fulfilling this need.
A key challenge for the industry though is sourcing a
sustainable, renewable high-protein ingredient. Cur-
rently, fishmeal, along with terrestrial plant meals, makes
up most of the protein content of diets. The production of
fishmeal cannot scale with the growth of the aquaculture
industry without having profound and lasting impacts on
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the health of the oceans, and often terrestrial plant meals
lack the EAAs required by many aquaculture species and
can contain antinutritional compounds, like phytic acid.
SCP-based protein meals have the potential to provide
the industry a sustainable, renewable feed ingredient to
make up for the deficiencies of plant-based meals and
reduce the need for fishmeal in diets. Long held as a
promising technology, SCP meals are actually now being
commercially produced by a number of companies and,
even more promising, is the ever increasing positive
feeding trial data on important aquaculture species,
including salmon, trout, and shrimp. These data clearly
show the positive effects of SCP inclusion in diets and
point to SCP meals become true commodity products.
While there are still challenges in scale-up, SCP proces-
sing, and the economics of a commodity product, the
progress made in the past several years, in regards to
finding new strains, developing new processes, and suc-
cessful testing on fish species, is highly encouraging for
SCP products.
Conflict of interest statement
All authors are full-time employees of White Dog Labs,
Inc., which is a biotechnology-based company that is
actively commercializing fermentation solutions for the
production of animal health and nutrition products.
CRediT authorship contribution statement
Shawn W Jones: Conceptualization, Data curation, Writ-
ing - original draft, Writing - review & editing, Supervi-
sion. Alon Karpol: Conceptualization, Data curation,
Writing - original draft, Writing - review & editing,
Supervision. Sivan Friedman: Conceptualization, Data
curation, Writing - original draft. Biniam T Maru:
Conceptualization, Data curation, Writing - original draft.
Bryan P Tracy: Conceptualization, Methodology, Data
curation, Writing - original draft, Project administration,
Funding acquisition.
Acknowledgements
This research did not receive any specific grant from funding agencies in
the public, commercial, or non-for-profit sectors.
References and recommended reading
Papers of particular interest, published within the period of review,
have been highlighted as:
 of special interest
 of outstanding interest
1. Ritchie H, Roser M: Meat and Seafood Production and
Consumption. 2019.
2. Tibbetts SM: The potential for’ Next-Generation’, microalgae-
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technical challenges of methane and oxygen fermentation, which led to
the pivot decision towards methanol fermentation, which was fully
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Current Opinion in Biotechnology 2020, 61:189–197
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isms to produce single-cell protein and related products from dry-grind
ethanol thin stillage concentrated solubles. This study demonstrates the
potential to further valorize some of the world’s largest, point source
feedstocks, which will continue to be an emerging opportunity for the
global ethanol industry.
28. Saejung C, Thammaratana T: Biomass recovery during
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 Single cell protein advances for aquaculture use Jones et al. 197
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38. Tomas-Almenar C, Larran ?am?, de Mercado E, Sanz-Calvo MA,
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39. Gamboa-Delgado J, Navarro YIM, Nieto-Lopez MG, Villarreal-
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juvenile Atlantic salmon (Salmo salar L.) diets containing
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A good example of considering additional post-processing requirements
to optimize nutrient availability of single-cell protein biomass for feed
applications.
41. Teuling E, Wierenga PA, Agboola JO, Gruppen H, Schrama JW:
Cell wall disruption increases bioavailability of
Nannochloropsis gaditana nutrients for juvenile Nile tilapia
(Oreochromis niloticus). Aquaculture 2019, 499:269-282.
42. Lee AK, Lewis DM, Ashman PJ: Disruption of microalgal cells for
the extraction of lipids for biofuels: processes and specific
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distiller’s dried grain product to replace corn protein concentrate
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our knowledge) genetically engineered SCP product to obtain GRAS
(generally regarded as safe) recognition by the US Food and Drug
Administration. This sets a significant precedent for the advancement
of additional, genetically engineered feed ingredients for aquaculture.
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60. Arnold S, Smullen R, Briggs M, West M, Glencross B: The
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65. Rasouli Z, Valverde-Perez B, D’Este M, De Francisci D,
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66. Khoshnevisan B, Tsapekos P, Zhang Y, Valverde-Perez B,
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A good example of a multifermentation approach with well conducted
research to combine multiple modalities of fermentation for the produc-
tion of single-cell protein.
Current Opinion in Biotechnology 2020, 61:189–197