An Overview of Microalgae Biomass As A Sustainable

BIOENGINEERED
2022, VOL. 13, NO. 4, 9521–9547

https://doi.org/10.1080/21655979.2022.2061148
REVIEW
An overview of microalgae biomass as a sustainable aquaculture feed

ingredient: food security and circular economy
Ashfaq Ahmad , Shadi W. Hassan , and Fawzi Banat
Department of Chemical Engineering, Khalifa University of Science and Technology, Abu Dhabi, United Arab Emirates
ABSTRACT ARTICLE HISTORY

Sustainable management of natural resources is critical to food security. The shrimp feed and Received 18 February 2022
fishery sector is expanding rapidly, necessitating the development of alternative sustainable Revised 24 March 2022
components. Several factors necessitate the exploration of a new source of environmentally Accepted 25 March 2022
friendly and nutrient-rich fish feed ingredients. Microalgal biomass has the potential to support KEYWORDS
the growth of fish and shrimp aquaculture for global food security in the bio-economy. Algal Aquaculture; microalgae;
biorefineries must valorize the whole crop to develop a viable microalgae-based economy. aquafeed; biochemical
Microalgae have the potential to replace fish meal and fish oil in aquaculture and ensure composition; sustainability;
sustainability standards. Microalgae biomasses provide essential amino acids, valuable triglycer bio-economy
ides such as lipids, vitamins, and pigments, making them suitable as nutritional supplements in
livestock feed formulations. Fish and microalgae have similar nutritional profiles, and digestibility
is a critical aspect of the aquafeed formulation. A highly digestible feed reduces production costs,
feed waste, and the risk of eutrophication. Due to low input costs, low carbon footprint, waste
water treatment benefits, and carbon credits from industrial CO2 conversion, microalgae-based
fish and shrimp feeds have the potential to provide significant economic benefits. However,
several challenges must be addressed before microalgal biomass and bioproducts may be used
as fish feeds, including heavy metal bioaccumulation, poor algal biomass digestion, and antinu
trient effects. Knowledge of biochemical composition is limited and diverse, and information on
nutritional value is scattered or contradictory. This review article presents alternative approaches
that could be used in aquaculture to make microalgal biomass a viable alternative to fish meal.
1. Introduction
food consumption [1,2]. Fish meals, which are
Aquaculture has become increasingly important high in protein, are an excellent source of nutri
for food security in the 21st century. By 2050, the ents for fish and shrimp in aquaculture systems
global population is projected to increase from 7.6 [3,4]. However, volatility in the global market for
to 9.8 billion, resulting in a 60 to 100% increase in fish meals has harmed long-term revenue and
CONTACT Ashfaq Ahmad ashfaq.ahmad@ku.ac.ae;fawzi.banat@ku.ac.ae; Fawzi Banat Department of Chemical Engineering, Khalifa University of
Science and Technology, Abu Dhabi 127788, United Arab Emirate
© 2022 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted
use, distribution, and reproduction in any medium, provided the original work is properly cited.
9522 A. AHMAD ET AL.
security in the aquaculture sector [3,5]. In recent PUFA) was observed in fish fed 5% Rosamin
years, the gradual exhaustion of marine fisheries [12]. Therefore, a plant-based fish meal has
resources has posed a serious problem for fish become an increasingly popular option in recent
meal production [6]. In China, for example, mar years.
ine capture increased more than 20-fold from 0.6 Improved fish nutrition can reduce feed
million tons in 1950 to 13.6 million tons in 2011. waste, resulting in enhanced financial sustain
Additionally, periodic closures and efforts to man ability. A diet rich in omega-3 fatty acids, anti
age fishing capacity in the offshore ocean have oxidants, and prebiotics has been shown to
significantly reduced fisheries productivity, increase the production, duration, and quality
thereby limiting fish meal production [7]. Finally, of farmed fish [13]. Furthermore, supplementing
industrialization and urbanization-related marine animal feed with algae improves growth and
pollution contaminate fish products and affect the weight gain, decreases feed consumption,
safety of fish meals [8]. Aquaculture is one of the increases immunological response, resistance to
fastest-growing sectors of the food industry. In illness, antibacterial and antiviral activity, and
2019, the aquaculture industry was expected to enriches livestock products with bioactive com
be valued at US$ 31.94 billion [9]. The aquaculture ponents [14]. Microalgae-related biotechnologies
industry is expected to grow at a rate of more than and bioproducts have been rapidly developed in
7.1% between 2020 and 2027. Increased human recent years. Research in microalgae has pre
consumption and commercial acceptability are viously focused on improving biomass harvest
now driving the growth of the aquaculture ing efficiency and producing specific high-value
industry. compounds in algal cells [15,16]. The rapid
In recent years, several new species have been growth of the algal bio-economy has been driven
introduced into the industry. Aquaculture has by significant advances in algal biotechnology
grown faster than other major food industries in that have turned algae into an efficient ‘cell
recent years due to increased fish consumption. It factory’ for food production [14]. The cost of
has been demonstrated that a combination of two microalgal feed remains higher than that of con
microalgal species, Nannochloropsis oculata, and ventional feed. The cost of microalgal feed must
Schizochytrium sp., can be used to feed Nile tilapia come down to be competitive. Algal biotechnol
(Oreochromis niloticus), the second-largest farmed ogy is closely related to the growth of the algal
fish in the world. This study demonstrated that bio-economy in terms of food and feed produc
microalgae-based feeds can enhance nutritional tion. Algal biotechnology focuses on increasing
quality and fish growth metrics [10]. In another algal productivity to reduce the cost of biomass
study, Nannochloropsis sp. and Isochrysis sp. were production. Several recent biotechnological
used to substitute fish meals and fish oil in the diet approaches have resulted in increased biomass
of rainbow trout (Oncorhynchus mykiss), a key production and accumulation of useful metabo
model species for salmonid aquaculture. lites . These include bioreactor design, produc
Compared to fish meal and fish oil, Isochrysis sp. tion of genetically modified strains, high-
was found to significantly increase apparent throughput screening, rapid sampling, and
digestibility coefficients for crude proteins, amino genetic and metabolic engineering. Microalgal
acids, lipids, and fatty acids [11]. Several commer biotechnology focuses on improving the produc
cially available algae products, including Verdemin tion of carbohydrates, proteins, polyunsaturated
(Ulva ohnoi) and Rosamin (Entomoneis spp.), were fatty acids, pigments, and other valuable nutri
evaluated as potential feed components for ents from microalgae through strain optimiza
Atlantic Salmon (Salmo salar). Verdemin and tion, carbon flux alterations, stress condition
Rosamin were found to have no significant effects modifications, and metabolic pathway predic
on the growth or feeding efficiency of Atlantic tion. In recent years, biotech and bioengineering
Salmon at doses of 2.5 and 5.0%, respectively. techniques have enabled algae to become more
However, a significant increase in long-chain efficient ‘cell factories’ for carbon sequestration
omega-3 polyunsaturated fatty acids (n-3 LC- and food production [14,17].
BIOENGINEERED 9523
Microalgae biomass has been proposed as a [20]. Microalgae cells, which contain a range of
high-value feed for fish and shrimp in sustainable negative charges, have been found to have a sub
aquaculture [18,19]. Table 1 outlines the benefits stantial potential for the adsorption and accumu
and drawbacks of a variety of fish-based alterna lation of heavy metals (HMs) [21]. As a
tives to traditional meals. The idea of employing consequence of the food chain, these harmful
microalgae biomass and bio-products as aquafeed components will concentrate on aquatic animals,
for fish and shrimp growth is novel; nevertheless, endangering the health of humans who consume
various obstacles must be overcome before the fish and shrimp. The majority of previous research
concept can be successfully applied. Several issues focused on the benefits and downsides of micro
need to be resolved, including potential safety algae biomass in aquafeed. Aquafeed accounts for
issues, antinutritional factors (ANFs), limited at least 75%-90% of aquaculture<apos;>s opera
digestibility, and others. There is a significant tional expenses. New feed additives are needed,
amount of cellulose in microalgal cells, which can as traditional feed materials such as fish meal,
affect the digestion of algal biomass in fish diets fish oil, and soybean meal have become
Table 1. Presents the benefits and drawbacks of an alternative fish diet.

Alternate
Feed Benefits Drawbacks Ref.
Guar meal ● Guar meal may be substituted for soy meal ● This product includes antinutritional and anti-digestive sub [22,23]
without harming growth in certain fish. stances such as residual gum, saponin, phytate, and protease
inhibitor tannin. Gastrointestinal evacuation is slow.
● Indigestible amino acids.
● Oil production and guar gum usage affect the availability of
guar meals.
Macroalgae ● Bioactive compounds from macroalgae can ● Complex polysaccharides are indigestible [24]
help farmed fish. ● Excess heavy metals
● The probable presence of phlorotannins, lectins, phyto acids,
trypsins, and amylase inhibiting substances
Yeast ● Lignocellulosic waste can grow ● Yeast protein is low in sulfur-containing amino acids such as [25,26]
● Yeast protein is beneficial for fish, except for methionine and cysteine.
its low methionine content.
● Due to a rapid increase
Insects ● Food waste may be used as a source of ● Most insect diets were lacking in methionine and cysteine. [27]
nutrients ● Antinutritional chitin is reported.
Blood meal ● The protein content is high. ● Methionine deficiency [28,29]

(cow ● Lysine-rich ● Protein digestibility is greatly influenced by heat sensitivity
blood) and drying conditions.
Hydrolyzed ● Hydrolyzed feather meal is rich in cystine ● Not easily digested [30,31]
feather (74–61%) and protein. ● Low in lysine and methionine (2 % of the crude protein) (1 %
meal crude protein)
Wheat ● Protein content is low (11%). ● Wheat<apos;>s high starch content makes it a primary [32,33]
source of energy (usually more than 70%).
● There is a deficiency of lysine.
Microalgae ● Microalgae<apos;>s rapid rate of growth ● Formulated feeds have a high production cost. [34,35,40]
and Algal ● A broad selection of species is available, ● Microalgae with rigid cell walls are difficult to digest.
oil each with a unique set of characteristics. ω-3
fatty acid-rich
● Contains antioxidants, colorants, and has a
probiotic impact
unsustainable. Aquafeed made from microalgae is consider when formulating it. A highly digestible
not only ecologically beneficial, but with appropri feed can help reduce production costs, waste, and
ate optimization, it may also be commercially fea the risk of eutrophication in the environment. This
sible. Microalgae also have a nutritional profile article discusses the digestibility of numerous
that is comparable to that of many fish. The microalgae in fish, as well as approaches to
digestibility of the feed is an important issue to enhance microalgal digestibility. Figure 1
Figure 1. Technology process lineup for the production of beneficial fish-derived food by using an algae-based feed.
Figure 2. Metabolites produced by microalgae during their photosynthetic activity.

BIOENGINEERED 9525
represents the technology process lineup for the abundant in microalgae, unlike plant-based com
production of beneficial meals derived from fish ponents such as Chlorella and Chlamydomonas
using the algae-based feed. [43]. The proportion of starch in microalgal spe
cies varies from 7 to 45% [44]. Other microalgae
contain less starch (30–49%) than Tetraselmis
2. Microalgae as fish meal
subcordiformis, C. rheinhardtii, and C. vul
Microalgae are photosynthetic microorganisms garis [44].
that utilize atmospheric carbon dioxide (CO2) Next-generation microalgae-based feeds have
and sunlight energy to produce a variety of pro the potential to provide a sustainable source of
teins, carbohydrates, lipids, minerals, vitamins, aquaculture food. In addition to providing essen
polyphenols, flavonoids, and carotenoids, as tial nutrients, microalgae are an essential food
shown in Figure 2. Microalgae-based products source for zooplankton and lower trophic fish,
can be used in a variety of industries, including which in turn provide food for fish at higher
food and beverages, animal feed, cosmetics, che trophic levels. Microalgae can contain up to 60%
micals, and biofuels. By 2028, the market for protein, 60% carbohydrates, or 70% oil, depend
microalgae-based products is expected to grow ing on the species of algae and its growing con
from US$ 1,547.23 million in 2020 to US$ ditions. Secondary metabolites generated by
2,811.10 million. It is expected to grow at a com microalgae, such as pigments, growth-promoting
pound annual growth rate (CAGR) of 7.9% compounds, and hormones, have intrinsic anti
between 2021 and 2028 [36]. To date, the micro oxidants, antibacterial, anti-inflammatory, and
algae sector has focused mainly on species that are immune-stimulant properties that benefit both
associated with food and cosmetics. Microalgae marine and freshwater species [19,45–47].
species, including Spirulina sp., Dunaliella sp., Furthermore, numerous species can produce
Isochrysis sp., Pavlova sp., and others, are also eicosapentaenoic acid (EPA), docosahexaenoic
used as larvae feed by fish hatcheries, although acid (DHA), and colors (e.g., carotenoids) from
these are not normally grown on a large scale. the ground up, proving their adaptability. In the
However, in recent decades, microalgae have future, the cultivation of microalgae on non-ara
been explored as a possible bulk-feeding ingredi ble land or along coastlines may greatly increase
ent for fingerlings and adult fish [37]. Microalgae the global photosynthetic primary production by
have been suggested as a substitute for fish food lowering water demand and recycling nutrients
for several reasons. Microalgae have the highest by using wastewater and seawater, as well as con
net biomass productivity compared to any terres verting atmospheric CO2 into nutrient-dense feed
trial plant or animal [38]. and animal health products. In this sense, a cir
Microalgae, unlike land-based plants, do not cular aquaculture firm could arise as part of the
need fertile soil to grow. Microalgae can even larger circular bio-economy [19,48–50]. In addi
grow in seawater or wastewater [39]. Current tion, the contents of EPA and DHA in microalgal
land-use patterns do not require large-scale lipids are significantly higher and also less con
microalgae cultivation on non-arable land or taminated than fish oil. PUFA-rich microalgae
non-potable water (or practices). Unlike insects include species of Schizochytrium,
and bacteria, microalgae have minimal nutri Crypthecodinium, Nannochloropsis, Isochrysis,
tional requirements. In a biorefinery, microalgae Nitzschia, Diacronema, Porphyridium, and
might be utilized to produce fish feed [40–42]. Desmodesmus, which produce EPA and DHA,
The promise of microalgae is based on its pro respectively [51,52]. For example,
tein, lipid, carbohydrate, and pigment composi Nannochloropsis sp. and Phaeodactylum tricornu
tion, which is ideal for fish health. Some tum have about 39% EPA and 30% DHA of total
economically relevant microalgal species have omega-3 fatty acids, while Schizochytrium sp. and
chemical compositions that are equivalent to Thraustochytrium have about 40% DHA and 22%
feed components utilized in the aquafeed indus EPA, respectively [53]. Furthermore, microalgae
try (Table 2). Methionine, for example, is produce antioxidant pigments, and some
Table 2. Nutrient content of several microalgae species.

Composition (%) Ref.
Microalgal strains Lipids Protein Carbohydrates
Haematococcus pluvialis 15 48 27 [103]
Dunaliella 25–75 50–80 10–25 [61,62]
Botryococcus braunii 33 39.61 2.38 [63]
Nannochloropsis sp. 22–31 33–44 8–14 [155]
Botryococcus 25–75 3–10 17–21 [64,65]
Scenedesmus quadricauda 1.9 47 21–52 [156]
Chlamydomonas 20–25 47–50 15–20 [66,67]
Synechococcus sp. 11 63 15 [68]
C. vulgaris 14–22 12–17 1–58 [69]
Arthrospira platensis (Spirulina) 7–23 57–65 20–30 [70,71]
Isochrysis galbana 12–14 50–56 10–17 [155]
Porphyridium cruentum 5.78–7.55 27.7–40.8 22.8–39.3 [72]
Spirulina maxima 6–7 60–71 13–16 [155]
Tetraselmis maculata 3 52 15 [156]
Nannochloropsis granulata (CCMP-535) 33.5 23.6 36.2 [73–75,182]
Phaeodactylum tricornutum (CCMP-1327) 18.2 39.6 25.2
Arthrospira platensis 14.2 55.8 22.2
Tetraselmis chuii (PLY-429) 12.3 46.5 25.0
microalgae contain vitamins and immunostimu molecules. Although microalgae may have oil con
lants that are beneficial to aquatic organisms tents exceeding 60% by weight of dry biomass, the
[13,54]. most common oil contents are in the range of 20–
Microalgae have the potential to control food 50% [76–78]. Polyunsaturated fatty acids present
production and pollution by assimilating nutrients in microalgal lipids include arachidonic acid
from water and wastewater [55,56]. Wet effluents (ARA) and DHA. Cryptothecodinium and
from markets and slaughterhouses contain large Schizochytrium contain DHA, Phaeodactylum,
amounts of nitrogen and phosphorus [57]. Nitzschia, Isochrysis, and Diacronema contain
Washing fruits and vegetables at public markets or EPA, and Cryptothecodinium and Schizochytrium
slaughterhouses and washing poultry and fish pro contain ARA. Microalgal species can produce EPA
duces wet market wastewater [58], which has higher concentrations ranging from 7 to 34% fatty acids.
levels of nitrogen, phosphate, chemical oxygen These fatty acids are rare and difficult to synthe
demand, biological oxygen demand, fats, solid parti size in a lab. These components are currently
cles, oils, and greases than domestic wastewater [59]. derived from fish oil and are restricted in vegetable
Algae biomass derived from nutrient recycling of wet oils, including palm, soybean, rapeseed, and
market and slaughterhouse effluents can be used as a canola, in aquafeed [79]. Microalgae may produce
fish meal. A biorefinery strategy may result in high amounts of lipids and have a nutritionally
resource-efficient, environmentally friendly value advantageous fatty acid composition when grown
chains with a low carbon footprint through the co- under stress conditions. Polyunsaturated fatty
production of bio-based and biodegradable products acids (PUFAs), such as EPA (20:5, −3), and DHA
when algal generation systems are integrated into (22:6, −3) are the important constituents of micro
aquaculture operations (a biorefinery approach). algal lipids. Microalgae species, such as S. limaci
This strategy may also benefit the aquaculture indus num, P. tricornutum, and Nannochloropsis sp.,
try and the general public in other ways, such as comprise 30–40% of ω-3 fatty acids in their total
healthier diets and ecosystem services [48,60]. content [53]. LC-PUFAs such as ω-3 and 6 must
be consumed regularly because humans and many
animals cannot synthesize them [80]. PUFAs are
2.1 Microalgae fatty acids
also necessary ingredients for fish growth because
Lipids play a dynamic role in the formation of they cannot be formed from saturated and mono
membranes and are important energy storage unsaturated fatty acids [81]. Fish have high
BIOENGINEERED 9527
tropical levels in aquatic environments due to their variety of fish [93,94]. When S. Platensis is used at
ability to metabolize PUFA to produce LC-PUFA, a 10% concentration, it has been proven to dra
which strongly contributes to tropical upgrading. matically increase the production of white blood
It is critical to understand how farmed fish create cells, red blood cells, hemoglobin, albumin, and
and store LC-PUFAs to substitute terrestrial plant total protein in rainbow trout [95].
lipids for fish oil in commercial aquafeeds. LC-PUFAs are beneficial to animals and
Through enhanced synthesis and selective use of humans as they provide biologically active com
dietary fatty acids, it is possible to increase the pounds such as prostaglandins and thromboxanes,
body<apos;>s storage of LC-PUFA. Data synthesis which are crucial for the formation of cholesterol
was performed to determine optimal fatty acid and triglycerides in the blood, as well as the pre
levels that improve the generation and storage of vention of certain diseases such as arthritis and
omega-3 polyunsaturated fatty acids in edible por rheumatoid arthritis [96]. DHA (22:6) and EPA
tions of salmonids [82]. (20:5), two of the most beneficial LC-PUFAs,
Microalgae improved the fatty acid profile of stand out for their many health benefits. DHA
fish and shrimp by improving the ω-3/ ω-6 ratio, improves brain health, which maintains neurons,
increasing PUFA content, and enriching long- boosts short- and long-term memory, and aids in
chain PUFAs [83,84]. Aquatic animals have higher the treatment of brain disorders such as memory
nutritional value when their fatty acid profile is loss and cognitive decline. By reducing oxidative
improved, which benefits consumers. Additionally, stress and plasma triglycerides, as well as provid
supplementing aquatic animals’ diets with micro ing benefits for the treatment of inflammation,
algal PUFAs can increase their growth and immu arrhythmia, and cardiovascular disease [97,98].
nity It has been reported that Nile tilapia can Microalgae also produce a wide range of fatty
digest Schizochytrium sp. lipids with 98% efficiency acids that are useful for the food and feed industry,
[85]. Lipid digestibility in juvenile European sea including gamma-linolenic acid (GLA), linoleic
bass (Dicentrarchus labrax), Nile tilapia with C. acid (LA), alpha-linolenic acid (ALA), and ARA
vulgaris, Schizochytrium sp. Spirulina sp., and [99]. Wound healing and regeneration, as well as
Chlorella sp., and African catfish (Clarus gariepi the eradication of invading microorganisms, may
nus) with C. vulgaris and S. maxima has been be aided by these essential fatty acids [100]. The
shown to be more than 80% [85–87]. Lower lipid content of C. vulgaris has been reported to be
digestibility has been noted under certain condi 35–40% by weight, with a content of 27% linolenic
tions, such as for N. gaditana in juvenile African and 24% linoleic acid [101]. Spirulina sp. has been
catfish [88]. A microalgae-based feed might also promoted as an inexpensive source of GLA [100].
improve fish survival due to its functional proper Recent research has indicated that microalgae such
ties, including probiotics, prebiotics, immunosti as Chlorella sp. and Schizochytrium sp. are more
mulants, antivirals, antibacterials, etc. Bacteria or attractive than other autotrophic species due to
microbes that are probiotics are believed to con their nutritional qualities and ability to be effec
tribute to a healthy gut when consumed. For tively consumed by aquaculture species. Thanks to
example, microalgae are probiotics for fish. The the well-developed technology for mass cultivation
microbiome in the colon digests algal cells and of Schizochytrium and Chlorella sp. in aquafeeds,
generates probiotics that inhibit the growth of the use of fish oil can be significantly reduced in
infectious agents [89,90]. Consumption of the future. To achieve sustainable substitution of
Tetraselmis suecica live cells significantly reduced fish oil in aquaculture, research must focus on the
the number of harmful bacteria in the stomach of use of microalgal species.
white shrimp (Fenneropenaeus indicus) as com
pared to a control group [91]. The addition of
2.2 Proteins and amino acids derived from
Schizochytrium sp. meal to the diet at 1.2% signif
microalgae
icantly improved Nile tilapia health [92]. Previous
studies have found that Spirulina can induce non- Proteins have been recognized as the building
specific immune responses against infections in a components that are responsible for individual
growth. Proteins are constructed from peptide European seabass and Atlantic salmon [87,111].
bonds that link amino acid units [102]. In terms Tetraselmis sp. exhibited significantly lower mino
of quality and amino acid composition, microalgae acid digestibility than juvenile European seabass.
protein is a great alternative to fish meals. The Pretreatment would break down larger proteins
protein concentration of algae has been reported into peptides and individual amino acids, which
to range from 40 to 60 wt/wt % [103]. Another increases amino acid digestibility [110].
study discovered that C. vulgaris has between 51 Threonine, isoleucine, lysine, leucine, methio
and 58% protein, while Spirulina sp. comprises nine, valine, and histidine are essential amino
between 60% and 71%. Additionally, Arthrosphira acids that the body cannot produce itself.
platensis has a protein content of 70% by weight Therefore, it is important to consume them
[104]. Protein is so abundant in some microalgae through foods that contain EAAs, such as tofu,
species that it accounts for more than half of their eggs, and fish [112]. Vegans and vegetarians have
biomass. Most Spirulina strains, as well as a few few options since the majority of plant-based pro
Chlorella and Nannochloropsis strains, have a pro teins do not meet the EAA profile. To overcome
tein content of 40 to 65% [105]. Microalgae can this problem, an alternative source with a balanced
synthesize all amino acid molecules; therefore, protein profile and low cost is required [113].
algae-derived amino acids are preferred over Protein digestibility of the microalgal protein (S.
other protein-rich foods [106]. Microalgae can platensis protein concentration) ranges from 87.5
synthesize several protein compounds faster than to 97.8% [114]. However, certain algae (510–710 g/
traditional protein sources. Table 3 summarizes kg) have more protein than eggs or soybeans (132–
the current research on microalgal biomass used 370 g/kg) and have fairly comparable EAAs [115].
as a replacement for fishmeal and fish oil. Due to their high content of EAA, microalgae are
Kim et al. [107] discovered that parrotfish considered one of the best vegan protein sources.
(Oplegnathus fasciatus) fed 5% Arthrospira had It is well known that microalgae contain EAAs and
significantly higher weight, protein efficiency, and non-NEAAs, both of which have health benefits
feed consumption compared to the control group [113]. NEAAs include amino acids, proline, argi
fed fish meal. Fish need meals with 30% to 55% nine, glutamic acid, glycine, aspartic acid, tyrosine,
crude protein and amino acids tailored to their cysteine, serine, and glutamic acid are a few exam
individual nutritional needs to achieve maximum ples. The amino acid profile of C. vulgaris and H.
growth [108]. Sørensen et al. [109], demonstrated pluvialis, the proportion of NEAAs is around 51%
that Phaeodactylum tricornutum can replace up to and 49%, respectively [116]. A healthy immune
6% of fish meal in the diet of Atlantic salmon system is influenced by these chemicals, as well
(Salmo salar) without affecting digestibility, utili as gene expression, antioxidant responses, and
zation, or growth performance. Protein digestibil cell signaling [117]. Many studies indicate that
ity of microalgae varies from 50% to 94% in the amount of microalgal meal that should be
different fish species. Protein digestibility of rain added to aquafeed varies depending on the type
bow trout, Nile tilapia, European seabass, and of algae used and the aquaculture species being
African catfish has been reported to exceed 80% fed. It would be beneficial to study the growth
[110]. In African catfish and Nile tilapia, bead capacity of microalgae and identify the variables
milling improved N. gaditana protein digestibility that influence their effectiveness.
by 16 and 17 %, respectively [86]. Enzymatically
processed microalgae digest protein 6% faster than
whole-cell Nannochloropsis oceanica [40]. Protein 2.3 Microalgae-based pigments
digestion was enhanced in a diet containing The color of microalgae is one of its most distin
Schizochytrium sp. when organic minerals were guishing properties, which is determined by pig
added [87]. N. oceanica and C. vulgaris provided ments. Microalgae pigments are critical for their
amino acids with higher digestibility than 90% for nutritional performance in aquaculture. In
BIOENGINEERED 9529
Table 3. Microalgal biomass as an alternative or supplement to fishmeal and fish oil.

Fish oil/fish meal/
dietary inclusion level
Microalgae species Aquaculture species replacement Effects of algae biomass Ref.
Schizochytrium Pacific white shrimp ● 4% inclusion in ● Although shrimp survival, digestive enzyme activity, [118]
(Litopenaeus the diet and fatty acid content were not affected, their specific
vannamei) growth rate was much higher than in the control
group.
Dunaliella salina Giant tiger prawn ● 5–10% incor ● The immune system and antioxidants (superoxide [119]
(Penaeus monodon) poration in feed dismutase and catalase) improved significantly and
the survival rate was significantly boosted.
Phaeodactylum Atlantic salmon (Salmo ● 6% replacement ● There is no negative impact on growth, protein, [109]
tricornutum salar) of fish meal lipids, energy, ash, growth performance, etc., in the
feed that is used.
Nannochloropsis sp. and Juvenile Atlantic cod ● 15% fish meal ● Increased feed intake and fish growth. Survival, feed [120]
Isochrysis sp. (Gadus morhua) protein conversion ratios, and muscle ω-3 and ω-6 fatty acid
replacement levels did not differ between the treatment groups.
Schizochytrium sp. Tilapia ● 100% replace ● However, the survival rate did not alter substantially. [85]
(Oreochromis niloticus) ment of fish oil
Nannochloropsis gaditana, Gilthead seabream ● 0.5 and 1% ● Increased defensive activity [121]
T. chuii, and (Sparus aurata) inclusion in feed
Phaeodactylum
tricornutum
Chlorella vulgaris Giant freshwater prawn ● 6–8% fish meal ● M. rosenbergii postlarvae had a faster growth rate, a [122]
(Macrobrachium substitute better immunological response (total haemocyte
rosenbergii) count and prophenoloxidase activity), and were
resistant to Aeromonas hydrophila infection.
Arthrospira sp. Golden barb (Puntius ● 20% fishmeal ● Fish growth rates have increased significantly. [123]
gelius) substitute
Pavlova viridis European sea bass ● Fish oil replace ● In terms of growth performance and nutrient con [124]
Nannochloropsis sp. (Dicentrarchus labrax) ment 50–100% sumption, there are no detrimental consequences on
fish.
Nanofrustulum sp. Atlantic salmon (Salmo ● fish meal repla ● Algal meal outperformed fish meal in terms of [125]
salar), common carp cement 5 or 10% growth and feed intake, indicating that it may be used
(Cyprinus carpio) in place of fish meal.
Tetraselmis sp. Pacific white shrimp
(Litopenaeus
vannamei)
Arthrospira platensis Nile tilapia (Oreochromis ● 0.5–2% inclu ● Enhanced fish health through tissue protection and [126]
niloticus) sion in feed antioxidant effects
Arthrospira sp. Tilapia (Oreochromis ● Replacing fish ● Unlike corn-gluten meal control, there was no dele [242]
niloticus) meal by up to terious impact on growth or feed consumption.
43%
addition to chlorophyll, microalgae include caro chlorophylls, and phycobilin are the three primary
tenoids and phycobiliproteins. The types of microalgal photosynthetic pigments
Nannochloropsis genus contains pigments such as [127,128]. Microalgae pigments are eye-catching
chlorophyll and astaxanthin. Photosynthesis in natural colors that include high-value components
algae is facilitated by pigments, which are brightly with health-promoting qualities that include anti
colored chemical compounds. Carotenoids, oxidants, vitamin precursors, neuroprotective, and
immunological boosters [129]. These pigments [135]. Phaeodactylum tricornutum produces large
may address the increased demand for natural quantities of fucoxanthin, which has been shown
colors due to health concerns about the adverse to contribute to the golden yellow coloration of
effects of synthetic pigments [127,130]. gilthead seabreams [136]. Carotenoids are found
Aquaculture uses a high concentration of carote in a wide variety of products, including natural
noids, such as β-carotene and astaxanthin, due to feed colors, food supplements, vitamin supple
their vibrant color and antioxidant effects. These ments, and health foods. The high concentration
molecules have the potential to improve the qual of carotenoids in D. saline makes it the most
ity and value of farmed fish, such as salmon and popular species for large-scale production (up to
Asian tiger shrimp (Penaeus monodon) [131]. 14% dry weight) [137]. Microalgae strains that are
Phytochemicals such as astaxanthin and β-caro commercially feasible for pigment synthesis must
tene are abundantly generated by the microalgae meet a series of criteria, including improved nutri
Haematococcus pluvialis and Dunaliella salina (3– tional components, non-toxicity, and the presence
7% wt/wt) in natural abundance [132,133]. Table 4 of digestible cell walls for nutrient absorption
shows the pigment compositions of numerous [129]. Phycobiliproteins, β-carotene, and astax
algae species, as well as the health benefits asso anthin are used mainly as colorants, pharmaceuti
ciated with them. cals, aquaculture, and nutraceuticals.
Microalgal biomass has been shown to affect Chlorococcum sp. (Astaxanthin, lutein, β-caro
fish pigmentation. H. pluvialis is the most com tene), D. salina (β-carotene, zeaxanthin, chloro
monly used microalgae specifically for color phylls a, b), H. pluvialis (astaxanthin,
enhancement. In the aquaculture industry, whole canthaxanthin, lutein), Spirulina sp. (β-carotene,
cells and extracts of H. pluvialis extracts are used zeaxanthin, phycocyanin, allophycocyanin),
as feed additives (1.5–1.7%) [134]. Several algae Porphyridium sp. (phycoerythrin) [138–143].
species are used as pigments in fish feed. The Natural feed pigments, feed additives, nutri
Haematococcus produces Astaxanthin, which ents, and health food products are commonly
gives salmon its pink hue [134]. Additionally, made from carotenoids. Carotenoids found in
Spirulina contains additional carotenoids that abundance in D. salina make it the species
ornamental koi and other fish can convert to most often exploited for large-scale production
astaxanthin and other brightly colored pigments [144]. Carotenes from Dunaliella species were
shown to improve the health of L. vannamei
Table 4. Algal pigment compositions and their health benefits. shrimp given high-carotene diets. Coloration
Microalgal and market acceptability can be achieved by
species Pigments Health benefits Ref. supplementing Red tilapia diets with A. platensis,
Haematococcus astaxanthin Pink colored pigment, [134] a source of pigmentation [145,146]. According
Antioxidant, Improved
disease resistance, faster to these studies, low amounts of Arthrospira or
growth other microalgae can enhance the color and fla
Spirulina β-carotene, Yellow, orange, and red- [135]
astaxanthin colored pigment,
vor of numerous fish species, such as tilapia.
antioxidants, improved Therefore, additional studies are required to
disease resistance, faster evaluate the impact of various microalgal pig
growth
Phaeodactylum fucoxanthin Golden and yellow [136] ments on commercial aquaculture.
tricornutum coloration, Antioxidant,
Anti-inflammatory
D. saline carotenoids Photo-protection, [137]. 2.4 Microalgae-based vitamins
camouflage, and
signaling enhance Microalgae are high in vitamins, and vitamin B has
immune system
Chlorella fucoxanthin, Yellow-colored pigment, [182] been shown to function as a cofactor for mito
vulgaris. zeaxanthin, antioxidant, anti- chondrial enzymes, reducing oxidative breakdown
and lutein inflammatory
Scenedesmus lutein Greens and orange-yellow, [75]
and improving metabolism [147,148]. Microalgae,
sp. Antioxidant, reduce including Spirulina sp., have more Vitamin B12
inflammation (127–244 g/g) than plant or animal-based foods.
BIOENGINEERED 9531
This vitamin helps prevent megaloblastic anemia, Table 5. Algal monosaccharide compositions.
which causes fatigue and weakness [149]. Another Monosaccharide composition (%)
study found a significant amount of vitamin E (3.7 Microalgal strains Arabinose Glucose Galactose Xylose Ref.
mg/g) in the Euglena gracilis microalgae. Vitamin S. platensis 1.4 24.1 16.4 6.1 [165]
Arthrospira platensis – 38.3 36.4 0.7 [166]
E has been reported to reduce the risk of cancer, Chlorella sp. 34 20 41 – [167]
eye disease, heart disease, and other diseases [150]. Porphyra ochotensis – 5.3 30.4 1.2 [168]
A high amount of vitamin C content (3.44 mg/g) C. vulgaris 1.6 54.9 – 2.3 [169]
JSC-6
has been found in the Eisenia arborea brown C. marina 37.6 30.3 10.0 – [166]
microalgae, which is equivalent to that of man
darin oranges [151]. This antioxidant vitamin is
necessary for immune system function, tissue for higher carbohydrate digestibility (greater than
mation, and repair [152]. 70%) in Nile tilapia [86]. Microalgae contain
starch-like carbohydrates that can be easily
digested. In vitro research revealed that C. soro
2.5 Carbohydrates derived from microalgae kiniana, Klamath, and N. sphaeroides showed
Microalgae are rich in carbohydrates, and polysac greater carbohydrate digestibility [161]. Studies
charides are readily found in both their cytoplasm conducted on Nile tilapia, African catfish, and in
and chloroplast [153]. Microalgae carbohydrates vitro studies have shown that C. vulgaris has a
are used for several reasons, including energy sto higher carbohydrate digestibility value than other
rage and structural components in cell walls [154]. algal species [86]. Certain microalgal species, such
Due to the high photoconversion efficiency, as Spirulina sp., Chlorella sp., and Schizochytrium
macroalgae such as P. cruentum contain carbohy sp., have been demonstrated to have excellent fiber
drates (40–57 wt% dry weight), Prymnesium par digestibility [162]. Isochrysis sp. and
vum (30–33 wt% dry weight), and S. quadricauda Nannochloropsis sp. fiber digestibility in rainbow
(21–52 wt% dry weight) [155,156]. For each spe trout was determined to be 96% and 38%, respec
cies of the algal genus, there is a distinct variation tively [11]. Even though Isochrysis sp. contained
in glucose metabolism and content [156,157]. more fiber than Nannochloropsis sp., the latter had
Algae with high carbohydrate yield and sugar con a higher fiber digestibility, suggesting that the type
tent are suitable for human consumption. The of fiber (soluble or insoluble) is important for
culture system and environmental conditions can digestion. Compared to other nutrients, starch is
impact algal production and glucose content [153]. a readily digestible food for fish and crusta
Microalgae contain about 10–25% carbohydrates ceans [163].
and their amount varies with culture age and The antiviral and antibacterial properties of β-
growth conditions [158]. A variety of starches, glucan have been shown in humans, and it has also
cellulose, sugars and other polysaccharides are been shown in fish to have high antibacterial and
found in microalgae. Bacteria and fungi naturally immune-stimulating properties [164].
produce the polysaccharide β-glucan, which is Furthermore, the types of carbon sources used
made of D-glucose. These polysaccharides can be and the metabolic mechanism employed are
found in large quantities in the Chlorella sp., other important variables that affect the sugar
microalgae [78,159]. Table 5 displays the mono concentration in microalgae [115]. The use of
saccharide compositions of several microalgae. light has been shown to influence both algal
Microalgal carbohydrates are digestible accord growth and biomass composition during algal cul
ing to the type and quantity of carbohydrates in ture since light is an important energy source for
biomass, as well as the kind of fish that consumes photosynthetic activity [78,153]. For algal cultiva
microalgal carbohydrates [160]. The carbohydrate tion, the light intensity is normally between 200
digestibility of microalgal species varies between and 400 mol photons m2/s. Nutritional restriction
22% and 83%, depending on fish species [110]. S. may be used to alter the metabolic process of
maxima and C. vulgaris were found to have a microalgae and cause glucose accumulation [44].
3. Nutrients digestibility been supported by any other study. Therefore,

more research is needed to link fiber content
It is important to understand how digestible feed
with nutrient digestion [161].
components are used to calculate their nutritional
In rainbow trout, Isochrysis sp. absorption of
value. Microalgal products are assessed for their
nutrients was shown to be superior to
nutritional digestibility before they can be used in
Nannochloropsis sp [11]. Fiber and other anti-
aquafeeds. Many different types of microalgal
nutrients also reduce proteolytic and amylase
organisms, from prokaryotes to eukaryotes, con
activity and digestibility [180,181]. Other variables
tribute to the wide variety. There has also been
can impact the digestion of a microalgae diet.
evidence of intra-species diversity in metabolic
Exopolysaccharides can form stable complexes
profiles. The nutritional and energy digestibility
with proteins, inhibiting proteolysis [161].
of various microalgal species varies due to differ
Exopolysaccharides are exopolysaccharides that
ences in chemical composition and physical struc
are released or remain attached to cells in an
ture. Fish digestion of microalgae is influenced by
algal culture [161]. Phenolic chemicals found in
the composition and rigidity of the cell [161]. In
plants and seaweed can precipitate proteins.
the prokaryotic (cyanobacterial) microalgae, pepti
Although microalgae have a modest phenolic con
doglycan layers are present in the cell wall, while
tent (0–20 mg GA/g-DW), plant phenolic sub
in eukaryotic microalgae, cellulosic layers occur
stances in the diet could impact microalgal
[86,170,171]. Fish prefer microalgae with peptido
protein digestion [182–184]. The digestibility of
glycan (murein) layered cell walls over cellulose
amino acids may be impaired. Furthermore, dif
layered microalgae [86]. The rigidity of the cell
ferences in physiology between fish species cause
wall also affects digestion. Thick-walled microalgae
different digestibilities for the same microalgae
are less digestible than species with thin or no cell
[86]. Fish species vary in their digestive enzyme
walls. Desmodesmus, Nannochloropsis,
profiles, in addition to their physiology. Only a few
Haemotococcus and Chlorella, microalgae are with
fish, such as the Rohu (Labeo rohita), have the
thick cell walls, while I. galbana, Porphyridium
enzymes required to break down cellulose [185].
cruentum, and D. salina are with thin cell walls
[85,110,172].
However, certain proteins may still alter the
3.1 Bioaccumulation of heavy metals
digestion of microalgae. Unwanted trypsin inhibi
tor, an enzyme that inhibits proteolytic enzymes, During cultivation, microalgae cells can absorb
may cause poor digestion of Nannochloropsis sp. and accumulate substantial amounts of heavy
Some marine microalgae include lipase inhibitors metals (HMs) [78,186–188]. Several studies have
that may affect lipid digestion [87,173,174]. examined the bioaccumulation of HMs in micro
Microalgae are made up of non-starch polysac algal biomass due to the existence of negatively
charides and fibers, making them difficult to digest charged functional groups on the surface of micro
[11]. Polysaccharides that are not made of starch, algal cells [189–191]. HMs absorbed by microalgae
such as cellulose, gums, pectins, and hemicellu include predominantly arsenic (As), cadmium
loses, are usually difficult to break down (Cd), copper (Cu), chromium (Cr), lead (Pb),
[170,175,176]. Digestive enzymes are absent in mercury (Hg), nickel (Ni), and lanthanum (La).
certain fish species, such as Nile tilapia, which In fact, both living and non-living microalgae are
cannot break down the beta glycosidic bonds effective in absorbing HMs from the environment
found in non-starch polysaccharides [177]. and water [21,191]. Non-living microalgae can
Undigested carbohydrates are transported quickly accumulate Cd, Cr, Hg, and Pb at 59, 98, 36, and
by the digestive tract, absorb proteins, and reduce 131 mg/g, respectively. Non-living algal biomass
protein digestibility [178,179]. Fiber concentra has a strong capacity to adsorb HMs primarily due
tions were shown to have a negative relationship to their surface functional groups [192].
with organic matter, protein, and carbohydrate When microalgal biomass containing concen
digestibility [147,161]. This hypothesis has not trated HMs is fed to fish, harmful compounds
BIOENGINEERED 9533
can migrate up the food chain. Certain strategies omnivore fish digest carbohydrates are better than
can be employed to avoid bioaccumulation of carnivorous fish, although they exhibit significant
HMs in the food chain when microalgae are used anatomical and physiological differences in their
as aquafeed. The amount of HMs in the culture digestive systems [198,199]. Another study found
media should be monitored regularly during that omnivorous fish have various digestive capa
microalgae culture. If HMs concentrations exceed cities [20]. Rather than raw biomass, defatted
the EPA<apos;>s guidelines, the culture medium microalgae biomass is often used as a supplement
must be treated or relieved. This method may to fish feed in the fishing sector. After oil extrac
avoid the bio-accumulation of HMs in the food tion, the starch content in the biomass could be
chain. Before using microalgae as fish food, it is increased. The microalgae biomass must be effec
possible to eliminate HMs from them. HMs deso tively managed to avoid an excessive starch con
rption may occur through pH-induced desorption tent. The apparent digestibility coefficients (ADCs)
or metal-chelating agent treatment. The pH of of macronutrients, amino acids, and fatty acids
algal cells must be lowered to the isoelectric points were studied in freshwater (Arthrospira,
of functional groups on their surfaces to neutralize Chlorella) and marine microalgal
their surface charge [193]. Heavy metals adsorbed (Schizochytrium) components in Nile tilapia [85].
on the surface of algae might be combined with Compared to chlorella, Arthrospira exhibited sig
metal chelating agents to facilitate their deso nificantly higher ADCs of crude protein and all
rption. HMs have been successfully desorbed EAAs (86%), which corresponded well with the
from microalgae biomass using these two reported values for fishmeal and plant feeds.
approaches [193–196]. When HMs are attached Schizochytrium had the highest DHA content, as
to the surface of algal cells rather than within well as the highest ADCs for lipids (total PUFA
algal cells, desorption techniques may be used to 98%), ω-3 (98%), and ω-6 (92.4%), as well as the
remove them from the biomass of microalgae. In maximum digestibility of DHA. Spirulina and
contrast to the accumulation of HMs inside algal Schizochytrium were shown to be effective protein
cells, microalgal cells can be treated by desorption substitutes for tilapia diets, while Schizochytrium
to remove HMs adhering to the surface. Therefore, was discovered to be a good LC-PUFA supple
after desorption, HMs content in algal cells should ment [200].
be evaluated to determine whether microalgae can For microalgal cells to be preserved, they usually
serve as aquafeed. need to be spray-dried after they have been
removed from the growing reactor. Furthermore,
microalgal biomass is dried in a variety of ways,
3.2 Deficiency in digestibility
such as sun drying, drum drying, and oven drying.
Since microalgae have a high starch concentration, However, the microalgae cell wall remains intact
replacing the biomass of microalgae with a fish in these circumstances, indicating that digestion is
meal reduces digestibility. Furthermore, aquatic restricted [87,201,202]. A cellulose-rich cell wall
animals must dissolve the cellulose-rich cell walls causes poor digestion of microalgae-supplemented
of algae before consuming their nutritional com fish diets [203]. The cell wall protects the algal cell
ponents. For these two challenges, meals supple and its intracellular components during growth. It
mented with microalgae- in fish and shrimp is necessary to disrupt the cellulose structure of the
culture are poorly absorbed [197]. Dietary wheat cell wall in order to access the intracellular con
starch levels of 20% in juvenile largemouth bass tents. Due to a lack of intestinal flora, several
(Micropterus salmoides) not only impeded weight species of fish are unable to digest non-starch
gain but also produced oxidative stress and carbohydrates effectively [204,205]. Therefore, cel
impaired innate immunity. Fish offered 5% and lulose cannot be digested by fish, and the structure
10% starch diets showed significant weight gain, of the cell walls of microalgae cannot be effectively
growth rate, protein efficiency ratio, and feed con dissolved by many species of fish. As a conse
version ratio when compared to fish-fed 20% quence, aquatic organisms will not be able to con
starch diets [198]. It is generally recognized that sume nutrients found in algal cells. To improve the
digestibility of microalgal biomass for use in fish/ 3.3 Anti-nutritional factors

shrimp aquaculture, certain pretreatment techni
ques could be applied. Microalgae, for starters, Anti-nutritional factors (ANFs) are biological
contain starch, which may be separated before components that affect gastrointestinal and meta
using algal biomass as aquafeed. Second, destroy bolic function in animals and humans. These
ing the cell wall may cause the nutritional compo ANFs were detected in vegetable soybean and pea
nents of the microalgae to be released [206]. nut proteins together with phytic, lectin, and tan
Enzymatic digestion and physical treatment have nic acids. Due to the increased nature of ANFs,
been shown to destroy microalgae cell walls effec protein meals or peanut meals cannot be used in
tively [206,207]. Microalgae cell walls are made up aquafeed [213]. Microalgae have been shown to
of pectin, hemicellulose, cellulose, and glycopro contain a variety of ANFs, such as Tetradesmus
teins. In algae, enzymes such as glucosidase, cellu obliquus, Kirchneriella lunaris, and
lase, hemicellulase, xylanase, and exoglucanase Pseudokirchneriella subcapitata [214,215].
break down the cell wall [206]. Previous studies indicate that the tannic acid levels
To improve nutrient digestibility, biomass can of S. maxima (6.86 mg/g) and C. vulgaris (1.44
be pretreated/processed using pasteurization, mg/g) were comparable [216]. High absorption of
freeze-drying, bead milling, high-pressure homo microalgae may negatively affect fish growth
genization, pulse electric field, microwave, chemi because of antinutritional components in micro
cal, and enzymatic treatments [208]. During algae. ANF-supplemented aquafeed reduced liver
biomass processing or pretreatment, the rigid cell enzyme activities and reduced intestinal brush
wall of algae is broken down, releasing internal border enzymes in juvenile Japanese seabass
nutrients that can be digested and absorbed by (Lateolabrax japonicus) [217].
fish. According to studies, the processing of speci It would be beneficial to discover and study
fic microalgae is linked to improved digestibility in anti-nutritional components in microalgae to
certain fish [87,209,210]. Physical treatment pro increase their incorporation into fish diets. A pre
cesses such as sonication, the beating of beads, and treatment strategy for microalgae should also
freezing can be used to damage the microalgal cell include the scavenging of anti-nutritional factors.
wall. The cost of the aforementioned methodolo Many studies have demonstrated effective ways to
gies should be used to determine their viability in a eliminate the anti-nutritional components from
real-world application [4]. The grinding of soybean meals [218,219]. Antinutritional chemi
Tetraselmis sp. beads, for example, increased pro cals may be removed by specific microorganisms
tein digestibility in European seabass by 20% com through their activities. When Aspergillus sojae
pared to untreated cells [87]. Pretreatment with and/or Aspergillus ficuum are used in conjunction
bead milled microalgae rather than whole biomass with solid-state fermentation, the phytic acid was
increased the digestibility of amino acids such as reduced by 53.27% to 73.16%. Enzymes produced
phenylalanine and aspartic acid in European sea from microorganisms may also be used to remove
bass, but not the digestibility of essential amino certain anti-nutritional substances [220]. Another
acids [87]. Nannochloropsis and Chlorella cell walls study found a maximum tannin removal rate of
were broken by bead milling for 10 minutes, which 73%. If antinutritional components are identified
could release nutrients and improve digestibility in microalgae, different physical treatment meth
[86,209,211]. For European seabass, enzyme pro ods, including extraction, frying, blanching, and
cessing improved the protein digestion of soaking, may be utilized to minimize their con
Nannochloropsis sp. and Chlorella sp., as well as centration. The pretreatment treatments described
energy digestion of Nannochloropsis sp. above have been shown to significantly decrease
Tetraselmis sp., and Chlorella sp., by 14%, 11%, ANFs in a variety of vegetable proteins [221–224].
and 40%, respectively [87]. Due to the fact that For example, extrusion cooking with optimal bar
the degree of cell rupture differs between species, rel temperature, extruder speed, and moisture con
nutritional accessibility can vary despite equal pro tent removed 61.25 % of tannin in linseed meals
cessing conditions [212]. [224]. Eventually, these technologies may be
BIOENGINEERED 9535
utilized to pre-retreat microalgae to eliminate anti- aquaculture system. Zooplankton is the main
nutritional components and surge algal biomass food source for fish and shrimp in this environ
assimilation into fish diets, which would be bene ment. In addition to breaking down animal waste,
ficial in the long run. microalgae can also transform it into valuable
components [15,231]. Therefore, harvesting and
drying can be avoided, and microalgae can be
3.4 Challenges in microalgae biomass
used to improve aquaculture water quality.
harvesting and processing
Therefore, this unique approach to microalgae
Most aquafeed substitutes microalgae biomass for aquaculture is more economically feasible while
fish meal. Microalgae cultivated in a culture med still addressing the objectives of a circular
ium should be harvested and dried, and the economy.
moisture adjusted before being used to make
algae biomass-supplemented fish feed pellets.
4. Economic and environmental feasibility
Centrifugation, sedimentation, and filtration are
all processes used to collect suspended algae cells To develop algae-based products as a possible
in a culture medium. The collection of algae source of food or feed for humans and animals,
biomass accounts for 30% of microalgae produc their economic feasibility and long-term viability
tion costs. In addition to that, flocculation har must be improved. The Techno-economic assess
vesting incorporates aluminum into biomass, ment (TEA) and life cycle assessment (LCA) meth
making microalgae-based aquafeed based on ods can analyze both the manufacturing route and
microalgae undesirable [225]. Drying wet micro the technical process of R&D activities to achieve
algae (with a moisture content of 70–90 %) is commercial and environmental viability [232].
required before the algae can be delivered to the Based on six potential alternative sites, a techno-
feed factory. This is an energy-intensive, time- economic study was conducted on the entire life
consuming, and costly technique [226]. cycle of a 100-hectare microalgal production plant.
Managing the moisture content of dehydrated Due to improved photosynthetic efficiency and
microalgae biomass is important throughout the photobioreactors with shorter light paths, it has
pelletization process. been determined that the cost of algal production
Certain alternatives have been proposed and in Spain is approximated at 3.4 euros per kilogram
thoroughly explored to overcome the aforemen of dry biomass with a predicted decrease to 0.5
tioned concerns. The development of more cost- euros per kilogram in ten years. The production of
effective biomass harvesting methods is an essen high-value metabolites (such as pigments) could
tial first step. Co-cultivation of a filamentous fun generate 657 million euros in profits over the next
gus with microalgae previously resulted in fungal- 15 years (or more) [233].
algal pellets [15,227]. As a consequence, biomass Biotechnology can be used to create bio-based
collection has become a passive process that does products, bioenergy, food, and feed while improv
not require human interaction. Several fungal ing ethical and environmental sustainability, opti
strains, including Mucor circinelloides and mizing production processes, and lowering costs.
Aspergillus oryzae, produce high-value compo Furthermore, to generate biomass, certain micro
nents when cultured with microalgae, which can algal species have been shown to thrive in com
improve the nutritional content of biomass plex organic waste streams (digestate, wastewater,
[228,229]. Microalgae immobilization techniques etc.) and to remove nutrient pollutants (P, N, and
have also been implemented to minimize collec other toxins) [234–237]. As a consequence, inte
tion costs. After growing in the substratum, a grated biorefineries based on microalgal biotech
biofilm<apos;>s biomass may be scraped off nology could recover some agricultural by-
using scrapers. Immobilized microalgae are more products while reducing disposal costs.
economical to harvest than suspended microalgae Compared to monoculture cultivation, hetero
[230]. Microalgae can be used to make an eco- trophic cultivation is cost-effective since it takes
friendly closed or semi-closed food chain less land and investment, consumes more energy
and carbon, and has a lower cost of downstream [244,245]. The lack of substantial amounts of
processing [237,238]. Microalgae protein has a microalgal biomass may hinder the growth of the
significant environmental impact because the dry aquafeed sector. Massive amounts of microalgal
ing process is so energy-intensive. In contrast to biomass for the aquafeed industry require success
beef and pork, the LCA in autotrophic and het ful large-scale algal growth of commercially rele
erotrophic algae cultures resulted in more ecolo vant microalgae species [246].
gically sustainable products derived from
heterotrophic cultivation [239]. When using
6. Perspective and future direction
hydrolyzed food waste to obtain carbon in micro
algal production, the environmental benefit could The cell walls of microalgae vary in content and
be 4.5 times greater, making it much more eco- structure. This emphasizes the need to screen for
friendly among protein sources. commercial strains that can be readily handled for
cell disruption. Only a few microalgal species have
been digestible tested. Several microalgae and cya
5. Current concerns with microalgae in
nobacteria species, including Anabaena sp. and
aquafeed
Nostoc sp., have not been tested for digestibility
The high cost of microalgae production continues by fish or other aquatic species. D. salina, for
to be a barrier to aquaculture. Microalgae have the example, does not have a cell wall, which can
potential to be used economically by cutting the improve digestion of cellular metabolites. It is
costs of production and distribution [85,240]. also critical to explore the impact of environmen
Microalgae are difficult to dry and pelletize, and tal variables and stress on the chemical composi
incorrect drying can alter their nutritional and tion of microalgae. For example, nitrogen
physical qualities, which in turn reduces their use deficiency could cause a buildup of PUFA and
as feed. Some microalgae (such as Chlorella) have starch, which fish can consume. The metabolic
thick cell walls that hinder nutrient uptake. Some response of microalgae to stress, such as nitrogen
microalgae (e.g., D. tertiolecta) have extracellular deficiency, differs between species. In certain
polysaccharides that might interfere with nutri strains, nitrogen deficiency lowers protein levels
tional absorption. Poor digestibility and significant while increasing carbohydrate levels [247].
salt buildup in marine microalgae species used as Optimal stress management and species selection
fish feed might cause problems. Algae can only are required to achieve a desirable biochemical
provide 10–15% of dietary protein requirements profile and digestibility.
in test diets without affecting development or In addition to the availability of microalgal bio
food consumption. Several microalgae have a mass at a reasonable price, microalgal and aqua
high carbohydrate and low protein content feed producers must address substantial variability
because their tough cell walls inhibit the digestion in the proximate composition, digestibility, and
of fats and proteins [163]. growth conditions. Aquaculture requires a diverse
Due to the high concentrations of trace ele range of microalgal species. Microalgal species
ments and toxins found in microalgae biomass, it with better nutritional or growth properties may
is not recommended for use in aquafeeds. Protein be more efficient. The nutritional value of various
left behind after fat is extracted for the generation microalgae should be investigated. This means that
of biofuels is often proposed for use in animal feed the cell wall must be broken down to make the
[241]. It is possible that microalgae used to pro algal elements available to digestive enzymes
duce biofuels are not suitable for feeding and that [147,211,248]. However, further processing stages
the demand for low-cost fuel production could may increase costs. To improve fish health, micro
result in toxic protein residues [242,243]. It is algal material must be tested for harmful chemicals
preferable to first utilize algal biomass for higher- prior to commercialization. In addition to nutri
value products, such as aquafeed, and then use any tional content and digestibility, the processing
remaining chemicals for biofuel production. This requirements of industrial production lines for
means a high-value product-first philosophy compound feed must also be considered. The
BIOENGINEERED 9537
composition of extruded fish feed is acknowledged Astaxanthin, polyunsaturated fatty acids

to be one of the most critical elements that impact (PUFAs), and phycocyanin are just a few of the
physical quality [249]. Microalgae were examined high-quality components found in microalgae.
in a recent study to physically determine their There is little doubt that these high-quality con
effects on extruded fish feed, but further research stituents can improve an animal<apos;>s immu
is necessary [208]. nological response, increase its survival rate, and
Due to the explosive growth of the aquacul help it gain weight. The biosynthesis of astax
ture industry, it is important to study the digest anthin, PUFAs, and phycocyanin in microalgal
ibility of microalgae. So far, digestibility has cells is technically feasible to produce high-quality
been examined only for salmonids, tilapia, sea biochemical products. Thanks to significant bioen
bass, and African catfish. During feed screening gineering technology advances, microalgal biomass
trials, microalgae should be evaluated for anti- and bioproducts such as aquafeed are no longer
nutritional factors, such as digestion enzyme just a theory. In addition, the culture medium
inhibitors (e.g., caulerpenyene, a terpene) and should be regulated to avoid the accumulation of
nutrient bioavailability factors. Studies on HMs in the biomass. To make microalgae suitable
macronutrients, such as lipids, carbohydrates, for aquafeeds, the cell wall of the algae must be
protein, and energy, have previously focused on broken down to release intracellular nutrients and
microalgae. However, specific classes of these scavenge nutrient-hostile chemicals. A co-culture
macromolecules have been shown to impact of microalgae with filamentous fungi is being
digestion. Therefore, it is essential to determine developed to facilitate harvesting and minimize
the digestibility and composition of microalgae microalgal biomass costs. Eco-friendly aquaculture
for certain species of fish. systems based on microalgae, zooplankton, and
The energy-intensive harvesting stage of fish/shrimp are also gaining importance in the
microalgae biomass results in high production circular bio-economy. A critical analysis revealed
costs. Microalgal biorefinery approaches might that bioengineering tools and strategies could sig
reduce microalgal feed component manufactur nificantly improve the development of aquacul
ing costs. This approach extracts high-value bio ture-based food production. Additionally, the
chemical components from biomass, such as study also provides new insights into the chal
lenges and potential breakthroughs regarding
lipids and carotenoids, which are then used in
microalgae biomass production and bioproducts
fish and animal feed. Supercritical fluid extrac
as sustainable ingredients in the future.
tion and organic solvent extraction in combina
tion are effective for defatting biomass. Although
defatted biomass from the aforementioned bior
Acknowledgements
efinery processes has been studied, no reports on
saponified biomass have been published. The We would like to express our gratitude to Khalifa University,
saponification-based biorefinery produces caro Abu Dhabi, United Arab Emirates, for their assistance via the
grant CIRA-2018-27.
tenoids and other value-added microalgae pro
ducts. The residual biomass may be used as fish
feed. Therefore, the residual biomass of biorefin
ery methods must be examined for digestion in Disclosure statement
fish. No potential conflict of interest was reported by the author
(s).
7. Conclusions
Due to the depletion and pollution of marine Funding
resources, aquaculture is increasingly turning to The author(s) reported there is no funding associated with
microalgae as a substitute for fishmeal. the work featured in this article.
ORCID juvenile Atlantic salmon. PLoS One. 2015;10(4):

e0124042. DOI:10.1371/journal.pone.0124042.
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Shadi W. Hassan http://orcid.org/0000-0002-7269-094X microalgae as a sustainable feed ingredient for aqua
Fawzi Banat http://orcid.org/0000-0002-7646-5918 culture. J Biotechnol. 2021;341:1–20.
[14] Kusmayadi A, Leong YK, Yen H-W, et al. Microalgae
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An Overview of Microalgae Biomass As A Sustainable

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BIOENGINEERED

2022, VOL. 13, NO. 4, 9521–9547

An overview of microalgae biomass as a sustainable aquaculture feed

ABSTRACT ARTICLE HISTORY

Table 1. Presents the benefits and drawbacks of an alternative fish diet.

Blood meal ● The protein content is high. ● Methionine deficiency [28,29]

Figure 2. Metabolites produced by microalgae during their photosynthetic activity.

Table 2. Nutrient content of several microalgae species.

Table 3. Microalgal biomass as an alternative or supplement to fishmeal and fish oil.

3. Nutrients digestibility been supported by any other study. Therefore,

digestibility of microalgal biomass for use in fish/ 3.3 Anti-nutritional factors

composition of extruded fish feed is acknowledged Astaxanthin, polyunsaturated fatty acids

ORCID juvenile Atlantic salmon. PLoS One. 2015;10(4):

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