applied

sciences
Review
Current Status and Economic Prospects of Alternative Protein
Sources for the Food Industry
Fábio Medeiros 1 , Ricardo S. Aleman 2 , Lucia Gabríny 3 , Seung Woon You 1 , Roberta Targino Hoskin 1
and Marvin Moncada 1, *

1 Plants for Human Health Institute, Department of Food, Bioprocessing and Nutrition Sciences,
North Carolina State University, 600 Laureate Way, Kannapolis, NC 28081, USA
2 School of Nutrition and Food Sciences, Louisiana State University Agricultural Center,
Baton Rouge, LA 70803, USA
3 AgroBioTech Research Centre, Slovak University of Agriculture, Tr. A. Hlinku 2, 949 76 Nitra, Slovakia;
[email protected]
* Correspondence: [email protected]

Abstract: The rising demand for novel and alternative protein (AP) sources has transformed both the
marketplace and the food industry. This solid trend is driven by social awareness about environmental
sustainability, fair food production practices, affordability, and pursuit of high-quality nutritional
sources. This short review provides an overview of key aspects of promising AP sources (plants, algae,
insects, fungi and cultured protein) as well as the economic potential, prospects, and operational
challenges of this market. The low environmental performance of livestock production, associated
with high GHG emissions and land use, can be overcome by less resource-intensive AP production.
However, despite the forecasted expansion and improved economic viability, key challenges such as
regulatory concerns, consumer acceptance and product functionality still need to be addressed. While
the consumption and production of plant-based products are relatively well established, research and
development efforts are needed to remediate the main commercialization and manufacturing issues
of unprecedented protein sources such as cultured protein and the emerging edible insects sector.

Keywords: global food chain; food production; market trends; sustainability; protein-rich products
Citation: Medeiros, F.; Aleman, R.S.;
Gabríny, L.; You, S.W.; Hoskin, R.T.;
Moncada, M. Current Status and
Economic Prospects of Alternative
Protein Sources for the Food Industry. 1. Introduction
Appl. Sci. 2024, 14, 3733. https:// Protein intake is commonly associated with animal-derived foods such as meat, shell-
doi.org/10.3390/app14093733 fish, fish, dairy products, and eggs. These traditional protein sources cover general nutri-
Academic Editors: Yiying Zhao,
tional requirements and have been the most produced and consumed throughout history
Wei Wang, Lei Zhou and Chu Zhang (Aiking & de Boer, 2020) [1]. However, there is an increasing social awareness about food
production, adequate nutrition, and environmental sustainability. The greenhouse gases
Received: 28 March 2024 (GHG) emissions associated to agriculture, forestry and other land uses have doubled in the
Revised: 22 April 2024 last 50 years (Deprá et al., 2022; Shabir et al., 2023) [2,3], but the current resource-intensive
Accepted: 25 April 2024
food production model still does not provide adequate nutrition to around 3.37 billion
Published: 27 April 2024
people (Amato et al., 2023; Cucurachi et al., 2019; UN, 2023a) [4–6]. In addition, solid
evidence has shown that high intake of animal protein is associated with higher risk of
cardiovascular disease, diabetes, and mortality (Hemler & Hu, 2019) [7]. Moreover, ethical
Copyright: © 2024 by the authors.
concerns towards animal production could also stifle demand in the Western world (Grahl
Licensee MDPI, Basel, Switzerland.
et al., 2020) [8]. Pressing issues about the sustainability of food production are expressed in
This article is an open access article the United Nations Sustainable Development Goals (UN SDG), which highlights concerns
distributed under the terms and related to food security, clean water supply, responsible consumption, and production,
conditions of the Creative Commons among others (UN, 2023b) [9].
Attribution (CC BY) license (https:// This scenario stimulates consumer interest for alternative protein (AP) sources as a
creativecommons.org/licenses/by/ way of providing sustainable, affordable, and nutritional options to replace or partially
4.0/). substitute for current mainstream food products (Amato et al., 2023; Bashi et al., 2019;

Appl. Sci. 2024, 14, 3733. https://doi.org/10.3390/app14093733 https://www.mdpi.com/journal/applsci

Appl. Sci. 2024, 14, x FOR PEER REVIEW 2 of 16

Appl. Sci. 2024, 14, 3733 This scenario stimulates consumer interest for alternative protein (AP) sources2 of as15a
way of providing sustainable, affordable, and nutritional options to replace or partially
substitute for current mainstream food products (Amato et al., 2023; Bashi et al., 2019;
Grossmann &
Grossmann & Weiss,
Weiss, 2021)
2021) [4,10,11].
[4,10,11]. Also, a significant
significant portion
portion of thethe population
population is is now
now
more inclined
more inclined to adopt vegetarian,
vegetarian, flexitarian
flexitarian andand vegan
vegandiets
diets(Kent
(Kentetetal.,
al.,2022)
2022)[12].
[12].AsAsa
aresult, this
result, shift
this in consumer
shift in consumer demand
demandis the
is biggest traction
the biggest in thein
traction market now. The
the market now.current
The
AP market,
current estimated
AP market, to be around
estimated USD 2.2
to be around USDbn,2.2
is expected to continue
bn, is expected growing
to continue to reach
growing to
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reach size ofsize
a market more ofthan
moreUSDthan150 bn by
USD 1502030
bn by(Figure
2030 1). The steep
(Figure sales
1). The growth
steep salestendency
growth
of meat-free
tendency products products
of meat-free and on market
and onvalue
marketof plant-based food companies
value of plant-based point to apoint
food companies con-
to a consumer
sumer market
market that that is to
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cross tothecross
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bridge from traditional
traditional animal-protein
animal-protein oriented
oriented
diets intodiets into diets
AP-rich AP-rich diets et
(Talwar (Talwar et al.,
al., 2024) 2024) [13].
[13].

Figure 1.
Figure 1. Projection
Projection of
of alternative
alternative protein
protein market
market growth.
growth. Adapted
Adapted from
from EY
EY Parthenon
Parthenon Analysis
Analysis
(2021) [14].
(2021) [14].

Althoughopportunities
Although opportunitiesfor forAP
AP sources
sourcessuch
such asas plants,
plants, insects
insects and
and other
other novel
novel protein
protein
sources have
sources have skyrocketed
skyrocketed in in both
both developing
developing and advanced
advanced markets (Bashi et al., 2019;
Lähteenmäki-Uutela
Lähteenmäki-Uutela et al., 2021) [10,15], studies studies have
have demonstrated
demonstrated the resistance
resistance ofof aa
significant
significantshare
shareofofthe population
the population to AP sources
to AP sources(Balfany et al.,et2023)
(Balfany [16]. Food
al., 2023) [16]. neophobia
Food neo-
(the unwillingness
phobia to try new
(the unwillingness foods),
to try newand the and
foods), sensethe
of sense
prejudiced disgust towards
of prejudiced specific
disgust towards
alternative protein sectors,
specific alternative proteinsuch as insect-based
sectors, and cultured
such as insect-based andproteins,
culturedhave been have
proteins, identified
been
as the two as
identified main
the factors
two main hindering market growth
factors hindering market (Siegrist
growth&(Siegrist
Hartmann, 2023; Wood
& Hartmann, &
2023;
Tavan, 2022) [17,18]. Undesirable organoleptic attributes such as off-flavors
Wood & Tavan, 2022) [17,18]. Undesirable organoleptic attributes such as off-flavors and and unpleas-
ant texture, texture,
unpleasant have also been
have alsoraised
been as obstacles
raised for thefor
as obstacles larger acceptance
the larger of APof(Malek
acceptance AP (Ma-&
Umberger, 2023; Mancini & Antonioli, 2019) [19,20]. In this sense, the
lek & Umberger, 2023; Mancini & Antonioli, 2019) [19,20]. In this sense, the market interest market interest has
driven research
has driven and and
research development
development towards overcoming
towards overcoming these challenges
these challengesandand
warranting
warrant-
wide access
ing wide and and
access acceptability of AP.
acceptability of AP.
Therefore,
Therefore, thisthis short
short review
reviewbrings
bringsan anoverview
overview ofofthethe fundamental
fundamental aspects
aspects about
about the
the composition and applications of promising AP sources (plants,
composition and applications of promising AP sources (plants, algae, insects, fungi algae, insects, fungi
and
and cultured),
cultured), as well
as well as the
as the economic
economic potentialand
potential andperspectives
perspectivesofofthis this emerging
emerging protein
protein
market.
market. It also identifies key aspects of currently available technologies and
It also identifies key aspects of currently available technologies and foreseeable
foreseeable
challenges
challenges in in the
the future
future of
of alternative
alternative proteins.
proteins.
2. Alternative Protein Sources
2. Alternative Protein Sources
2.1. Plant-Based
2.1. Plant-Based
Plant-based (PB) proteins are derived from vegetable sources, commonly legumes,
Plant-based
grains, nuts, and (PB)
seedsproteins
(Malekare derived from
& Umberger, vegetable
2023) sources,
[19]. Among commonly
these, legumes,
soybeans, peas,
grains, nuts, and seeds (Malek & Umberger, 2023) [19]. Among these, soybeans,
chickpeas, beans, rice, lentils and almonds are the most widely used (Gomes & Sobral, peas,
chickpeas, beans, rice, lentils and almonds are the most widely used (Gomes &
2022) [21]. In addition, oilseeds are also regarded as potential sources since protein-Sobral,
rich meals (press cakes from oil extraction) are low-cost by-products from oil production
(sunflower, rapeseed and sesame, for example), which can potentially be upcycled for
Appl. Sci. 2024, 14, 3733 3 of 15

protein production (Nicholson, 2022) [22]. PB proteins have rapidly become popular as the
most readily available and easily accessible alternative to animal-based protein. Although
animal-sources are usually perceived as the most common protein source, plant proteins
account for an average of 60% of the global protein supply, with a significant geographic
dependence, ranging from as low as 35% in Northern America, to as high as 84% in Western
Africa (Roser et al., 2023) [23].
Protein profile and content are highly dependent not only on the PB source, but also
on factors associated with plant growth conditions. Protein content in plant sources vary
widely from as low as 2.9 g/100 g fresh weight (FW), for spinach and other green leaves,
to 13.0–23 g/100 g FW, for common pulse sources such as peas and chickpeas (Balfany
et al., 2023) [16]. Although proteins are a key part of human diet, the amount of protein
ingested is not the only determinant influencing biological functions. Protein profile and
digestibility also play a major role in the nutritional effects derived from protein intake.
While animal-based protein sources, such as meat and eggs, have a complete profile of
essential amino acids (histidine, isoleucine, leucine, lysine, methionine, phenylalanine,
threonine, tryptophan and valine), PB proteins usually lack select essential amino acids,
which compromises their biological utilization (Maestri et al., 2016; Mariotti & Gardner,
2019; McCusker et al., 2014) [24–26]. Meat and eggs rank between 95–98% for digestibility,
with a protein digestibility corrected amino acid score (PDCAAS) around 96–100%, while
the digestibility of PB proteins is 97%, 89% and 64% for soybean, chickpea and kidney
beans, with PDCAAS of 100%, 78% and 68%, respectively (Jiménez-Munoz et al., 2021;
Kent et al., 2022) [12,27].
On the other hand, plant-based proteins have been associated with reduced allergenic-
ity (when compared to dairy and eggs), and with biologically active peptides linked to
enhancement of gastrointestinal health, and anti-hypertensive and anti-microbial activ-
ities (Balfany et al., 2023) [16]. For instance, buckwheat, beans and peas have bioactive
peptides with in vitro anti-hypertensive activity, while potent antioxidant activity (both
metal chelating and radical scavenging) has been attributed to cocoa, walnuts, rye and
maize (McCusker et al., 2014; Samarakoon & Jeon, 2012; B. P. Singh et al., 2014) [26,28,29].
The combination of different sources of plant proteins and the use of PB products (flours,
concentrates and isolates) is an efficient strategy to meet the recommended daily allowance
(RDA) of protein and essential amino acid intake in animal-free diets.
Functionality and sensory performance of plant-based proteins are decisive factors
for the use of AP for product development. Indeed, technological attributes such as
emulsification, foaming and gelation capacities are essential aspects for processing and
product development, although flavor and texture are still the biggest drivers for consumer
acceptance of novel PB products (Ma et al., 2022) [30]. Functional attributes are related to
physical and chemical properties that affect their interaction with other food components
during product development, processing and storage (Akharume et al., 2021; Grossmann &
Weiss, 2021; Jiménez-Munoz et al., 2021) [11,27,31]. Both intrinsic (protein content, amino
acid profile, overall net charge, isoelectric point) and extrinsic (temperature, pressure, pH)
factors govern the interaction between proteins and other molecules in food models (Pérez-
Vila et al., 2022) [32]. Off-flavors, such as beany or grassy tastes, are usually perceived in
PB proteins derived from pulses and green leaves, even after hydrolysis and concentration
(Adámek et al., 2018; Mariotti & Gardner, 2019) [25,33]. While studies have shown that
off-flavors might be related either with the binding of proteins to flavor-inducing molecules
under processing conditions (usually pH and temperature) during isolation/concentration,
or with the oxidation of polyunsaturated fatty-acids from the plant sources (Pérez-Vila
et al., 2022) [32], product development strategies are usually necessary to either reduce or
mask inherited flavors. Texture, on the other hand, is dependent not only on the protein
technological attributes, but also on the overall food model characteristics. Wettability,
water/oil adsorption, foaming, gelation and emulsification capacities, as well as the ability
of the protein to maintain these attributes during storage, shape the textural properties of
the proteins in food model systems. Studies have shown that process conditions such as
Appl. Sci. 2024, 14, 3733 4 of 15

temperature and pH have major impact on plant proteins solubility and, consequently, on
their ability to interact with oil droplets, air bubbles and create stable films, for instance
(Ma et al., 2022) [30].
The life cycle of plant-based proteins; however, is highly geographically dependent and
influenced by national agricultural and transportation policies (Cucurachi et al., 2019) [5].
The technological development of high yielding crops, such as soybean and lentils, for
example, is crucial for more efficient land use and water consumption (Shabir et al., 2023) [3].
Furthermore, technological development of crops and enhanced agricultural practices are
expected to drive a significant change in the PB protein supply chain (Aimutis, 2022; Amato
et al., 2023; Aschemann-Witzel et al., 2020) [34,35].

2.2. Algae
Algae-based protein sources are associated with seaweeds and microalgae and have
gained increased attention in the food industry over the last decade (Gkarane et al.,
2020) [36]. Both seaweeds and microalgae are photosynthetic, oxygen-producing organisms,
but seaweeds are macroscopic, multicellular, marine species of algae, while microalgae
are unicellular, microscopic forms. Algae species used for protein production are known
to contain protein concentrations similar to animal-based sources such as meat, egg and
milk (McCusker et al., 2014) [26], while delivering 4–30 times higher protein productivity,
when compared to plant-based protein sources such as soybeans and pulses, and requiring
far less resources (Thiviya et al., 2022) [37]. In fact, marine seaweeds, for example, do not
require fresh water or arable land to grow (Birch et al., 2019; Rawiwan et al., 2022) [38,39].
Among seaweeds, brown, green and red algae are commonly used for human con-
sumption, especially in East Asia. According to the Food and Agriculture Organization
of the United Nations (FAO, 2023) [40], the Republic of Korea is the largest consumer
market of algae, followed by China and Japan. High-protein seaweed species such as
Undaria pinnatifida (brown algae, 24 g protein/100 g dry weight), Ulva lactuca (green algae,
32.7 g/100 g DW) and Porphyra spp. (red algae, 50 g/100 g DW) are considered promising
candidates for alternative protein production. Among microalgae, Arthrospira platensis, also
known as Spirulina, and Chlorella vulgaris are the most commonly cultivated species, with
multiple applications as functional foods and generally regarded as safe (GRAS) both by
the Food and Drug Administration (FDA) and the European Food Safety Authority (EFSA)
(Lucakova et al., 2022; Thiviya et al., 2022; Wang et al., 2021) [37,41,42]. In addition to the
relatively high protein content, algae-based protein is considered a favorable source of
essential amino acids, comparable to eggs and soybeans (Eilam et al., 2023; Rawiwan et al.,
2022) [39,43].
Given that the amino acid profile is highly dependent on species, growth parameters
and phase, algae-based protein generally meets the essential amino acids standards. On the
other hand, it has been associated with lower digestibility, when compared to traditional
sources of plant-based and animal-based proteins (McCusker et al., 2014; Samarakoon &
Jeon, 2012) [26,28]. De Bhowmick and Hayes (2022) [44] evaluated the in vitro digestibility
of different species of seaweed and reported PDCAAS levels ranging from 8%, for Fucus
vesiculosus (brown algae), to 69%, for Palmaria palmata (red algae). Among microalgae, the
two most common species, Spirulina and Chlorella spp., are reported to present PDCAAS
of around 77–84% (De Bhowmick & Hayes, 2022; Wang et al., 2021) [42,44]. However, the
digestibility of algae protein is still poorly described in literature, since the shift of algae
utilization as a source of proteins is relatively recent.
The technological attributes of algae-based proteins are comparable to traditional
plant-based proteins, which contributes to their introduction in traditional food model
systems (López-Pedrouso et al., 2020; Moura et al., 2022) [45,46]. Solubility, foaming,
emulsification and gelation capacities, for example, have been found to be pH-dependent,
which is also the case for most proteins from plant and animal sources, with a lower
performance under acidic conditions closer to the isoelectric point, and favored under
alkaline conditions, where the proximity to the pKa of amine groups favors solubility and
Appl. Sci. 2024, 14, 3733 5 of 15

decreases hydrophobic repulsion (Bleakley & Hayes, 2017; Gómez et al., 2019; Strauch &
Lila, 2021) [47–49]. On the other hand, the presence of di- and trivalent cations, such as
Ca2+ and Al3+ , can negatively affect the performance of both microalgae and algae-based
proteins in food applications, since they can promote flocculation by ion bridging the
usually negatively charged surfaces of algae-derived products (Gkarane et al., 2020; Gómez
et al., 2019) [36,48].

2.3. Edible Insects

Insects have been historically consumed as part of the human diet for centuries, es-
pecially by African and Asian cultures (De Castro et al., 2018) [50]. Insect proteins have
recently received increasing attention from an environmental perspective, especially for
their efficient energy conversion to biomass that exceeds many livestock, which makes
them produce protein more efficiently (Chéreau et al., 2016; Yi et al., 2013) [51,52]. Cur-
rently, over 2000 insect species have been identified as edible (Mishyna et al., 2021) [53] and
commonly studied edible insects include mealworms (Tenebrio molitor), superworm (Zo-
phobas morio), lesser mealworms (Alphitobius diaperinus), house crickets (Acheta domesticus),
dubia roaches (Blaptica dubia), black soldier flies (Hermetia ilucens) and western honey bees
(Apis mellifera). Edible insects show high protein content with suitable amino acid profiles
and studies confirm that their nutritional profile is suitable according to FAO standards
(FAO/WHO/UNU, 2007) [54]. On average, they have 40–75% protein by weight, on a dry
basis (Verkerk et al., 2007) [55], including A. domesticus (32.6%), T. molitor (38.3%), beetles
(42.2%) (Huang et al., 2019) [56], grasshoppers (61–77%) and butterflies (15–60%) (Verkerk
et al., 2007) [55]. H. ilucens has a high content of essential aromatic amino acids (pheny-
lalanine and tyrosine) and sulfur essential amino acids (methionine and cysteine). On the
other hand, T. molitor, Z. morio, A. diapernius, A. domesticus and B. dubia were reported to
be deficient in tryptophan, valine, isoleucine and threonine, three of which are essential
amino acids (Yi et al., 2013) [52].
Industrial processing of edible insects mostly follows three main steps: pre-treatment,
drying and formulation (Liang et al., 2024) [57]. Following harvest, pre-treatments generally
intend to decrease microbial contamination and enzyme activity for the purpose of ensuring
food safety and avoiding possible lipid oxidation (Laroche et al., 2019) [58]. Blanching and
freezing are the most cited pre-treatment options applied to common insect protein sources
(Hermetia illucens, Tenebrio molitor, Acheta domesticus), with blanching being featured as the
most cost-effective (Singh et al., 2020; Yongkang et al., 2020) [59,60]. Drying is an important
step for increasing the shelf-life of edible insects. Traditional solar drying has been replaced
by industrially preferred convection oven drying. Freeze drying and microwave drying
have also been investigated, but the high costs associated with equipment and operation
hinder their wide implementation (Liang et al., 2024) [57]. Finally, the formulation step
varies with the intended application of the insect-based product and it goes from simple
grinding and bleaching, for the production of protein-rich insect flours, to alkaline protein
extraction (protein-rich ingredients) or solvent-based solid-liquid extraction (insect oil or
chitin production) (Liang et al., 2024) [57].
Reports on functionality of insect proteins vary with species and are scattered. Defat-
ting is a common pre-treatment, and the choice of defatting solvent, as well as the choice of
extraction procedure affect the solubility and functionality of insect protein (Queiroz et al.,
2023) [61]. Mishyna et al. (2021) [53] demonstrated that protein concentrates of mealworm
(T. molitor), cricket (Gryllodes sigillatus), and locust (Schistocerca gregaria) had water-holding
capacity and oil-holding capacity up to 300% greater than their respective raw flours. H.
ilucens was reported to have a great foaming capacity, increasing up to 1080% after thermal
treatment (Queiroz et al., 2023) [61], while A. mellifera (Mishyna et al., 2019) [62], A. domesti-
cus, T. molitor, Z. morio, A. diaperinus, B. dubia exhibited poor foaming properties (Yi et al.,
2013) [52]. On the other hand, A. domesticus exhibited far better gelling properties than
other insects (Queiroz et al., 2023) [61]. Due to the wide range of functionalities between
Appl. Sci. 2024, 14, 3733 6 of 15

species, insect protein needs to be screened for their functionalities before their use in
food formulations.
Currently, there are two major challenges for the insect protein industry. Despite the
history of consumption, insects are commonly deemed as dirty and unpleasant, which is a
serious impediment for their commercialization (Alhujaili et al., 2023; Caparros Megido
et al., 2016; De Castro et al., 2018) [50,63,64]. In addition, insect protein has been reported
to negatively affect the sensory properties and certain physical properties when used
as ingredients in either carbohydrate-based foods (pasta, cereal, bread) or animal food
analogues (ice cream, sausage, jerky) (Kim et al., 2022) [65]. For example, current problems
with T. molitor protein are the particle size and salt content, which affects the appearance,
texture, and perceived coarseness (Wendin et al., 2019) [66] of the final product. Addition
of T. molitor in biscuits led to negative sensory scores (Biró et al., 2020) [67]. Similarly,
the use of cricket powder in protein bars and energy bars were associated with lower
liking scores (Adámek et al., 2018) [33]. However, insect protein hydrolysates have shown
promising results. The addition of cricket and locust protein hydrolysates significantly
improved the sensory quality, microbiological characteristics and antioxidant properties
of aged cheese and goat or sheep meat emulsions (Lone, Bhat, Aït-Kaddour, et al., 2023;
Lone, Bhat, Kumar, et al., 2023; Singh et al., 2023) [68–70]. Reports on the incorporation of
insect protein into food products are still incipient, but there is a pressing need to invest
in research and development strategies to enable the offer of appealing insect-derived
products in the market.

2.4. Cultured (Cell-Based) Protein

Cultured protein, also known as cell-based, in vitro or lab-grown protein, refers to
protein products obtained from the cultivation of mammalian cells under controlled condi-
tions (Mancini & Antonioli, 2019) [20]. Developed by the biopharmaceutical industry and
usually implemented for monoclonal antibodies and vaccine production, the technology
for mammalian cell cultivation has secured significant investments both from industry and
academia, aiming to a rapid technological advance on protein production. However, this
protein segment is still in its infancy, as the first patents and technical reports addressed
this subject around the year 2000. In 2013, the first cultured beef was tasted in London
(Hadi & Brightwell, 2021) [71].
Significant improvements in product quality and affordability have occurred along the
last decade, from around USD 1.2 million per pound, in 2013, to around USD 50 per pound,
in 2021 (Chriki & Hocquette, 2020; Specht, 2020) [72,73]. The process of producing cultured
protein can be divided into four main steps: (1) The mammalian stem cells are harvested
from an animal via biopsy, followed by mechanical and enzymatic digestion for isolation;
(2) The cells are proliferated in a culture medium to increase the number/concentration of
viable cells; (3) The cells are differentiated into skeletal muscle cells and fibers; (4) The cells
can be assembled in scaffolding materials to deliver a final meat product (Post, 2014) [74].
Two main types of stem cells are considered suitable for protein production: skeletal
muscle cells, also known as satellite cells, and embryonic stem cells (Bogliotti et al., 2018;
Post et al., 2020) [75,76]. When satellite cells are used, the final product is composed only
of skeletal muscle fibers and do not resemble the complexity of animal-based protein,
which includes other components such as fat and connective tissue, leading to a decrease
in consumer appeal and overall quality (Bryant & Barnett, 2020; Bryant & Sanctorum,
2021) [77,78]. A highly complex meat structure can be achieved with embryonic stem cells,
which are able to be further differentiated into different tissues including intramuscular fat
and connective tissues, via adipogenesis and fibrogenesis, respectively, and microvascular
network, via vascularization (Ben-Arye & Levenberg, 2019) [79].
Considering that they come from mammalian cells, cultured protein is expected to
deliver the same structure, taste, texture, and functionality of animal-based protein (Chriki
& Hocquette, 2020; Mancini & Antonioli, 2019; Onwezen et al., 2021) [20,72,80]. Studies
have shown that cultured protein digestibility scores high in the PDCAAS test (around
Appl. Sci. 2024, 14, 3733 7 of 15

92%), with an amino acid balance comparable to beef. However, there is little information
on the presence and bioavailability of minerals, such as iron, zinc and selenium, and
vitamins, such as A, B12, D and E, commonly found in animal-sourced proteins (Hurrell &
Egli, 2010) [81]. Consumer acceptance of cultured protein is still hampered by taste and
texture, which are still far from being considered optimal, when directly compared to them
by consumers (Bryant & Sanctorum, 2021) [78]. The food industry has targeted comminuted
products such as burgers, minced beef and nuggets, for example, in order to overcome
texture issues (Mancini & Antonioli, 2019; Onwezen et al., 2021) [20,80]. Furthermore,
consumer preference studies have shown that the perception of “unnatural” production,
coupled with concerns with food safety and proper production regulations, can generally
lead to a sense of disgust and neophobia that increases consumer resistance to cultured
meat proteins (Bryant & Barnett, 2020) [77].
Scalability and environmental performance remain the biggest challenges in the im-
plementation of cultured protein (Specht, 2020) [73]. Because it involves unprecedented
and yet to be established techniques, there is a lack of previous operational data and
economically efficient cultivation protocols on a larger scale. Moreover, the high demand
of resources, especially electricity and water use, influence both the production and the
consumer acceptance, and still require improvement before cultured proteins get to the
commercial level (Bashi et al., 2019) [10].

2.5. Fungi
Mushrooms have received renewed interest due to their nutritional properties and high
protein content but especially for their sensory profile which is acceptable for their use in
meat analogues (You et al., 2022) [82]. Their relatively simple production protocol using low
value materials such as sawdust and branches (Okuda, 2022) [83] justifies why fungi-based
proteins are a steep-growing trend in the food sector. Indeed, their estimated consumption
exceeds 12.7 million tons (El-Ramady et al., 2022) [84]. The cost of industrial mushroom
production can be as low as USD 0.026 per fresh kg, or even lower USD 0.014 cents per fresh
kg, with optimized heating and cooling systems (Beghi et al., 2020) [85]. The growth period,
depending on the species, can be as short as 42 days to produce 25,000 kg of fresh white
button mushrooms (A. bisporus). The biological efficiency of converting growth substrate to
mushroom biomass can be as high as 250%, which is an impressive efficiency compared to
other crops where 80–90% of biomass is left unused after harvest (Chiu et al., 2000) [86].
Their high protein content (average 27.5% on a dry basis) and complete amino acid
profile by FAO standards (Kalač, 2013) [87] differentiate them from most non-animal
proteins. For instance, individual chemical scores of nine essential amino acids of three
commonly consumed mushrooms (Agaricus bisporus, Pleurotus ostreatus and Lentinus edodes)
satisfy the FAO standards (FAO/WHO/UNU, 2007) [54]. Additionally, evidence shows
that mushroom protein has antioxidant activity (Goswami et al., 2021; Khongdetch et al.,
2022) [88,89] and therapeutical properties (Bovi et al., 2013; De Mejía & Prisecaru, 2005;
Ditamo et al., 2016) [90–92].
Despite these advantages, the incorporation of mushroom biomass or protein into food
matrices, and their processing into protein concentrates and food ingredients are recent.
Mushroom processing is generally divided into three main categories: (1) post-harvest
processing for fresh market, (2) processing into convenient canned, sauces or pickled
foods, and (3) extraction of high-value ingredients for functional and pharmaceutical
applications (Zhang et al., 2021) [93]. While post-harvest processing focuses on drying and
irradiation techniques to extend the shelf-life of fresh mushrooms (Barzee et al., 2021) [94],
the use of mushroom-derived ingredients is more developed for high-end pharmaceuti-
cal applications.
As AP source, the use of mushrooms in meat analogues such as nuggets or patties is
limited, but sensory acceptance has been positive when mushrooms were used as partial
replacement (You et al., 2022) [82]. Alkaline extraction followed by isoelectric precipitation
has been the only method tested so far for the extraction of mushroom protein aiming at the
 the use of mushroom-derived ingredients is more developed for high-end pharmaceutical
applications.
As AP source, the use of mushrooms in meat analogues such as nuggets or patties is
limited, but sensory acceptance has been positive when mushrooms were used as partial
replacement (You et al., 2022) [82]. Alkaline extraction followed by isoelectric precipitation
Appl. Sci. 2024, 14, 3733 8 of 15
has been the only method tested so far for the extraction of mushroom protein aiming at
the production of fungi-based protein concentrates. One interesting study analyzed pro-
teins of 50 different mushrooms
production under the
of fungi-based Osborne
protein classification,
concentrates. based on protein
One interesting solu- proteins of
study analyzed
50 different2001)
bility (Bauer Petrovska, mushrooms under theculinary
[95]. Common Osbornemushrooms
classification, based
such on protein
as white buttonsolubility (Bauer
mushrooms (A. Petrovska,
bisporus),2001)
shitake [95].mushrooms
Common culinary mushrooms
(L. edodes), and oystersuch as white (P.
mushrooms button
os- mushrooms
treatus) contain bisporus),
(A.only 20–30%shitake
globulin mushrooms (L. edodes),
(salt-soluble) fraction.and
Thisoyster mushrooms
contrasts with plant ostreatus) contain
(P.pro-
teins that largely contain storage globulin proteins (Chéreau et al., 2016) [51], that favor that largely
only 20–30% globulin (salt-soluble) fraction. This contrasts with plant proteins
containthrough
protein production storage isoelectric
globulin proteins (Chéreau
precipitation. Onetthe
al., contrary,
2016) [51],protein
that favor protein production
fractions
such as albuminthrough
(water isoelectric
soluble) precipitation. On the contrary,
are left unrecovered protein
in the liquid phasefractions
during such as albumin (water
protein
soluble)
production (Yang et al.,are left [96].
2022) unrecovered in the liquid phase during protein production (Yang et al.,
2022) [96].
3. Economic Potential and Market Perspectives of Alternative Protein
3. Economic Potential and Market Perspectives of Alternative Protein
3.1. Global Market
3.1. Global Market
New productsNew withproducts
protein-related claims (“high claims
with protein-related protein” or “source
(“high protein”of protein”)
or “sourceare of protein”) are
increasing in Africa/Middle East (+41%), Europe (+26%), Asia (+15%),
increasing in Africa/Middle East (+41%), Europe (+26%), Asia (+15%), Australasia (+12%),
Australasia (+12%),
Latin AmericaLatin
(+8%), and North
America (+8%), America
and North(+2%), between
America 2018 between
(+2%), and 20222018 (Research
and 2022 and(Research and
Markets, 2021a) [97]. The market value for AP reached USD 77.0 bn
Markets, 2021a) [97]. The market value for AP reached USD 77.0 bn globally, globally, in 2022, andin 2022, and it
it is expected to increase to USD 193.75 bn, by 2028. In the United States alone,
is expected to increase to USD 193.75 bn, by 2028. In the United States alone, the market the market
value corresponded to about USD
value corresponded to 1.1 bn,USD
about in 2020 (Vegconomist,
1.1 bn, 2021) [98]2021)
in 2020 (Vegconomist, and it[98]is and
ex- it is expected
pected to reach USD 10.1 bn, by 2027, with an annual growth rate of around
to reach USD 10.1 bn, by 2027, with an annual growth rate of around 9.7% (Research and 9.7% (Re-
search and Markets,
Markets, 2021a)
2021a) [97].
[97].Indeed,
Indeed,the theNorth
NorthAmerican
Americanmarketmarketisisthethebiggest
biggestsales
sales share on this
share on this protein segment, followed by Europe (Figure 2A).
protein segment, followed by Europe (Figure 2A). Other relevant Other relevant APAP markets
markets are China,
are China, Japan, andand
Japan, Canada,
Canada, withwitha growth forecast
a growth of 9.5%
forecast of (to
9.5%reach
(to USd
reach 351.4
USdmillion),
351.4 million), 7.1%,
7.1%, and 7.4%,and respectively, from 2020
7.4%, respectively, fromto 2020
2027.to In2027.
Europe, the German
In Europe, AP market
the German is ex- is expected
AP market
pected to grow to approximately
grow approximately 7.3%, while7.3%,the restthe
while of the
restEuropean market should
of the European marketreachshould reach USd
USd 351.4 million,
351.4by 2027 (Albrecht,
million, 2021) [99]. In
by 2027 (Albrecht, terms
2021) of AP
[99]. source,ofPB
In terms AP accounts
source, should
PB accounts should
continue dominating
continue the market up to
dominating the2029, followed
market up toby fungi-based
2029, followedand byalgae-based
fungi-basedpro- and algae-based
teins (Figure 2B).
proteins (Figure 2B).

Figure 2. Global alternative

Figure protein
2. Global market
alternative by (A)
protein region
market byand (B) protein
(A) region source.
and (B) Adapted
protein source. from
Adapted from MMR
MMR (2023) [100].
(2023) [100].

The revolutionThe revolution

resulting fromresulting
the growth from ofthe
APgrowth
market of AP market
could lead to could lead to the generation
the generation
of agriculture.
of novel jobs in novel jobs inSubstitute
agriculture. Substitute
foods foods might
might experience experience
a new growthawave
new growth
with wave with
technological advances such as data analytics, artificial intelligence,
technological advances such as data analytics, artificial intelligence, and biotechnology and biotechnology that
enable the creation of new foods using 3D printers and other
that enable the creation of new foods using 3D printers and other enhanced techniques enhanced techniques (Knorr
& Augustin,
(Knorr & Augustin, 2024;
2024; R. SinghR. Singh et al., [70,101].
et al., 2023) 2023) [70,101].

3.2. Plant-Based Protein

Different from other segments in the AP ecosystem, PB proteins are well known in the
marketplace. Due to its historical presence, the sector lies on well-established regulatory
grounds, and it is generally well accepted by consumers. In fact, the market demand for soy
and wheat protein precedes the recent renewed interest for plant-based proteins. Together
with rice, peas and chickpeas, they are now mainstream plant-based protein sources offered
in the global marketplace (Aimutis, 2022) [34].
 Different from other segments in the AP ecosystem, PB proteins are well known in
the marketplace. Due to its historical presence, the sector lies on well-established regula-
tory grounds, and it is generally well accepted by consumers. In fact, the market demand
for soy and wheat protein precedes the recent renewed interest for plant-based proteins.
Appl. Sci. 2024, 14, 3733 9 ofprotein
15
Together with rice, peas and chickpeas, they are now mainstream plant-based
sources offered in the global marketplace (Aimutis, 2022) [34].
The market growth depends on the producers’ ability to develop tasty, competitive,
The market growth
and versatile depends
products. on thefoods
Functional producers’ ability
and sports to develop
nutrition tasty, competitive,
are major segments of the
and PB
versatile products. Functional foods and sports nutrition are major
protein market share, and protein-rich powders, bars and ready-to-drink segments of the
beverages
PB protein market share, and protein-rich powders, bars and ready-to-drink beverages
are popular products in the United States (Figure 3). Moreover, the use of chickpeas, are
popular products in the United States (Figure 3). Moreover, the use of chickpeas, brown
brown rice, fava bean and mung bean protein is growing significantly as egg substitutes. rice,
fava Lentil,
bean and mung
wheat andbean protein
flaxseed is growing
proteins significantly
are gaining as egginsubstitutes.
popularity Lentil, wheat
products directed to athletic
and performance.
flaxseed proteins are gaining popularity in products directed to athletic performance.
Plant-based versions of much-loved animal-based products are now availa-
Plant-based versions of much-loved animal-based products are now available in grocery
ble in grocery stores. Food manufacturers, both small and large food companies world-
stores. Food manufacturers, both small and large food companies worldwide, are investing
wide, are investing in plant-based milk, eggs and meat substitutes. They are increasingly
in plant-based milk, eggs and meat substitutes. They are increasingly competitive among
competitive among animal-derived products in terms of taste, price, and accessibility. AT
animal-derived products in terms of taste, price, and accessibility. AT Kearney predicts that
Kearney predicts that by 2040, 60% of the consumed meat will come from alternative-
by 2040, 60% of the consumed meat will come from alternative-sourced meat substitutes,
sourced meat substitutes, 25% of which will be plant-based (AT Kearney, 2019) [102].
25% of which will be plant-based (AT Kearney, 2019) [102].

Figure 3. (A)3.US
Figure Plant-based
(A) proteinprotein
US Plant-based supplement market,market,
supplement (B) Global
(B)plant-based protein supplement
Global plant-based protein supple-
ment market share. Adapted from Grand View Research
market share. Adapted from Grand View Research (2023) [103]. (2023) [103].

The The
plant-based milkmilk
plant-based segment
segmentis theis most popular
the most category
popular among
category amongPB substitutes.
PB substitutes.
The The
market associated with this segment is around USD 2.5 bn, and
market associated with this segment is around USD 2.5 bn, and plant-based milkplant-based milksub-
substitutes alone correspond to 35% of the total market size of PB foods.
stitutes alone correspond to 35% of the total market size of PB foods. Refrigerated non- Refrigerated
non-dairy
dairy milk
milk makes
makesup upmost
mostsales
salesininthis
thissegment,
segment, ledled
by by
almond, oat,oat,
almond, andand
soy soy
milk.milk.
Oat Oat
milk,milk,
for example, has experienced
for example, has experienced massive
massivegrowth in recent
growth years,
in recent going
years, fromfrom
going a small
a small
market segment in 2018 to becoming the second most demanded plant-based
market segment in 2018 to becoming the second most demanded plant-based dairy prod- dairy product
in 2020
uct(GFI, 2021)
in 2020 [104].
(GFI, Although
2021) being a well-established
[104]. Although market, some
being a well-established novel
market, alternative
some novel alter-
milknative
products are now found on supermarket shelves such as
milk products are now found on supermarket shelves such as bananabanana milk and products
milk and
produced
productswithproduced
plant blends.
with plant blends.
The The
growth of plant-based
growth milkmilk
of plant-based substitutes, which
substitutes, is currently
which purchased
is currently purchasedby 39% of of
by 39%
American
Americanhouseholds has stimulated
households has stimulated the growth
the growthof other plant-based
of other dairy
plant-based substitutes,
dairy substitutes,
suchsuch
as ice cream, yogurt, butter, cheese, and eggs. Soy protein, one of
as ice cream, yogurt, butter, cheese, and eggs. Soy protein, one of the first the first leading
leading
ingredients heavily
ingredients usedused
heavily for alternative
for alternativeproducts, has decreased
products, at a compound
has decreased at a compound annual
annual
growth rate (CAGR) of 6% from 2004 to 2019. Conversely, interest in pea protein grew at
growth rate (CAGR) of 6% from 2004 to 2019. Conversely, interest in pea protein grew at
a CAGR of 30% during the same period (McKinsey, 2020) [10]. This trend is partly due
a CAGR of 30% during the same period (McKinsey, 2020) [10]. This trend is partly due to
to health concerns related to soy protein, such as allergenicity and estrogenic concerns.
health concerns related to soy protein, such as allergenicity and estrogenic concerns. The
The total market value associated with these alternative products amounted to USD 1.9 bn,
in 2020, and represents a combined sales growth of 28%, compared to 2019, and 55%,
compared to 2018.
In 2020, PB meat substitutes reached a market value of USD 1.4 bn, representing an
increase above USD 430 million in sales compared to the previous year and a sales growth
of 45%, between 2019 and 2020. The best-selling format of PB meat substitutes are burgers,
sausages, nuggets, tenders, and chops. PB refrigerated meat substitutes have been more
available to the public in animal-based meat sections in supermarkets, rather than in aisles
Appl. Sci. 2024, 14, 3733 10 of 15

dedicated to vegan, organic, or special dietary needs. This alone reflects significant changes
in both product innovation and marketing strategies.
Some interesting trends have been observed. For example, data shows that in the
United States, 98% of people who buy plant-based meat substitutes also buy conventional
meat (GFI, 2021) [104]. Products that try to replicate and emulate taste, texture, and
appearance of animal meat dominate sales in this segment, in contrast to non-similar
products such as black bean burgers and vegetarian hamburgers, whose sales are much
lower. PB beef is the best-selling substitute in the meat substitute subcategory, followed
by pork and chicken substitutes, but PB seafood (fish and shellfish) still needs competitive
proposals in the United States market (GFI, 2021) [104].

3.3. Insect Protein

Entomophagy as a possible solution for food insecurity and sustainable diets has
been studied and discussed for many years, but its actual implementation in Western
societies is still incipient. On the contrary, this type of protein source has been present
in Asian and African markets for a long time. The global market value for insect protein
represented around US$ 144 million, in 2021, and it is expected to reach USD 1.33 bn by
2026, with a growth rate of 45.7% per year (Market Data Forecast, 2021) [105]. Other market
forecasts; however, have more optimistic predictions. Barclays estimates that the insect
protein industry could be worth USD 8.0 bn by 2030 (Barclays, 2021) [106]. World-leading
food manufacturers such as Nestlé, PepsiCo and Tyson have started operations in this
sector (PitchBook, 2021) [107], and restaurants, supermarkets, and even the Seattle Baseball
Stadium are incorporating insect-based snacks.
Insect protein farming is less expensive than traditional animal protein sources (Market
Research Future, 2021) [108]. Besides pursuing technological advances to enable a more
efficient and appealing way to incorporate insect ingredients in traditional foods, and
addressing organoleptic and functional issues, a lot needs to be done in the legislation
arena. To take full advantage of the potential benefits of insect protein and expand markets,
regulatory efforts are crucial. Extensive revisions of the food regulations are ongoing at
this moment in several countries, and the future of the insect protein industry is highly
dependent on this (Sogari et al., 2023) [109].

3.4. Cultured Meats

Actual cultured meat products are still not marketed in the United States and several
other countries, which makes it difficult to build reliable market predictions. However,
some agencies expect a market size of around USD 140 million, by 2030, and USD 630 mil-
lion, by 2040, including 35% of the world’s demand for meat (AT Kearney, 2019) [102].
Cultured meat is expected to be in restaurants in the next years as this production method
is supposed to become cost-competitive in the next decade (Rabb, 2021) [110].
Optimistic growth prospects rely on the similarity of cultured meat to animal-sourced
meat regarding sensory and nutritional attributes (Jeffries, 2019) [111]. On the other
hand, high production costs, disputable consumer acceptance of cultured and genetically
modified foods, and sanitary regulations are some of the reasons why any realistic market
growth might only occur after 2030 (McKinsey, 2020) [10]. However, the recent marketing
authorization obtained by Eat Just in Singapore for cultured chicken meat after a lengthy
regulatory process could favor the approval of this type of meat in other countries (Poinski,
2021) [112]. In addition to meat grown for human consumption, there are companies
dedicated to developing farmed seafood products and pet food.

3.5. Fungi Proteins

Mycofermentation has become the third technology pillar of the AP revolution in
2020. Fermentation has been used to produce foods and drinks for thousands of years, but
2020 became a landmark for mycoprotein production, due to the notable increased use of
mycelia and mycoproteins (GFI, 2021) [104]. The global market for fermentation-based
Appl. Sci. 2024, 14, 3733 11 of 15

AP is expected to reach USD 422.26 million, by 2026, with an annual growth rate of 5%
between 2021 and 2026 (Research and Markets, 2021b) [113]. It is anticipated that the main
factors driving the growth of this market are food allergies and intolerances associated to
dairy protein, which have increased in recent years.
Fermented mycoproteins have a longer shelf life and environmental advantages,
such as reduced organic waste production (Research and Markets, 2021b) [113], and fast
processing time, since fermented protein can be obtained within a month through industrial
fermentation (Research and Markets, 2021b) [113]. Furthermore, fungi fermentation costs
are very moderate. To date, companies that have developed promising mycofermentation
technologies are very young and need more financial and personal resources. On the other
hand, a factor that could negatively affect the growth of mycoproteins is the regulatory
requirements for manufacturers to label these products, including the term “mold”, which
does not have a positive connotation among consumers (GFI, 2021) [104].

4. Conclusions and Outlook

The high demand for AP sources has motivated both the mainstream food industry
sector as well as new entrepreneurs to venture into this promising market. Current evi-
dence suggests that they are a viable alternative to animal proteins as they confer health
benefits, and use fewer natural resources, and comparably more environmentally friendly
processing techniques. The PB protein market is relatively well-established, but new supply
sources have been added to the industry portfolio to satisfy consumers avidness for novelty,
convenience, and sustainability, without compromising sensory attributes and affordabil-
ity. The forecast for the AP market predicts significant expansion and more competitive
products in the near future. Algae protein-based products perform well in specific regional
markets and show desirable high productivity with low resource input, making them
promising candidates for market expansion. However, despite the optimistic scenario and
large investment in this sector, key challenges still need to be addressed. Emerging protein
sources such as insect protein and cultured meats need advanced research and development
to improve product attributes and consumer acceptance. Moreover, regulatory issues and
allergenicity concerns have yet to be solved. The food industry needs to be aware of the
trends, challenges and opportunities taking place in the protein production arena. The
AP market not only promises economic prosperity but also leads to a broad societal shift
towards sustainable and ethical food production and consumption, and plays a pivotal role
in the future of the global food industry.

Funding: This publication was supported by the Operational Program Integrated Infrastructure
within the project “Demand-driven Research for the Sustainable and Innovative Food”, Drive4SIFood
313011V336, co-financed by the European Regional Development Fund.
Conflicts of Interest: The authors declare no conflicts of interest.

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