Annual Review of Food Science and Technology

Insects as an Alternative
Protein Source
Andrea M. Liceaga,1 José Eleazar Aguilar-Toalá,2
Belinda Vallejo-Cordoba,3 Aarón F. González-Córdova,3
and Adrián Hernández-Mendoza3
1
Protein Chemistry and Bioactive Peptides Laboratory, Department of Food Science, Purdue
University, West Lafayette, Indiana, USA; email: [email protected]
2
Laboratorio de Procesos de Transformación y Tecnologías Emergentes de Alimentos,
Departamento de Ingeniería y Tecnología, Facultad de Estudios Superiores Cuautitlán,
Universidad Nacional Autónoma de México, San Sebastián Xhala, Cuautitlán Izcalli, Estado de
México, México
3
Laboratorio de Calidad, Autenticidad y Trazabilidad de los Alimentos, Centro de Investigación
en Alimentación y Desarrollo A.C. (CIAD, A.C.), Hermosillo, Sonora, México

Annu. Rev. Food Sci. Technol. 2022. 13:19–34 Keywords

First published as a Review in Advance on
entomophagy, food security, insect protein, processing technologies, safety
October 26, 2021

The Annual Review of Food Science and Technology is Abstract

online at food.annualreviews.org
The recent COVID-19 pandemic drastically affected food supply chains
https://doi.org/10.1146/annurev-food-052720-
worldwide, showing the vulnerability of food security. Efforts to develop al-
112443
ternative protein sources that are sustainable and can help alleviate global
Copyright © 2022 by Annual Reviews.
food shortage problems should be prioritized. Insects have been part of our
All rights reserved
diet for thousands of years and still are today, and market trends show a
global increase in the number of food-grade insect producers. The global
market for edible insects has been forecasted to reach US$8 billion by the
year 2030. Insects are highly nutritious and have bioactive peptides with po-
tential therapeutic effects. This review provides an overview of the consump-
tion of insects from ancient to modern times, discusses the rationale for using
insects as alternative protein sources, and presents a summary of the major
insects consumed worldwide as well as a brief description of the traditional
and novel technologies currently used to process insects and/or extract their
nutritional components.

19
 INSECT PROTEIN FROM PAST TO PRESENT
Insect consumption (entomophagy) dates back thousands of years, before people, particularly
those living in warmer regions of the planet who had access to a year-round variety of insects,
had tools for hunting or farming (Imathiu 2020, Kouřimská & Adámková 2016). This is perhaps
one of the main reasons why insects are now primarily consumed in tropical countries in Africa,
Asia, and Latin America (Garofalo et al. 2019). In these countries, insects are an important compo-
nent of cultures, gastronomies, and staple diets. The countries with the highest reported number
of edible insect species include Mexico, China, Thailand, and India (Baiano 2020). One of the first
documented records of entomophagy dates back to Aristotle’s (384–322 BCE) Historia Animalium,
where he writes about the delicacy of eating cicadas; in Ancient Rome, cossus, a dish consisting of
larvae from the longhorn beetle, was highly coveted by its citizens (van Huis et al. 2013). Modern-
day historical records show that in 1557, Spanish Conquistadores noted that many insect species
were considered delicacies by the Aztecs. For example, the ahuauhtli, which are eggs deposited by
flies on the surface of stagnant waters, were highly prized by the Aztec Emperor Moctezuma II
(van Huis et al. 2013) and are still widely consumed in public markets in Mexico. In the United
States, it is estimated that 25–50% of Native American communities consume insects. For exam-
ple, the Kitanemuk of the western Mojave are known to eat red ants and caterpillars, and the Utah
Paiute are known to calendarize insect harvesting around the life cycles of larvae (Anderson 2006).
The Onondaga tribe used ants to add a citrus taste to dishes, and the Tlicho tribe allowed larvae
to grow on the surface of meat prior to cooking (Nicholas 2018). For a more detailed overview of
the culture, evolution, and sustainability of entomophagy, the reader is referred to Ramos-Elorduy
(2009).
Despite the strong evidence of entomophagy in ancient and modern times, the consumption
of insects in Western countries is uncommon or even considered culturally unacceptable (Baiano
2020, Shockley & Dossey 2014). In these countries, insects are mostly associated with plagues
or decaying matter. Additionally, the social and cultural characteristics such as neophobia and
lack of familiarity (e.g., to the taste and texture) by Western consumers are the main reasons
for the low acceptability of insects as food (Wendin & Nyberg 2021). Nevertheless, as efforts
increase to generate sustainable protein sources to meet the growing global demand for animal
protein, the consumption of edible insects has gained increased attention worldwide (Imathiu
2020). This growing interest in edible insects can be demonstrated by the increasing number of
scientific publications on the subject. In this context, Baiano (2020) utilized a major abstract and
citation database (Scopus) to find that 90% of the literature has been published from 2010, and
an online Google Scholar search for “edible insects” shows more than 147,000 results, with more
than 20,000 since the year 2017. Insects’ remarkable nutritional value (e.g., high-quality protein,
unsaturated fats, vitamins, minerals), comparable to, or even higher than, that of conventional
livestock, has placed them in the spotlight of the food and feed industry sectors. In the past 10 years,
several edible insect startup companies in Europe, North America, and South Asia have emerged.
Melgar-Lalanne et al. (2019) list more than 130 edible insect companies and/or commercial brands
worldwide.

RATIONALE FOR THE USE OF INSECTS AS A PROTEIN SOURCE

As stated above, entomophagy is not a new concept; evidence shows that humans have consumed
insects for thousands of years, and at present there are approximately 2 billion people in 113
countries who eat insects (Lesnik 2018). In fact, recent market trends show a global increase in the
number of food-grade insect producers, forecasting the global market for edible insects to reach
US$8 billion by the year 2030 (Glob. Mark. Insights 2020). In Europe and North America, market

20 Liceaga et al.
 Antioxidant High protein quality

Antiobesogenic Essential amino acids

ies

Nu
e propert

tritio
Edible insects

nal benefi
Antihypertensive Fiber

activ
Bio

ts
Anti-inflammatory

Mono- and
Antimicrobial polyunsaturated fats

Immunomodulatory Vitamins and minerals

Sustainability

↓ Impact natural resources High feed efficiency conversion ↓ Greenhouse gas emissions
↓ Water and land High fecundity ↓ CH4, CO2
and short life cycles
Figure 1
Rationale for using insects as a protein source. Edible insects are considered more sustainable protein sources and provide nutritional
(e.g., protein, fiber, fat, vitamins, and minerals) and health (e.g., antioxidant, antihypertensive, anti-inflammatory, etc.) benefits when
consumed. Spiders, including tarantulas, are classified as arachnids (Arachnida), not insects (Insecta). Both Arachnida and Insecta
belong to the phylum Arthropoda. Images sourced from depositphotos.com.

demand is predicted to grow more than 43% by the year 2024, and in the Asia Pacific market it
is estimated that the market will surpass US$270 million by 2024 (Baiano 2020). In modern-day
culture, the sustainability factors and nutritional and, more recently, health benefits of insects are
some of the main reasons for the gaining interest in insects as alternative sources for food and
feed production (Figure 1). The recent COVID-19 pandemic in 2020 drastically affected food
supply chains worldwide, showing the vulnerability of food security. Doi et al. (2021) highlight the
merits of modern entomophagy in a post-COVID-19 world, particularly due to insects’ low risk
for zoonotic disease transmission and high production rate and industrial output. These authors
elucidate how boosting consumption of edible insects could play an important role in establishing
and rebuilding global food security.

Sustainability
In 2009, several international organizations, including the World Bank, the Food and Agricul-
ture Organization (FAO), and the US Department of Agriculture (USDA), listed the increase of
food production without a further rise in greenhouse gas (GHG) emissions as a major critical
challenge to ensure food security (FAO 2006, Ingram et al. 2012). On July 17, 2019, the World
Research Institute released the World Resources Report: Creating a Sustainable Food Future, indicating
that world food demand will increase by 50% by 2050, with the demand for animal-based foods

www.annualreviews.org • Insects as a Protein Source 21

 increasing by 70%. To meet the rise in food demand, farmers must produce ∼60% more crop
calories (7,400 trillion calories) and increase land use by 593 million hectares (twice the size of
India) (Searchinger et al. 2019). In the United States, beef and dairy consumption remain one of
the primary dietary protein sources, with livestock using ∼43% of the agrarian land (more than
3.9 billion hectares) and responsible for ∼37% of methane gas emissions (Aarnink et al. 1995,
Goodland 2013, Losey & Vaughan 2006). These data confirm the need to develop an alterna-
tive agriculture system that considers the rising population and looks beyond conventional food
sources. Scientists have proposed that substituting at least 25% of protein from livestock with al-
ternative (more sustainable) proteins would allow the reforestation of agrarian land and reduce
4% or more of agricultural GHG emissions, equivalent to 23 million metric tons per year (EPA
2017, Searchinger et al. 2019).
Extensive research shows that edible insects require limited land and water use and produce
lower GHG emissions during their production (de Carvalho et al. 2020, Jantzen da Silva Lucas
et al. 2020). For example, crickets use 15 m2 of land to produce 1 kg of protein, whereas pigs
use 50 m2 and cattle use 200 m2 (Goodland 2013, van Huis et al. 2013). Furthermore, crickets
use 2,000 L of water to produce 1 kg of protein, whereas chicken, pork, and beef require 2,300;
3,500; and 20,000 L, respectively (Gahukar 2016). The efficiency conversion of ingested foods
(ECI) is a quantifiable metric of a farm’s productivity defined as the weight gained by an animal
per pound of food intake. The overall ECI of insects is higher than livestock. For example, house
crickets (Acheta domesticus) require on average 1.7 kg of feed to produce 1 kg of protein, whereas
5 kg and 10 kg of feed are required for pork and beef, respectively (Nakagaki & Defoliart 1991).
Other positive aspects of insect farming/production are their high fecundity (e.g., large numbers
of progeny per individual), rapid growth rates, and short life cycles as well as their low cost of
production compared to traditional animal protein sources (Shockley & Dossey 2014, Wendin &
Nyberg 2021).

Benefits of Insect Protein: Nutritional Quality and Therapeutic Applications

Insects’ nutritional composition consists mainly of protein, including all essential amino acids (20–
76% of dry matter); lipids (2–50% of dry matter); chitin (2.7 mg–49.8 mg/ kg of fresh matter);
and minerals such as calcium, phosphorous, potassium, and magnesium (Womeni et al. 2012).
Table 1 summarizes the nutrient composition of the major edible insect orders. A detailed nu-
tritional composition for 236 edible insects has been provided by Rumpold & Schlüter (2013). In
terms of protein quality, insect protein contains all the essential amino acids and has suitable di-
gestibility. The protein digestibility determined by true fecal nitrogen levels in rat models report
values between 86% and 90% for insects, which are higher than that of many plant-based pro-
teins but lower than those reported for other animal proteins (egg, 95%; beef, 98%; casein, 99%)
(Finke 2013). The slightly lower protein digestibility results from some of the protein bound to
the insects’ exoskeleton that makes it unavailable when consumed. In addition, studies have re-
ported that amino acids from insects such as silkworm larvae and crickets exceed the amino acid
requirements established by the FAO and World Health Organization (Hall et al. 2017, Wu et al.
2011). However, it is important to state that the variability of the protein profile depends on the
species, life cycle stage, diet composition, and preparation or processing (e.g., thermal and me-
chanical treatments) (Baiano 2020, van Huis 2013). For example, Eri silkworms (Samia ricinii) dis-
played a similar nutritional profile irrespective of their developmental stages (i.e., pre-pupae and
pupae) and feed source (Longvah et al. 2011). Conversely, mealworms (Tenebrio molitor) showed
differences in their protein and fatty acid profiles between larvae and adult stages (Finke 2002,
Ravzanaadii et al. 2012).

22 Liceaga et al.
 Along with their excellent nutritional quality, several studies report that through enzymatic
proteolysis, insect protein can be an excellent source of bioactive peptides. Zielińska et al. (2017)
reported the antioxidant activity of various edible insects (e.g., Blaptica dubia, Gromphadorhina por-
tentosa, Locusta migratoria, Amphiacusta annulipes, Zophobas morio) produced by enzymatic proteol-
ysis using pepsin and pancreatin. The authors found that peptides with the highest antioxidant
activity were those derived from A. annulipes (IC50 = 19.1 μg/mL, DPPH method) and Z. morio
(IC50 = 4.6 μg/mL, ABTS method). Hall et al. (2020) identified antihypertensive, antiglycemic,
and anti-inflammatory peptides derived from the gastrointestinal digests of cricket (Gryllodes sig-
illatus) peptides. They found that the cationic peptide fraction (<0.5 kDa) significantly decreased
(p < 0.05) α-amylase, α-glucosidase, and angiotensin-converting enzyme (ACE) activity in vitro
and inhibited the expression of NF-κB in RAW 264.7 macrophage cells. In addition, two identi-
fied pentapeptides (PHGAP and VGPPQ) exhibited a higher degree of noncovalent interactions
with the enzyme active site residues and binding energies. Tao et al. (2017) purified and identified
the peptide GNPWM from silkworm (Bombyx mori) pupa protein hydrolysates, which exhibited
ACE inhibitory activity (IC50 = 21.70 μM). Using molecular docking analysis, the authors es-
tablished that the inhibition mechanism was the interaction of the peptide with the active site of
ACE. There are a few in vivo studies on bioactive peptides derived from edible insects. For exam-
ple, antioxidant (Yuan et al. 2012), antihypertensive (Yuan et al. 2012), and antidiabetic (Han et al.
2016, Jung et al. 2010) effects of silkworm fibroin peptides have been reported in animal models.
Edible insect proteins are a promising sustainable source of bioactive peptides. Research suggests
that bioactive peptides could be used as components of functional or therapeutic food applications
and may contribute to health promotion by improving specific physiological functions.

TYPES OF INSECTS USED FOR HUMAN CONSUMPTION

There are approximately 1,900 identified edible insect species consumed worldwide. These in-
clude beetles, caterpillars, wasps, ants, bees, grasshoppers, locusts, crickets, cicadas, leafhoppers,
termites, dragonflies, and flies, among others. In Mexico alone, more than 500 species of edible
insects have been recorded (Ramos-Elorduy 2009). The majority of edible insects are classified
into eight orders (Table 1), with most of them being wild-harvested and only a smaller number

Table 1 List of major edible insect orders, their nutritional content, and number of species consumed worldwide
Number of species
Insect Order Examples % Protein % Fat % Fiber consumed worldwide
Blattodea Cockroaches, 35.3 29.9 5.3 96
termitesa
Coleoptera Beetles, grubs 40.7 33.4 10.7 659
Diptera Flies 49.5 22.8 13.6 37
Hemiptera Cicadas, stink bugs 48.3 30.3 12.4 237
Hymenoptera Bees, wasps, ants 46.5 25.1 5.7 321
Lepidoptera Butterflies, moths 45.4 27.7 6.6 362
Odonata Dragonflies, 55.2 19.8 11.8 61
damselflies
Orthoptera Grasshoppers, 61.3 13.4 9.6 278
locusts, crickets

a
Termites were previously classified in the order Isoptera.
Protein, fat, and fiber are percent values on a dry basis.
Data from Jongerma (2017) and Rumpold & Schlüter (2013).

www.annualreviews.org • Insects as a Protein Source 23

 of species currently commercially reared for human consumption. The order Coleoptera consti-
tutes 350,000 species of beetles, making them the single largest order of animals on the planet;
of these, there are 659 species of beetles that are consumed as food primarily in their larval stage.
Orthoptera refers to the order of insects commonly known as crickets, locusts, and grasshoppers,
among others. Within the Western market, these insects have been instrumental in the intro-
duction of insects for human consumption due to their familiarity in Western culture. Bees, silk-
worms, house crickets, and cochineals are the main species of insects that are considered to be
fully domesticated (Resh & Cardé 2009). However, because of their commercial rearing success
in recent years, house crickets (A. domesticus), tropical banded crickets (G. sigillatus), and yellow
mealworms (T. molitor) are the most widely farmed insects for human consumption worldwide
(Melgar-Lalanne et al. 2019). In the North American market, cricket species are the major insect
consumed, whereas in the European market (primarily Germany and France) the most used insect
species for food and feed are mealworms, crickets, and silkworms. Thailand and China typically
consume grasshoppers, crickets, locusts, and beetles (Baiano 2020).

PROCESSING, CHARACTERIZATION, AND TECHNOLOGICAL

APPLICATIONS OF INSECT PROTEIN
Although the sustainability of insects can be a driving factor in the consumption of insects as
food, it is not a sufficient reason for wide public acceptance, mainly because of the need to over-
come the aversion associated with consuming them. Studies indicate that low acceptability of ed-
ible insects is associated with emotional factors (e.g., disgust) and unfamiliar tastes and textures;
therefore, marketing exposure and incorporating insects as part of an ingredient in a familiar food
could be potential gateways to increasing acceptance by consumers (Wendin & Nyberg 2021).
Baiano (2020) and Liceaga (2019) suggest that processing insects into unrecognizable forms, such
as protein powders and textured insect proteins in beverages, meat analogs, dairy replacements,
and snack and bakery products (Luna et al. 2021, Shockley et al. 2018), may increase the ac-
ceptability of edible insect consumption. Farming and transforming insects into different insect-
based consumable products (i.e., food products and industrial or medical ingredients) is a new
evolving practice in some countries (Reverberi 2020). Accordingly, insects are being consumed as
whole insects (recognizable form), processed whole insects (nonrecognizable form), and extracts
(Figure 2).
Edible insects are traditionally eaten whole (recognizable as such); however, in some cases, re-
moval of unpalatable body parts (e.g., legs, wings, head, antenna, and guts) is required, especially
when insects are consumed raw. For instance, it is recommended to cut the back end of the palm
weevil (Rhynchophorus ferrugineus) larvae before eating them to reduce the pungent taste that is
produced, perhaps, by the action of the digestive juices of the larvae (Cartay 2018). Similarly, be-
cause edible stinkbugs’ release a noxious scent, they are first beheaded and carefully squeezed; thus,
chemicals and scent-producing secretion glands are discarded (Dzerefos et al. 2013). It should be
highlighted that eating raw insects could pose not only sensory drawbacks but also certain health
risks. Hence, to improve the innocuous consumption of insects and increase their acceptability
(i.e., overcome the disgust factor), several processing techniques have been used. Although the
number of such existing processes is highly variable, the elemental operations include cleaning,
heating, and drying (Nyangena et al. 2020).
The cleaning process usually includes sieving or washing the insects several times in cold or
tepid water to eliminate frass and other related matter. Insect euthanasia is then usually performed
by decreasing the temperature (freezing for 24 h at −18 °C) or by heating (blanching 4–7 min,
at 80–100 °C), although other methods have also been reported, such as pithing, microwaving,

24 Liceaga et al.
 INSECTS

Desired clean parts

Whole insect Whole insect

Processed
Insect extracts
Recognizable as such

Raw Cooked Insect proteins

Maceration or drying
and milling/extrusion
Insect fat/oil
Blanching
Granular/paste/powder Insect colorants
Steaming, roasting, smoking, formats
frying, boiling, curing, and/or Insect biopolymers
a combination thereof

Human Health Animal Industrial/medical

Human consumption food product feed applications

Figure 2
Schematic representation of insect processing for human consumption as whole insects (steaming, roasting, smoking, etc.), in processed
form (granular, paste, powders, etc.), and as extracts (protein, oils, colorants, etc.). Figure adapted from images created with
BioRender.com.

use of infrared tunnels, and asphyxiation (Hernández-Álvarez et al. 2021). Thereafter, the indi-
gestible parts of the insect are separated by, e.g., removing hard hairs and spiny parts to not only
avoid unpleasant sensations in the mouth but also reduce the risk of intestinal constipation. As a
slaughtering method, freezing can facilitate the removal of certain body parts (Fraqueza & da Silva
Coutinho Patarata 2017, van Huis et al. 2013). Furthermore, the insect’s gut may also be removed
due to the presence of spoilage-causing bacteria and potential pathogens. However, because it is
not possible to completely remove the gut in most edible insects, a fasting (∼24 h) step may be
applied to reduce the content of such gut-dwelling microorganisms. It should be mentioned that
the opposite of degutting (gut-loading) has also been practiced for commercially reared insects to
increase the levels of desired nutrients (e.g., the calcium:protein ratio and vitamin A content) in
the insect gut (van Huis et al. 2013).
In practice, whole (recognizable) insects are often consumed cooked as roasted/fried snacks
or ready-to-eat dishes. Hence, some popular heat processing methods have been applied, which
include steaming, boiling (blanching), roasting, frying (pan and deep), smoking, and stewing, or
a combination thereof. These traditional processing methods improve overall microbial safety by
reducing and/or eliminating spoilage and pathogenic bacteria, yeasts, and mold. Heating meth-
ods have also been shown to cause a decrease in the immunoreactivity of the para-allergen
tropomyosin (Hall et al. 2018). Processing methods are known to inactivate enzymes and in-
crease the nutritional composition, palatability, and digestibility of insects. These methods can
alter their chemical and nutritional composition by inducing inter- and intra-biochemical reac-
tions, proteolysis, lipid oxidation, and solubilization of vitamins and minerals (Murefu et al. 2019).

www.annualreviews.org • Insects as a Protein Source 25

 In this regard, it has been reported that boiling, toasting, and drying lead to significant increases
in protein and decreases in fat content on a dry-matter basis (Dobermann et al. 2019, Nyangena
et al. 2020). For example, during blanching, sorption properties of insect proteins may be altered
because some proteins undergo structural changes, including denaturation, crosslinking, and in-
teraction with lipids and carbohydrates. Furthermore, color parameters may also be altered by the
promotion of certain secondary reactions, including lipid oxidation and nonenzymatic browning
(Melgar-Lalanne et al. 2019). Similarly, other measured components, such as vitamins (e.g., ri-
boflavin, niacin, pyridoxine, folate, ascorbic acid), cholesterol, fiber, and micronutrients (e.g., Ca,
Na, Fe, Mg, Zn, Co, K), as well as their extractabilities (potential bioavailability), are also affected
(Mutungi et al. 2019). Attempts have also been made to apply fermentation technology to in-
sects to develop fermented edible insect products (e.g., sauces, powders, and paste) containing
secondary metabolites able to impart nutritional and antimicrobial benefits (Castro-López et al.
2020, Kewuyemi et al. 2020).
Despite cooking being a well-known way to inactivate most microorganisms and enzymes
present in insects, it is difficult to store cooked insects without drying due to the high mois-
ture content present. Therefore, it is suggested that insects should be quickly subjected to re-
frigeration to avoid the multiplication of survival microorganisms and/or germination of spores
during cooling; as such, it has been recommended to avoid storing insects at room temperature
(Fraqueza & da Silva Coutinho Patarata 2017). Additionally, different drying technologies may be
used for whole edible insects, including traditional methods such as sun-drying and oven-drying
(Hernández-Álvarez et al. 2021). The use of high temperatures during drying allows for a greater
decrease in microbial populations; however, it may also result in the loss of their original color
and nutritional value and changes in the aroma and flavor. Hence, freeze-drying has gained at-
tention because due to the low temperature used and the resulting water sublimation, organic
molecules such as proteins, lipids, carbohydrates, and vitamins and sensory properties (e.g., color,
aroma, and flavor compounds) will hardly change (Melgar-Lalanne et al 2019). However, freeze-
drying has low throughput and a high cost compared to most other drying technologies, which
currently limits its application on a large scale. Other promising alternatives such as microwave-,
radio frequency–, infrared radiation–, ultrasound-, and pulsed electric field–assisted drying have
also been explored (Hernández-Álvarez et al. 2021).
Although the cooking and drying methods described above are frequently used to prepare
more palatable insects, they remain whole (recognizable), which makes the idea of eating insects
repellent for many people. Hence, whole insects are being processed into nonrecognizable forms
to increase consumer acceptability (Hall et al. 2017). In this regard, insects are often ground to an
appropriate size and sifted into powder or flour that can be added to a wide variety of product for-
mulations such as health products, animal feed and foods, and gluten-free and high-protein/low-
fat products (van Huis et al. 2013). Particularly, food extrusion has been used in the production
of many common cereal-based foods (e.g., bakery products and pasta) to be used as a carrier for
the introduction of different amounts of insect flours and to study the products’ quality (Carcea
2020, Luna et al. 2021). It should be noted that insect flours have a high fat content (8–50%) and
are therefore prone to lipid oxidation. For this reason, insect flours are usually defatted to reduce
their lipid content (Hernández-Álvarez et al. 2021). Additionally, minced products may also be
prepared by mincing cooked insects to adequate particle size and combining them with other in-
gredients to obtain foods similar to those prepared with meat, such as hamburgers, meatballs, and
sausages (Elhassan et al. 2019, Fraqueza & da Silva Coutinho Patarata 2017). Efforts to produce
protein supplements, beverages, and energy bars based on insect powder have also been reported
(Mutungi et al. 2019).

26 Liceaga et al.
 The nutrients and other components in insects can also be extracted. For instance, proteins
can be extracted using water, organic solvents, and enzymes. The extraction rate and the physic-
ochemical, functional, and bioactive properties of extracted proteins are influenced by the in-
sect species, solvent used, drying treatment, and/or extraction temperature (Melgar-Lalanne et al.
2019). Protein concentrates (65–90% protein dry basis) or isolates (>90% dry basis) are pro-
duced from extracted proteins via the use of different methods, including enzyme-assisted extrac-
tion, reverse micelle precipitation, two-phase aqueous extraction, electrodialysis ultrafiltration,
ultrasound-assisted extraction, supercritical CO2 extraction, high pressure, and alkaline extraction
followed by isoelectric precipitation (Hernández-Álvarez et al. 2021, van Huis et al. 2013). The
general process for obtaining concentrates and isolates from insect proteins can be summarized
using certain key steps (Figure 3).
Before supplementing products (e.g., meat products and analogs, pasta, bread, snacks) with
protein concentrates and isolates, additional research regarding functional properties, including
thermal stability, solubility, gelling, water-holding capacity, foaming, oil absorption capacity, and
emulsifying capacity, is required (Gravel & Doyen 2020). Extracted proteins can be further pro-
cessed into protein hydrolysates, which can potentially be used for the development of insect-based
ingredients with improved functional and bioactive properties (Hall et al. 2017).
Among different components of insects, chitin is a significant biopolymer that can also be ex-
tracted from the exoskeletons of a variety of edible insects. There are numerous methods used to
extract chitin from insects, which include five main steps: defatting, demineralization, decoloriza-
tion, deproteinization, and deacetylation (Malm & Liceaga 2021, Mohan et al. 2020). Potential
applications of chitin involve its use as a nutraceutical or pharmaceutical product (i.e., as a drug
vehicle), in the water industry, in biodegradable materials for plastic alternatives, as an antimicro-
bial agent in food, and in cosmetics (Malm & Liceaga 2021, Mutungi et al. 2019).
Solvent-based methods (i.e., ethanol and n-hexane) have been widely applied to a variety
of insects for lipid extraction; however, other alternatives have also been implemented, such as
ultrasound (as pretreatment to Soxhlet, Folch, and aqueous extractions), supercritical CO2 ex-
traction, and high hydrostatic pressure extraction (Choi et al. 2017, Ugur et al. 2020). Mariod
(2013) reported that blending sunflower kernel oil with melon bug oil resulted in the im-
proved oxidative stability of the former and that transesterification of two insect oils using
methanol or ethanol in the presence of sulfuric acid yielded insect biodiesel. Insect oil has also
been suggested as a replacement for palm oil in broiler chicken nutrition (Benzertiha et al.
2019).
It should be noted that natural lacquers, dyes, and various types of varnishes have been obtained
from some insects. A few species from the superfamily Coccoidea (Hemiptera) have been used as
a source of pigments. Probably the best known is the carmine dye obtained from the cochineal
(Dactylopius coccus), which is widely used as a natural colorant in cosmetics, foods, pharmaceuticals,
textiles, and plastics (Borges et al. 2012). The traditional extraction process is a multistep pro-
cedure involving the treatment of dried, adult cochineal females with organic solvents and then
alkaline extraction, solid–liquid separation (i.e., flocculation and filtration), recovery of the precip-
itate, resolubilization of carminic acid, concentration, and drying into a powder form (Borges et al.
2012). Another example is the giant Margarodidae scale insect (Llaveia axin), commonly known as
niij in Mexico, which produces an internal body substance used as lacquer for woodwork, ceramic,
metal, and many other substrates. For the extraction procedure, adult females are boiled and then
mashed through fine cloth to produce a crude aqueous extract. After a prolonged mixing of the
extract, the fat globules separate from the other components and coalesce into bright yellow globs
that are washed thoroughly with water, shaped into a cylindrical form, and stored (MacVean 2008).

www.annualreviews.org • Insects as a Protein Source 27

 Insects

Pretreatment
Freeze-drying/oven-cooking
Ground and sieve

Fine powder

Defatting
Organic and nonorganic solvents
Three-phase partitioning Protein concentrates and isolates
Mechanical pressing
Supercritical CO2

Defatted protein solution/flour Drying

Oven-drying
Fluidized bed-drying Solid samples
Protein extraction Microwave-drying
Acid extraction (pH <2) Spray-drying method (?) Liquid samples
Salt extraction (salting-in, 0.5–1M)
Freeze-drying Wet solid/liquid samples
Aqueous extraction (pH 6–7)
Alkaline extraction (pH >7)

Soluble protein Purified protein

Protein recovery
Isoelectric precipitation Protein purification
Salting-out method Ion exchange
Precipitated protein Reversed phase
Thermolabile proteins Affinity chromatography
Freeze-drying Gel filtration
Ultrafiltration/diafiltration

Figure 3
Representative processing steps for obtaining protein concentrates and isolates that include defatting, protein extraction, recovery, and
purification. Adapted from Gravel & Doyen (2020). Figure adapted from images created with BioRender.com.

LEGISLATION AND SAFETY ASPECTS REGARDING THE USE

OF INSECT PROTEIN
Legislative restrictions (e.g., standards, labeling, and other regulatory instruments) and safety
aspects are important considerations for edible insects to become part of the worldwide food

28 Liceaga et al.
market (Belluco et al. 2017, Imathiu 2020). Until a few years ago, in Western countries, insects
were not considered a food or food source (Belluco et al. 2017). For example, the Codex Alimen-
tarius, which is one of the main organizations that regulate the trade of food and animal products
in the world, did not consider edible insects as food and classified them instead as “impurities”
(Petkov 2019). In the United States, edible insects would be regulated by the United States Food
and Drug Administration (FDA) with the Food Safety Modernization Act (FSMA) instating the
Preventative Controls for Human Food (PCHF) rule. Under this Act, insect farms are exempt
unless they change the raw agricultural commodities into a processed food. Although there are
no specific standards for edible insects, the US Food, Drug and Cosmetic Act, states that “insects
are considered food if that is the intended use” (Sec. 201(f )); therefore, wild-crafted insects and
those reared for animal/pet feed cannot be used for human food. Edible insect farms must have a
good manufacturing practice certification accompanied by pertinent microbial tests. Additionally,
in the United States, insect-containing foods require allergen-warning labels given significant
evidence that shellfish-allergic persons may also be allergic to insects (Baiano 2020, Liceaga 2019).
According to European regulations, farmed insects are defined as “farmed animals” and there-
fore can be reared only with qualifying substrates/feed materials. The European Food Safety Au-
thority considers whole insects and their parts as “novel foods” (EC 2015/2283). However, cur-
rently, edible insects represent a challenge for food legislation, as they need proper scientific back-
ing that addresses their safety concerns before obtaining market permission. Furthermore, there
is a need to create legislative rules regulating their production and commercialization (Belluco
et al. 2017, Loveday 2019).
Insect consumption occurs worldwide and is a part of the diets in many Asian, African, and
South American countries, either consumed as whole insects or incorporated in various food
products (Imathiu 2020, Kim et al. 2019). However, there are some safety aspects regarding the
use of insect protein related to the fact that, like other foods or food ingredients derived from
plants or animals, insects can accumulate contaminants such as microbial pathogens (Kouřimská
& Adámková 2016) and may possess some antinutritive factors (Ekop et al. 2010). Commonly
used preservation methods (e.g., heat treatments) applied during the cooking or processing of ed-
ible insects can decrease the microbial load and antinutritive content. For example, Klunder et al.
(2012) found a low microbial count of Enterobacteriaceae (<2.7 log CFU/g) in different cooked (i.e.,
boiled, roasted, or stir-fried) edible insects, including mealworm larvae (T. molitor), small crickets
(A. domesticus), and large crickets (Brachytrupus spp.). Authors reported that some methods such as
blanching and boiling were more effective at eliminating Enterobacteriaceae counts (<1 log CFU/g)
in all the insects analyzed. Ali et al. (2010) found that fried grasshoppers showed low counts of sev-
eral microbial populations compared with their fresh counterparts. Frying considerably decreased
the microbial loads of Escherichia coli (∼3.6 log CFU/g reduction), Salmonella (∼4.5 log CFU/g
reduction), fecal coliforms (∼5.9 log CFU/g reduction), and total coliforms (∼4.3 log CFU/g re-
duction). Other microorganisms or indicators of their presence, such as lactic acid bacteria, yeast,
and sulfite-reducing clostridia (indicator for Clostridium sp.), were not detected in fried grasshop-
pers. In contrast, Mujuru et al. (2014) found that the combination of boiling and pan-roasting
methods was the most effective at decreasing the microbial load in Mopani worms (Gonimbrasia
belinak) compared with other methods (i.e., ash-drying, drum-roasting, and sun-drying).
Antinutritive risks need to be evaluated on a case-by-case basis, as they seem to be species
dependent. For example, pupae of African silkworm (Anaphe spp.) are known to contain heat-
resistant thiaminase, which was responsible for several cases of thiamin deficiency in Nigeria
(Rumpold & Schlüter 2013). However, other studies on emperor moth larvae (Cirina forda), yam
beetle (Heteroligus meles), and palm weevil (Rhynchophorus phoenicis) found trace amounts of phytic
acid and oxalate, far below the nutritionally accepted values, including those found in common

www.annualreviews.org • Insects as a Protein Source 29

 legumes ( Jonathan 2012, Omotoso 2006). The impact of processing conditions on these antinu-
tritive factors needs to be further explored, as such conditions could assist in removing or inacti-
vating the antinutrients. Boiling and drying methods were able to decrease the content of three
antinutritive factors (oxalates, tannins, and alkaloids) in a stinkbug (Encosternum delegorguei) con-
sumed in Zimbabwe (Musundire et al. 2014).
In terms of chitin content, this long-chain polymer of N-acetylglucosamine is widely found
in the exoskeleton of insects. Although chitin can be digested to some extent by humans, con-
cerns exist about urinary stone formation and immunologic responses to chitin in some individuals
(Yhoung-Aree 2008). Additionally, chitin is believed to interfere with the protein functionality of
insect-based flours. Conversely, chitin has been suggested to improve human gut microbiota due
to its prebiotic potential (Selenius et al. 2018, Stull et al. 2018). Overall, based on the literature
available, edible insects do not seem to represent critical microbiological and antinutritive hazards
to humans. Further studies on the impact of processing on their safety are required.

CONCLUSION AND FUTURE DIRECTIONS

The consumption of insects by Western cultures is an emerging trend that will require changes in
our perception of insects. History has shown us that large-scale behavioral changes are possible,
especially if we consider, e.g., how lobster was perceived in the nineteenth century as nuisance
garbage in New England (USA); regulations prohibited feeding lobsters to prisoners more than
twice a week, as it was seen as inhumane treatment (Berger et al. 2018). Today, lobster has become
a luxury food widely accepted by consumers, suggesting that a similar trend for edible insects is
not implausible.
Consumer research suggests that utilitarian claims related to eating insects (e.g., their environ-
mental and health benefits) have successfully increased self-awareness of entomophagy. However,
consumers will respond more favorably to utilitarian claims when the hedonic response is already
in place (Berger et al. 2018). This suggests that a significant shift to what are being called “hedonic
claims” (e.g., tasty, delicious, savory) will help increase insect consumption across the United States
and Europe, with sensory appeal being the key factor for the acceptance of insects as pleasurable
components of food products.
As more countries begin to acknowledge the potential of insects as a source of protein, clear
regulations are needed to help insect farmers and industry commit to their application in food.
Food safety hazards, including allergens, pesticides, and microbial pathogens, need to be con-
sidered during the farming, harvesting, processing, and distribution stages. Edible insects are a
promising alternative to address the future protein demand by an ever-increasing world popula-
tion, especially in situations where food security is affected such as during the COVID-19 global
pandemic in 2020. In Western markets, insects are most likely to be consumed if they are not in a
recognizable form, e.g., if they are part of a food and/or formulated into a conventional food prod-
uct. In this context, food scientists need to explore mechanisms to improve the functionality and
acceptability of insect-derived protein to make it widely applicable in food and beverage formu-
lations. In addition, many of the other components of insects such as chitin, lipids, and colorants
increase the potential of fully utilizing edible insects as sustainable resources.

DISCLOSURE STATEMENT
The authors are not aware of any affiliations, memberships, funding, or financial holdings that
might be perceived as affecting the objectivity of this review.

30 Liceaga et al.
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