Trends in Food Science & Technology 118 (2021) 207–229

Contents lists available at ScienceDirect

Trends in Food Science & Technology

journal homepage: www.elsevier.com/locate/tifs

The science of plant-based foods: Approaches to create nutritious and

sustainable plant-based cheese analogs
Lutz Grossmann *, David Julian McClements
Department of Food Science, University of Massachusetts, Amherst, MA, 01003, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Background: There has been a growing interest in consumers around the world in adopting a more plant-based
vegan Cheese diet for health, sustainability, and ethical reasons. Many commercially successful products have already been
Cheese alternative developed, including plant-based meat and milk analogs. However, the production of plant-based cheese analogs
Imitation cheese
that consumers find desirable and acceptable has proved extremely challenging. This is mainly due to the
Healthy
Dairy-free
compositional and structural complexity of real cheese products, which is difficult to mimic using plant-derived
Emulsion gel ingredients.
alternative proteins Scope and approach: In this review article, we start by providing a brief overview of the production and properties
dairy-free of real dairy cheese. We then describe the plant-based ingredients and processing operations that can be used to
assemble cheese analogs that mimic the composition, structure, physicochemical properties, sensory, and
nutritional attributes of real cheese. We also consider in this review the potential impact of switching from
animal-based to plant-based cheese on the environment and human health.
Key findings and conclusions: Plant-based cheeses can be produced from plant proteins obtained using fraction­
ation or tissue disruption routes. These products are typically complex colloidal dispersions consisting of lipid
droplets embedded within a viscoelastic polysaccharide and/or protein network. These plant-based cheeses are
likely to be more environmentally sustainable and better for animal welfare than their regular counterparts.
More research is needed to identify appropriate ingredients and processing methods, including understanding the
changes in texture and flavor as well as creating appropriate melting behaviors. Moreover, further research is
required to improve the nutritional profile and test the health effects of plant-based cheeses.

1. Introduction
“I’d had no idea that cheese was a miracle.” (Holm, 2014).
Cheese is an extremely complex food. By one estimate there are more
than 1,000 cheese varieties that can be produced from the milk from
different animals, as well as other raw materials, using a variety of
production and aging operations (McSweeney, Ottogalli, & Fox, 2017).
The great variety reflects the popularity of regular cheese with around 6
and 10 million tons being produced in the USA and EU, respectively,
which is around three-quarters of the world’s production (USDA, 2020).
Cheese is generally acknowledged to be a nutritious food that is rich in
macronutrients, especially fat and protein, as well as micronutrients
such as vitamin A, vitamin B2 and B12, calcium, phosphorous, selenium,
and zinc (SELF Nutrition Data, 2021). Moreover, cheese has a long
shelf-life with some varieties lasting several years, which facilitates its
storage, distribution, and sale.
Recently, however, there has been a growing trend towards the
production of cheese analogs manufactured from plant-based in­
gredients. For instance, the Good Food Institute, a nonprofit organiza­
tion based in Washington D.C., reported a growth in plant-based cheese
analog sales of around 42% from 2019 to 2020, with $270 million being
sold in the USA alone in 2020 (Fig. 1). This trend in plant-based cheese
sales is surprising because around a decade ago these products were
regarded as cheap and less nutritious imitations of the cheeses produced
from animal milk. Producers of such products were accused of deceiving
customers by selling a low-quality product designed to mimic a high-
quality one. For example, the BBC documentary “Food Factory” from
2012 had an episode about “fake cheese”, which is “[…] quicker and
cheaper to produce than the real thing.” (BBC One, 2012).
However, this attitude has changed drastically over the past few
years with more and more consumers seeking out and purchasing plant-
based foods. The trend in plant-based cheese sales is driven by several

* Corresponding author.
E-mail address: lkgrossmann@umass.edu (L. Grossmann).

https://doi.org/10.1016/j.tifs.2021.10.004
Received 27 April 2021; Received in revised form 30 September 2021; Accepted 3 October 2021
Available online 5 October 2021
0924-2244/Published by Elsevier Ltd.
L. Grossmann and D.J. McClements
Fig. 1. Annual plant-based cheese sales in the USA according to (GFI, 2020).
factors and is part of a larger change in consumption habits directed
towards a general increase in plant-based food consumption (Asche­
mann-Witzel et al., 2020). The reasons for this change are numerous but
include food intolerances, environmental sustainability, health, and
animal welfare considerations, as well as social trends. Moreover, the
number and quality of cheese analog products on the market are
continuing to increase, which is increasing the acceptance and adoption
of these products.
It is interesting to address the question: why should a new food
product category (such as a plant-based cheese) be designed to mimic an
existing product (such as real cheese)? In principle, it is possible to use
modern technology to create entirely new food products that have not
been encountered before. This is also true for plant-based cheeses and
conclusions about why these products are helpful to guide dietary
transitions can be drawn from plant-based meats. While there is no
consumer research on plant-based cheese yet, some studies on plant-
based meats are helpful to understand some general mechanisms
involved. In general, it is accepted that a diet transition is eased by
decreasing barriers to change (e.g., food neophobia, adverse taste, price
differences, lack of familiarity, general beliefs about diet, etc.) and
supporting positive drivers of change (e.g., health and environmental
benefits, ethical concerns) (Szejda et al., 2020). For example, a UK study
showed that around 40% of the surveyed consumers were considered to
have some degree of food neophobia (Hoek et al., 2011), which limits
the adoption of new products by consumers, such as plant-based cheese.
This might be overcome by several factors, e.g., by efficient product
marketing to decrease the barrier and by focusing on social impacts.
For example, a study has indicated that meat-eaters are more willing
to shift to a plant-based meat product, if the product closely resembles
that of real meat in taste and texture (Michel et al., 2021) and this result
may also be applicable for plant-based cheese. This study also showed
that the acceptance of meat analogs is strongly influenced by social
factors, which are not directly related to the product itself but are based
on social patterns (e.g., it was found that meat analogs are less likely to
be consumed during a family dinner which commonly follows strict
social patterns) and social interactions (e.g., experienced judgmental
behavior of others based on own diet or negative experience with people
who follow a vegan diet). However, such results most likely have a
strong regional connotation and may vary from country to country but
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Trends in Food Science & Technology 118 (2021) 207–229
this research shows that plant-based cheese manufacturers should
emphasize the positive impacts of plant-based consumption habits while
positively emphasizing social patterns and trying to meet the taste and
price of real cheese, which have a strong influence on purchasing habits
(Szejda et al., 2020).
However, designing plant-based products that can accurately mimic
the full range of physicochemical, sensorial, and nutritional attributes of
real cheese is extremely challenging due to their compositional and
structural complexity. For the sake of clarity, the term “plant-based
cheese” will be used in this article but it has to be kept in mind that
regulations in many countries currently ban the use of dairy-related
descriptors for plant-based product names (CVRIA, 2017). In this re­
view, we mainly focus on processing approaches, health, and sustain­
ability considerations, whereas the sensory aspects of plant-based
cheeses have recently been reviewed by Short et al. (2021).
2. Cheese derived from animal milk
The production of cheese from animal milk is most likely one of the
oldest processes to preserve milk and the first cheese products were
already manufactured around 8000 years ago (Fox & McSweeney,
2017). Typically, cheese is classified according to its type (rennet
cheese, fresh cheese, processed cheese), its textural properties (soft,
semi-soft, medium-hard, semi-hard, and hard), and its origin (e.g., cow,
buffalo, or goat) but many more regional cheese classifications are
known.
Cheeses are produced from milk, which is the nutrient-rich fluid
produced by the mammary glands of mammals to feed their young (Fox
& McSweeney, 2017). The main aim of cheese processing is to obtain a
viscoelastic solid material from fluid milk by inducing a sol-gel transi­
tion and collecting the curd formed, which usually has around a 10-fold
higher protein content than the original milk. The agglomeration of the
caseins is induced by acidification and/or enzyme addition leading to
the formation of a curd, which is then separated from the
non-aggregated whey part (Johnson, 2017). The main processing routes
used for cheese production are described in various references (Kam­
merlehner, 2009; Kessler, 2002; McSweeney, Fox, et al., 2017), but
commonly involves a series of processes that may include: Pasteurization
→ Standardization → Acidification → Coagulation → Curd processing→
Ripening. This process is varied to obtain different types of cheeses with
various characteristics. However, some general cheese properties are
also important for plant-based formulations, which are reviewed below.
2.1. Texture and melting
All cheeses are viscoelastic materials, ranging from soft to hard
depending on cheese type. The texture is governed by the composition of
the milk (standardization), pretreatments such as homogenization (only
for some cheeses, such as blue cheese) and thermal processing, and
operating variables such as rennet-, calcium-, and sodium chloride-
concentration, as well as pH, and amount of whey removed.
However, the main transformation from a springy texture to a ho­
mogenous soft or hard texture occurs during the ripening phase (Iru­
dayaraj, 1999). During ripening, the casein network is partly broken
down, which releases peptides and functional groups. Depending on the
pH and the water-to-casein ratio, the cheese may soften or harden during
ripening (Lawrence et al., 1987). Typically, cheddar cheese hardens as a
result of proteolysis because the released functional groups bind more
water. Here, the relatively low water-to-casein ratio fosters the forma­
tion of a more rigid structure (Irudayaraj, 1999). In contrast, the aging of
camembert gives rise to a more creamy texture through the same
mechanism but the relatively high water-to-casein ratio fosters the for­
mation of a more viscous structure (Schlesser et al., 1992).
The pH of cheese also plays an important role in determining its
texture because it alters the protein-protein interactions and calcium
solubility. Typically, a less hard structure is obtained for cheeses that
L. Grossmann and D.J. McClements
have a pH close to the isoelectric point (pI) of the caseins, because more
calcium is solubilized and so the casein network is weakened (Pastorino
et al., 2003). This effect is especially relevant for acid-coagulated cheese
but has also been demonstrated for rennet-coagulated cheeses, e.g., for
camembert cheeses produced with different pH values during draining.
Indeed, a low draining pH (e.g., pH 4.6) results in considerable calcium
loss in the whey fraction. As the pH during ripening increases, the curd
liquifies because the curd is low in calcium bridges, which results in an
increase in protein solubility with increasing pH and as a consequence a
drop in viscosity during the ripening phase (Batty et al., 2019).
One key material property of cheese is its melting behavior, which is
critical for many applications, such as pizza toppings and cheese slices.
Cheese melting involves converting a predominantly elastic-like solid
cheese into a more viscous-like material by inducing a gel-sol transition
(Schenkel et al., 2013a). Most cheeses soften when heated and if the
temperature is further increased, the cheese begins to flow. The melting
behavior of cheese is commonly attributed to the melting of fat until 40
◦
C, but it is also strongly influenced by changes in casein-casein in­
teractions. Hydrophobic interactions strengthen as the temperature is
raised, resulting in a contraction of the individual casein molecules,
thereby weakening their interactions with other casein molecules,
which leads to the formation of a weaker gel structure (Lucey et al.,
2003). In general, any process that weakens casein-casein interactions
tends to increase the meltability of cheeses by, e.g., washing the curd to
remove calcium or other processes that increase calcium solubility.
Commonly, most cheeses melt in the temperature range from 30 to 75 ◦ C
(Karoui et al., 2003; Ray et al., 2016; Schenkel et al., 2013b,a) but some
do not melt at all. Cheeses that do not melt are typically prepared by
utilizing milk that was heated longer than the common
high-temperature/short-time treatment followed by pH reduction below
5. This results in the formation of covalent bonds that are less affected by
heating and thus no melting behavior is observed (Lucey et al., 2003).
The meltability of cheeses is also influenced by additives known as
“emulsifying salts”, such as citrates (e.g., sodium citrate) and phosphates
(e.g., sodium phosphate), which are added during the manufacturing of
processed cheese types. This cheese variety is produced by melting
rennet-induced medium-hard to hard cheese types in the presence of
emulsifying salts (and other additives if required), followed by a cooling
step. The effect of the emulsifying salts is threefold: (i) they sequester
calcium and thereby weaken the casein network; (ii) they increase the
pH, which increases protein solubility; and (iii) they exert a salting-in
effect. These effects improve the melting properties and foster the for­
mation of a stable network during the melting and cooling process
without fat and protein destabilization (Guinee, 2017).
2.2. Aroma
Ultimately, the aroma of a cheese is governed by the complex
mixture of volatile molecules produced during the cheese
manufacturing process. This aroma is a result of the composition of the
milk used, the type of processing operations employed, and the break­
down of the components within the cheese during the ripening phase.
First, cheese aroma is influenced by the species of animal the milk comes
from (e.g., cow, sheep, goat, or buffalo), but also by factors such as an­
imal feed (fresh grass, silage, hay, or grain), age, and health status
(Faulkner et al., 2018). Second, the different processing steps influence
flavor development. For example, a study performed with different types
of French cheeses (Brie, Camembert, Saint Nectaire, etc.) concluded that
the use of raw milk for production led to a higher overall aroma intensity
and also different flavors, especially flavor notes like sharp, butyric,
bitter, goaty and moldy, in most of the cheeses studied compared to
those produced with pasteurized milk (Chambers et al., 2010). Last, the
ripening of the freshly prepared cheese has a profound influence on the
aroma profile. The degradation of lipids, proteins, and carbohydrates by
indigenous enzymes, exogenous enzymes (rennet), and the microflora
produce an abundance of volatile compounds including carboxylic
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Trends in Food Science & Technology 118 (2021) 207–229
acids, lactones, ketones, alcohols, and aldehydes (Ianni et al., 2020).
2.3. Shreddability
The shreddability of cheese is a key material property that is of
particular importance for some specific cheese products, such as pizza
cheese, because it facilitates their utilization and accelerates their
melting (Apostolopoulos & Marshall, 1994). The shreddability of cheese
is influenced by a set of material characteristics and is mainly controlled
by a fine balance of its viscoelastic behavior. Soft cheeses, which have
relatively high moisture contents, low storage moduli, and high surface
energies, exhibit strong blade adhesion during cutting, which decreases
their shreddability (Childs et al., 2007). In contrast, hard cheeses, which
have relatively low moisture contents and high storage moduli may be
too gummy and dry, which results in shattering into smaller particles
during the shredding process (Kindstedt, 1995).
3. Plant-based cheese analogs
Most likely the first plant-based cheese varieties were produced and
consumed in China as fermented tofu. Reports about fermented tofu date
back to the 17th century and were called “furu” which might be trans­
lated as “spoiled milk” (Shurtleff & Aoyagi, 2011). However, this already
brings up the main question: what is plant-based cheese? Why is tofu
mostly regarded as a meat replacer, whereas fermented tofu is typically
qualified as a cheese analog? In this review, we will define plant-based
cheese as follows:
“Plant-based cheese is an edible material prepared from plant in­
gredients that is designed to have a similar appearance, texture, and
flavor as animal-based cheeses.”
Following this definition, the design principle behind plant-based
cheese production is to match the physicochemical and sensory attri­
butes of a specific conventional cheese (such as cheddar or mozzarella),
which can be achieved by using a variety of different ingredients and
structuring techniques depending on the properties required. Ideally, it
would be advantageous to use similar processing operations as used to
commercially manufacture regular cheeses, as then the same equipment
and manufacturing facilitates could be utilized. In practice, this is
extremely difficult because plant-derived ingredients behave very
differently from those found in animal milk.
The main difference between regular and plant-based cheese pro­
duction is that there is only one main ingredient in regular cheese (milk
from different sources) and the different cheese varieties are produced
by utilizing different processing routes. In contrast, plant-based cheese
analogs are created from a range of different ingredients and so their
development requires both ingredient and process selection.
3.1. Raw materials
In this section, the main properties and compositions of the in­
gredients that can be used to assemble plant-based cheese analogs will
be summarized. Only a brief overview will be given here since more in-
depth discussions on the extraction, characterization, and properties of
plant-derived ingredients can be found in recent reviews and books
(BeMiller & Whistler, 2009; Day, 2013; Grossmann & Weiss, 2021;
Kyriakopoulou et al., 2021; McClements et al., 2021, 2019; Nadathur
et al., 2016; Zia-ud-Din et al., 2017).
3.1.1. Polysaccharides
Starches are one of the most commonly used polysaccharide-based
ingredients in plant cheese analogs (Table 1). They consist of long
chains of glucose units linked together by glycosidic bonds and come in
two major forms: amylose (linear, α-1–4 bond) and amylopectin
(branched, α-1–4 and α-1–6 bonds). The precise molecular
 L. Grossmann and D.J. McClements
Table 1
Selected plant-based cheese analogs and regular cow milk cheeses on the market and their composition in g per 100 g sorted by their main ingredient into two different processing routes. The websites were accessed in April
2021. Note that some products are combinations of the fractionation and tissue disruption route. Kcal = kilocalories; CHO = Carbohydrates; S = Saturated Fats.
Product Ingredients kcal Fat (S) CHO Protein Fiber Sodium Others Type Company Website

Material Fractionation Route

Field Roast: Filtered water, coconut oil, corn and potato starch, modified potato starch, 286 21 21 0 0 0.93 Calcium: Semi-hard, https://fieldroast.com/produc
Creamy Original: Chao fermented chao tofu (soybeans, water, salt, sesame oil, calcium sulfate), (21) 0% DV ripened flavor t/chao-block-creamy-original/
Block sea salt, natural flavor, olive extract (antioxidant used as a preservative),
beta carotene (color), powdered cellulose (to prevent caking).
Daiya: Filtered Water, Tapioca Starch, Coconut Oil, Vegan Natural Flavors, Pea 286 21 25 4 0 0.82 Calcium: Cheddar-style, https://daiyafoods.com/our
Medium Cheddar Style Protein, Expeller Pressed: Canola and/or Safflower Oil, Chicory Root (16) 0.48 semi-hard -foods/blocks/medium-cheddar
Block Fiber, Tricalcium Phosphate, Salt, Xanthan Gum, Lactic Acid (Vegan), Pea /
Starch, Vegan Enzyme, Cane Sugar, Annatto (Color), Yeast Extract,
Coconut Cream.
Follow Your Heart: Filtered Water, Coconut Oil, Modified Potato and Corn Starches, Potato 286 21 21 0 0 0.93 Calcium: Mozzarella-style, https://followyourheart.com/
Dairy-Free Mozzarella Starch, Sea Salt, Natural Flavor, Olive Extract, Beta Carotene for Color. (21) 0% DV semi-soft products/dairy-free-mozzarell
Blocks a-blocks/
Follow Your Heart: Filtered Water, Organic Palm Fruit Oil†, Modified Potato Starch, Expeller- 321 25 29 0 0 1.64 Calcium: Parmesan-style, https://followyourheart.com/
Dairy-Free Parmesan Pressed Canola Oil, Natural Flavors (Contains Autolyzed Yeast), Organic (13) 20% DV hard products/dairy-free-parmesan-
Shredded Vegetable Glycerin, Less than 2% of: Sea Salt, Calcium Phosphate, Bamboo shredded/
Fiber, Sodium Phosphate, Carrageenan, Lactic Acid, Nutritional Yeast,
Organic Chickpea Miso (Organic Handmade Rice Koji, Organic Whole
Chickpeas, Sea Salt, Water, Koji Spores), Sunflower Lecithin, Citric Acid,
Annatto.
Follow your heart: Filtered Water, Organic Expeller-Pressed Soybean Oil, Organic Soymilk 286 29 (5) 7 4 4 0.34 Calcium: Mozzarella-style, https://followyourheart.co
Dairy-Free Mozzarella Powder (Organic Soybeans), Natural Flavors (Plant Sources), Inulin 0% DV soft m/products/dairy-free-mozzare
(Chicory Root Extract), Agar Agar, Sea Salt, Organic Soy Protein, Lactic lla-block-soy/
210

Acid (Vegetable Source)

Violife: Filtered Water, Coconut Oil, Potato Starch, Salt (Sea Salt), Glucono Delta 321 29 11 0 0 0.68 Calcium: Feta-style, https://violifefoods.com/us/
Just like Feta block Lactone, Flavor (vegan sources), Olive Extract, Vitamin B12. (25) 0.0 product/just-like-feta-block/
Vit. B12:
30% DV
Violife: Filtered Water, Coconut Oil, Potato Starch, Salt (Sea Salt), Glucono-Delta- 233 23 7 0 0 0.47 Calcium: Cream-cheese https://violifefoods.com/us
Just Like Cream Cheese Lactone, Flavor (vegan sources), Olive Extract, Vitamin B12. (20) 0.0 style /product/just-like-cream-
Original Vit. B12: cheese-original/
30% DV
Bute Island Foods: Water, Coconut Oil (21%), Modified Potato Starch, Maize Starch, Gluten 292 23 19 0.5 5.1 1.7 (as Calcium: Gouda-style, https://www.buteisland.
Sheese Block Gouda Style Free Oat Fibre, Modified Maize Starch, Thickeners (Carrageenan, Guar (19) NaCl) 0.15 semi-hard com/products/sheese-bloc
Gum), Salt, Natural Flavourings, Acidity Regulators (Lactic Acid, Sodium ks/gouda-style/
Lactate), Yeast Extract, Color (Carotenes).

Trends in Food Science & Technology 118 (2021) 207–229

Simply V: 53% cashew nut preparation (drinking water, 2% roasted cashews), 272 19 24 <0.5 2 (as Semi-hard https://www.simply-v.de/
Fein cremige coconut oil, modified starch, starch, salt, potato protein, flavor, coloring (18) NaCl)
Genieβerscheiben food (carrot and apple concentrate), antioxidant: sodium ascorbate.
OATzarella: Original Mini water, organic steel cut oats, organic extra virgin olive oil, organic tapioca 179 16 11 0 0 0.68 Mozzarella-style, https://oatzarella.com/cheeses
Cheese Wheel flour, natural flavors, sea salt, organic agar agar, lactic acid (vegan). soft

Tissue Disruption Route

Fermented Tofu Tofu (98%) (soya beans, water, magnesium chloride, calcium sulfate), sea 116 8 5.15 8.15 2.87 Calcium: https://fdc.nal.usda.gov/
salt, vegan yogurt culture: Str. Thermophilus. (depending on manufacturer) (1.16) 1.23 fdc-app.html#/food-details/
174305/nutrients
Kite Hill: Almond Milk (Water, Almonds), Salt, Enzymes, Tartaric Acid, Cultures. 246 21 (2) 9 9 4 0.40 Calcium: Ricotta-style, https://www.kite-hill.com/our-
Almond Milk Ricotta 0.1 semisoft food/artisanal-delicacy/
Alternative
(continued on next page)
 L. Grossmann and D.J. McClements
Table 1 (continued )
Product Ingredients kcal Fat (S) CHO Protein Fiber Sodium Others Type Company Website

Parmela Creamery: Nutmilk (Water, Cashews), Coconut Oil, Modified Food Starch, Potato 286 25 21 7 0 0.91 Calcium: Monterey Jack- https://www.parmelac
Nut Cheese Pepper Jack Starch, Sea Salt, Peppers (Jalapeno, Habanero), Natural Flavor, Annatto, (18) 2% DV Style, semi-hard reamery.com/products/slices
Slices Yeast Extract, Cultures
Catalyst Creamery: Hemp milk (filtered water, hemp hearts), organic apple cider vinegar, Not available Muenster-style, https://www.catalystcreamery.
Muenster Style H*mp Seed organic coconut oil, tapioca, nutritional yeast, kappa carrageenan, sea semi-soft com/cheesecave/muenster
Cheese salt, cultured cane sugar, organic mustard, organic onion powder, organic
smoked paprika
Cheeze & Thank You: Organic tofu (water, organic whole soybeans, calcium sulfate, magnesium 214 16 4 7 0 1.46 Calcium: Feta-Style https://www.cheezeandt
Artisanal Herbed Feta chloride), organic refined coconut oil, white wine vinegar, sea salt, less (11) 15% DV hankyou.com/cheezes
than 2% of spice, vegan source lactic acid
Cheeze & Thank You: Organic soy milk (soybeans, water), organic refined coconut oil, tapioca 393 36 11 4 0 0.68 Calcium: Mozzarella-style https://www.cheezeandt
Artisanal Mozzarella Capri flour, less than 2% of garlic, sea salt, spice, kalamata olives, sundried (29) 2% DV hankyou.com/cheezes
tomato, kappa carrageenan, vegan source lactic acid
Treeline Cheese: Cashew nuts, filtered water, vegan lactic acid, vegan L. Acidophilus, 500 39 (7) 18 18 4 0.45 Calcium: Soft-ripened https://www.treelinecheese.
Classic hickory smoked sea salt. 0.0 com/pages/classic
Treeline Cheese: Cashew nuts, filtered water, sea salt, lemon juice, vegan L. Acidophilus. 321 25 (4) 14 7 0 0.46 Calcium: Cream-cheese https://www.treelinecheese.
Plain 0.07 com/pages/plain
Happy Cheeze: Cashews (64%), water, salt, vegan fermentation and edible mold cultures. 359 27.3 13.3 13.9 1.2 (as Camembert- https://happy-cheeze.
Happy White (5.9) NaCl) style, soft- com/products/vegane-came
ripened mbert-kaese-alternative-dr-ma
nnahs
The Frauxmagerie: Raw cashews, nutritional yeast, vegan probiotics, salt, bacterial culture, 440 36 (4) 12 16 4 0.26 Calcium: Blue-cheese https://thefrauxmagerie.
Botanic True Blue filtered water 4% DV style, soft- com/product/botanic-true-blu
ripened e/
Miyoko’s Creamery: Organic Cashew Milk (Organic Cashews, Filtered Water), Organic 393 29 (5) 25 14 4 0.71 Calcium: Farmstead https://miyokos.com/pro
211

Aged Sharp English Chickpea Miso (Organic Rice Koji (Organic Rice, Koji Spores), Organic 0.04 cheese style ducts/vegan-cheese-wheel
Farmhouse Cashew Milk Whole Chickpeas, Sea Salt, Water), Nutritional Yeast, Sea Salt, Natural
Cheese Flavors (derived from Oregano, Plum, Flaxseed), Cultures
Miyoko’s Creamery: Organic Cashew Milk (Filtered Water, Organic Cashews), Organic Coconut 214 18 4 4 0 0.75 Calcium: Mozzarella-style, https://miyokos.com/
Organic Cashew Milk Oil, Organic Tapioca Starch, Sea Salt, Organic Agar, Mushroom Extract, (11) 0.04 soft products/fresh-vegan-mozza
Mozzarella Organic Konjac, Cultures. rella-cheese
Miyoko’s Creamery: Miyoko’s Cultured Vegan Milk (Oat Milk (Filtered Water, Organic Oats), 250 16 18 11 0 0.96 Calcium: Farmstead-style, https://miyokos.com/prod
Cultured Vegan Cheddar Navy Beans, Organic Garbanzo Beans, Cultures), Filtered Water, Organic (13) 0.52 semi-hard ucts/cultured-vegan-farmhous
Cheese Block Coconut Oil, Faba Bean Protein, Potato Starch, Organic Tapioca Starch, e-cheddar-chunk
Contains Less Than 2% Of Sea Salt, Calcium Sulfate, Natural Flavors,
Organic Yeast Extract, Organic Annatto, Organic Cultured Dextrose,
Konjac, Organic Locust Bean Gum

Trends in Food Science & Technology 118 (2021) 207–229

Nuts for Cheese: Organic cashews, organic coconut oil, organic coconut milk, water, 467 43 17 10 3 0.67 Calcium: Brie-style, soft- https://nutsforcheese.com/veg
Organic Un-Brie-Lievable organic quinoa rejuvelac (water, organic quinoa), sea salt, nutritional (20) 0.03 ripened an-cheese-products/
Wedge yeast, organic chickpea miso (organic rice, organic chickpeas, sea salt,
water, koji spores), fermented organic oregano extract (water, organic
oregano, organic raw cane sugar, active cultures)

Regular cheese from cow milk

Camembert Pasteurized Milk, Salt, Starter Cultures, Non-animal Rennet. (depending 300 24 0.5 20 0 0.84 Calcium: Soft-ripened USDA (2021)
on manufacturer) (15) 0.39
Vit. A: 241
μg
Vit. B12:
1.3 μg
(continued on next page)
L. Grossmann and D.J. McClements Trends in Food Science & Technology 118 (2021) 207–229

characteristics of the amylose and amylopectin (such as molecular

weight and degree of branching), as well as their ratio, depend on the
source of the starch. Most commonly, tapioca, potato, and corn starches
are used in plant-based cheese formulations (Table 1). Tapioca starch is
Company Website
extracted from the root of Manihot esculenta, commonly called cassava or
USDA (2021) manioc, by a series of processing steps to obtain isolated starch granules

USDA (2021)

USDA (2021)
with diameters of around 4–35 μm. The roots are washed, chopped into a
slurry, and the starch granules are separated by a sequence of screens
with decreasing pore size, hydrocyclones, and decanter centrifuges,
followed by a drying step (Breuninger et al., 2009). Similarly, potato
starch is produced by grinding potatoes with subsequent separation of
the starch granules and fibers from the potato juice by decanter centri­
fuges. The fibers (80–500 μm) are removed from the granules (1–120
Semi-hard

μm) by a series of centrifugal sieves, followed by refinement in centrif­

Hard
Type

Soft

ugal separators and hydrocyclones and a drying step to produce the

starch powder (Grommers & van der Krogt, 2009). Corn starch is ob­
tained by soaking the corn in an aqueous solution, followed by a series of
Vit. A: 316

Vit. A: 203
Calcium:

Calcium:

Calcium:
Vit. B12:

Vit. B12:

1071 IU wet milling, cyclone, screening, centrifuging, and filtration steps to

Others

Vit. A:
1.1 μg

1.7 μg
0.71

0.69

1.25

release the starch from the kernel, mill the endosperm, and separate the
μg

μg

granules from the germ, proteins, and fibers (Eckhoff & Watson, 2009).
Starches are used in plant-based cheeses because of their ability to
Sodium

form a viscous paste or gel upon heating (gelatinization) and cooling

0.65

0.68
0.7

(retrogradation/setback), which entraps fluids and other ingredients

within the 3D-polysaccharide network formed (Kasprzak et al., 2018).
Fiber

Starch gelatinization occurs when the starch granules are heated in the
0

presence of water. Initially, the starch granules absorb water and swell,
which leads to an increase in viscosity. However, above a particular
Protein

temperature, the starch granules are ruptured, which results in the

23

24

32

release of starch molecules (mainly amylose). Here, the pasting tem­

perature (maximum viscosity during heating) depends on the starch
CHO

2.4

4.4

type, being 63 ◦ C for tapioca, 64 ◦ C for potato, and 80 ◦ C for corn, and is
0

influenced by the amylose-to-amylopectin ratio (Taggart & Mitchell,

Fat (S)

2009). Upon cooling, the solubilized amylose may form a gel due to the
(19)

(12)

(21)
34

20

29

formation of hydrogen bonds between helical regions on their back­

bones, which is known as retrogradation or setback. Amylose solutions
kcal

408

298

393

have a low viscosity but are able to form strong irreversible gels due to a
high retrogradation rate, whereas amylopectin has a strong effect on the
viscosity but only forms weak and reversible gels upon heating and
cooling because of a low retrogradation rate (Schirmer et al., 2015).
Milk, Salt, Cultures, Enzyme (Rennet). (depending on manufacturer)

Pasteurized Cows Milk, Salt, Non-animal rennet, Starter cultures.

However, the overall viscosity is not only influenced by the amylose to

amylopectin ratio, but also by the granule size with larger granules (as in
potato) typically having high thickening powers, especially during
Milk, Salt, Animal Rennet. (depending on manufacturer)

heating (Schirmer et al., 2013). For plant-based cheese production, the

starches should gelatinize into a stretchy and formable mass and build a
viscoelastic gel upon cooling via partial retrogradation. Tapioca starch
has recently been shown to exhibit these desirable attributes (Mattice &
Marangoni, 2020).
Commonly, different starch types are combined to obtain an opti­
mum degree of retrogradation, softening temperature, and final gel
(depending on manufacturer)

strengths as well as viscoelastic properties which ultimately relates to

the amylose:amylopectin ratio, the corn size, and also the modifications
of the starch (Breuninger et al., 2009; Schirmer et al., 2015). For
example, corn starch has a high tendency of retrogradation, whereas
potato and tapioca starch have a much lower one, which influences the
Ingredients

final texture of the plant-based cheese (Jackson, 2003). Waxy starches (i.
e., high amylopectin content) are often used in plant-based cheese for­
mulations most likely because of their low tendency for retrogradation,
which enables the formation of a more soft texture. For example, waxy
potato and rice starches but also tapioca starches have been described as
exhibiting excellent melting properties, especially when combined with
Parmigiano Reggiano
Table 1 (continued )

plant proteins such as proteins from soy (Bergsma, 2017). Moreover, the
starches used in plant-based cheeses are modified to various degrees to
alter their physicochemical and functional properties. For example, acid
Mozzarella

hydrolysis is carried out to decrease the paste viscosity during gelati­

Cheddar
Product

nization and increase the gel strength upon retrogradation, octenyl

succinate derivatization is carried out to minimize oiling-off effects, and

212
L. Grossmann and D.J. McClements
cross-linking increases the heat and shear stability of the starch granules
(Klemaszewski et al., 2016; Taggart & Mitchell, 2009).
3.1.2. Proteins
The proteins used in plant-based cheeses are mainly derived from
pea, soy, lupin, potato, nuts, and corn. The proteins need to fulfill
different physicochemical and functional attributes with emulsification,
gelation, water holding, and flavor pre-cursor properties being the most
important ones. Some of the most critical properties that are important
to know about plant proteins are their solubility at different pH- and
ionic strengths, their interfacial activity and stabilization behavior, their
thermal denaturation temperature, their ability to form networks that
can entrap water and oil, and their susceptibility to enzymatic modifi­
cations such as proteolysis and crosslinking. In the remainder of this
section, we briefly discuss some of these properties but the reader is
referred to other articles that have focused on plant proteins for more
details (Day, 2013; McClements & Grossmann, 2021; Nadathur et al.,
2016). Moreover, the emulsifying and gelation properties of plant pro­
teins have been reviewed recently (Grossmann & Weiss, 2021).
• Legume proteins: Pea, soy, and lupin belong to the family of Fabaceae,
also known as legumes, and the proteins are mainly (50–90%)
comprised of salt-soluble globulins (Day, 2013). When being
dispersed at low ionic strengths, the 7S globulins have a minimum
solubility at pH 4.5–7.0 and a high solubility in more acidic and
alkaline regions. The 11S globulins have a broader minimum solu­
bility at pH 4.0–7.5 at low ionic strengths. In contrast, the 7S glob­
ulins have a solubility >90% at pH 3–9, while the solubility of most
11S globulins is low up to a pH of 5–6 (only soy 11S has no clear
minimum solubility and a high solubility (>80%) above a pH of 3–4)
at high ionic strengths (Grossmann & Weiss, 2021; Kimura et al.,
2008; Ruiz & Hove, 1976). Soy glycinin (11S) denatures at 78–94 ◦ C
at pH 7.6, depending on the ionic strength, whereas β-conglycinin
denatures between 67–87 ◦ C (Lakemond et al., 2000; Murphy, 2008).
Pea legumin denatures at around 87 ◦ C, whereas the total pea
globulin (legumin and vicilin/convicilin) fraction was reported to
have a denaturation temperature of 76–82 ◦ C depending on pH, ionic
strength, and extraction conditions (Chihi et al., 2016; Mession et al.,
2013; O’Kane et al., 2004). Finally, lupin globulins were reported to
denature in the region of 94–114 ◦ C (Sousa et al., 1995). Commonly,
pea, soy, and lupin proteins are susceptible to transglutaminase
cross-linking but adverse processing during protein extraction may
impair their crosslinking potential (Ceresino et al., 2021; Ionescu
et al., 2009; Sun & Arntfield, 2011; Yasir et al., 2007). Lastly, the oil
binding capacity is commonly reported to be highest for soy followed
by lupin and pea protein (Tömösközi et al., 2001).
• Potato proteins: Potato protein is composed of the glycoprotein
patatin (88-kDa dimer) and other proteins including various protease
inhibitors (5–25 kDa) (Ralet & Guéguen, 2000; Zhang et al., 2017).
Patatin has a minimum solubility at a pH of 3.5 and I = 200 mM, but
it can be enhanced at low ionic strengths (I = 0.15 mM). In contrast, a
high solubility under acidic conditions and at neutral to alkaline
conditions has been reported for this fraction (Ralet & Guéguen,
2000; van Koningsveld et al., 2001). Interestingly, the 16–25 kDa
fraction was reported to have a very high solubility across a broad
pH-range from 2.5 to 12 (Ralet & Guéguen, 2000). Patatin has a low
thermal denaturation temperature of around 59–60 ◦ C at I = 10
mmol L− 1 and a pH of 7, whereas the protease inhibitor fraction has a
slightly higher denaturation temperature of 66–68 ◦ C (Creusot et al.,
2011; Koningsveld, 2001; van Koningsveld et al., 2001). Potato
proteins have been shown to be susceptible to a range of
213
Trends in Food Science & Technology 118 (2021) 207–229
cross-linking enzymes such as transglutaminase, peroxidase, laccase,
and tyrosinase (Gui et al., 2020).
• Nut and seed proteins: Generally, nuts are seeds that consist of a hard
shell and an edible kernel. Thus, strictly speaking, almonds and
cashews should not be considered to be nuts but for the sake of
clarity, they will be discussed in this section. In general, much less
information is available for protein derived from nuts as for some
other protein sources, and more research is needed to fill this gap.
Proteins derived from cashew kernels have a minimum solubility in
the pH range from 4.0 to 5.0 (Neto et al., 2001; Ogunwolu et al.,
2009). Moreover, it has been shown that cashew proteins undergo a
sol-gel transition after heat treatment at 100 ◦ C with a least gelling
concentration of 6.5–13.5% (Ogunwolu et al., 2009). Typically, nut
proteins have an oil absorption capacity in the region of 3–4 g oil per
g of protein (Jitngarmkusol et al., 2008; Ogunwolu et al., 2009;
Sharma et al., 2010; Sze-Tao & Sathe, 2000). This value might be
higher or lower compared to soy protein, depending on the soy raw
material. However, most nut proteins have been reported to already
gel at lower concentrations than soy proteins (typically between 4
and 14%) while especially proteins found in cashew kernels possess
high thickening powers (Sathe & Sze, 1997; Sharma et al., 2010).
• Zein: Zein is the major prolamin fraction (soluble in aqueous ethanol
solutions) in the endosperm of corn with a concentration of around 60%
of the total protein. It is extracted from milled corn with different sol­
vents that solubilize the α-zein, β-zein, γ-zein, and δ-zein, with α-zein
being the most abundant fraction (Anderson & Lamsal, 2011). The
α-zein (molecular weight 21–26 kDa) is a hydrophobic protein that
forms a triple helix with a hydrophobic core, which can be used to
solubilize and stabilize hydrophobic bioactives, such as lutein
(Anderson & Lamsal, 2011; Momany et al., 2006). Zein has also been
employed to entrap other hydrophobic substances such as vitamins and
nutraceuticals, and zein particles have been used to stabilize
the oil-water interfaces in emulsions through the Pickering effect
(Fathi et al., 2018). Zein has a glass transition temperature of 139 ◦ C in a
dried state but water plasticizes the protein, which enables the for­
mation of rubbery stretchable textures at sufficiently low temperatures
(Madeka & Kokini, 1996; Mattice & Marangoni, 2020). However, there
is currently a lack of data about how zein could stabilize an oil-phase in
plant-based cheeses, whereas it is known that zein is not susceptible
to enzymatic cross-linking with transglutaminase but to tyrosinase
(Glusac et al., 2018; Mattice & Marangoni, 2020).
3.1.3. Fats
The fats in real cheese play an important role in determining their
formation, texture, sensory quality, and nutritional profile. Triglycerides
are by far the most common fats found in milk, which are comprised of
three fatty acids attached to a glycerol via ester bonds (Brady, 2013). The
most common fatty acids in milk are saturated (around 70%), mono­
unsaturated (around 25%), and polyunsaturated (around 2.3%), but
short-chain fatty acids (such as butyric and caprylic acid) are also present,
which make an important contribution to the flavor profile of cheese after
ripening (Macedo & Malcata, 1996; Månsson, 2008). However, this
composition may change depending on the season, region, and breed. The
relatively high levels of long-chain saturated fatty acids in milk fat
(>25%) mean that it tends to be partially crystalline at temperatures
around ambient and below, which is important for the formation of
solid-like structures in cheese. Indeed, the fat crystals tend to interact
with each other to form a 3D-network that contributes to the mechanical
properties of real cheese. The variation of the solid fat content (SFC) of
milk fat with temperature is therefore an important functional attribute,
which is desirable to mimic using plant-based fats (Fig. 7).
L. Grossmann and D.J. McClements
Plant-based fats and oils tend to exhibit a wide range of different
fatty acid compositions depending on their botanical origin and the
methods used to isolate them, which leads to different melting charac­
teristics. In general, the melting point of edible fats tends to increase as
their chain lengths increase and the number of double bonds decreases.
Many common plant-based oils (such as sunflower, corn, and canola
oils) contain substantial amounts of unsaturated fatty acids so they tend
to be liquid at ambient temperature and cannot form fat crystal net­
works. Historically, these liquid oils were often converted into solid fats
using hydrogenation, since this reduced the number of double bonds
present. However, this process is less popular now because it converts
healthy unsaturated fats into unhealthy trans- and saturated ones if the
process is not carefully controlled, which may be associated with an
increased risk of cardiovascular disease (Hu et al., 2001). Instead,
manufacturers of cheese analogs often blend natural sources of
plant-based solid fats (e.g., coconut oils) with liquid oils to obtain the
required SFC versus temperature profile (Fig. 7) and hence obtain the
required textural and melting characteristics. It should be noted, how­
ever, that these oils also have relatively high amounts of saturated fats
that might also have adverse effects on human health (Ludwig et al.,
2018).
It should also be noted that the fatty acid profile of plant-based fats
also influences their nutritional attributes and chemical stability
(McClements & Decker, 2017, chap. 5). Unsaturated fats, particularly
polyunsaturated ones (like flaxseed or algal oils), may have beneficial
health effects (Saini & Keum, 2018; Shahidi & Ambigaipalan, 2018). On
the other hand, they tend to be liquid at room temperature and are
highly susceptible to lipid oxidation, which can reduce the acceptability
of a product due to the creation of rancid odors (Arab-Tehrany et al.,
2012; McClements & Decker, 2017, chap. 5; Nogueira et al., 2019). A
number of different approaches have therefore been developed to try to
overcome this problem, such as minimizing the exposure of the unsat­
urated lipids to oxygen, heat, or light by controlling the storage condi­
tions or packaging; minimizing pro-oxidant contamination (such as
transition metal ions); utilizing antioxidants or chelators (such as
Fig. 2. Flow chart comparing the overall processing steps from the raw material to the final regular cheese and plant-based cheese (fractionation and tissue
disruption route) into a solid viscoelastic material based on phase transitions.
214
Trends in Food Science & Technology 118 (2021) 207–229
phenolic substances and organic acids); and by employing structural
design principles (such as interfacial engineering) (Jacobsen, 2015;
Jacobsen et al., 2013; McClements & Decker, 2018).
A wide variety of plant-derived fats have been used to formulate
plant-based foods, such as avocado, canola, cocoa, coconut, corn, palm,
vegetable, safflower, sesame, soybean, and sunflower oils (Sha & Xiong,
2020). Only a few of these are solidified at room temperature (cocoa,
coconut, and palm oils) and so can be used to generate desirable textural
attributes in cheese analogs. The SFC versus temperature profile and
textural attributes of the cheese analogs can be optimized for particular
products by blending these solid fats with liquid oils, as mentioned
earlier.
3.2. Processing routes
An important goal in the successful production of plant-based cheese
analogs is to induce a sol-gel transition that generates a viscoelastic solid
with textural characteristics similar to those found in regular cheese.
This can be achieved by utilizing different ingredients and processing
routes.
Fig. 2 compares the overall principles of regular and plant-based
cheese production by starting from the initial energy source and
describing the main phase transitions induced during production. In
principle, the value chain begins for all cheese types by the "harvesting"
of light energy by plants and it is important to realize that photons are
the primary energy source for all cheese types (just like as for many
other foods). For regular cheese, sunlight drives the growth of the feed
material (e.g., grass or cereals) through photosynthesis, which is then
consumed by the cow. The cow then breaks down the feed material
through its digestive system operating in symbiosis with the gut mi­
crobes residing there. This process releases energy and the building
blocks that the cow can absorb and use to assemble milk within the cells
in its mammary glands. After being expressed from the cow, the liquid
milk is then further treated using a variety of processing operations, such
as pasteurization, standardization, acidification, coagulation, curd
L. Grossmann and D.J. McClements Trends in Food Science & Technology 118 (2021) 207–229

Table 2
Overview of studies that investigated the formation of plant-based cheese analogs.
Ingredients Pretreatment Sol-gel transition Properties Reference

Fractionation Route: Starch or protein-based emulsion gels (oil/fat crystal dispersed phase)

waxy non-modified potato starch, potato gelatinization → setback, semi-hard, melting Bergsma (2017)
protein, sunflower oil retrogradation
modified, pregelatinized high-amylose corn gelatinization, melting → setback, semi-soft, good shredding, poor meltability Zwiercan et al.
starch, shortening, buffering salts retrogradation, crystallization (1987)
different starches, fat, carrageenan, xanthan, gelatinization, melting → setback, Atapattu and Fannon
guar gum, buffering salts, and others retrogradation, crystallization (2014)
Zein, high oleic sunflower oil, coconut oil, self-assembly, plasticizing, melting, high stretchability, weaker structure at Mattice and
starch, xanthan gelatinization → plasticized protein elevated T Marangoni (2020)
network

Fractionation Route: Protein-based emulsion gel (filled/particulate)

Zein, potato protein, olive oil Enzymatic crosslinking (Tyrosinase) shear-thinning behavior, crumbly texture Glusac et al. (2018)
Zein, pea protein, corn oil Enzymatic crosslinking (Tyrosinase) paste-like structure Glusac et al. (2019)
Soy tofu, soy protein isolate, maltodextrin, Enzymatic crosslinking Cream-cheese style Lim et al. (2011)
palm oil (Transglutaminase)
Pea-globulins, sunflower cream fraction, Enzymatic crosslinking Holz-Schietinger
glucose (Transglutaminase) + Acidification by et al. (2014)
starter cultures
Soy globulins, sunflower oil/moong bean 8S, Heat Heat-aggregation/acidification with Melting, stretching properties Holz-Schietinger
palm oil, buffering salts/soy globulins, pea treatment starter cultures et al. (2014)
globulins, pea prolamins, xanthan

Tissue Disruption Route: Protein-based emulsion gel (filled/particulate)

Cashew-, soy milk analog Heat- Ion gelation with multivalent cations Semi-hard, protein content in dry matter up Oyeyinka et al.
treatment to 64% (2019)
Lupine paste, oil, emulsifying salts Heat aggregation Protein content up to 14.9%, increased oil Awad et al. (2014)
separation, firmness, and a lower
springiness compared to control
Soy milk analog Heat- Acidification with starter cultures Further processing to a soft cheese analog Matias et al. (2014)
treatment
Soy milk analog Heat- Acidification with starter cultures, ion Semi-hard, hardness higher compared to Chumchuere et al.
treatment gelation (CaSO4) commercial sample (2000)
Soy milk analog, coconut oil, carrageenan, Heat- Acidification with starter cultures/ Cream-cheese-type, protein content up to Li et al. (2013)
emulsifying salts treatment glucono-delta-lacton 19.5%
Soy milk, soy oil, okara Heat- Acidification with starter cultures Cream-cheese-type, protein content up to Giri et al. (2018)
treatment 17.6%
Soy milk Heat Acidification with starter cultures L. rhamnosus ferments soy oligosaccharides Liu et al. (2006)
treatment
Soy milk, pigeon pea milk Heat Acidification with starter cultures, ion Up to 58.7% protein in dry matter Verma et al. (2005)
treatment gelation with multivalent cations
(CaCl2)
Soy milk Heat Acidification with starter cultures, ion Hard-cheese type, fat content of 11.80% (El-Ella, 1980)
treatment gelation with multivalent cations (Ca2+- after 3 months ripening
lactate)
Soy milk Heat Ion gelation with multivalent cations Cream-cheese type after blending with palm Zulkurnain et al.
treatment (CaSO4), acidification with citric acid oil and polysaccharides (2008)
Soy milk Heat Ion gelation with multivalent cations Chikpah et al. (2015)
treatment (alum)
Macadamia-almond nut milk analog Heat Acidification with starter cultures, Fresh-, salted-, soft-ripened cheese types Holz-Schietinger
treatment enzymatic crosslinking et al. (2014)
(transglutaminase)
Soy milk Heat Enzyme-rich Moringa oleifera extract soft-white cheese Sánchez-Muñoz et al.
treatment (2017)
Soy milk Heat Plant and microbial proteases Only curd production Murata et al. (1987)
treatment
Cashew kernels Acidification with fermented quinoa brie, red, herb, cheddar, and blue cheese- Chen et al. (2020)
dispersion like types

formation, and ripening. Thus, the two main phase transitions that occur
during regular cheese production are solid (feed) → liquid (milk) → solid
(cheese). In contrast, more phase transitions are typically required to
create plant-based cheeses with two main processing approaches that
can be distinguished being:
• Fractionation Route: The raw plant-based material is treated to isolate
and purify refined functional ingredients, such as polysaccharides,
proteins, and fats. Proteins are commonly extracted from soybeans,
peas, beans, lupines, and potatoes. Starches are isolated from corn,
pea, tapioca, and potato. Other functional polysaccharide-based
hydrocolloids may be isolated from terrestrial plants (e.g., pectin,
guar gum, locust bean gum, or cellulose), seaweed (e.g., agar, algi­
nate, and carrageenan), or microbes (e.g., xanthan gum). Plant-based
fats, which may be liquid or semi-solid at ambient temperature, are
often obtained from coconut, palm, cocoa, soy, sunflower, or rape­
seed. Proteins and polysaccharides are usually isolated using a liquid
extraction step followed by a drying step to yield the final ingredient
(with the exception of dry fractionated proteins). Thus, the solid raw
plant material is dispersed in an aqueous solution to extract and
dissolve the desired ingredients, followed by a drying step to convert
them into a powdered form. The fractionated protein and/or poly­
saccharide ingredients are then often solubilized in water and
blended with oil to create a plant-based emulsion. Finally, this

215
L. Grossmann and D.J. McClements
mixture may be solidified using a variety of approaches, including
heating, chilling, pH adjustment, enzyme addition, or salt addition.
During these various processes, proteins are often solubilized, de­
natured, and/or aggregated, starches are often gelatinized, fats are
emulsified, melted, and/or crystallized, and gums may undergo coil-
helix transitions, with the precise processes occurring depending on
the nature of the ingredients and processes used. Thus, this pro­
cessing route employs numerous phase transitions to yield the final
cheese analog: solid (plant raw material) → liquid (extraction/frac­
tionation) → solid (dried ingredients) → liquid (emulsion) → solid
(plant-based cheese) (Fig. 2).
• Tissue Disruption Route: In this route, a whole plant-based raw ma­
terial serves as a basis for plant-based cheese production. For
example, soybeans, peas, or nuts are employed. The plant-based raw
materials are soaked to soften them and then broken down to form a
colloidal dispersion, which may contain oil bodies, plant tissue
fragments, dissolved biopolymers, sugars, and salts. This dispersion
can then be converted into a cheese analog by inducing a sol-gel
transition using the same methods described for the fractionation
route. Thus, this processing route only employs two phase transitions
to yield the final cheese analog: solid (plant raw material) → liquid
(emulsion) → solid (plant-based cheese) (Fig. 2). It should therefore
be more energy efficient than the fractionation method, which was
also highlighted in a recent study that compared different strategies
for the production of protein-rich ingredients from plants and found
the highest impact for highly refined ingredients (Lie-Piang et al.,
2021). However, actual data for specific plant-based cheese products
is currently missing, which highlights the need for more research in
this area.
Fig. 3. Flow process chart comparing the different processing routes for plant-based cheese production for the material fractionation route based on extracted
polysaccharides (left) or proteins (right) from different raw materials.
216
Trends in Food Science & Technology 118 (2021) 207–229
These two main processing routes will be described in more detail in
the following sections. It is important to note that these two processing
routes are distinguished by the primary actor that induces the sol-gel
transition. The sol-gel transition in the “fractionation route” is facili­
tated by fractionated and mostly isolated plant ingredients coming from
different origins that are recombined in the process, whereas the sol-gel
transition in the “tissue-disruption route” is mainly induced by the in­
gredients coming from the initial raw material that is used as a whole (e.
g., nut paste) or has undergone only a limited number of separation steps
(i.e., a plant-based milk). The curds obtained can be classified depending
on the coagulation process employed: self-associated-, enzymatic-, acid-,
ionic-, and thermal-curd, as well as combinations of these (this is similar
to the classifications used for conventional cheese products, such as
acid- or rennet-curd).
Studies that have employed these processing routes to obtain plant-
based cheeses will be discussed in the following sections and are sum­
marized in Table 2.
3.2.1. Processes based on the fractionation route
In the fractionation route, a selection of functional ingredients (such
as isolated proteins, polysaccharides, and fats) is blended in an appro­
priate ratio to form an oil-in-water emulsion, and then a sol-gel transi­
tion is promoted using one of various means to form a semi-solid cheese
analog (Fig. 3). Two of the most common ingredients that have been
used to form a three-dimensional gel network in commercial products
are starches and proteins (Table 1).
Polysaccharide-based cheese analogs: Starch is the most common
polysaccharide-based ingredient used to form gelled structures within
cheese analogs, but other polysaccharides can also be used for this
purpose, including alginate, carrageenan, or guar gum. There is
L. Grossmann and D.J. McClements Trends in Food Science & Technology 118 (2021) 207–229

Fig. 4. Different viscoelastic emulsion gels

are formed during plant-based cheese pro­
duction. Starch structures are produced from
the fractionation route, whereas protein-
based may be produced from the fraction­
ation and tissue disruption route. Depending
on the process, protein-based structures can
be distinguished as coming from a diluted
colloidal dispersion (i.e., a plant-based milk)
process that involves separation of insoluble
particles or coming from a concentrated
colloidal dispersion (i.e., a plant-based paste
for example ground nuts) using the whole
seed material.

Fig. 5. Flow process chart comparing the different processing routes for plant-based cheese production for the tissue disruption route based on concentrated (i.e., a
seed paste often based on nuts; left) or diluted (i.e., a plant-based milk; right) colloidal dispersions.

217
L. Grossmann and D.J. McClements Trends in Food Science & Technology 118 (2021) 207–229

Fig. 6. Schematic representation of the interaction

potential between two proteins modeled as colloidal
spherical particles showing the primary minimum
(1st Min), secondary minimum (2nd Min). For
protein-based plant-based cheese production, the
energy barrier has to be lowered by reducing elec­
trostatic repulsion (ionic screening, pH near iso­
electric point) and/or increasing hydrophobic
interactions (thermal treatment) to induce aggre­
gation or facilitate enzymatic/ionic crosslinking by
bringing molecules in close proximity.

cheese types can be described as polysaccharide-based viscoelastic

emulsion gels with the polysaccharide forming the 3-dimensional
network that entraps water and oils. The oils might partially coalesce
or when fats being used (e.g., coconut oil) it partly or fully crystallizes
with partial coalescence upon cooling, giving the plant-based cheese its
characteristic texture (Fig. 4A and B).
In one patent, the main ingredients used were 15–22% waxy non-
modified potato starch, 0.5–8% potato protein, 15–35% fat (short­
ening), and 35–74.5% water (Bergsma, 2017). These ingredients were
then heated at 70–90 ◦ C while mixing to induce starch gelatinization.
The overcooked starch emulsion obtained was then poured into a mold
and cooled to induce a sol-gel transition, leading to the formation of a
semi-solid cheese analog. In another patent, pregelatinized modified
high amylose corn starch was utilized as a plant-based gelling agent
(Zwiercan et al., 1987). In this case, the cheese analogs were prepared by
mixing starch (>20%), fat (22.8%), buffering salts, and water at a
temperature at which the fat was melted, followed by cooling to 4 ◦ C to
solidify the system. Another patent used a similar procedure but pro­
posed a variety of different kinds of starches as gelling agents, including
native starch, acid-treated corn, octenyl succinate-modified starch, and
hydroxypropyl distarch phosphate (Atapattu & Fannon, 2014). More­
over, the mixture also contained hydrocolloids (such as carrageenan and
guar gum) to provide texture and buffering salts to control the pH at
around 5 to 6. The plant-based cheese analog was produced using a fat
with a melting point between 30 and 52 ◦ C. The fat, water, and various
functional ingredients were mixed to form an oil-in-water emulsion that
Fig. 7. The solid fat content (SFC)-temperature profile of edible fats depends on
was heated to around 83 ◦ C to melt the fat and gelatinize the starch. This
the fatty acid composition, which is determined by their biological origin.
emulsion was then cooled, which promoted the formation of a semi-solid
Plant-based fats should mimic the SFC profiles of animal-based ones.
texture mainly due to fat crystallization and starch crosslinking.
Depending on the plant-cheese type required, it was recommended that
currently a lack of research published in the scientific literature that use
the water content could range from 20 to 80% water and the fat content
starch as the main gelling agent to form plant-based cheese analogs but
from 15 to 30%.
several patents have been filed that describe this process (Atapattu &
Protein-based cheese analogs: Dairy proteins, especially caseins, play a
Fannon, 2014; Bergsma, 2017; Klemaszewski et al., 2016; Schelle et al.,
critical role in the production and the formation of the characteristic
2020; Zwiercan et al., 1987).
properties of regular cheeses. For this reason, there has been great in­
In general, the production of plant-based cheeses with starches as the
terest in the utilization of plant proteins to provide the required func­
main gelling ingredient is based on starch gelatinization at elevated
tional, sensorial, and nutritional attributes in cheese analogs. However,
temperatures (i.e., a thermal curd) followed by a sol-gel transition dur­
it is extremely challenging to find plant proteins that can accurately
ing cooling through entanglement, setback, and retrogradation of the
mimic the structure and behavior of casein micelles. This is mainly
starch molecules (Taggart & Mitchell, 2009). The structure of these
because casein molecules, and the micelles they form, are highly

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L. Grossmann and D.J. McClements
unusual structures in nature (Lamichhane et al., 2018). The individual
casein molecules have an open structure with a high degree of flexibility,
which is different from the tight compact globular structure of most
plant proteins. Moreover, the casein molecules have regions with
numerous phosphate, sugar (κ-casein contains glycated residues that can
consist of mixtures of N-acetylgalactosamine, galactose, N-acetylglu­
cosamine, N-acetyl-neuraminic acid (O’Riordan et al., 2014)), or
non-polar groups attached, which provides them with unique func­
tionalities, such as the ability to bind calcium, to associate into clusters,
or to generate repulsive interactions. Casein micelles typically have an
average diameter of around 200 nm but actually exhibit a broad range of
sizes ranging from around 50 to 600 nm. In contrast, individual plant
proteins are typically much smaller than this. For example, the 7S and
11S soy globulin protein fractions have dimensions below 6 nm (radius
of gyration) (Glantz et al., 2010; Guo et al., 2012). However, it is known
that these globular plant proteins can be made to form colloidal particles
with dimensions of around 200 nm similar to casein micelles by pro­
moting their controlled aggregation, e.g., by heating under specific pH
and ionic strength conditions. The drawback is that these plant protein
aggregates typically behave differently from casein micelles because of
their different surface chemistries, which leads to different kinds of
aggregation behavior and gelation properties (Chen et al., 2019).
Nevertheless, current research shows that the dimensions of plant pro­
tein aggregates have an important influence on their gelling properties.
For instance, the water holding capacity of gels prepared from soy
protein aggregates produced by controlled heat treatment has been re­
ported to depend on their size, as well as the crosslinking agent used to
promote their gelation (Wang et al., 2017; Wu et al., 2019). The impact
of crosslinker type is likely to be related to the nature of the functional
groups exposed on the surfaces of the proteins (Ni et al., 2015).
Consequently, it is important to be able to control the dimensions and
surface chemistry of any plant protein aggregates that are going to be
used to mimic the behavior of casein in cheese. It should also be noted,
that plant protein aggregates have been reported to evoke a gritty
mouthfeel that is undesirable to consumers, especially in fresh-cheese
analogs (Li et al., 2013). Therefore, it is also important to be able to
design casein-micelle mimetics that do not exhibit this undesirable
sensory attribute.
Several researchers have tried to mimic the properties of real cheeses
using different plant proteins and processing methods following the
fractionation route based on knowledge of casein micelles and the curds
they form when they aggregate. The overall processing scheme is shown
in Fig. 3.
In general, gels with different properties can be produced from the
same protein using different gelation techniques, as demonstrated by
Ben-Harb et al. (2018). The authors used enzymatic (chymosin and
transglutaminase), acid (glucono-delta-lactone), and thermal treatments
to induce gelation of a 15% pea protein dispersion. They concluded that
all treatments resulted in a sol-gel transition but the elasticity was
highest in the acid-induced gels and the highest resistance to strain was
in the enzymatic-induced gels. Thus, by carefully controlling the gela­
tion conditions, different structures ranging from soft to hard
plant-based cheeses might be obtained. However, the structures formed
as described below are typically known as semisolid emulsion gels, that
consist of a continuous protein network that entraps liquids (oil/fat,
water) and other ingredients, such as polysaccharides (Fig. 4C).
Some researchers have utilized the self-association tendency of
hydrophobic proteins when dispersed in aqueous solvents, such as zein
(the prolamin of maize), to mimic the aggregation behavior of casein
micelles upon the addition of proteolytic enzymes (Glusac et al., 2018,
2019; Mattice & Marangoni, 2020). For instance, Mattice and Mar­
angoni (2020) employed a 30% zein dispersion mixed with 1.5% fat,
2.8% starch, and 0.7% xanthan. The dispersion was mixed and heated to
80 ◦ C for 5 min to allow for plastification, followed by cooling. The
resulting plant-based cheese analogs had similar properties to regular
cheddar cheese with comparable values obtained through texture profile
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Trends in Food Science & Technology 118 (2021) 207–229
analysis and stretchability tests. However, it has to be noted that zein
has a relatively poor nutritional value (PDCAAS score) because of its low
level of bioavailable essential amino acids (Boye et al., 2012). To
overcome this shortcoming, other researchers have mixed zein with
other proteins, such as those from potato and pea, to form plant-based
cheese analogs.
In these studies, emulsification and enzymatic crosslinking were
employed to form plant-based cheese analogs (Glusac et al., 2018,
2019). The protein-coated oil droplets formed by homogenization were
crosslinked by tyrosinase to simulate the aggregation processes that
occur in milk after rennet is added. Tyrosinase was used rather than
transglutaminase, because it was better able to crosslink the zein mol­
ecules (Mattice & Marangoni, 2021). These emulsions were fabricated
by homogenizing an aqueous phase containing pea or potato protein
with an oil phase containing zein and then incubating them together
with tyrosinase, which led to a final emulsion with an oil concentration
of 40% and zein concentration of 0.8% (Glusac et al., 2018, 2019). The
results showed that zein increased the elastic properties of the emulsions
gels, which were further enhanced by the crosslinking action of the
tyrosinase. The mixture of pea and zein protein induced a crumbly solid
texture, whereas the combination of potato and zein protein yielded a
viscous paste. A similar cross-linking approach has been utilized for soy
proteins. In this study, a plant-based cream cheese analog was produced
when soy tofu was recombined with around 10–20% of soy protein
isolate and maltodextrin followed by incubation with transglutaminase
for 24 h (Lim et al., 2011). The soy-based cream cheese produced was
more solid-like at room temperature, but less elastic at refrigerated
temperature than real cream cheese. The enzyme crosslinking approach
coupled with acidification has also been utilized by Holz-Schietinger
et al. (2014) to produce plant-based cheese analogs from pea proteins.
These authors used 4% pea globulins, 20% of a sunflower cream frac­
tion, and a standardized starter culture. After 1 h of incubation, trans­
glutaminase was added and the dispersion was incubated for 12 h to
reach a final pH of 4.2 and to induce the sol-gel transition. After the
removal of the plant-whey fraction, the plant-based cheese produced
was transferred into molds and stored at 4 ◦ C.
A heat-treatment approach for creating plant-based cheese analogs
was described in the patent of Holz-Schietinger et al. (2014). Melting
plant-based cheese curds were obtained by a simple heat treatment
applied to a homogenized mixture of 6% soy protein globulins and 20%
sunflower oil to 95 ◦ C. Another approach combined acidification with
thermal treatment resulting in a reversibly melting cheese by using a
mixture of 4% moong 8S proteins at pH 7.4 and 50 mM sodium chloride,
mixed with 20% palm oil and inoculated with commercial starter cul­
tures and 3% sodium citrate after heating to 95 ◦ C. Similarly, a stretchy
cheese was obtained by utilizing a mixture of soy globulins (4%), pea
globulins (2%), and pea prolamins (2%) after heating to 95 ◦ C and
inoculation with starter cultures after heating. The authors related the
stretchy and melting behavior of the cheese analogs especially to the
presence of the prolamins.
In summary, a number of different fractionated plant-based in­
gredients, used in isolation or in combination, have been shown to form
cheese-like structures. The creation of gelled textures can be achieved
using a variety of physicochemical mechanisms, including gelatinization
of starch and controlled aggregation of proteins through physical,
chemical, and enzymatic crosslinking methods. The limited number of
studies that have been carried out so far show that it is possible to create
materials with cheese-like textures by carefully controlling the types and
amounts of functional plant ingredients used, as well as the processing
operations employed to induce structure formation. Nevertheless,
knowledge about some important processing operations, sensory prop­
erties, and nutritional aspects of these materials is still lacking. Espe­
cially there is a lack of studies carried out on ripening conditions and
processes that enhance the food safety, such as pasteurization. More­
over, there is a lack of data published about what kind of structures are
formed. Most likely, viscoelastic emulsion gels with continuous starch or
L. Grossmann and D.J. McClements
protein networks are formed as shown in Fig. 4, but it is unclear to what
extend filled or particulate (aggregated droplets) gels are formed (Far­
jami & Madadlou, 2019).
3.2.2. Tissue disruption route
The previous section outlined approaches to generate cheese-like
structures from fractionated plant ingredients that were recombined to
generate specific compositions. Another approach is to employ non-
fractionated plant-based ingredients. In this case, whole plant mate­
rials (such as seeds) are broken down into a colloidal dispersion of small
particles and dissolved molecules (a “plant-based milk” or a “plant-
based paste”), which is then converted into a cheese analog by pro­
moting a sol-gel transition (McClements et al., 2019). There are a
number of commercial cheese analog products that are produced using
the tissue disruption route that generate a plant-based milk first
(Table 1). For instance, soy, pea, lupin, oat, and nut milk have all been
utilized as starting materials to form this kind of product. Afterwards, a
number of different processing steps are required to convert these
plant-based milks into cheese analogs. Some of the most commonly used
manufacturing operations employed to promote the sol-gel transition
are heat treatment, acidification, enzymatic crosslinking, and
salting-out, either alone or in combination.
The formed structures are emulsion gels with a continuous protein
network, which entraps the liquids and other particles. In principal,
these structures are obtained from either a diluted (i.e., a plant-based
milk, Fig. 4C) or a concentrated (i.e., a plant-based paste, Fig. 4D)
colloidal dispersion that is transformed into an emulsion gel by different
principles as outlined below and in Fig. 5. Here, the main difference is
that the diluted dispersion is characterized by the removal of insoluble
particles, such as larger protein fragments or insoluble polysaccharides
such as cell wall material, whereas the concentrated dispersion is pro­
duced from the whole seeds (e.g., nuts) without separating any com­
pounds and thus insoluble materials and particles remain within the
plant-based cheese and commonly no whey is separated. However,
overall this tissue disruption processing approach is more closely related
to the production of cheese from regular milk.
The sol-gel transition is induced by various means and foremost it
has to be mentioned that most studies use a combination of heat treat­
ment combined with other techniques that induce the formation of a 3-
dimensional protein network (Table 2). As described earlier, globular
proteins become more susceptible to aggregation after heat denatur­
ation because this increases the attraction between them by exposing
more hydrophobic and other functional groups (Fig. 6) (Ni et al., 2015).
These unfolded proteins are more susceptible for interactions and to
form a gel by the addition of calcium or other salts (electrostatic
screening and calcium bridging of carboxyl groups) or by lowering the
pH close to the isoelectric point (reduction of electrostatic repulsion)
(Zhang et al., 2018; Zheng et al., 2020). Thus, a heat-treatment
commonly “activates” the proteins by unfolding and exposing more
functional groups but does not induce enough attractive potential to
induce a sol-gel transition, if carried out carefully. By implementing
another step, such as adding ions, or altering the pH of the colloidal
dispersion, the attractive interaction becomes strong enough for the
proteins to interact. Such behavior may be modeled in a first approxi­
mation by describing the interaction potential between two spherical
colloidal particles that are influenced by van der Waals, electrostatic,
steric, and hydrophobic interactions as shown in Fig. 6. Nonetheless, the
heat treatment step is commonly part of the dispersion preparation and
is also used to inactive enzymes like lipoxygenases to minimize the
production of off-flavors and it is not well understood what the effect of
the heat treatment on the structure formation in plant-based cheeses
actually is.
However, a heat treatment approach without any salt addition has
been successfully used in the study of Awad et al. (2014) who used lupin
seeds as a starting material for the plant-based cheese. Initially, lupin
seeds were transformed into a colloidal dispersion by soaking and
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Trends in Food Science & Technology 118 (2021) 207–229
grinding them. This dispersion was then mixed with oil (17.5%),
emulsifying salts, and water (11.8%) and heated at 85–90 ◦ C for 10 min.
The resulting plant-based curd was hot molded and stored for up to 3
months. Compared to regular cheese, the lupin-cheese exhibited
increased oil separation, increased firmness, and lower springiness. The
plant-based cheese had also acceptable sensory scores but these were
lower than those found for regular cheese.
In contrast, the most well-known plant-based cheese is fermented
tofu, which is produced from heat-treated soymilk combined with an
ion-gelation or an acidification approach. Commonly, soy milk is first
heated to 65–95 ◦ C to induce denaturation of the globular proteins and
exposure of the hydrophobic and other functional groups as explained
before. After the heat treatment, a sol-gel transition is induced by the
addition of either calcium ions (to promote ion bridging of the soy
globulins and charge screening) and/or by lowering the pH (to promote
isoelectric precipitation of the soy globulins) (Zhang et al., 2018; Zheng
et al., 2020). Despite the fact that soy proteins have many sulfhydryl
groups, disulfide bridges are not thought to be important in stabilizing
tofu structures (Kohyama et al., 1995). The resulting curd is pressed and
either soft or hard tofu can be produced depending on the properties of
the gel network and the amount of soy whey removal (Kao et al., 2003).
After pressing, the tofu is typically diced into cubes and then aged for up
to several months with various strains of Actinomucor spp., Rhizopus
spp., or mold Mucor spp. The fermentation induces proteolysis of the soy
protein (mainly the 7S and 11S globulins in tofu) by different enzymes or
proteases from microorganisms, followed by secondary proteolysis and
enzymatic or chemical conversion of amino acids into derivatives (Liu
et al., 2018). This process is therefore analogous to the structural
breakdown and volatile generation that occurs due to microbial
fermentation in regular cheese. However, the final textures and aromas
produced are often quite different from those in cheese made from an­
imal milk. For this reason, there is still an interest in creating plant-based
cheese analogs using other processing routes and starting materials so as
to more closely mimic real cheese. For example, fermentation has been
carried out using other kinds of plant proteins, as well as other types of
microbes. Other studies demonstrated that such ion-gelation ap­
proaches used in tofu production are also useful for converting other
plant-based milk into plant-based cheeses. For example, a combination
of soy- and cashew nut milk with the aim of improving the nutritional
value over traditional soy-based products has been used (Oyeyinka
et al., 2019). The plant-milks were obtained by a typical soaking, mill­
ing, and filtration process scheme. An optimum sensory acceptance was
reported for cheese analogs produced from 60% soy milk and 40%
cashew nut milk. The sol-gel transition was achieved by heating the milk
at boiling temperature, followed by the addition of ammonium
aluminum sulfate to induce coagulation (Table 2). The curd produced by
this method was filtered, pressed, and cooked again with salts and
spices. The resulting plant-based cheese had a protein content of 64%
and a lipid content of 6% in the dry matter. The authors concluded that
mixing two kinds of plant-based milk might be a feasible option for the
sensory acceptance and nutritional profile of soy-based cheeses.
Some studies have used a lactic acid fermentation (i.e., acid curd
preparation) approach to produce cheese analogs from colloidal dis­
persions of plant materials. For instance, a soy milk has been used as a
starting material and the overall process was similar to tofu production
but the sol-gel transition was achieved by using starter cultures (Matias
et al., 2014) and not glucono-delta-lactone or calcium addition, as
described before (Zheng et al., 2020). Similarly, the soy milk was heated
to 95 ◦ C for 10 min and incubated with strains of Bifidobacterium animalis
subsp. lactis Bb-12, L. acidophilus La-5, and the starter S. thermophilus.
The authors obtained two different structures, depending on the final
pH. A quark-type cheese structure was produced when the pH was
lowered to 5.7 and a soy-cheese was obtained at 4.8 after the removal of
the plant-whey fraction (Matias et al., 2014). This reveals that the type
of structure formed is influenced by the pH and thus the magnitude of
the electrostatic repulsion interactions between the protein moieties. In
L. Grossmann and D.J. McClements
this study, the soy-cheese was further processed with soy cream and
hydrocolloids to obtain a product similar to petit-suisse, a cream cheese.
In another study, Streptococcus thermophilus and Lactobacillus fermentum
were employed to obtain a semi-hard cheese based on soy milk fer­
mented to a pH of 4.5 after heat treatment to 63 ◦ C for 30 min and
pressing. While the plant-based cheese had similar textural properties as
its dairy counterpart, the sensory scores were lower (Chumchuere et al.,
2000). Similarly, spreadable and firm plant-based cheeses were pro­
duced by fermenting soy milk with Lactobacillus casei ssp. casei to reach a
final pH of 6.3 or by utilizing a combination of Lactobacillus acidophilus
and Bifidobacterium animalis subsp. lactis together with
glucono-delta-lactone to reach a final pH between 4.7 and 7.2 (Giri
et al., 2018; Li et al., 2013). Giri et al. (2018) achieved spreadability by
mixing the curd with soybean oil to achieve a fat content of around 25%
and a protein content of 18%. Moreover, it was demonstrated that
papain could be employed to improve the sensory scores of soy-based
cheese spreads by lowering the colloidal protein particle size and
therefore most likely reduce the grittiness (Li et al., 2013). Last, a
soy-based plant cheese was produced by inoculating soy milk with
Lactobacillus rhamnosus and fermenting the plant-based milk to a pH
between 4.7 and 5.2. The curd was cut, pressed, and salted for further
analyses (Liu et al., 2006). On a final note, the production of plant-based
cheese by lactic acid fermentation has been mainly reported for
soy-based products but should be expanded also for other raw materials.
New research could leverage from findings on the fermentation of
plant-based milks derived from lupin and others (Canon et al., 2020;
Tangyu et al., 2019).
The interactions and structural organization of the molecules in
plant-based cheese analogs obtained using the heating → pH-
precipitation processing sequence are typically considerably different
from that present in regular cheeses. For instance, the structures in
cheeses obtained from rennet curd are stabilized by calcium bridges,
hydrogen bonds, and hydrophobic interactions (Paula Vilela et al.,
2020), whereas those in such plant-based cheese analogs as described in
the previous section are mainly stabilized by hydrophobic attraction
(Fig. 6). Thus, other studies have utilized either the addition of multi­
valent cations described above for the tofu production (Chikpah et al.,
2015) or a combination of acid- and ion-gelation (i.e., a combination of
lowering the pH and the addition of calcium) to prompt the sol-gel
transition (Table 2) (El-Ella, 1980; Verma et al., 2005; Zulkurnain
et al., 2008). In this case, the amount of calcium bound to the soy pro­
teins depends on the pH used and is typically decreasing as the pH is
lowered because then there are fewer ionized carboxyl groups (-COO-)
present (Canabady-Rochelle et al., 2009). One early study used
heat-treated soy milk that was subsequently inoculated with Lactococcus
lactis (El-Ella, 1980). After the initial fermentation, the sol-gel transition
was induced by the addition of calcium lactate. The curd was pressed,
salted, and ripened for up to three months. Results showed that the
plant-based cheese had a flat flavor after 1 month of ripening, which
turned to a slightly acid and cheesy flavor after 2–3 months. In another
study, pigeon pea milk and soy milk were used to produce a plant-based
cheese. The milk was prepared by grinding, followed by filtration and
inoculation with Streptococcus thermophilus and Lactobacillus delbrueckii
subsp. bulgaricus. The plant-based curd was obtained through coagula­
tion of the fermented milk by the addition of calcium (0.02% CaCl2) and
a heat treatment (up to 95 ◦ C, 15 min), followed by filtration and
pressing of the curd (Verma et al., 2005). The authors carried out ex­
periments to identify the optimum processing conditions to promote
good curd production. This study used a combination of heat denatur­
ation, pH reduction, and calcium addition to produce a plant-based
cheese curd. However, other studies carried out with tofu have
showed that the breaking energy and storage moduli are fairly similar
for curds obtained with glucono-delta-lactone and calcium sulfate
(Kohyama et al., 1995). Thus, the question remains how specific mo­
lecular and colloidal interactions influence curd formation and proper­
ties and how these can be manipulated to create cheese analogs with the
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Trends in Food Science & Technology 118 (2021) 207–229
desirable properties required.
Enzymatic cross-linking has been employed in a series of ap­
proaches to produce plant-based cheeses as described in different pat­
ents and studies (Brown et al., 2013; Holz-Schietinger et al., 2014;
Murata et al., 1987; Sánchez-Muñoz et al., 2017). In the patents, mac­
adamia and almond nuts were soaked in boiling water for 30 s, followed
by 16 h hydration, and blending to disrupt the plant tissue structure
(Brown et al., 2013; Holz-Schietinger et al., 2014). The emulsified
slurries were then centrifuged to remove insoluble solids and the
resulting skim and cream phases were recombined at different ratios,
depending on the cheese type. The plant-milk was subsequently
pasteurized and cooled before further processing. The plant-based
cheese types prepared with this milk were for example fresh,
soft-ripened, and salted (cheddar-like) plant cheeses. The curds for these
three plant cheese types were prepared following the same overall
procedure. The milk was first fermented with mesophilic starter cultures
(and Geotrichum candidum, Penicillium candidum, and Debaromyces han­
senii for soft-ripened cheese). The curds formed had a pH of 5.6 after 1 h
and a pH of 4.4 after 12 h of fermentation. The crosslinking enzyme
transglutaminase was then added and the system was incubated at room
temperature for another 12 h to induce the sol-gel transition. The
resulting curd was cut into diced pieces of around 1.3 cm and around
40–50% of plant-whey was removed followed by a whisking step for 10
min. Depending on the cheese type, the cheese was molded with addi­
tional pressure (fresh cheese and salted cheese) or without pressure
(soft-ripened cheese) to obtain the necessary weight loss ratio. The
cheeses were subsequently brined for 30 min at 10 ◦ C. The fresh cheese
was packed without ripening after draining, whereas the soft-ripened
cheese was aged for 17 days at temperatures between 10 and 16 ◦ C
and relative humidities between 75 and 90%. The salted cheese was
aged for 3 weeks at 13 ◦ C and 55% relative humidity followed by
smoking or waxing. The authors of this patent also highlighted that the
addition of exogenous proteases and lipases to the milk or the curd
improved the textural and sensorial attributes, most likely by similar
processes as during the ripening stage of regular cheeses. Moreover, they
evaluated the effect of fermentation on the production of different vol­
atile compounds typically attributed to cheese, such as fermenting with
Staphylococcus xylosus and Brevibacterium to generate 3-methyl butanoic
acid and dimethyl trisulfide, respectively (Holz-Schietinger et al., 2014).
Another enzymatic approach was taken by Murata et al. (1987). Instead
of inducing covalent crosslinks via enzymes, the authors studied
different proteases to induce a sol-gel transition by proteolysis. Thus,
this approach is more similar to regular milk curdling and the study
revealed that a proteolysis induced a sol-gel transition in soy milk at pH
6.1 for most proteases used, except for Aspergillus saitoi proteases,
rennin, and pepsin.
Finally, some commercial products use fermented nut paste (i.e.,
acid curd preparation), often cashew nuts as a raw material, for pro­
ducing plant-based cheese analogs, typically camembert-style cheeses.
This approach has the advantage that ingredients that are known to have
a positive impact on food structure and stability are not separated. For
example, phospholipids and fibers are retained which may improve
emulsifying and oil-holding properties (Nikiforidis, 2019;
Sánchez-Zapata et al., 2009). The main difference in this approach
compared to the previous ones is that the whole kernels are used to form
a concentrated dispersion, whereas the aforementioned approaches
have typically removed the solids (insoluble proteins and poly­
saccharides) during the plant-based milk production step, which yields a
diluted colloidal dispersion (Fig. 4C and D). Here, the nuts are soaked in
boiling water for several hours to ease the blending and reduce the
microbial load of the kernels. The nuts are then blended into a
concentrated colloidal dispersion with the addition of water and inoc­
ulated with mesophilic lactic acid bacteria and with fungus, such as
Penicillium camemberti, Lactococcus lactis subsp. lactis, and Lactococcus
lactis subsp. cremoris (Fig. 5). The cheese curd is fermented at optimum
conditions for mesophilic bacteria to grow for around 24 h, formed, and
L. Grossmann and D.J. McClements
further fermented for several weeks at a temperature from around 12 to
18 ◦ C and a relative humidity >80% until the required characteristic
flavor profile is developed and the surface is covered with mold. For
example, a recent study prepared different cheese types based on
grounded cashew kernels that were inoculated with a fermented quinoa
dispersion and chickpea miso, depending on the final plant-based cheese
type (Chen et al., 2020). The authors concluded that the ‘brie’ and ‘blue’
cashew plant-based cheeses became dominated by Lactococcus, Ped­
iococcus, and Weissella genera. Moreover, the allergenicity decreased
with fermentation. However, it is still unclear what kind of structure is
formed upon acidification in such plant-based cheeses prepared from
concentrated dispersions. Most likely, a viscoelastic filled emulsion gel is
formed in which the fat droplets are embedded in an aggregated protein
matrix that is further enriched with polysaccharides. However, the
advantage of this approach is that no whey fraction is generated and all
the nutrients in the original nuts are incorporated into the cheese
analog. Thus, this plant-based cheese is rich in proteins and fats
(Table 1).
In conclusion, it should be mentioned that there is currently a lack of
published data on the important physicochemical parameters of most
plant proteins (i.e., denaturation state, aggregation state, surface
chemistry, and impurities), which prevents fine-tuning of processing
parameters to create the structures desired in plant-based cheeses. The
few examples presented here show that there should be more research
on every stage of the plant-based cheese analog process to establish the
molecular, physicochemical, material, and process parameters required
to generate a blueprint for the structuring process as well as to generate
the desired flavor. Currently, there is a lack of knowledge of the mo­
lecular events occurring during processing and structures formed, which
limits further improvement. In the future, the structure and texture of
the cheese analogs could be further tuned by carefully controlling the
colloidal interactions through balancing the degree of denaturation, pH,
ionic strength, or by the addition of crosslinking enzymes.
4. Products on the market
There is a growing market for plant-based cheeses, which is expected
to increase appreciably in the future as their quality improves and more
consumers are willing to purchase them (Fig. 1). The products currently
available have been created using both processing approaches outlined
in this article: the fractionation route and the tissue disruption route.
Some current products sorted by their main ingredient into two different
processing routes are given in Table 1. While this table provides a broad
overview of products on the market at the time of writing this review, it
is certainly not complete. Even so, it does provide a good overview of
cheese analog types, ingredients, and nutrient compositions. A variety of
plant-based cheese analogs has already been developed and marketed,
including cheddar-, gouda-, mozzarella-, camembert-, provolone-, and
parmesan-style cheeses. Overall, however, there is still a lack of diversity
in the product range compared to regular cheese, which might change in
the future with emerging processing approaches and ingredients being
developed. This is in line with a study carried out on commercial plant-
based cheeses in the UK that identified 109 products on the market with
the majority of 74% of them being coconut-oil based, 10% of them nut-
based, 6% of them palm-oil based, 5% rice-based, 3% soy-based, and 2%
sunflower oil-based (Nicolás Saraco & Blaxland, 2020).
Products following the fractionation route are mainly based on
starches and coconut or palm oil (Table 1). It has to be emphasized that
these products may successfully mimic the functional and textural at­
tributes of regular cheese but they do not deliver the same nutrients as
regular cheese, which will be discussed in more detail in the following
section. Because these products are designed with high amounts of co­
conut and palm oil, they also contain high amounts of saturated fatty
acids. Moreover, because isolated ingredients are used in these formu­
lations, thickening agents (e.g., methylcellulose, carrageenan, or agar),
colorants, and flavors are commonly added (Nakamura et al., 2016). By
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the addition of such additives to isolated ingredients, the manufacturers
can specifically tune and design the functional, textural, and sensorial
attributes of the plant-based cheese. However, consumers often prefer
not to have long ingredient lists and such formulations might not be in
line with clean-labeling efforts.
Plant-based cheeses produced following the tissue disruption route are
typically based on soy- and hemp milk or are produced using whole
cashew nuts (Table 1). While there are some plant-based cheeses that
only contain a limited number of ingredients (e.g., cashews, water,
starter cultures, and salt), other products also contain coconut oil,
thickening agents, and flours to optimize the textural attributes that
have been produced by employing the tissue disruption approach. Thus
these products are produced by a combination of the fractionation and
disruption routes. However, the protein contents are typically higher
compared to products based on starches and are responsible for forming
the continuous 3-D network in many products. Nonetheless, the overall
protein content is still lower compared to real cheeses. In contrast, plant-
based cheeses based on cashew nuts often have lower contents of satu­
rated fats and salts, which will be discussed in more detail in the
following section.
5. Sustainability, health, and ethical considerations
Current estimates indicated that reducing the proportion of animal-
derived food products in the human diet would have a positive impact
on many environmental sustainability factors, including a reduction in
greenhouse gas emissions and pollution (Clark et al., 2019; Willett et al.,
2019). Current studies estimate that 34% of total greenhouse gas
emissions are related to the food system (Crippa et al., 2021). It should
be noted that animal-based foods can be produced on marginal lands
that are unsuitable for growing arable crops and that they can utilize
side streams from food production, thereby converting waste materials
into food (Cassidy et al., 2013; Salter, 2017; Shahid & Al-Shankiti, 2013;
van Zanten et al., 2016). Consequently, there are sustainability argu­
ments for maintaining some level of animal production for food. A
limited number of studies exist that have investigated production ca­
pacities using such measures and these studies report that every day
between 7 and 27 g per animal-based protein per capita could be pro­
duced using for example marginal land or byproducts of food production
(Hal et al., 2016, p. 137). Nonetheless, recent studies indicate that there
are substantial opportunity costs for using pastures for ruminant meat
and dairy production, because less carbon is sequestered through
ecosystem restoration when used as feedstock (Hayek et al., 2021).
However, the increase in awareness of the relationship between the food
system and environmental sustainability as well as health are major
drivers for the development and release of plant-based cheese products.
While it is beyond the scope of this review to outline the research carried
out on the sustainability of the food system and healthy diets over the
past decades, we want to briefly discuss some important recent de­
velopments. In addition, we briefly highlight some of the ethical issues
that are also important to some consumers.
5.1. Global warming
Plant-based cheeses are typically designed to mimic a regular cheese
type and thus the environmental sustainability should be compared to
regular cheese. As cow cheese is the most often consumed cheese type,
we will focus on this type but results may vary by the source of the milk.
In general, the global warming potential of regular cheese is mainly
influenced by the production of milk, which typically accounts for
around 65–98% of CO2-eq emissions (average 82%) of the cheese,
depending on the type of cheese and the boundary conditions used (Bava
et al., 2018; Finnegan et al., 2018; González-García et al., 2013; van
Middelaar et al., 2011; Üçtuğ, 2019). The environmental impact of the
production and retail stage was mainly influenced by the type of energy
source, wastewater treatment, packaging, and logistics used (Bava et al.,
L. Grossmann and D.J. McClements
2018; González-García et al., 2013; Dalla Riva et al., 2017; Tar­
ighaleslami et al., 2020).
The raw materials used are likely to have the greatest impact on the
environmental impact of plant-based cheeses compared to regular
cheeses. Typically, cow’s milk has a global warming potential of around
1.39 kg CO2-eq per L (min: 0.54; max: 7.50 kg CO2-eq per L) (Clune
et al., 2017). It has to be noted that this value highly depends on farm
management practices, geographical region, study design, allocation of
other products, and other issues (Baldini et al., 2017). However, because
1 kg of cheese requires commonly 4–10 L of milk (depending on cheese
type) the impact of the raw milk per kg of cheese is in the range of 1.62
(fresh cheese) to 8.3 (semi-hard cheese) kg CO2-eq per kg of cheese,
resulting in total average emissions of 8.86 (min: 5.33; max: 16.35) kg
CO2-eq per kg of cheese (Clune et al., 2017; Finnegan et al., 2018).
The comparison of these values to plant-based cheeses is challenging
at the moment because there is currently a lack of life cycle assessments
(LCAs) on plant-based cheese production, probably because of the lack
of producers. However, because the raw material has the greatest impact
on the environmental sustainability of regular cheeses (Bava et al., 2018;
Finnegan et al., 2018; González-García et al., 2013; van Middelaar et al.,
2011; Üçtuğ, 2019), a reasonable comparison might be drawn based on
the life cycle assessments (LCAs) carried out on the raw materials used
for plant-based cheese production and further studies are needed to
compare the impact of processing on the greenhouse gas emissions of
plant-based cheese products.
For some raw materials mentioned in this review, the global warm­
ing potential in kg CO2-eq per kg or L has been reported as: almond and
coconut milk 0.42 (min: 0.39; max: 0.44), soy milk 0.88 (min: 0.66; max:
1.40), soy protein isolate 2.4 per kg protein, pea milk 0.387, cassava starch
0.594, tree nuts 1.42 (min; 0.43; max: 3.77), palm oil 1.4–2.0, and sun­
flower oil 0.8 (Braun et al., 2016; Clune et al., 2017; Henderson &
Unnasch, 2017; Schmidt, 2015; Usubharatana & Phungrassami, 2015).
It has to be noted that these findings may be somewhat limited by the
functional unit employed and the number of published studies. For
example, almond and coconut milk have lower nutritional values (e.g.,
protein) and therefore a conclusive comparison based on volume or
mass is not always applicable. Moreover, the conversion ratios of
plant-based milk to plant-based cheese have not been reported yet, i.e.,
how much plant-based milk is needed to produce one kg of plant-based
cheese, which may limit the conclusions at the time of writing this
review.
However, based on the data reported, it may be concluded that the
global warming potential of plant-based cheese ingredients is lower than
regular cow’s cheese. Assuming a similar conversion ratio of milk to
cheese, plant-based cheeses may have lower CO2 emissions than regular
cheese. For example, cheeses based on nuts may have lower CO2 emis­
sions because the whole kernels are used, enabling a direct comparison
between these two products. Therefore, the presented data may already
show that there is a potential to reduce the CO2 emissions by employing
nuts as a base ingredient for plant-based cheeses: 1.42 kg CO2-eq for 1 kg
of nuts vs. 6.5–8.3 kg CO2-eq for milk produced for soft to semi-hard cow
cheeses (Clune et al., 2017; Finnegan et al., 2018).
The assumption that plant-based cheeses have a lower global
warming potential is further reinforced by life cycle assessments carried
out on soy-based tofu, which has comparable processing operations
compared to regular cheese and has a reported global warming potential
of 0.98 kg CO2-eq per kg tofu (which equals to 9.8 kg CO2-eq per kg
protein) using a farm to factory gate approach (Mejia et al., 2018). As
already mentioned, this value is lower than commonly reported values
of cheeses with a mean of 8.86 (min: 5.33; max: 16.35) kg CO2-eq per kg
cheese (Clune et al., 2017). However, the difference might be smaller
when calculated based on the same product category (e.g., semi-hard
cheese types) and on actual nutrient values such as protein or the con­
tent of essential amino acids, which has been shown to have a strong
influence on data interpretation (Tessari et al., 2016). In conclusion, life
cycle assessment studies on plant-based cheeses are needed to draw a
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Trends in Food Science & Technology 118 (2021) 207–229
final conclusion on these preliminary assumptions.
5.2. Health aspects
Plant-based cheeses may become an important part of diets in the
future for consumers who follow a plant-based or mainly plant-based
diet or try to reduce their intake of animal-based products. With more
and more people switching to an increase in consumption of plant-based
foods, plant-based cheeses may become a more important factor in
people’s diets on a daily basis. Thus, we want to briefly elucidate the
possible effects of such diet shifts. We begin by briefly discussing some
general findings regarding plant-based diets that have been published in
the last few years, followed by a review of the health impacts of cheese
consumption. Lastly, we briefly elaborate on the possible health effects
of the consumption of plant-based cheeses based on the ingredients
used.
In general, an increase in consumption of healthy plant-based foods
(e.g., whole-grains, fruits, vegetables, nuts, legumes, tea, and coffee
(Kim et al., 2018)) has been associated with positive health outcomes in
observational studies, such as a decrease in all-cause and cardiovascular
disease mortality whereas an unhealthy plant-based diet (e.g., refined
grains, fruit juice, sweets, sugar sweetened beverages) seems to have no
such positive effects (Kim et al., 2018, 2019; Satija et al., 2017).
Moreover, a comprehensive meta-analysis of observational studies
concluded that a vegetarian diet is associated with a lower incidence
and/or mortality from ischemic heart disease and incidence from total
cancer but there was no correlation with all-cause mortality and mor­
tality from cancer (Dinu et al., 2017). The same effects might be true for
vegan diets, but comprehensive meta-analyses are still missing due to a
lack of studies.
However, these reported positive effects are based on some studies
that concluded a lower risk for specific causes of death, the occurrence of
diseases, and also all-cause mortality in vegetarians compared to meat-
eaters but some other studies could not replicate these findings in
population-based studies and some even found enhanced risks for some
diseases such as stroke (Appleby et al., 2016; Mihrshahi et al., 2017; Mj
et al., 2013; Tong et al., 2019). These contradicting results are
commonly explained by the fact that early studies did not distinguish
between healthy and unhealthy vegetarian/vegan diets and that the
foods eaten in vegetarian diets have turned more towards unhealthy
foods in recent years (Magkos et al., 2020; Mihrshahi et al., 2017). For
example, a diet based on starchy and sugar-rich plant-based foods is still
considered vegetarian/vegan but may have adverse health effects. These
findings are especially relevant for processed plant-based foods. Some
plant-based cheese types mainly contain processed ingredients such as
starch and oils (Table 1) and further studies need to elucidate their ef­
fects on human health in this food matrix. Such methodological gaps
have been closed by using plant-based diet indices (Kim et al., 2018,
2019; Satija et al., 2017). In these studies, the food intake of different
cohorts was evaluated based on plant-based diet indices that group
certain foods as either plant- or animal-based (= plant-based diet index)
and additionally as “healthy” or “unhealthy” (= healthy/unhealthy
plant-based diet index). It has to be noted that in these studies even
participants in the group with the highest consumption of plant-based
foods consumed on average some animal products such as meat. How­
ever, in these studies, all animal-based foods are categorized in one
group whereas it is known that not all animal-based foods have the same
association with negative health outcomes, such as dairy foods (Fardet &
Boirie, 2014; Mazidi et al., 2019; Schwingshackl et al., 2017). Thus, it
was hypothesized that well-controlled omnivore diets with a lower
amount of animal-based foods (e.g., the Mediterranean diet) may have
similar effects (Magkos et al., 2020).
The case of cheese is similarly not straightforward. While most
observational studies indicate that there is no link between cheese
consumption and higher mortality or higher risk of cardiovascular dis­
ease (de Goede et al., 2016; Farvid et al., 2017; Guo et al., 2017;
L. Grossmann and D.J. McClements
Hjerpsted & Tholstrup, 2016; Mazidi et al., 2019; Pala et al., 2019),
some authors note that controlled studies are missing to support this
statement (Sacks et al., 2017). The fact that most observational studies
could not find a link is surprising because cheese has a high content of
saturated fatty acids (e.g., 100 g of gouda contains around 18 g of
saturated fat), which has been linked to higher all-cause mortality and
an increased risk for cardiovascular diseases due to its LDL
cholesterol-raising effect especially when consumed instead of fats rich
in mono- and polyunsaturated fatty acids but not when replaced with
mostly refined carbohydrates (Sacks et al., 2017). However, there is still
an ongoing debate about whether saturated fat is actually detrimental to
human health and which fatty acids provoke these effects (Heileson,
2020; Lawrence, 2021). Nonetheless, it has been supposed that
food-matrix effects may impact the influence of certain ingredients on
human health. For example, cheese contains high levels of calcium,
protein, some beneficial fatty acids, such as conjugated linoleic acid, and
many more compounds that all together may have a totally different
effect on the body (Hjerpsted & Tholstrup, 2016; Magkos et al., 2020).
Most likely, more well-controlled observational and clinical studies are
needed to finally answer these questions.
Now, the question remains about what the health consequences of
increased plant-based cheese consumption would be. First, it has to be
noted that there are no studies as of the writing of this review that would
allow drawing a definite conclusion based on actual data for plant-based
cheeses. However, based on the ingredients used in plant-based cheeses
(Table 1), some assumptions can be made that need to be further eval­
uated in future studies. Animal-based cheese is a food rich in nutrients
such as protein, fats, calcium, phosphorus, vitamin A, and vitamin B12
(Górska-Warsewicz et al., 2019). These compounds are highly
bioavailable to the human body. For example, milk has a high protein
quality with caseins having a digestible indispensable amino acid score
(DIAAS) of 1.29, which is considerably higher than proteins from plant
sources, which is partly because of the absence of protein inhibiting
antinutrients such as protease inhibitors found in plants (Guillin et al.,
2021; Mathai et al., 2017). Additionally, a relatively high fraction of the
calcium in dairy foods (around 30%) is absorbed, whereas a lower
fraction is absorbed in plant-based foods, which is mainly due to the
presence of antinutrients such as oxalates and phytates (Yang et al.,
2012).
Plant-based cheese that is mainly based on starch and oils may fail to
deliver the same beneficial nutrients as regular cheese, namely calcium,
protein, vitamin A, and vitamin B12. Moreover, cheese analogs often
contain relatively high amounts of saturated fatty acids and refined
carbohydrates, which might be detrimental to human health if eaten on
a regular basis. There is also some debate about whether the oils used in
cheese analogs, such as coconut oils, have the same detrimental effects
on LDL levels as other fats that contain high levels of saturated fatty
acids (Eyres et al., 2016; Hewlings, 2020).
Nevertheless, plant-based cheeses based on proteins and unsaturated
fats may have a positive impact on human health. For example, nuts are
rich in nutrients and lower in saturated fatty acids and are associated
with positive health outcomes (Chen et al., 2017; Fardet & Boirie, 2014;
Schwingshackl et al., 2017). Commonly used cashew nuts are mainly
comprised of monounsaturated fatty acids (mostly oleic acid), along
with around 18% of protein and 3% of fibers. Moreover, they contain
important vitamins (B1, B5, B6, K) and minerals (e.g., zinc, iron, mag­
nesium, copper, selenium, phosphorus) (USDA, 2021). One strategy that
can be used to improve the nutritional profile of plant-based cheeses and
match the profile with their animal counterpart is to fortify them with
the micronutrients normally found in real cheese, such as calcium,
vitamin A, and vitamin B12. In addition, they could be fortified with
other health-promoting ingredients, such as omega-3 fatty acids or
nutraceuticals. However, since most plant-based cheeses are calorie-rich
foods, they should still only be consumed in limited quantities.
Another micronutrient that is rich in most regular cheeses is salt.
Especially, cheeses such as cheddar, feta, gouda, camembert, and
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Parmigiano-Reggiano contain high amounts of sodium (typically
600–1500 mg of sodium per 100 g of cheese) while swiss cheese and
ricotta contain less (typically 100–200 mg of sodium per 100 g of
cheese) (USDA, 2021). It is well accepted that increased sodium intakes
can increase blood pressure, which in turn can lead to higher rates of
coronary heart disease, stroke, heart failure, and kidney failure (Cho­
banian & Hill, 2000; Farquhar et al., 2015; Wang et al., 2020). However,
this notion came again under discussion recently especially the effects of
very low sodium intakes (Adedinsewo et al., 2021; Messerli et al., 2020).
However, the blood pressure raising effect is especially pronounced in
people grouped as “salt-sensitive”, which affects 30–50% of the hyper­
tensive and 25% of the normotensive subjects (Balafa & Kalaitzidis,
2021; Gholami et al., 2020). Thus, a decrease in salt intake (around 2.3 g
for adults per day) is part of many international and national initiatives
(Santos et al., 2021). The plant-based cheeses shown in Table 1 also
contain significant concentrations of salt, except for some formulations.
This indicates that future plant-based cheese formulations should also
focus on decreasing the sodium content in their products.
6. Future directions
Plant-based cheeses are a promising future food that has the poten­
tial to serve as a nutritious and sustainable alternative for regular
cheese. To further improve the quality and expand the variety of plant-
based cheese types, while ensuring a nutritious and sustainable food
product, a few important future directions for research and development
are highlighted here.
• Structuring: This review outlined the main structuring approaches
currently utilized to produce plant-based cheese structures. Howev­
er, there is still a lack of targeted approaches to extract and structure
plant proteins that would enable a rational-design based formulation
approach. Current techniques are mainly based on trial-and-error
attempts but a more in-depth understanding is needed to design
structures that mimic all cheese classes, ranging from fresh soft
cheese to matured hard cheese. At the moment there is no blueprint
available that would enable scientists and manufacturers to set up
processes based on plant protein properties, which prevents the
rational design of processes to structure proteins. For example, there
is a lack of understanding of how molecular and colloidal in­
teractions influence the texture of plant-based cheese analogs
created by different structuring approaches, which is holding back
the design and production of higher quality and more diverse
products.
• Aging and sensory analysis: Cheeses are aged to obtain the desired
structure and flavor. At the moment there is a lack of data on how
plant-based cheeses could be fermented to obtain desirable flavors
and structures. Some preliminary data shows that the inoculation
with the same microbial communities results in totally different
sensorial attributes, depending on the composition of the gel (e.g.,
pea vs. milk) (Ben-Harb et al., 2020). Moreover, some cheese flavors
have been produced using soy milk by employing specific microor­
ganisms (Holz-Schietinger et al., 2014) and it is also known that
fermentation can reduce off-flavors, such as beany and earthy notes,
and the antinutrients found in many plant-based foods (Tangyu et al.,
2019). However, there have only been a few comprehensive studies
on the sensory attributes of plant-based cheese products, which have
been reviewed recently by Short et al. (2021). It is very likely that
different starter cultures will be needed to obtain the desired
different flavor profiles in plant-based cheeses. This should be a focus
of future research because there is currently very little data on the
impact of different starter cultures on the processing, physicochem­
ical properties, and sensory attributes of these products.
• Melting: Melting is a key quality attribute of regular cheese. A survey
of commercial plant-based cheeses found a lower meltability than
regular cheese, which was attributed to the use of thermo-
L. Grossmann and D.J. McClements
irreversible starch as a gelling agent (Nicolás Saraco & Blaxland,
2020). Thus, more work is needed to overcome this hurdle. A study
suggests that zein can enhance the melting properties of plant-based
cheeses because it plasticizes at higher temperatures by a decrease in
non-covalent bond strengths, similar to casein networks (Mattice &
Marangoni, 2020). Moreover, some limited data suggests that by
gelling soy, rubisco, moong bean, or mixtures of pea protein and
xanthan yields thermo-reversible gels but more research about the
underlying mechanisms is needed (Holz-Schietinger et al., 2014).
• Nutrition and sustainability: It will be important to design healthy
plant-based cheeses to produce more healthy choices for consumers.
Thus, scientists and product developers are encouraged to use less
saturated fats, salts, and starches in their plant-based cheeses, as
discussed in this review. This will be challenging due to the chemical
stability and structuring features of saturated fats and more research
is necessary to find alternative structuring approaches to incorporate
more healthy fats in plant-based cheeses. Moreover, it will be
necessary to assess the nutritional and sustainable properties of
plant-based cheeses to ensure that there are no adverse effects on the
human body and the environment by consuming these foods.
Advanced encapsulation technologies, such as nanoemulsions, may
prove useful for fortifying the next generation of cheese analogs with
health-promoting ingredients, such as vitamins, omega-3 oils, or
nutraceuticals.
• Food safety: There have been very few food safety studies carried out
on plant-based cheeses as of the writing of this review. This gap
needs to be filled to guarantee safe plant-based cheese products for
consumers.
7. Conclusions
This review has outlined some of the most important advances that
have been made on the design and development of plant-based cheeses.
The use of more environmentally sustainable ingredients compared to
animal milk will most likely lead to more sustainable cheese analogs.
There is still a substantial opportunity for researchers and developers to
understand how plant-based ingredients need to be extracted and pro­
cessed to obtain the desired structures and flavors needed to further
increase the market penetration of plant-based cheeses.
Declaration of competing interest
The authors have no conflicts of interest. Brand names appearing in
this review article are for product identification purposes only and all
information presented is available to the general public. No endorse­
ment or criticism was intended, nor was endorsement or criticism
implied of products not mentioned.
Acknowledgments
The authors thank the Good Food Institute for partial support of this
work.
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