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Citation: Genet, B.M.L.; Sedó Molina,
G.E.; Wätjen, A.P.; Barone, G.;
Albersten, K.; Ahrné, L.M.; Hansen,
E.B.; Bang-Berthelsen, C.H. Hybrid
Cheeses—Supplementation of
Cheese with Plant-Based Ingredients
for a Tasty, Nutritious and
Sustainable Food Transition.
Fermentation 2023,9, 667.
https://doi.org/10.3390/
fermentation9070667
Academic Editor: Thomas Bintsis
Received: 30 June 2023
Revised: 13 July 2023
Accepted: 14 July 2023
Published: 15 July 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
fermentation
Review
Hybrid Cheeses—Supplementation of Cheese with Plant-Based
Ingredients for a Tasty, Nutritious and Sustainable
Food Transition
Blandine M. L. Genet 1, * , Guillermo Eduardo SedóMolina 1, Anders Peter Wätjen 1, Giovanni Barone 2,
Kristian Albersten 3, Lilia M. Ahrné2, † , Egon Bech Hansen 4, † and Claus H. Bang-Berthelsen 1, *,†
1Research Group for Microbial Biotechnology and Biorefining, National Food Institute, Technical University
Denmark, Kemitorvet, Building 202, 2800 Kongens Lyngby, Denmark
2Department of Food Science, University of Copenhagen, Rolighedsvej 30, 1958 Frederiksberg, Denmark;
lilia@food.ku.dk (L.M.A.)
3Thise Dairy, Sundsørevej 62, Thise, 7870 Roslev, Denmark
4Research Group for Gut, Microbes and Health, National Food Institute, Technical University Denmark,
Kemitorvet, Building 202, 2800 Kongens Lyngby, Denmark
*Correspondence: blagen@food.dtu.dk (B.M.L.G.); claban@food.dtu.dk (C.H.B.-B.)
†Last author shared.
Abstract:
With increasing awareness of the impact of food on the climate, consumers are gradually
changing their dietary habits towards a more plant-based diet. While acceptable products have
been developed in meat analogues and non-fermented dairy products, alternative fermented dairy
products such as yogurt and particularly ripened hard and semi-soft cheese products are not yet
satisfactory. Since the cheese category has such a broad range of flavors and applications, it has
proven complicated to find plant-based sources able to mimic them in terms of texture, meltability,
ripening and flavor. Moreover, plant-based dairy alternatives do not provide the same nutritional
supply. New technological approaches are needed to make cheese production more sustainable,
which should be integrated in the already existing conventional cheese production to ensure a fast
and cost-efficient transition. This can be tackled by incorporating plant-based components into the
milk matrix, creating so-called “hybrid cheeses”. This review will discuss the challenges of both
animal- and plant-based cheese products and highlight how the combination of both matrices can
associate the best properties of these two worlds in a hybrid product, reviewing current knowledge
and development on the matter. Emphasis will be drawn to the selection and pre-processing of raw
materials. Furthermore, the key challenges of removing the off-flavors and creating a desirable cheese
flavor through fermentation will be discussed.
Keywords:
cheese; plant based; sustainability; hybrid cheese; fermentation; lactic acid bacteria;
alternative dairy
1. Introduction
The production of food from animal sources has a significant impact on the climate
and on the destruction of ecosystems. Dairy, and particularly cheese, is not an exception.
Indeed, when ranking food categories for their greenhouse gas (GHG) emissions, cheese
comes fifth, after beef herds, lamb and mutton, beef from dairy herds and crustaceans, and
before pig, fish, poultry, eggs and milk [
1
]. Even though the dairy sector has become more
efficient in the past 50 years [
2
], there is a limit to how sustainable animal farming can
become; therefore, drastic changes towards more plants for food are needed in the food
system to maintain global warming below 2
◦
C [
3
]. Moreover, the Eat-Lancet commission
highlighted in their report the urgency to reduce the consumption of animal-based food and
to increase the proportion of plant-based items in the diet, both for health and sustainability
reasons [
4
]. Consumers are increasingly concerned about sustainability and the actions they
Fermentation 2023,9, 667. https://doi.org/10.3390/fermentation9070667 https://www.mdpi.com/journal/fermentation
Fermentation 2023,9, 667 2 of 17
can undertake to reduce their impact. Changing diet towards a more plant-based one is an
obvious choice that many are undertaking. For example, in Denmark, 3% of the population
declared themselves vegetarians in 2022, in comparison to 1.8% in 2017. In the age range
of 18 to 34 years old, the numbers rise to 7.4% [
5
]. This can be seen in the development
of new plant-based analogues, such as meat substitutes [
6
,
7
], milk analogues [
8
], and
yogurt [
9
]. Other plant-based products are in the early stages of development, such as
alternatives to eggs, ice cream or fish [
10
]. The main drivers for consumers to change to a
more plant-based diet are mainly health concerns for flexitarians and ethical reasons for
vegans and vegetarians [
11
]. However, changing habits is difficult and dietary habits are
not an exception. For new and more sustainable products to be adopted as a regular part
of the diet, the product must be pleasant to consume; otherwise, the curiosity purchase
will not transform into a regular purchase. Plant-based alternatives (PBA) are mainly
developed with the aim of replicating the properties of existing animal products. Due to
the different functionalities of the used materials, the perfect match is difficult to reach,
which generally results in low acceptance [
10
]. Plant-based cheese alternatives (PBCA)
are massively arriving on the market, with solutions for many different types of cheeses
such as cream cheese, brine cheese (mozzarella, feta), blue cheese or longer-ripened cheeses
such as edam or gouda. They are mostly made with coconut oil, starches, nuts or soy [
12
].
However, the products currently available have not yet reached satisfying quality in several
aspects such as sensory, nutrition and sustainability, depending on the base used, which
will be reviewed in the next paragraphs.
First, the sensory properties do not live up to the expectations of consumers. A major
sensory issue with soy-based cheese alternatives (CA) is the beany flavor and grittiness of
the mouthfeel, which could be alleviated via fermentation, processing or blending different
plant matrices together [13]. Falkeisen et al. [14] measured sensory aspects and emotional
response to five different plant-based (PB) shredded-cheddar-type products. They found
that even though some products performed better than others, they were overall not liked,
and even participants regularly consuming PBCA assigned it low liking scores. Also, they
reported the conflict of participants wanting to consume PBCA but not doing so because
the taste was not satisfactory [
14
]. Giacalone et al. provided a detailed review of the
different sensory and consumer behavior aspects leading to the different acceptance levels
of consumers towards PBA [
15
]. One of the highlights is that mimicking the exact properties
of conventional products is counterproductive with increasing familiarity to plant-based
alternatives: the more familiar consumers are with alternatives, the less they want them
to resemble real animal products [
16
]. This is an opportunity to create products with new
tastes and to embrace the plant-based aspect. However, overall liking of the product is the
crucial criterion for omnivores and flexitarians to consume PBA. In contrast to vegetarians
and vegans, they will not compromise on taste and might have less information about the
environmental impact of different food products [
17
]. The greatest impact can be reached
by convincing most consumers to switch to a more plant-based diet, and therefore the focus
when developing a new sustainable product should be the sensory aspects [16].
Secondly, changing the matrix from animal milk to plant-based dairy alternatives
(PBDA) does not provide the same nutritional properties. This can lead to severe deficien-
cies of which consumers are not always aware. Particularly concerned are young children,
elderly people and women in the reproducing stage of their life [
18
]. Dairy products, and
especially cheese, are important sources of macronutrients (protein and fats) and micronu-
trients (calcium, vitamin B2, vitamin B12, vitamin D and iodine) [
19
,
20
]. Few studies about
the nutritional value of plant-based cheeses and comparison with animal counterparts have
been conducted so far [
18
,
21
–
23
], while efforts were mostly concentrated on alternative
milk products. These point out that the main nutritional challenges for PBCA are the lack
of protein, calcium, vitamins (B2, B12 and D) and iodine and high amount of saturated
fatty acids due to the use of coconut fat as a main ingredient [
18
,
21
–
23
]. Supplementation
in micronutrients has proven to be an efficient way to achieve nutritional adequacy [
18
].
From the perspective of hybrid products, calcium can be fortified using the right propor-
Fermentation 2023,9, 667 3 of 17
tion of soluble and insoluble calcium salts to avoid effects on protein structure and salt
precipitation [
24
]. Iodine could be fortified in the salting and brining steps, as has been
shown by Wechsler et al. [25] in conventional cheese. On the macronutrient level, produc-
ing a nutritionally adequate product should be the primary concern when formulating
PBCA, prioritizing protein-rich ingredients and not introducing high amounts of saturated
fatty acids.
While the literature about the environmental impact of conventional dairy products is
increasing in quantity, quality and accuracy, there are still very few studies on the impact
of PBCA as such. A major barrier to creating these kind of data is the diversity of products,
ingredients and processes involved, while traditional cheesemaking only requires milk
from cows and, in a minority of cases, from other animals. In the review of 16 life cycle
assessment (LCA) studies, Finnegan et al. [
26
] found that the overall environmental impact
of cheese production is driven by the production of raw milk, accounting for 79 to 95% of
the total global warming potential (GWP) of cheese, while the processing of raw milk into
cheese only accounts for 2 to 18% of the GWP. Transportation and packaging appeared to
be negligeable. Therefore, rethinking the source of raw materials is crucial to reduce the
overall impact of cheese. Few studies have been conducted in the comparison between con-
ventional dairy and PBDA, which was reviewed by Carlsson Kanyama et al. [
27
]. Overall,
they found that PBDA had a much lower environmental impact than their conventional
dairy counterparts. However, these types of studies are still in their infancy and many
parameters are limiting, calling for the harmonization of the methods used, units used
and information collected. Even if they seem clearly more sustainable, PBCA should also
be questioned, depending on the choice of raw materials as a base and on their origin.
Indeed, nuts have a high environmental and social impact, and even though the Eat-Lancet
commission [
4
] recommends an increase in the consumption of nuts and seeds, they are
precious ingredients and care should be taken to not lose their nutritional aspect during
processing [
28
]. In their thorough assessment of the impact of nuts, Cap et al. [
28
] identified
cashews and almonds as the worst-performing nuts. They are, however, very prominent as
ingredients in PBCA. Moreover, the mostly manual post-harvest processing of cashew nuts
has a disastrous impact on the health of the workers, who are mostly uneducated women
whose voices are rarely heard in mainstream attention [
29
]. Therefore, to achieve a new
cheese alternatives which can truly be considered sustainable, the ingredients for PBDA
should be carefully selected in the perspectives of low environmental and social impact
as well as local production. The use of industrial side streams is a promising source of
material, allowing for both locality and circularity, which should be considered for hybrid
cheese formulations, as has been investigated in yogurt alternatives with brewer’s spent
grain [30].
Animal cheese is an important source of nutrients and has a long history of traditional
use, making it a category with wide diversity which is difficult to mimic and replace.
However, the current consumption of cheese is not sustainable, and alternatives should
be found. Even though numerous new PBCAs are brought to the market, their nutritional
profile and sensory properties are not satisfactory. We hypothesize that creating a hybrid
product combining the nutrition and functional properties of milk and milk proteins with
the sustainability of plant-based components is a solution to lower the environmental
impact of cheese consumption, while providing a high-quality product with both high
nutritional profile and great flavor. In this literature review, we will investigate the current
knowledge about mixing milk and plant-based proteins to create a gel network, summarize
prior achievements in creating cheese analogue products by mixing both animal and plant-
based milks as well as how can the raw ingredients be pre-processed to make them more
functional. Finally, we will review how fermentation, which is a process commonly applied
to cheese, is impacted by the change in matrix, and which opportunities it creates to
improve the sensory and nutritional profile of products containing plant-based ingredients.
Fermentation 2023,9, 667 4 of 17
2. Recent Developments in Hybrid Cheeses
We define hybrid cheese as a cheese product made from milk and plant-based ingredi-
ents, where both components are retained in the final product to various concentrations.
To the best of our knowledge, this terminology is not yet widely used, and only one pub-
lication was found using it [
31
]. Other terminologies are “mixed dairy and plant-based
alternatives” or “dairy supplemented with plant-based ingredients”.
Guyomarc’h et al. [
32
] reviewed the evolution of animal/plant mixed products, in
the context of mixing milk, eggs and plant bases. They also summarize nutrition and
digestibility aspects of such mixed foods. Mixing fast-digestible animal protein with slow-
digestible plant protein is a good approach to fight against malnutrition and build up
muscle mass [
32
]. Except for butter spreads, where parts of the fat are replaced with plant
oils, hybrid products do not seem to have arrived on the European consumer markets yet.
This might be mostly due to a lacking consumer base. However, to launch such products,
it is important to evaluate consumer perception. Drigon et al. [
33
] observed the attitude
towards mixed dairy products (milk, yogurt, solid dairy/cheese-like) in an online survey
on French consumers. They found three types of consumers: the first are the ones who
already consume dairy alternatives and have a negative opinion on milk (for health and
environmental reasons), and have a relatively low opinion of such products because they
already found their plant-based alternatives. The second type are consumers strongly
attached to milk, who do not think that dairy is bad for the environment and even if, would
not consider environment as a criterion to choose food. They have a bad perception of
soy and are hard to convince if the taste is not spot on. The last category is made up
of mostly women, self-declared omnivores, making altruistic and self-centered choices.
They would change to mixes because they have fewer calories and a lower environmental
impact, but the emotional properties would still be conserved because the product would
be close to traditional use and familiarity [
33
]. There are, therefore, good incentives towards
developing such hybrid products.
Milk-based cheese has two main components: protein and fat, with minimal amounts
of carbohydrates and essential micronutrients. Therefore, plant-based replacements can
be found for either the fat or the protein component, partially or fully, and in different
proportions. A key challenge in hybrid cheese production is forming a protein network
to obtain the desired structure, which will be discussed in the next section. Analogue
cheese products appeared in the 1980s, and were developed to create cheap replacements
for the ready-meal industry, such as pizza cheese toppings. Cost reduction was achieved
by removing the milk fat, to be sold as butter, and replacing it with vegetable oils and
fats. Such products permitted more flexibility in terms of functionality, were more con-
sistent throughout seasons and had longer shelf-life, which made them easier to handle
logistically [
34
]. Nowadays, consumers are more concerned about health aspects, and the
replacement of dairy fat with plant fat can be an advantage. Furthermore, there is a general
trend towards creating high-quality food products made for hedonic purposes to reduce
environmental and health impacts.
Some research has been conducted in the direction of hybrid cheese, but it seems
quite sporadic, and the methods used to assess the success of the formulation differ from
study to study. Key factors for successful formulations are texture and sensory profiles.
While the former can be conducted on machines with standardized protocols allowing for
reproducibility, sensory analysis is not always conducted, and when applied, the robustness
of the investigation is sometimes questionable. To significantly improve the quality of newly
developed products, sensory analysis should receive more attention and be conducted in a
more standardized way [
13
]. Table 1summarizes the studies found to formulate hybrid
cheeses by replacing either parts of the casein or fat with plant-based ingredients. It is
notable that diverse types of cheese and plant-based replacements are being investigated.
Fermentation is traditionally applied in cheesemaking, as lactic acid bacteria (LAB)
are essential for the acidification of the milk, enabling curd formation and increasing shelf
life. Moreover, secondary cultures are also important for the development of specific
Fermentation 2023,9, 667 5 of 17
cheese aromas. The potential of fermentation for plant-based dairy alternatives will be
discussed further down. However, while reports exist about fermented PBA in yogurt
and beverages [
30
,
35
,
36
], to the best of the authors’ knowledge, no studies have been
conducted specifically investigating the effect of fermentation on hybrid cheese. In the
papers listed here, only half of them used a starter culture in cheesemaking for acidification
purposes. One of the studies viewed their hybrid product as a potential probiotic and
recorded the evolution of the number of LAB during a two-week storage period, proving
that the number did not decrease below 7.0 log CFU/g, thus confirming the probiotic
potential [
37
]. Since fermentation with bacteria and fungi has such an importance for the
traditional cheesemaking process and to improve the functionality of plant proteins [
38
],
research conducted in this field should include these aspects as well.
Overall, these studies show that there is a low threshold for the proportion of plant
bases that can replace animal milk. Indeed, most studies report a negative impact on the
taste or the texture when adding more than 15%, and only a few reach values as high as
20% plant-based ingredients. Above this quantity, negative effects can be found such as
collapsing of the texture, grittiness, or off-flavors. When developing products to reduce
the environmental impact of cheese, such low proportions of plant ingredients will not
be sufficient, and development efforts should aim for higher standards. Moreover, some
studies have aimed to replace protein with starches. While the effort to include more fiber
in the diet is an honorable goal, one should not forget that the main macronutrient provided
by cheese is protein, and that consumers will expect alternative cheese products to be a
source thereof. The formulation of hybrid cheese should therefore aim for protein contents
at least as high as those of conventional cheese. Protein is a useful tool in food formulation,
as it plays important roles as an emulsifier, foaming and gelling agent. However, milk
and plant proteins are different in their composition and structure, making them difficult
to interchange.
Table 1. Summary of studies investigating hybrid cheese from complex materials.
Cheese Type Plant-Base Fermentation Main Results Reference
Mozzarella
Hydroxypropylated
barley starches No
Replacement of 15% of rennet casein with
starches provides acceptable textures with
improved meltability
Mehfooz et al.,
2021 [39]
Mozzarella Soy milk
Thermophilic Y 082
D (Clerici Sacco
International Srl,
Cadorago Como,
Italy): Lactobacillus
bulgaricus,
Streptococcus
thermophilus
Using 10 to 20% soy milk is acceptable.
Higher proportion decreases meltability but
increases nutritional profile
Jeewanthi et al.,
2014 [40]
Feta Lab-made lentil
milk, inulin No
Adding too much lentil protein disrupts the
structure, but 10% is acceptable. Inulin
increases likeability as a fat replacer
Moradi et al.,
2021 [41]
Cream cheese Lab-made soy
protein concentrate No
From 5 to 15 g/L soy protein concentrate
added to partially skim milk; addition of this
amount of soy protein did not impact sensory
experience too strongly. The texture was still
acceptable, taste was slightly negatively
impacted. Products were stable
Rinaldoni et al.,
2014 [42]
Cream cheese
Pea protein, lupin
protein or oats
protein isolates
No
All emulsions created were stable; caseins
and whey proteins are primarily adsorbed at
the oil/water interface
Grasberger
et al., 2021 [43]
Cheddar Soy milk
Streptococcus lactis
(now Lactococcus
lactis)
Cow’s milk can be replaced with soy milk up
to 15% without impairing sensory qualities
(but sensory experience was made by
untrained lab personal)
Rani and Verna,
1995 [44]
Fermentation 2023,9, 667 6 of 17
Table 1. Cont.
Cheese Type Plant-Base Fermentation Main Results Reference
Cheddar Soy protein isolate
(SPI)
S. thermophilus,
Lactobacillus
delbrueckii ssp.
bulgaricus
Up to 7% soy protein, no adverse effect on
taste, microstructure was less compact with
soy protein. Advise a max 5% soy protein
Atia et al., 2004
[45]
Yogurt cheese Lab-made soy milk
Thermophilic
Y332A (Clerici
Sacco International
Srl, Cadorago
Como, Italy), L.
bulgaricus and S.
thermophilus
Higher protein and lower fat content in
cheeses supplemented with up to 20% soy
milk; no significant difference in the rheology
character between control and soy
supplemented
Lee et al., 2015
[37]
3. Achieving Texture through Raw Material Selection and Pre-Processing
Food materials, including ingredients, are complex as they can be heterogeneous,
amorphous and hygroscopic while having different structural properties (e.g., morphol-
ogy) [
46
]. The processing of food materials is often associated with modification at all
levels such as micro, meso and molecular [
47
]. Also, the concomitant phenomena of mass
transfer with thermal treatment during food processing can result in physicochemical
changes. Phase transitions from liquid to solid during food processing are often encoun-
tered in systems dominant in proteins or carbohydrates due to aggregation and gelation,
respectively [
48
]. Understanding protein physicochemical properties, modifications and
interactions during processing can represent a powerful tool for modifying and tailoring
physicochemical properties, appearance and texture to develop formulated new products
such as plant–dairy hybrid food. The gelation of milk proteins is well understood and exten-
sively described [
49
–
51
]. There are also extensive studies on the gelation properties of plant
proteins, such as the heat-induced gelation of plant globulin [
52
–
54
], acid induction [
55
] or
combinations of treatments [
56
,
57
]. As a flexitarian diet is becoming increasingly common
among consumers due to health, environmental and sustainability concerns [
58
], scientific
reports investigating plant–dairy systems for their physicochemical properties, as those
influenced by processing conditions are emerging [
59
,
60
], with some relevant reports being
summarized in Table 2.
Early studies were primarily focused on combining SPI with whey protein isolate, or
micellar casein isolate, at neutral pH (i.e., 6.7 to 7.0) using mixing (or shear mixing) and heat
treatment. Although these early studies used ideal conditions such as purified ingredients,
there was virtually no interference of other components (e.g., lipids or carbohydrates) and
standardized pH; it was fundamentally established that the addition of dairy proteins,
especially whey proteins, to soy protein was modulating the rheological properties of such
hybrid systems towards low viscosity, and that the heat load applied during processing
influenced the formation and properties of the aggregates (soluble or insoluble).
Most recently, pea proteins and their fractions (i.e., legumin and vicilin) in combination
with different sources of dairy proteins (whey, casein and a combination thereof) have been
receiving a lot of attention from researchers. This draw of attention towards pea is ascrib-
able to pea being considered safer than soy, in terms of allergenicity, regulations (GMO)
and sustainability, as pea requires less water than soy to grow [
61
,
62
]. Most of the work con-
ducted on pea protein and dairy have focused on either pea protein concentrate or isolate
along with relatively simple dairy sources such as whey proteins or micellar casein proteins
and skim milk [
54
,
55
,
63
–
68
]. Most of these studies processed dairy and plant proteins using
simple mixing with ultra-pure water and typical standardized heat treatment processing for
achieving gelation (80 to 95
◦
C, 20 to 60 min). In addition, some of these works also studied
the influence of pH (either from acid addition or fermentation) or enzymatic treatment
for providing insights relative to gelation characteristics and protein–protein interactions.
At a general level, the different extents of processing (e.g., heat treatment, acidification,
Fermentation 2023,9, 667 7 of 17
etc.), plant-to-dairy protein ratio and total protein concentration significantly influenced
the following, but not limited to, properties: (a) onset gelling temperature decreasing with
a high proportion of plant protein; (b) plant protein produces a gel network independent
of dairy protein; (c) rheological properties of hybrid gel (especially storage module) was
influenced by the relative proportion of plant proteins; (d) acidification of the hybrid system
was accelerated when plant proteins were dominant whilst decreasing gel stiffness and (e)
inclusion of dairy protein increased functionality, especially when fat components were
included in the hybrid system. However, there are limitations for the comparison between
studies, since the technical properties of plant protein vary strongly depending on the
extraction method [
52
], as commercial protein isolates are usually denaturated during the
extraction process, which lowers solubility [53,64].
Overall, one should point out that limited reports have investigated other plant protein
sources, apart from pea or soy proteins, in the presence of dairy proteins. Additionally,
most of the work was conducted in a protein-dominant system without including other
necessary components for a fermentable hybrid cheese perspective, such as lipids and
fermentable carbohydrates sourced from plant or dairy (e.g., lactose). Fat is one of the major
macrochemical components of conventional full-fat dairy cheese (e.g., cheddar, Parmigiano
Reggiano and gouda), which is crucial for taste, texture and flavor, with an average protein-
to-fat ratio ranging from 0.70 to 1.02 [
69
]. It is, therefore, necessary to incorporate fat
components into hybrid cheese matrices; however, only a few studies, to the authors’
knowledge, can be considered fundamental from a hybrid cheese perspective, as seen
above. From a plant–dairy hybrid perspective, the work of Grasberger et al. [
43
], carried
out on fat content similar to conventional dairy cheese (i.e., 20 to 21.4%) but at a lower
protein content (i.e., 8%) than regular medium-hard cheeses, is a good example. They used
different plant proteins such as pea, oat and lupin in conjunction with fresh milk and whey
protein in an extensive range of plant–dairy ratios (see Table 2). The formulations were
processed very similarly to what is commonly carried out at the industrial level, such as the
reconstitution of the ingredients and pre-homogenization, followed by homogenization at
250 bar, with a thermal treatment (85
◦
C for 5 min) to generate protein gelation. Interestingly,
dairy protein inclusion was crucial to stabilizing the oil droplet at the interface, while plant
protein modulated the gel’s final characteristics, with lupin and pea resulting in creamy
spreadable-like cheese in contrast to oat protein, which resulted in the gel having pasta-
filata-like properties. Similarly, a very interesting work by Canon et al. [
36
] focused on
hybrid set-type yogurts fermented using LAB strains (Lactiplantibacillus plantarum and
Enterococcus faecalis). The yogurts’ formulation was relatively higher in protein content
(6.6%) than their conventional dairy counterparts. The proteins were supplied from lupin
protein isolate and whey protein or skim milk powder as dairy ingredients along with
the inclusion of milk and plant fat components (milk fat solids and coconut at 1.5%). It
was shown that pre-treatment such as homogenization (two steps, total 300 bar) and pre-
heating (95
◦
C for 10 min) prior to fermentation of the protein suspension can improve the
texture with a higher content of lupin, further modulating toward a high apparent viscosity.
Pre-treatment of the protein ingredient solution prior to the mixing stage produces hybrid
mixtures with different textures and gel structures than those not being pre-heat treated, a
finding established also in other published reports [
70
–
72
]. However, a milk/lupin protein
ratio of 67:33 was more acceptable compared to the 50:50 ones, as coconut oil did not
negatively impact the overall sensory properties. Fermentation also improved acceptability
by producing more and different aroma compounds and a more pleasant texture due to
acidity levels close to the isoelectric point of dairy–plant proteins.
Although there is an incremental number of studies on understanding the overall
properties of plant–dairy hybrid systems in terms of physicochemical properties and struc-
ture whilst correlating these properties to processing and pre-processing, there is limited
evidence on mimicking or reproducing plant–dairy hybrid systems that can resemble
conventional dairy products. Most of these works were carried out in model conditions
(composed only of protein-based ingredients) without being emulsified (no fat included).
Fermentation 2023,9, 667 8 of 17
However, the advantage of combining the pre-processing of the plant and dairy ingredi-
ents alone or in combination thereof, followed by fermentation using a combination of
dairy and plant carbohydrate, appears promising for improving and making plant-dairy
hybrid systems close to dairy. Although more reports in this area are necessary, this may
represent a very significant advancement toward developing plant–dairy hybrid products
(e.g., cheeses and spreadable creams) for sustainable diets aiming for a gradual and full
transition towards plant-based diets.
Table 2.
The relevant literature on main effects and properties of different plant–dairy hybrid systems.
Ingredients and
Concentrations Plant to Dairy Ratio Processing
conditions Highlights Reference
SPI, WPI—6% (w/v)
protein content
0:100, 30:70, 50:50,
70:30, 100:0
Mixed; 90 ◦C 60 min;
pH 7.0
While soy protein in isolation formed
large, soluble aggregates, the addition
to whey protein significantly reduced
the amount of soluble aggregate and
induced precipitation. 7S and the
basic subunit of 11S were present in
the precipitate after heating with WPI.
Roesch and
Corredig (2005)
[73]
SPI, MCI—10 and
15% (w/w) protein
content
50:50
High shear mixing;
40, 60 and 95 ◦C
15 min; native pH
(6.74–6.86)
Temperature above the denaturation
of soy glycinin induced aggregation
and gelling via disulfide bonding in
both soy and casein; below soy protein
critical gelling concentration, mixtures
resulted in a Newtonian liquid with
lower viscosity and improved storage
stability compared to non-denaturing
heat treatment.
Cosmin and
Moraru (2013)
[74]
PPC, WPC—10, 16
and 22% (w/v)
protein content
80:20, 50:50, 20:80 Mixed, 92 ◦C 30 min;
pH 4.0, 6.0, 8.0
Higher synergistic enhancement at
20:80 pea/whey was observed for gels
at pH 6.0; 50:50 and 20:80 ratios both
had synergistic viscosity enhancement
after heat treatment.
Wong et al.
(2013) [63]
PPC (legumin and
vicilin fractions),
Purified casein
micelles—1.8%
(w/v) protein
content
50:50 Mixed; 85 ◦C 60 min;
pH 7.10
Casein micelles stabilized pea proteins
against heat-induced unfolding;
vicilin, in presence of casein micelles,
produced heat-induced soluble
aggregates, while legumin produced
insoluble aggregates.
Mession et al.
(2017) [54]
PPI, SMP—14.8%
(w/w) protein
content
50:50
Mixed, (a) heat
treated 90 ◦C 60 min;
(b) acidified with
glucono delta-lactone
2% (w/v) and (c)
Enzymatic treatment
CaSO4 (0.3%),
chymosin (0.5%), and
TGase (0.3%)
Pea gels induced by acid or enzyme
had a higher storage modulus (G0).
Thermal treatment induced covalent
bonds. Enzyme-induced gels
produced coarse aggregates with a
more excellent resistance to strain.
Ben-Harb et al.
(2018) [64]
MPC, WPI, PPI,
SPI—12% (w/w)
Protein content
50:50, 58:41, 66:33,
75:25, 83:16, 90:10,
100:0
Stir mixed; 90 ◦C 1 h;
pH 7.00
Gelation temperature increased with
plant protein, with soy being more
effective than pea protein.
Silva et al.
(2018) [65]
PPI (globulin
fraction); WPI
(β-lactoglobulin
fraction)—2% (w/v)
Protein content
0:100, 30:70,
50:50,70:30, 100:0
Stir mixed; 85 ◦C
60 min; pH 7.20
Synergistic effect for increased gels
elasticity and water holding capacity
compared to gels containing pure
aggregates of Glob or mixtures of
Glob and βlg aggregates.
Mohamed–
Lazhar Chihia
et al. (2018) [66]
Fermentation 2023,9, 667 9 of 17
Table 2. Cont.
Ingredients and
Concentrations Plant to Dairy Ratio Processing
conditions Highlights Reference
RPI, WPI—0.0 to
20% (w/w) Protein
content;
90:10, 80:20, 70:30,
60:40, 50:50, 40:60,
30:70, 20:80, 10:90
Mixed, 80 to 95 ◦C
30 min; pH 7.00
Independent gel network formation:
higher plant to dairy ratio produced a
stronger gel than single standalone
ingredient.
Ainis et al.
(2019) [75]
OPC, OPI,
SMP—12.3, 13.8
and 15.3% (w/w)
total solids
60:40
Homogenized
one-pass 200 bar;
80 ◦C 20 min; native
pH
OPC good replacer of SMP; good
functionality due to oat starch.
Brückner-
Gühmann et al.
(2019) [76]
PPI, OPC, LPI, WPI,
SM—8.0% (w/v)
Protein content; 20
to 21.4% (w/v) fat
content
33:66, 32:67, 26:73,
67:32, 67:32, 68:31,
70:30,
Mixed, homogenized
one-step 250 bar;
85 ◦C 5 min; pH 6.1
to 6.6
Emulsion gel oil droplets were
stabilized by dairy proteins; LPI and
PPI induced low onset gelation
temperature.
Grasberger et al.
(2021) [43]
LPI, SMP, WPI,
milk fat and
coconut oil—6.6%
(w/w) Protein
content; 1.5% (w/w)
fat content
50:50, 67:33
Stirred and
homogenized
two-step 250/50 bar;
heated 95 ◦C 10 min;
pH native; LAB
fermented
Fermentation improved the texture
and reduced off-flavor of
lupin-dominant formulation.
Canon etl al.
(2022) [36]
SPI, WPI—4% (w/v)
protein content
100:0, 75:25, 50:50,
25:75, 0:100
Mixed; 95 ◦C 30 min;
pH 7.00 then acidified
SPI inclusion decreased stiffness (low
G0) and stretchability (lower γc) of
acid-induced gels; hybrid gels
displayed a relatively more elastic
response in the nonlinear viscoelastic
regime with a plastic behavior.
Xia et al. (2022)
[67]
PPI; SMP—5.0, 7.0,
9.0, and 11% (w/w)
Protein content
27:73, 33:66, 42:57,
60:40
Stir mixed; 85 ◦C
5 min; pH 6.30 to 6.80
The presence of pea proteins
accelerates acid-induced gelation but
weakens the structure of mixed gels.
Oliveira et al.
(2022) [68]
Dairy ingredients:
Whey protein concentrate (WPC); whey protein isolate (WPI); fresh skim milk (SM); skim
milk powder (SMP); milk protein concentrate (MPC) and micellar casein isolate (MCI).
Plant ingredients:
Soy
protein isolate (SPI); pea protein concentrate (PPC); pea protein isolate (PPI); oat protein concentrate (OPC); oat
protein isolate (OPI); lupin protein isolate (LPI) and rapeseed protein isolate (RPI). 4. Flavors, off-flavors and
antinutritional factors in hybrid cheese.
For the development of hybrid cheeses, the combination of plant-based ingredients
with milk may lead to the generation of nutritional, technological, and sensorial challenges
when aiming to mimic their dairy-based counterparts [
13
,
77
]. Implementing new raw
materials will inevitably demand the application of new methods to reach a convincing
flavor profile. In a fully milk-based cheese production, many parameters already affect
the flavor profile of the resulting cheese. Animal origin [
78
], diet [
79
], physical treatment
(i.e., pasteurization, cooking temperature and salting) [
80
] and ripening conditions (i.e.,
microbiota, storage temperature and maturation time) [
81
] have all been thoroughly studied
since they have a significant effect on the final product. The introduction of plant-based
ingredients will equally be influenced by these steps, and one will need to carefully study
how to maintain favored properties, while avoiding unwanted inputs like off-flavors and
antinutritional factors (Figure 1).
Fermentation 2023,9, 667 10 of 17
Fermentation 2023, 9, 667 9 of 17
21.4% (w/v) fat con-
tent
LPI, SMP, WPI, milk
fat and coconut oil—
6.6% (w/w) Protein
content; 1.5% (w/w)
fat content
50:50, 67:33
Stirred and homogenized
two-step 250/50 bar; heated
95 °C 10 min; pH native;
LAB fermented
Fermentation improved the texture and reduced
off-flavor of lupin-dominant formulation.
Canon etl al.
(2022) [36]
SPI, WPI—4% (w/v)
protein content
100:0, 75:25,
50:50, 25:75, 0:100
Mixed; 95 °C 30 min; pH
7.00 then acidified
SPI inclusion decreased stiffness (low G′) and
stretchability (lower γc) of acid-induced gels; hy-
brid gels displayed a relatively more elastic re-
sponse in the nonlinear viscoelastic regime with a
plastic behavior.
Xia et al.
(2022) [67]
PPI; SMP—5.0, 7.0,
9.0, and 11% (w/w)
Protein content
27:73, 33:66,
42:57, 60:40
Stir mixed; 85 °C
,
5 min; pH
6.30 to 6.80
The presence of pea proteins accelerates acid-in-
duced gelation but weakens the structure of
mixed gels .
Oliveira et
al. (2022)
[68]
Dairy ingredients: Whey protein concentrate (WPC); whey protein isolate (WPI); fresh skim milk
(SM); skim milk powder (SMP); milk protein concentrate (MPC) and micellar casein isolate (MCI).
Plant ingredients: Soy protein isolate (SPI); pea protein concentrate (PPC); pea protein isolate (PPI);
oat protein concentrate (OPC); oat protein isolate (OPI); lupin protein isolate (LPI) and rapeseed
protein isolate (RPI). 4. Flavors, off-flavors and antinutritional factors in hybrid cheese.
For the development of hybrid cheeses, the combination of plant-based ingredients
with milk may lead to the generation of nutritional, technological, and sensorial chal-
lenges when aiming to mimic their dairy-based counterparts [13,77]. Implementing new
raw materials will inevitably demand the application of new methods to reach a convinc-
ing flavor profile. In a fully milk-based cheese production, many parameters already affect
the flavor profile of the resulting cheese. Animal origin [78], diet [79], physical treatment
(i.e., pasteurization, cooking temperature and salting) [80] and ripening conditions (i.e.,
microbiota, storage temperature and maturation time) [81] have all been thoroughly stud-
ied since they have a significant effect on the final product. The introduction of plant-
based ingredients will equally be influenced by these steps, and one will need to carefully
study how to maintain favored properties, while avoiding unwanted inputs like off-fla-
vors and antinutritional factors (Figure 1).
Figure 1.
Metabolic pathways of the production and utilization of desired or undesired compounds
by LAB, propionic acid bacteria, yeast and molds in the context of plant and milk fermentation.
Plant-based raw materials are characterized by containing antinutritional factors such
as phytates, enzymatic inhibitors and saponins that affect the absorption and bioavailability
of valuable compounds [
9
,
77
,
82
]. Phytates reduce the mineral bioavailability of metal
ions by forming complexes, while enzymatic inhibitors, such as trypsin inhibitors present
in legumes, decrease plant protein digestibility [
82
,
83
]. Furthermore, saponins affect the
absorption of specific vitamins due to their cross-interaction with fat-soluble vitamins that
are chemically similar to plant sterols [
82
]. Different antinutrients are found depending on
the plant-based raw material used. For instance, cereals and legumes contain flatulence-
causing oligosaccharides such as raffinose and stachyose.
Off-flavor molecules are classified into volatiles, such as aldehydes, alcohols and
ketones, and non-volatiles such as phenolic compounds, peptides and saponins [
84
]. The
combination of ingredients from animal and plant origin in hybrid cheeses can result
in off-flavor notes from both compounds, influencing the products’ sensorial properties
and consumer acceptance [
84
,
85
]. First, milk itself can provide off-flavor compounds,
which could originate from physical (milk pasteurization and light exposure)-, enzymatic
(oxidation and lipolization)-, non-enzymatic (Maillard reaction)- and microbial (protein,
amino acid, carbohydrate and lipid metabolisms)-driven reactions, which could impart
“sulfurous”, “bitter” and “rancid” notes [
86
]. However, plant-based dairy alternatives
are known for their typical off-flavors, often characterized as “green” or ”beany”, and
are therefore a significant source of undesired compounds [
77
,
84
,
87
]. The desirability of
cheese flavors strongly depends on the type of cheese and can even be considered a defect
when in the wrong cheese. Therefore, specific starter cultures and processing conditions
are optimized for each type [
88
]. For instance, “sulfurous” notes produced from sulfur
amino acids such as methionine are desired in camembert and cheddar cheeses [
88
], while
undesired in parmesan cheese. Nevertheless, the bitterness caused by peptides containing
hydrophobic amino acids is usually considered “unwanted” in most types of cheeses [
88
].
Volatile off-flavors that impart “green” notes are mainly caused by the degrada-
tion of polyunsaturated fatty acids through oxidation reactions, enzymatically and non-
enzymatically driven [
84
]. Enzymatically, they are mainly started by the action of lipooxy-
genases (LOX) that degrade plant fatty acids such as linoleic acids and transform them
into hydroperoxide intermediates, producing fatty acid aldehydes such as hexanal, pen-
Fermentation 2023,9, 667 11 of 17
tanal and nonanal [
84
,
85
,
89
]. Those are converted into their respective alcohol and ketone
compounds via enzymatic and non-enzymatic reactions, respectively [
84
]. Non-volatile
off-flavors are characterized by giving off astringent and bitter taste notes [
9
]. Phenolic
compounds such as ferulic, coumaric and gallic acids, saponins, alkaloids, peptides and
amino acids are some of the examples [
9
]. Also, it has been demonstrated that specific
combinations of volatile off-flavor molecules could enhance (or reduce) their perception by
synergistic (or antagonistic) effects with other volatiles [90].
Different approaches have been investigated to reduce the concentration of off-flavors
and antinutrients when developing plant-based fermented dairy alternatives [
87
]. Physical-
based technologies such as roasting, dehulling, soaking and blanching and milling have
been implemented in sesame, legumes, soy and peas and soy, respectively [
87
]. For instance,
roasting, milling and soaking have been applied to reduce LOX activity, reducing lipid
oxidation and off-flavor formation [
91
–
93
], while the dehulling of legumes has been used
to decrease phytate concentration [
91
]. Chemical-based technologies such as pH alterations
(alkalinization), chemical deodorization in pulses (pea, lentil and soy) and pseudocereals
(quinoa) have been applied for texture purposes as well as for reducing LOX activity and
off-flavor generation [
87
]. The pasteurization process of milk, preceding the initiation of
the regular cheesemaking process, also alters the flavor profile, depending on the degree
and type of pasteurization [94,95].
Finally, fermentation-based approaches have been investigated due to the fermentation
step necessary to produce dairy and hybrid cheeses [
87
]. In the following, we will elaborate
on the contribution from fermentation-based solutions in terms of flavor alterations, as it is
the most significant contributor in cheesemaking.
In traditional cheesemaking, large parts of the flavor formation happen during the
ripening stage, where the secondary metabolism of the present bacteria, yeasts and molds
create the main part of flavor molecules associated with cheese [
96
]. Understanding the
individual role, as well as the interplay of these organisms, is essential if fermentation
is applied in the production of a hybrid product. Since these products will have a new
substrate baseline, it will be important to tailor the use of specific strains to secure satisfying
alternatives. Specific LAB strains are extensively used in the production of dairy cheeses
as starter cultures, and are responsible for generating texture, flavor and other desired
properties in different types of cheeses [
97
]. In hybrid cheese production, the plant-based
substrate added will affect the metabolism and growth performance of the starter cultures
added, as well as the organoleptic properties of the final product. Therefore, dairy-adapted
microorganisms might not be optimal when aiming to alter the flavor profile and remove
antinutrients present in hybrid cheeses [
98
]. LAB strains with specific gene clusters and
enzymatic activities should be selected and added for that purpose [
98
–
100
]. As an example,
the high activity of LAB alcohol and aldehyde dehydrogenases has been correlated with the
removal of aldehyde molecules such as hexanal in plant-based dairy alternatives [
89
]. Also,
high activities of
β
-glucosidase,
α
-rhamnosidase and
β
-galactosidase have been related to
the degradation of the sugar moieties attached to saponins and
glycoalkaloids [101–103]
.
Depending on the plant-based substrate, different sugars are combined to their respec-
tive saponins [
104
]. For example, in soyasaponins, avenacosides (A and B),
α
-solanine
and
α
-chaconine, are bonded with dimers and trimers of glucose, galactose, arabinose
and/or rhamnose through different O-glycosidic bonds to each other and to their respective
saponins [104,105].
In a dairy cheese context, the flavor contribution from fermentation can be split into
three categories of origin: protein-, lipid-, and carbohydrate-derived flavor compounds [
96
].
The carbohydrate metabolism is almost exclusively linked to primary metabolism, generat-
ing compounds such as lactate, acetate, ethanol, CO
2
and the dairy-related diacetyl, acetoin
and butanediol [
90
], usually linked to fermented dairy products. From lipid catabolism
(lypolysis), the initial breakdown of triglycerides provides free fatty acids (FFA), the precur-
sors for flavor compounds such as lactones, esters, alcohols and methylketones [
106
,
107
].
Many of these impart fruity and herbal flavors and are rarely linked to LAB fermentation,
Fermentation 2023,9, 667 12 of 17
although there are studies showing the significant release of FFA from LAB [
108
]. Lipolysis
is instead often linked to Propionibacterium [
109
] and mold-ripened cheeses [
107
]. Flavors
related to the breakdown of proteins (proteolysis) are produced by both the starter culture
strains as well as commensal strains of bacteria, yeasts and molds [
110
–
112
]. However,
proteolysis is initiated by the addition of rennet and the activity of cell-envelope proteinases
and peptidases from the starter LAB [
112
]. The resulting release of small peptides and free
amino acids imparts a savory and salty flavor but is most importantly fed into the amino
acid catabolism, which produces sulfurous aromas and keto-acids [
111
]. The proteolytic
nature of many LAB is possibly linked to them being auxothropic in their amino acid
anabolism, hence needing to collect certain amino acids from their surroundings [
113
].
However, as most dairy-adapted LAB strains have been selected for their ability to break
down the caseins in milk, little focus has been drawn on finding strains suitable for the
proteolysis of plant proteins. Studies have been conducted on the extracellular microbial
proteases, specifically targeting plant proteins [
38
]. Even though there are some LAB
strains showing proteolytic effects on certain plant proteins, molds and yeasts are far more
capable [
38
]. Molds are essential for the ripening of certain cheeses such as camembert and
Roquefort, and many mold-ripened PBCAs have appeared on the market. However, the
safety of such molds should be carefully assessed when changing the growth substrate
from milk to plant bases, since it could induce the production of mycotoxins. This topic is
rarely addressed and should receive more attention as the development of fungi-fermented
foods is gaining momentum [114].
To conclude the above, metabolic capabilities of the starter cultures added to the fer-
mentation of hybrid cheeses, including LAB, yeasts and non-LAB strains such as Propioni-
bacterium freudenreichii, need to be investigated regarding the production of cheese-related
flavors, and the elimination of antinutrients and off-flavors in a hybrid cheese products.
Metabolic pathways and their actual application in this field are currently being studied for
each of the off-flavor and antinutrient molecules for the overall organoleptic and functional
improvement in hybrid cheese production.
4. Concluding Remarks and Perspectives
The food system plays a prominent role in the challenges that society is facing in
terms of climate change and its impacts on food security. These are clear warning signs
that the food industry has an urgent need to adapt to more sustainable food production
practices. The increase in market demand for PBA is an incentive towards increased
product development. While plant-based dairy alternatives have not yet been developed
to a satisfactory point in terms of taste and nutrition, this review shows the great potential
for developing hybrid cheese products. Key challenges in this domain are the interaction
of plant and milk protein as well as sensory properties. Since cheese is a key source of
protein in the diet, a hybrid product should have comparable or superior nutritional,
textural and sensorial profile. Therefore, research should aim to find processes to make
plant proteins more functional, so they can provide unique functionalities in the hybrid
cheese matrix. Furthermore, focus should be drawn to fermenting such products, as
fermentation represents a great opportunity to reduce the off-flavor characteristics for
plant-based ingredients, while adding desired cheese aromas and nutritional compounds
such as vitamins. More systematic investigations are needed, both in terms of protein
interactions to provide target textures and sensory analysis to satisfy consumer preferences
and succeed in a fast transition.
Author Contributions:
B.M.L.G. and C.H.B.-B. conceptualized the review framework. All authors
contributed to the design of the review. B.M.L.G. made the first draft of the manuscript with large
contributions from A.P.W., G.B. and G.E.S.M. Figures and tables were made by A.P.W., G.B., G.E.S.M.
and B.M.L.G. The draft was reviewed by L.M.A., K.A., E.B.H. and C.H.B.-B. All authors have read
and agreed to the published version of the manuscript.
Fermentation 2023,9, 667 13 of 17
Funding:
This review was supported by a Danish Dairy Research Foundation grant (SusCheese).
B.M.L.G. and G.B., G.E.S.M., E.B.H., L.M.A. and C.H.B.-B. were supported by an Innovation Fund
Denmark grant—Innomission AgriFoodTure called REPLANTED. A.P.W. was funded by a joint PhD
alliance grant between Technical University of Denmark (DTU) and University of Queensland (UQ).
The funding source was not involved in the design of the review, writing of the review or the decision
to submit the review for publication.
Conflicts of Interest:
The authors from DTU and UCPH affirm that the review was conducted
without any commercial or financial relationships that could be interpreted as possible conflicts of
interest. K.A. is an employee of the Danish dairy Thise.
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