Critical Reviews in Food Science and Nutrition

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Inflammatory, oxidative and DNA damage status

in vegetarians: is the future of human diet green?

Goran Gajski, Marko Gerić, Ivone Jakaša, Ines Peremin, Ana-Marija Domijan,
Marijana Vučić Lovrenčić, Sanja Kežić, Martina Bituh & Vanessa Moraes de
Andrade

To cite this article: Goran Gajski, Marko Gerić, Ivone Jakaša, Ines Peremin, Ana-Marija Domijan,
Marijana Vučić Lovrenčić, Sanja Kežić, Martina Bituh & Vanessa Moraes de Andrade (2021):
Inflammatory, oxidative and DNA damage status in vegetarians: is the future of human diet green?,
Critical Reviews in Food Science and Nutrition, DOI: 10.1080/10408398.2021.1986464

To link to this article: https://doi.org/10.1080/10408398.2021.1986464

Published online: 11 Oct 2021.

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Critical Reviews in Food Science and Nutrition
https://doi.org/10.1080/10408398.2021.1986464

Review

Inflammatory, oxidative and DNA damage status in vegetarians: is the

future of human diet green?
Goran Gajskia , Marko Gerića , Ivone Jakašab , Ines Pereminb , Ana-Marija Domijanc , Marijana
Vučić Lovrenčićd , Sanja Kežiće , Martina Bituhf and Vanessa Moraes de Andradeg
a
Mutagenesis Unit, Institute for Medical Research and Occupational Health, Zagreb, Croatia; bFaculty of Food Technology and Biotechnology,
Department of Chemistry and Biochemistry, Laboratory for Analytical Chemistry, University of Zagreb, Zagreb, Croatia; cFaculty of Pharmacy
and Biochemistry, Department of Pharmaceutical Botany, University of Zagreb, Zagreb, Croatia; dDepartment of Laboratory Medicine, Merkur
University Hospital, Zagreb, Croatia; eCoronel Institute of Occupational Health, Amsterdam UMC, University of Amsterdam, Amsterdam, The
Netherlands; fFaculty of Food Technology and Biotechnology, Department of Food Quality Control, Laboratory for Food Chemistry and
Biochemistry, University of Zagreb, Zagreb, Croatia; gHealth Sciences Unit, Translational Biomedicine Laboratory, University of Southern
Santa Catarina, Criciúma, Brazil

ABSTRACT KEYWORDS
The health benefit of a vegetarian diet is still under debate as it may result in a higher intake of Vegetarians;
some beneficial micronutrients, while others may be reduced, thus influencing various metabolic vegans;
pathways and health-related biomarkers. This scoping review discusses inflammatory, oxidative omnivores;
inflammation;
and DNA damage status in vegetarians and vegans compared to omnivores. Most of the reviewed
oxidative stress;
studies indicated favorable effects of a vegetarian diet on oxidative status compared to omnivores DNA damage
but did not clearly associate particular dietary habits to genome damage. The evidence on the
effect of vegetarian diet on the inflammatory and immunological biomarkers is poor, which could
at least partly be explained by methodological constraints such as small sample size, short duration
of vegetarianism and inconsistent definitions of the omnivorous diet. The only inflammatory
biomarker that seems to be associated with the vegetarian diet was inflammatory mediator
C-reactive protein, which in several studies showed lower values in vegetarians as compared to
omnivores. There were very few studies on immunological markers and the results on the difference
between vegetarians and omnivores were inconclusive. Although several biomarkers involved in
oxidative stress and inflammation showed a beneficial association with the vegetarian diet, further
research in well-defined and sufficiently sized cohorts is needed to provide more evidence.

Introduction Finally, the constraints include the inability to make food

Comparative anatomy, physiological processes, and evolu-
tionary data classify humans as omnivores, however all pri-
mates are specific types of omnivores whose diet is comprised
of a large proportion of vegetables (Andrews and Johnson
2020; Milton 2003). The earliest documents proving a switch
to a vegetarian diet in humans from Europe come from
Ancient times where Pythagoras and his way of life pro-
claimed an avoidance of meat consumption, and such an
approach was quite inspirational to many people up to the
19th century (Leitzmann 2014). To understand the phenom-
enon of turning to a vegetarian diet, Rosenfeld and Burrow
(2017) proposed a concept that presumes the interplay of
certain motivations, aversions, and constraints. Motivations
include improvements to personal health or animal welfare,
battling environmental degradation, upholding religious prin-
ciples, etc., while aversions are usually being disgusted by
or having a distaste for meat. Those aversions can even
strengthen one’s motivation to avoid meat eating, particularly
in people who turned to vegetarianism due to ethical issues.
choices freely, such as financial issues to purchase certain
food items, or social influences that would lead individuals
to contextual barriers (De Backer and Hudders 2014;
Hoffman et al. 2013; Ruby 2012). The reasons for choosing
a vegetarian diet often go beyond health and well-being
including economical, ecological and social concerns, which
leads to another sphere of nutritional ecology concerned
with sustainable life styles and human development
(Leitzmann 2005); however, Hargreaves et al. (2021) have
recently thoroughly reviewed domains affecting the quality
of life that might go in favor or against adopting a vege-
tarian diet. Some estimates suggest that there is 1.5 billion
vegetarians worldwide and 75 million of them are vegetar-
ians by choice (Leahy, Lyons, and Tol 2010), while the num-
ber of vegans is growing more rapidly than the number of
vegetarians (Leitzmann 2014).
There are several different plant-based diets and they
are categorized according to the degree to which prod-
ucts of animal origin are avoided (Figure 1). So let’s start

CONTACT Goran Gajski ggajski@imi.hr

© 2021 Taylor & Francis Group, LLC
2 G. GAJSKI ET AL.
Figure 1. Different plant-based diets categorized according to the degree of to
which products of animal origin are avoided. Plant-based diets inside the green
box should be considered vegetarian, since they avoid meat and seafood.
from the beginning. One of the most restrictive diets is
vegan, which excludes all animal originated products such
as meat, poultry, fish, seafood, eggs, and dairy while basing
its nutritional intake exclusively on vegetables, legumes, fruit,
nuts, whole grains, and seeds, as well as their derivatives.
Several even more restrictive subgroups of vegans include
raw vegans (raw food eaters) with the amount of uncooked
food that usually varies from 75% to 100% (Leitzmann
2005; Melina, Craig, and Levin 2016), and frutarians that
consume fruits only (Causso et al. 2010). The consump-
tion of honey is somewhat controversial so those vegans
who consume honey and bee products are called beegans.
When we talk about the vegetarian diet, which includes
the consumption of animal products except meat, it is also
divided into several groups. Ovo-vegetarians include eggs in
their diet, whereas lacto-vegetarians include dairy. Lacto-ovo
vegetarians include eggs and dairy as products of animal
origin, but exclude meat, poultry, fish and seafood. There
are also several diet types such as semi-vegetarianism, flex-
itarianism, pesco-vegetarianism, pollo-vegetarianism, even
vegans-before-6 pm and freegans that include occasional con-
sumption of fish and seafood, poultry, and red meat (Gaby
2013; Leitzmann 2005; Melina, Craig, and Levin 2016; Tuso
et al. 2013). These groups should not be considered vege-
tarians since they do consume animal meat although to a
lesser extent (EVU 2019).
The human diet, both vegetarian and omnivorous, if
not well-planned could become imbalanced particularly if
avoiding certain foods, either animal or plant, and may lead
to deficiencies or an excess of specific nutrients (Desmond
et al. 2021; Melina, Craig, and Levin 2016; O’Keefe et al.
2020; Sabaté 2003). Vegetarians, especially vegans, could
be at risk for deficiency of key nutrients that are typi-
cally provided by animal sources, specially vitamin B12,
high-quality proteins, iron, zinc, ω-3 fatty acids, vitamin
D, and calcium (Gaby 2013; Gajski, Gerić, Vučić Lovrenčić
et al. 2018; O’Keefe et al. 2018), so they are advised to
consume fortified foods and supplements for certain essen-
tial nutrients. Hence, there are several available guidelines
and recommendations for a well-balanced vegetarian diet
(Agnoli et al. 2017; Baroni et al. 2018; Gomes Silva et al.
2015; Lemale et al. 2019; Marsh et al. 2009; Melina, Craig,
and Levin 2016; Nakamoto et al. 2009; USDA and HHS
2015). Those who live in dominantly vegetarian regions
or are part of a traditionally vegetarian culture (such as
India where ∼35% of the population is vegetarian) or have
sought for a professional intervention to balance their diet
for optimal nutrition may be at less risk (Leitzmann 2014).
On the contrary, those who are new to a vegetarian diet
should carefully monitor their nutrition balance and incor-
porate fortified foods or supplements when appropriate to
avoid the possibility of malnutrition (Tuso et al. 2013). The
variety of plant-based diets and drivers leading to their
practice makes the recognition of a specific diet’s benefi-
cial properties to human health pretty complex. One note
about the terminology, where it has to be pointed out that
a plant-based diet is not a synonym for vegetarianism, as
well as about diet quality assessment, where plant-based
diets are not equivalent to high-quality diets. Plant-based
diets can also include highly processed, less healthful foods,
including savory snacks, pastries, and sugary fruit drinks
(Hemler and Hu 2019; Mayra, Ugarte, and Johnston 2019).
In general, vegetarianism as an alternative dietary pattern is
associated with many health benefits, as well as with some
health risks that rely on nutrient deficiencies, changes on
physiological and molecular level (Craig 2010; Leitzmann
2005; Lemale et al. 2019; Orlich et al. 2015; Oussalah et al.
2020; Tuso et al. 2013).
A growing body of evidence suggests that inflammation,
oxidative stress and DNA damage have a key role in the
pathogenesis of various diseases, including diabetes, obesity,
cancer, neurodegeneration, metabolic syndrome, cardiovascu-
lar disease, liver disease, and other chronic disorders. Elevated
levels of reactive species (reactive oxygen (ROS) and nitro-
gen (RNS) species), that result from the oxidant-antioxidant
imbalance, may lead to increased damage of biomolecules,
including DNA. Therefore, understanding the background
and molecular mechanisms underlying the mutual relation-
ship between inflammation, oxidative stress and DNA damage
can lead to the discovery of dietary strategies that would
favor prevention and provide treatment for some partic-
ular diseases. Although cells are equipped with numerous
non-enzymatic molecules and enzymatic scavengers of ROS
as well as DNA repair mechanisms, these defenses are not
always adequate to attenuate excess ROS production or DNA
damage. Hence, agents that can boost the antioxidant defense
system and prevent increased ROS generation can represent
an effective treatment for oxidative stress/inflammation-related
diseases and the food that we are consuming, and our dietary
habits can play a crucial role in such a strategy (Cena and
Calder 2020; Chatterjee 2016; Kawanishi et al. 2017; Kurutas
2016; Reuter et al. 2010; Willett et al. 1995). Today, there is
a body of evidence suggesting that different dietary habits

and patterns yield different, either positive or negative, effects
on inflammation processes, oxidative status and consequently
on genome integrity.
Therefore, in this scoping review we will give an extensive
overview on these three areas of research based on different
vegetarian dietary patterns compared to omnivores in healthy
populations using studies developed to assess the effects on
inflammatory, oxidative and DNA damage status. Moreover,
this review offers a complex insight into the differences
between selected biomarkers related to specific dietary pref-
erences, which could benefit clinicians and nutritionists in
patient counseling regarding nutrition and dietetics.
Search strategy
We conducted a literature search to identify relevant papers
using scientific databases. The databases included were
PubMed (www.pubmed.com) and Web of Knowledge (www.
webofknowledge.com) up to December 2020. We did not
use restrictions concerning publication language, country or
dates. The following search terms were used in our literature
search: (vegetarian* or vegan) and (inflammation OR inflam-
matory biomarker* OR inflammatory marker* OR proin-
flammatory biomarker* OR proinflammatory marker* OR
pro-inflammatory biomarker* OR pro-inflammatory marker*
OR inflammatory cytokine* OR proinflammatory cytokine*
OR pro-inflammatory cytokine* OR cytokine* OR IL-18 OR
interleukin-18 OR IL-10 OR interleukin-10 OR IL-6 OR
interleukin-6 OR IL-1α OR interleukin-1α OR IL-1β OR
interleukin-1β OR IL-1RA OR interleukin-1RA OR
TNF-alpha OR tumor necrosis factor alpha OR CRP OR
C-reactive protein OR hs-CRP OR high sensitivity C-reactive
protein OR ICAM/ICAM-1 OR intercellular adhesion mol-
ecule/ intercellular adhesion molecule-1 OR VCAM/VCAM-1
OR vascular adhesion molecule/ intercellular adhesion
molecule-1 OR MCP-1/CCL2 OR monocyte chemoattractant
protein 1 OR VEGF OR vascular endothelial growth factor
OR adiponectin OR E-selectin OR omentin/omentin-1 OR
resistin), for the impact of vegetarian diets on inflammatory
status, (vegetarian* or vegan) and (biomarker OR thiols OR
nitrotyrosine OR isolevuglandin OR isoprostanes OR gly-
coxidation OR hydroxynonenal OR peroxidase OR antioxi-
dants OR oxidative stress OR malondialdehyde OR carbonyls
OR glutathione OR catalase OR dismutase OR reactive oxy-
gen species) for oxidative status, and (vegetarian* or vegan)
and (biomarker OR DNA damage OR genome damage OR
micronucleus OR comet OR chromosome aberrations OR
H2AX OR sister chromatid exchange OR adducts OR epi-
genetics OR methylation OR acethylation OR phosphoryla-
tion OR ubiquitination OR miRNA) for DNA damage status.
Other relevant related original and review papers were also
checked in the reference lists of papers found in the search
and those papers have also been included in the pres-
ent review.
Our search retrieved 3695 articles for the impact of veg-
etarian diets on inflammatory, oxidative and DNA damage
status. For inclusion in this scoping review, we selected
altogether 115 papers following a screening of titles and
Critical Reviews in Food Science and Nutrition 3
abstracts. We omitted 3580 papers (exclusion criteria) with
duplicate results, conference abstracts, papers with no cor-
responding omnivore group, papers dealing with short-term
interventional dietary studies, papers dealing with
non-healthy subject, in vitro and animal studies, and papers
dealing with biomarkers beyond the scope of this review.
A detailed description of the selected studies can be found
in Tables 1–3 and the following chapters, whereas their
geographical distribution is shown in Figure 2.
Effect of vegetarian diets on inflammatory status
Inflammation is an important response of the body evolved
to protect the host from noxious stimuli, infections and
tissue injury. Next to acute stimuli, inflammation can be
triggered by a homeostatic imbalance of certain physiological
systems e.g., as a result of life-style factors like obesity or
malnutrition. Altered homeostasis may lead to sustained
adaptive changes that are of lower magnitude than by e.g.,
infections or tissue injury. Nevertheless, changes in inflam-
matory status even of lower intensity could shift the system
to different set points which may either be beneficial or
adverse for one’s health. The Western diet is characterized
by high consumption of foods associated with induced
inflammation such as fried foods, white bread, and meat,
while vegetarian diets generally contain higher amounts of
foods that are thought to be anti-inflammatory (Melina,
Craig, and Levin 2016). Furthermore, vegetarian diets are
generally associated with lower body weight (Berkow and
Barnard 2006; Spencer et al. 2003). Obesity is characterized
by a state of chronic, low-grade inflammation, and adipose
tissue is a significant source of hormones, adhesion mole-
cules, growth factors and cytokines (Engin 2017). Another
mechanism by which diet may influence inflammatory status
is the dynamic interaction between gut microbiota and
immune system, a pathway that is being increasingly studied
(Zheng, Liwinski, and Elinav 2020).
The inflammatory response is coordinated by a large
range of mediators which act as the effectors of inflamma-
tion (Chen et al. 2018). Many of these inflammatory medi-
ators have effects on the vasculature and recruitment of
leukocytes. Roughly, inflammatory mediators can be classi-
fied into seven groups based on the mode of action: vaso-
active amines, vasoactive peptides, fragments of complement
components, lipid mediators, cytokines, chemokines and
proteolytic enzymes. In this review we will focus on the
mediators involved in the inflammatory and immune (innate
as well as adaptive) response and have been investigated in
relation to the vegetarian and vegan diet.
The literature search resulted in a total of 37 articles
investigating the impact of the vegetarian diet on inflam-
mation (Table 1). The majority of studies measured the
inflammatory mediator C-reactive protein (CRP) as the end
point of the effect of a vegetarian diet, followed by a smaller
number of studies evaluating IL-6, tumor necrosis factor
alpha (TNF-α), adiponectin and IL-10. This is not surprising
as increased expression and activation of CRP is associated
with inflammatory processes extending into the vascular
Table 1. Characteristics of the reviewed studies regarding the association of the vegetarian and vegan diet with inflammatory biomarkers. 4

∼Age (years, Type of diet Biomarkers tested (in

No. of mean or Duration of relation to vegetarians/
Reference & country subjects range) Sex Vegetarian (vegan) Omnivores vegetarianism vegans) Main findings/comments
Suwannuruks et al. 1990 80 18–50 M&F Lacto-ovo – >1y fibrinogen Ø Data taken from Craddock et al. (2019).
(Thailand) Original article not available online
Pan et al. 1993 (Taiwan) 114 20–30 M&F Lacto & vegan Usual Chinese diet 2 y (min) fibrinogen Ø Women had a much higher concentration
G. GAJSKI ET AL.

of fibrinogen than did men, irrespective

Famodu et al. 1999
(Nigeria)
Haddad et al. 1999
(California, USA)
Mezzano et al. 1999
(Chile)
Sebeková et al. 2001
(Slovakia)
Szeto, Kwok, and Benzie
2004 (China)
Fontana et al. 2005
(Missouri, USA)
Krajcovicova-Kudlackova 270
and Blazicek 2005
(Slovakia)
Fontana, Klein, and
Holloszy 2006
(Missouri, USA)
of their diets.
76 18–22 & – Lacto-ovo & vegan Habitual African > 1 y, vegetarian fibrinogen ↓ (in both Results suggest that the vegetarian diet is
35–60 diet diet vegan and lacto-ovo a part of a healthier lifestyle resulting in
customarily vegetarians, also a more favorable status of hemostatic
followed in lower in vegans cardiovascular risk factors.
conjunction compared to
with principles lacto-ovo
recommended vegetarians)
by the 7th-Day
Adventist
Church
45 35 M&F Vegan All food categories 1 y (min) CRP Ø, IgG Ø, IgM Ø, It is not possible to determine from these
included IgA Ø, complement findings whether the immune status of
factor 3 ↓, vegans is compromised or enhanced
complement factor compared with non-vegetarians.
4 Ø, complement
factor 50 Ø
52 39 M&F Lacto-ovo, lacto & Mixed diet 1 y (min) CRP Ø, fibrinogen ↓ Considering the predictive value of
vegan inflammatory markers on cardiovascular
risk, the vegetarian diet does not seem
to offer an advantage at this level.
61 35 – Lacto-ovo, vegan & Traditional Western 8y CRP Ø and cystatin C Ø Plasma cystatin C and C-reactive protein
semi (plant food, mixed diet (for all vegetarian concentrations did not differ significantly
eggs, dairy and groups compared to between the groups.
fish) omnivores)
60 44 M&F Lacto-ovo & vegan – 5–55 y hs-CRP↓ Lower hs-CRP in the vegetarians indicates
that a long-term vegetarian diet may be
cardio-protective and anti-inflammatory.
36 54 (33–85) M&F Raw food vegetarian Typical American 3.6 (1.5–10) y hs-CRP ↓, leptin ↓, Circulating levels of CRP were significantly
(only diet IGF-1 ↓ lower in the raw food vegetarian group
plant-derived than in the control group, suggesting a
foods that have low level of systemic inflammation.
not been cooked,
processed or
otherwise altered
from their
natural state)
19–75 M&F Lacto-ovo & lacto Traditional mixed 10 y hs-CRP ↓ The mean concentration of hs-CRP in
diet vegetarians is significantly reduced as
compared to non-vegetarians. Moreover,
the hs-CRP concentrations decrease with
duration of vegetarianism.
42 53 M&F Raw food vegan Typical Western 2 y (min) hs-CRP ↓, IGF-I ↓, Plasma concentrations of inflammatory
(low-protein, mixed diet IGFBP-3 Ø, IGF-1/ markers were lower in subjects
low-calorie) IGFBP-3 ↓, fasting consuming a low-protein, low-calorie
C-peptide ↓, leptin diet than in non-obese, sedentary
↓ subjects who were consuming typical
Western diets.
Šebeková et al. 2006 136 37 M&F Lacto-ovo Standard Western 2–25 y hs-CRP Ø, cystatin C Ø Data suggest that a vegetarian diet may
(Slovakia) mixed diet postpone microinflammation associated
with the metabolic syndrome risk
factors.
Su et al. 2006 (Taiwan) 118 58 F Lacto-ovo – 5 y (min) sVCAM-1 ↑, sICAM-1 Ø In apparently healthy postmenopausal
vegetarians, elevation in homocysteine
and sVCAM-1 levels may attenuate the
potential beneficial effects of a
vegetarian diet on plasma lipids which
may possibly lower the development of
carotid atherosclerosis.
Chen et al. 2008 198 50 M&F Lacto-ovo – 1 y (min) hs-CRP ↓ Serum hs-CRP level was significantly lower
(Taiwan) in vegetarians than in omnivores as a
group, however non-significant trend
was seen when M & F were compared
separately.
Yen et al. 2010 (Taiwan) 98 2 age groups: M&F Lacto-ovo, lacto, ovo Mixed diet (plant 5&9y hs-CRP Ø No significant differences were found in
5 & 35 & vegan and animal serum hs-CRP between vegetarian
products) parents or their preschool children and
omnivorous controls.
Krajcovicova-Kudlackova 83 60–70 F Lacto-ovo & semi Traditional mixed 10 y CRP ↓ Low values of CRP in vegetarians are the
et al. 2011 (Slovakia) (white meat and diet consequence of regular and sufficient
fish ≤ 1 time per consumption of fruit and vegetables,
week) which are important sources of dietary
salicylates as well as other
anti-inflammatory compounds.
Yang et al. 2011 (China) 300 34 M Laco-ovo & vegan – 1 y (min) CRP Ø There were no differences in C-reactive
protein between vegetarians and
omnivores.
Su et al. 2011 (Taiwan) 90 58 F Laco-ovo & vegan – 5 y (min) hs-CRP Ø There were no differences in C-reactive
protein between vegetarians and
omnivores.
Ambroszkiewicz et al. 90 4–10 M&F Lacto-ovo, lacto, ovo – From birth leptin ↓, adiponectin Different kinds of diet might influence not
2011 (Poland) & vegan ↑, serum soluble only body mass and lipid profile but
leptin receptor ↑, also serum concentration of
leptin/sOB-R ↓ adipocytokines in prepubertal children.
Gorczyca, Prescha, 40 2–18 M&F – No food restrictions 1 y (min) IgA Ø, IgM Ø, IgG Ø Despite negligible differences in serum
Szeremeta, and immunoglobulin levels between
Jankowski et al. 2013 vegetarian and omnivore children, the
(Poland) impact of several nutrient intakes on
IgM and IgG levels differed between
groups.
Yu et al. 2014 (China) 195 36 M Vegetarian (no red Meat (at least 5 0.5 y (min) IL-1 Ø, IL-2 Ø, IL-6 ↑, Vegetarians have higher IL-6, but lower
meat, fish and times per week) IL-10 Ø, TNF-α Ø, LTB4, COX2 and PGE2 levels.
chicken) LTB4 ↓, COX2 ↓,
PGE2 ↓
(Continued)
Critical Reviews in Food Science and Nutrition
5
Table 1. (Continued) 6

∼Age (years, Type of diet Biomarkers tested (in

No. of mean or Duration of relation to vegetarians/
Reference & country subjects range) Sex Vegetarian (vegan) Omnivores vegetarianism vegans) Main findings/comments
Montalcini et al. 2015 52 32 M&F Lacto-ovo & vegan – 3 y (min) IL-1α Ø, IL-1β Ø, IL-2 Ø, Vegetarian diet could change the cytokine
(Italy) IL-4 Ø, IL-6 Ø, IL-8 profile, which in turn could influence
Ø, IL-10 Ø, IFN-γ Ø, resting energy expenditure.
G. GAJSKI ET AL.

TNF-α Ø, MCP-1 Ø,
VEGF Ø, EGF Ø
(log-transformed)
Chuang et al. 2016 4109 45 M&F Lacto-ovo, ovo & Mixed diet – CRP Ø Results suggest that lower inflammation
(Taiwan) vegan may not be a major mechanistic step
explaining the lower risk of hypertension
in vegetarians.
Ambroszkiewicz et al. 89 5–9 M&F Lacto-ovo Traditional mixed From birth CRP ↓ The median level of CRP was lower in
2017 (Poland) diet vegetarian children than omnivorous,
but CRP values were within the normal
range in all individuals.
Acosta-Navarro et al. 88 47 M Lacto-ovo, lacto & Meat (any type at 4 y (min) hs-CRP Ø Despite a better cardiovascular disease
2017 (Brazil) vegan least 5 servings biomarker profile, no differences were
per week) seen between omnivores and
vegetarians in CRP levels.
Franco-de-Moraes et al. 268 50 M&F Lacto-ovo & vegan Animal products – Median levels of CRP Observations of higher CRP and TNF-α/IL-10
2017 (Brazil) (more than once and TNF-α/IL-10 ratio in the omnivorous group reinforce
a month) increased gradually previous hypothesis that a saturated
from vegan toward fat-enriched diet could induce
lacto-ovo vegetarian inflammation.
and omnivores.
Pinto et al. 2017 (UK) 46 40–70 M&F Vegan Mixed diet (meat, 2 y (min) IL-6 Ø Finding of no significant differences in IL-6
fish, eggs and between vegans and omnivores does
dairy) not necessarily indicate that there are
no differences between groups in their
capacity to inhibit or resolve acute
inflammatory events due to limited
utility of circulationg cytokine
concentrations as biomarkers of
inflammation occurring in localized areas
of tissue.
Ambroszkiewicz et al. 117 5–10 M&F Lacto-ovo Traditional From birth Leptin/sOB-R ↓, HMW/ A well-planned vegetarian diet might
2018 (Poland) omnivorous diet total adiponectin Ø, beneficially affect the adipokine profile
resistin ↓, visfatin Ø, and inflammatory status expressed by
vaspin Ø, omentin Ø the ratios of anti-inflammatory to
pro-inflammatory adipokines in
prepubertal children.
Gajski, Gerić, Vučić 80 32 M&F Lacto-ovo & vegan Mixed diet 9y hs-CRP Ø It seems that meat consumption did not
Lovrenčić et al. 2018 (Mediterranean) affect the inflammatory status in
(Croatia) non-vegetarians.
Jaceldo-Siegl et al. 2018
(California, USA)
Singh et al. 2019
(California, USA)
Morgan-Bathke and
Jensen 2019 (USA)
Ganie et al. 2019 (India)
Grant et al. 2019
(Australia)
Menzel, Biemann et al.
2020 (Germany)
Cinegaglia et al. 2020
(Brazil)
Vučić Lovrenčić et al.
2020 (Croatia)
Yang et al. 2020
(Taiwan)
↑, significant increase; ↓, significant decrease; Ø, no effect; BMI, body mass index; COX2, cyclooxygenase-2; CRP, C-reactive protein; EGF, epidermal growth factor; F, female; HMW, high-molecular weight multimers;
hs-CRP; high-sensitivity C-reactive protein; ICAM-1 (sICAM-1), (soluble) intercellular adhesion molecule-1; sOB-R, soluble leptin receptor; IFN-γ, interferon gamma; IgA, immunoglobulin A ; IGF-1, insulin-like growth
factor 1; IGFBP-3, insulin-like growth factor-binding protein 3; IgG, immunoglobulin G ; IgM, immunoglobulin M; LTB4, leukotriene B4; M, male; MCP-1, monocyte chemoattractant protein-1 (chemokine (C-C motif )
ligand 2, CCL2); PGE2, prostaglandin E2; RA, receptor antagonist; TNF-α, tumor necrosis factor alpha; VCAM-1 (sVCAM-1), (soluble) vascular cell adhesion molecule-1; VEGF, vascular endothelial growth factor.
1371 59 M&F Vegan, lacto-ovo & Meat and fish (>1 – CRP ↓, significant The lower CRP and IL-6 concentrations
partial (semi and per week, no indirect associations among vegetarians may be mediated by
pesco vegetarians limits on dairy) of CRP and IL-6 with BMI.
included) vegetarian diet
mediated by BMI
no association
between IL-10 and
TNF-α with
vegetarian diet
74 51 M&F Vegan, lacto-ovo & Meat (quantity not – IL-6 Ø, CRP Ø There were no significant differences
pesco defined) between vegetarians and
non-vegetarians for log-transformed CRP
or log-transformed IL-6.
10 39 M&F Vegan, lacto-ovo & – 5 y (min) TNF Ø, IL-6 Ø Plasma TNF and IL-6 concentrations were
pesco not different between the omnivores
and vegetarians tested.
320 27 F Vegan Mixed diet (meat/ 1 y (min) TNF-α Ø, IL-6 ↑, IL-1β Pro-inflammatory markers were higher and
chicken/fish/egg Ø, hs-CRP ↑, resistin anti-inflammatory markers were lower
at least 5 days Ø, adiponectin ↓, among the healthy vegetarian women
per week) IL-10 Ø from Delhi.
688 15 M&F All vegetarians, Mixed diet (any – CRP Ø Although there was no statistically
regardless of the amount of meat, significant difference in the mean CRP
type, were poultry or fish) between the two diet groups, the
grouped in the prevalence of abnormal CRP
same category concentrations was somewhat high in
both groups.
72 38 M&F Vegan Mixed diet (at least 5y adiponectin Ø, hs-CRP No significant differences of the
3 portions of Ø, IL-18 Ø, inflammatory biomarkers were observed
meat per week omentin-1 Ø, resistin between vegans and omnivores.
or 2 portions of Ø, IL-1 RA Ø, ICAM-1
meat and 2 Ø
portions of
processed meat
per week)
88 46 M Lacto-ovo, lacto & Meat (any type of 4 y (min) hs-CRP Ø There was no significant difference in
vegan meat at least 5 hs-CRP level between vegetarians and
servings per omnivores.
week)
76 30 M&F Lacto-ovo & vegan Mixed diet 3–20 y hs-CRP Ø, adiponectin Serum hs-CRP and adiponectin levels did
(Mediterranean) Ø not differ between the vegetarian and
omnivore participants, however
adiponectin levels were significantly
higher in women than in men and in
female vegetarians compared to
omnivore controls.
75 38 M&F Lacto-ovo – <5 & >10 y CRP Ø There was no significant difference in CRP
level between vegetarians and
omnivores.
Critical Reviews in Food Science and Nutrition
7
Table 2. Characteristics of the reviewed studies regarding the association of the vegetarian and vegan diet with oxidative stress biomarkers. 8

∼Age (years, Type of diet

Subjects mean or Vegetarian Duration of Biomarkers tested (in relation
Reference & country (no.) range) Sex (vegan) Omnivores vegetarianism to vegetarians/vegans) Main findings/comments
Akesson and Ockerman 94 Different age M&F Lacto & vegan Common Swedish – Se ↓ No relation between dietary Se and urinary Se
1985 (Sweden) groups diets was observed.
Millet et al. 1989 106 40 M&F Semi, lacto-ovo – 12 y Vit A ↑Ø (sex dependent), Vit Differences in vitamin intake and level between
G. GAJSKI ET AL.

(France) & vegan (1–32) C ↑, Vit E ↑ vegetarians and omnivores.

Debski et al. 1989 38 29 F Lacto-ovo – – GPx ↑, Se ↑ Although data from the present study show that
(California, USA) Se in human milk can be modified by the
maternal diet, the increased Se content and
GPx activity in milk from vegetarians cannot
be explained by an increased Se intake.
Malter, Schriever, and 22 28–50 M – – – Vit A Ø, Vit E Ø, carotene ↑, Among the vitamins tested, only the level of
Eilber 1989 UA Ø carotene was significantly higher in
(Germany) vegetarians.
Flagg et al. 1993 12 18–61 M&F Lacto-ovo – – GSH ↑ Among Seventh-day Adventists, consumption of
(Georgia, USA) a vegetarian diet was associated with
increased plasma GSH concentration.
Pan et al. 1993 114 22 M&F – – 6y UA ↓ (sex dependent) Vegetarian women but not vegetarian men had
(Taiwan) lower uric acid than omnivores.
Krajcovicova-Kudlackova 109 24 M&F Lacto-ovo & – 3y CAT ↑, GPx Ø, CD Ø, Vit C ↑, Results indicate favorable consequences of
et al. 1994 (Slovakia) lacto Vit E Ø↑ vegetarianism with respect to pro-oxidative/
anti-oxidative parameters.
Rauma et al. 1995 40 45 M&F Vegan (raw) Matched 5y GPx Ø, SOD ↑, Vit C ↑, Vit E The present data indicate that the "living food
(Finland) omnivores ↑, β-carotene ↑ diet" provides significantly more dietary
antioxidants than does the cooked,
omnivorous diet, and that the long-term
adherents to this diet have a better
antioxidant status than do omnivorous control
subjects.
Fenech and Rinaldi 264 20–89 M&F Lacto-ovo Meat or meat 3 y (min) Vit C ↑, Vit E Ø Vegetarians were generally found to have
1995 (Australia) products significantly higher plasma levels of Vit C and
similar levels of Vit E compared with
non-vegetarian.
Kadrabová et al. 1995 88 35 M&F – – 1 y (min) GPx ↓, Se ↓, Zn ↓, Cu ↓ A vegetarian diet does not provide a sufficient
(Slovakia) supply of essential antioxidant trace elements.
Krajcovicova-Kudlackova 26 51 M&F Lacto-ovo & – 10 y Vit C ↑, Vit E ↑, β-carotene ↑ Favorable values of prooxidative-antioxidative
et al. 1995a lacto parameters demonstrated a reduced risk of
(Slovakia) lipoperoxidation in vegetarians.
Krajcovicova-Kudlackova 225 11–60 M&F Lacto-ovo & – 1.4 y (min) CAT ↑, CD Ø↑↓ (age and sex Results indicates positive effect of vegetarian
et al. 1995b lacto dependent), Vit C ↑ nutrition on efficiency of protection system.
(Slovakia)
Krajcovicova-Kudlackova 142 46 M&F Lacto-ovo & – 6y Vit A ↑, Vit C ↑, Vit E Ø, The results document a beneficial effect of
et al. 1995c lacto β-carotene ↑, Se ↑, Zn Ø, vegetarian nutritional habits on antioxidative
(Slovakia) Cu Ø parameters.
Krajcovicova-Kudlackova 162 20–40 M&F Lacto-ovo & – 3.5 y GPx Ø, CAT ↑, CD ↓, Vit C ↑, Results imply a higher protective effect against
et al. 1995d lacto Vit E ↑ lipid peroxidation.
(Slovakia)
Krajcovicova-Kudlackova 143 27 M&F Lacto-ovo & – 6y GPx ↑, CD ↓, Vit C ↑, Vit E Favorable values of prooxidative-antioxidative
et al. 1996a lacto Ø↑, β-carotene ↑, Se ↑ parameters demonstrated a reduced risk of
(Slovakia) lipoperoxidation in vegetarians.
Krajcovicova-Kudlackova 321 47 M&F Lacto-ovo & – 5y CD ↓, Vit A ↑, Vit C ↑, Vit E Results support the positive effect of
et al. 1996b lacto ↑, β-carotene ↑, Se ↑, Zn vegetarianism on the regulation of
(Slovakia) Ø, Cu Ø prooxidative processes.
Verhagen et al. 1996 40 45 F Vegan (living – 5y GPx Ø, SOD ↑, Vit C ↑, Vit E The present data indicate a few differences in
(Finland) food diet) Ø↑, β-carotene ↑ biomarkers of chemopreventive potential in
strict vegans vs matched omnivorous controls.
Meißner 1997 256 38 M&F – – – Se Ø There was no influence of vegetarian diet on
(Germany) blood and plasma selenium concentration.
Krajcovicova-Kudlackova 58 11–14 M&F Lacto-ovo & Mixed diet 3y TAS ↑, CD ↓, Vit A ↑, Vit C Favorable lipid and antioxidant parameters in
et al. 1997 (Slovakia) lacto ↑, Vit E ↑, β-carotene ↑ vegetarian children reflect the optimal
nutrition composition with respect to the
prevention of free radical diseases.
Kováčiková et al. 1998 196 35 M&F Semi, lacto-ovo – – GPx ↓, MDA ↓, Se ↓, Vit A Vit C was significantly higher, while the selenium
(Slovakia) & vegan ↑↓ (sex dependent), Vit C and the activity of GPx was lower both in
↑, Vit E ↑↓ (sex plasma and erythrocytes of vegetarians.
dependent)
Nagyová et al. 1998 38 19–56 M&F Lacto-ovo – 8y GSH Ø, TAS ↑, TBARS ↓, Vit A Although the resistance of LDL to in vitro
(Slovakia) Ø, Vit C ↑, Vit E Ø oxidation was not altered in vegetarians, the
effect of vegetarian nutritional habits
favorably affected TBARS levels in LDL and
total plasma antioxidant capacity.
Hänninen et al. 2000 40 – – Vegan (living Mixed food 14 y (min) Vit C ↑, Vit E ↑, α-carotene The subjects eating living food showed highly
(Finland) food diet) ↑, β-carotene ↑, lycopen increased levels of carotenes, lycopen, lutein
↑, lutein ↑ and vitamins.
Li et al. 2000 (Australia) 147 35 M Lacto-ovo & Moderate & high 0.5 y (min) Vit A ↓, Vit E Ø There are differences between the four diet
vegan meat eaters groups that have potential to affect the
subjects’ health and susceptibility to chronic
diseases.
Herrmann et al. 2001 104 23 M&F Lacto-ovo & High & low – TAS ↓ Vegetarians, especially the LME and vegan
(Germany) vegan meat-eaters groups, had a reduced TAS, whereas the TAS
of vegetarians was not different from that of
the HME group.
Manjari et al. 2001 74 31 M&F Lacto Consumed meat – MDA ↓, SOD ↑, CAT ↑, GPx Vegetarians have a higher concentrations of
(India) and Ø anti-oxidant enzymes and lower levels of
meat-based plasma lipid peroxides compared to
preparations at omnivores. Results suggest that vegetarians
least once in 3 have a higher anti-oxidant status.
days
Poornima et al. 2003 45 46 M Lacto Fish-based diet – MDA Ø, GSH Ø, Vit C ↑, UA Results show that the Vit C values were
(India) (occasionally Ø, ceruloplasmin Ø significantly higher in vegetarians.
other meats)
Krajcovicova-Kudlackova 211 36 M&F Lacto-ovo Traditional mixed 6y Vit C ↑, Vit E ↑, Se Ø, Zn Ø, A correct vegetarian nutrition or optimized mixed
et al. 2003 (Slovakia) diet Cu Ø, Fe ↓ diets may be an effective contribution to the
age-related chronic degenerative disease
prevention.
Krajcovicova-Kudlackova, 48 41 M&F Lacto-ovo Traditional mixed 10 y CD ↓, Vit C ↑, Vit E ↑, Results indicate that vegetarians have a better
Spustová, and diet β-carotene ↑, Fe ↓ antioxidative status as a consequence of
Pauková 2004 regular consumption of protective food.
(Slovakia)
(Continued)
Critical Reviews in Food Science and Nutrition
9
Table 2. (Continued) 10

∼Age (years, Type of diet

Subjects mean or Vegetarian Duration of Biomarkers tested (in relation
Reference & country (no.) range) Sex (vegan) Omnivores vegetarianism to vegetarians/vegans) Main findings/comments
Krajcovicova-Kudlackova, 48 41 M&F Lacto-ovo Traditional mixed 10 y Vit C ↑, Vit E ↑, β-carotene Higher antioxidative status in vegetarians indicate
Spustová, and diet ↑, oxidized purines ↓, that a correct vegetarian nutrition might
Pauková 2004 oxidized pyrimidines Ø represent an effective cancer prevention.
G. GAJSKI ET AL.

(Slovakia)
Kažimírová et al. 2004 48 41 M&F Lacto-ovo Traditional dietary 10 y TAC (FRAP) Ø, Oxidized bases Vegetarian diet can lead to a slight decrease in
(Slovakia) habit (Endo III Ø, Fpg ↑) oxidative DNA damage in lymphocytes. The
lowest level of DNA damage was found in
lymphocytes of lacto-vegetarians suggesting
that this diet provides some protection
against oxidative stress.
Szeto, Kwok, and 60 44 M&F Lacto-ovo – 22 y (5–55) MDA Ø, FRAP Ø, Vit C ↑, Vit Long-term vegetarians have a better antioxidant
Benzie 2004 (China) E ↓, UA ↓ status than do apparently healthy omnivores.
Tiahou et al. 2004 113 41 M&F Mostly Fish-based diet – MDA Ø, GPx ↓, AOPP Ø, Vit A The mountain region subjects have a higher
(Ivory Coast) vegetarian (coastal region, ↓, Vit E Ø, Se ↓, UA Ø, exposure risk to oxidative stress. However, the
(mountain predominantly tocotrienol ↑ absence of oxidative stress damage in
region, crude refined palm mountain subjects provides evidence of an
palm oil oil) effective protection. The long-term
diet) consumption of crude palm oil in mostly
vegetarian group could be considered as an
effective protective factor against oxidative
stress.
Cariappa et al. 2005 45 46 M Lacto Predominantly fish – MDA Ø Oxidant status does not vary significantly in the
(India) eaters two groups, suggesting that neither a
vegetarian diet nor a fish diet has a benefit
over the other.
Šebeková et al. 2006 136 37 M&F Lacto-ovo Standard Western 10 y (2–25) MDA Ø, SOD Ø, FRAP Ø, A balanced nutritional regimen renders
(Slovakia) mixed diet AOPPs ↑, AGEs Ø, Vit A Ø, comparable antioxidant defense in prevention
Vit C Ø, Vit E Ø, of plasma lipid peroxidation and advanced
β-carotene ↑ glycation.
Kažimírová et al. 2006 48 41 M&F Lacto-ovo & Traditional mixed 10 y FRAP Ø, Vit C ↑, Vit E Ø, Although the vegetarian lifestyle is supposedly
(Slovakia) lacto diet β-carotene ↑ healthy, it may lead to deficiency in some
micronutrients, which can influence the level
of certain biomarkers and partly counteract
the beneficial lifestyle of vegetarians.
Haldar et al. 2007 89 38 M&F Lacto-ovo & Did not have any 0.5 y (min) GSH Ø, GPx Ø, GST Ø, SOD Ø, The overall antioxidant status was similar
(Northern Ireland) vegan dietary FRAP Ø, UA Ø, Vit A Ø, Vit between vegetarians and omnivores.
(vegetarians restrictions C Ø, Vit E Ø, β-carotene Ø,
were defined α-carotene Ø, lutein ↑,
as those lycopene Ø,
who did not α-cryptoxanthin ↑,
consume β-cryptoxanthin Ø, Se ↓,
meat, Zn Ø
chicken, fish
or other
flesh foods,
any more
than six
times per
year)
Krajcovicova-Kudlackova 368 39 M&F Lacto-ovo & Traditional mixed 9y Vit C ↑ The data emphasize the role of Vit C in free
et al. 2007 (Slovakia) lacto diet radical disease prevention under the condition
of protective, antioxidative concentrations.
Krajcovicova-Kudlackova 161 45 (25 young F Semi & Traditional mixed 10 y Oxidative DNA damage The results suggest that increase of oxidative
et al. 2008 (Slovakia) & 65 older) lacto-ovo diet (oxidized purines ↓, damage in aging may be prevented by
oxidized pyrimidines ↓), vegetarian nutrition.
PC Ø, Vit A Ø, Vit C ↑, Vit
E Ø, β − carotene ↑ (Ø in
young group)
Hoeflich et al. 2010 107 28 M&F Lacto-ovo & Food of both 0.5 y (min) GPx Ø, Se ↓, SEPP Ø Whether German vegetarians and vegans need to
(Germany) vegans plant and be considered as a Se-deficient group
(vegetarians animal origin. depends on the biomarker chosen.
were defined
as those
who did not
consume
meat,
chicken, fish
or other
flesh foods,
any more
than six
times per
year)
Krajcovicova-Kudlackova 83 64 F Semi & Traditional mixed 10 y CD ↓, Vit C ↑, Vit E Ø↑, Favorable values of cardiovascular risk markers in
et al. 2011 (Slovakia) lacto-ovo diet β-carotene ↑ older vegetarian women document a
beneficial effect of vegetarian nutrition.
Somannavar and 100 45 M&F Lacto-ovo Consuming animal Since birth MDA ↓, GPx ↑, Vit A ↑, Vit E Results indicate that there was an increased lipid
Kodliwadmath 2012 products such ↑ peroxidation and a low antioxidant status in
(India) as meat, non-vegetarians compared to vegetarians.
poultry, fish
and other sea
foods at least
twice a week.
Kim, Cho, and Park 75 49 M&F Lacto-ovo – 25 y (17–35) SOD Ø, CAT Ø, GPx Ø, d-ROM Oxidative stress was lower in long-term
2012 (Korea) ↓, BAP Ø vegetarians compared to omnivores.
Ingenbleek and McCully 39 18–30 M Vegetarian Occasional – GSH ↓ Plasma levels GSH were significantly lower in the
2012 (Chad) consumption of study group.
bovine meat,
canned or
powdered milk
and smoked
fish.
Nadimi et al. 2013 40 37 M&F – – 3 (1–10) MDA Ø, PC Ø, Total Total anti-oxidant levels were higher in
(Iran) anti-oxidant ↑ vegetarians, carbonyl protein levels and MDA
levels showed no differences between the
groups.
Amirmozafari, 50 22–25 F – – 4–8 y SOD ↓, CAT ↓, peroxidase ↓ Data indicate a significant reduction in protecting
Pourghafar, and ability of saliva in vegetarian subjects.
Sariri 2013 (Iran)
(Continued)
Critical Reviews in Food Science and Nutrition
11
Table 2. (Continued) 12

∼Age (years, Type of diet

Subjects mean or Vegetarian Duration of Biomarkers tested (in relation
Reference & country (no.) range) Sex (vegan) Omnivores vegetarianism to vegetarians/vegans) Main findings/comments
Gorczyca, Prescha, 40 7 (2–18) M&F Semi & Reported meat 1 y (min) Fe Ø Children who follow a vegetarian diet may suffer
Szeremeta, and lacto-ovo, consumption from iron deficiency in spite of having a high
Jankowski et al. lacto Vit C intake.
G. GAJSKI ET AL.

2013 (Poland)
Schmidt et al. 2013 1693 45 M&F Lacto-ovo & Meat & fish eaters – UA ↑ Individuals consuming a vegan diet had the
(UK) vegans highest serum concentrations of UA compared
to meat eaters, fish eaters and vegetarians,
especially in men. Vegetarians and individuals
who eat fish but not meat had the lowest
concentrations of serum UA.
Boancă, Colosi, and 86 28 M&F Lacto-ovo – 2 to 29 y MDA ↓, SOD ↓ The SOD activity and MDA concentration were
Crăciun 2014 significantly lower in vegetarians. The duration
(Romania) of the vegetarian diet has a significant impact
only on the SOD activity.
Markussen et al. 2015 361 57 F Vegetarian Western, – Carotenoids ↑ (α-carotene, Vegetarian pattern is associated with a more
(Norway) continental & β-carotene, favorable profile of the plasma carotenoids.
high-protein β-cryptoxanthin, lutein,
diet lycopene, zeaxanthin)
Elorinne et al. 2016 41 34 M&F Lacto & vegan Omnivores 9 y (2–16) Vit E ↓, β-carotene ↓, Se ↓ Long-term consumption of a vegan diet was
(Finland) associated with some favorable laboratory
measures but also with lowered
concentrations of key nutrients compared to
reference values suggesting the need for
nutritional guidance to vegans.
Navarro et al. 2016 84 46 M Lacto-ovo, lacto Consuming any 4 y (min) MPO ↓ A vegetarian diet is associated with a healthier
(Brazil) & vegans type of meat at profile of cardiovascular biomarkers compared
≥5 servings per to omnivorous.
week.
Vanacore et al. 2018 30 29 M Lacto-ovo & All major types of 2 y (min) DPPH Ø, ABTSs Ø, FRAP Ø, The results demonstrated that restrictive vegan
(Italy) vegan food TBARS Ø, NO2- ↑, phenols diet could not prevent the onset of metabolic
Ø (↓ vegans) and cardiovascular diseases or protect from
oxidative damage.
Gajski, Gerić, Vučić 80 32 M&F Lacto-ovo & Mixed diet 9 y (3–20) MDA Ø, GSH ↓, UA Ø, Se Ø , The patterns of biomarkers were in favor of the
Lovrenčić et al. 2018 vegan (Mediterranean) Zn ↓, Cu ↓, Fe Ø consumption of both plant and animal
(Croatia) derived food.
Nebl et al. 2019 73 27 M&F Lacto-ovo & Both plant and 0.5 y (min) MDA ↑, nitrate ↑, nitrite ↑ The highest pre-exercise MDA and nitrate
(Germany) vegan animal foods concentrations were observed in vegans,
suggesting oxidative stress and NO synthesis
in this group.
Lin et al. 2019 (Taiwan) 63 53 M&F – – 13 y Vit A ↑ Data indicate that the vegetarians have higher
plasma Vit A then the omnivores.
Miles et al. 2019 840 59 M&F Semi, pesco, – – Vit A Ø, Vit E Ø, α-carotene Adventist Health Study participants following
(California, USA) lacto-ovo & ↑, β-carotene ↑, lutein ↑, certain vegetarian diet patterns have
vegan zeaxanthin Ø, significant differences in biomarkers associated
cryptoxanthin ↑, lycopene with dietary intakes compared to
Ø, total carotenoids ↑ non-vegetarians.
Weikert et al. 2020 72 38 M&F Vegan – 5 y (1–20) Vit A ↓, Fe Ø, Se Ø, Zn ↓ Lower SEPP P levels, but not selenium levels,
(Germany) indicate a reduced intake of selenium in
vegans compared to omnivores.
Cinegaglia et al. 2020 88 46 M&F Lacto-ovo & – 4–10 y, more HO-1 ↓ HO-1 induction in omnivorous may indicate a
(Brazil) vegan than 10 y pro-oxidative status since HO-1 is activated
under oxidative stress a state not seen in
vegetarians.
Björn Potthast et al. 71 27 M&F Lacto-ovo & – 0.5 to >3 y MDA Ø↑, SIRT1 Ø↓, SIRT3 In vegan participants, increased oxidative stress
2020 (Germany) vegan Ø↓, SIRT5 Ø↓ (exercise despite higher amounts of the antioxidative
dependent) substances in the diet was observed after
exercise.
↑, significant increase; ↓, significant decrease; Ø, no effect; AGEs, advanced glycation end products; AOPP, advanced oxidation protein products; BAP, biological antioxidant potential; CAT, catalase; CD, conjugated
dienes; Cu, copper; d-ROM, diacron reactive oxygen metabolites; F, female; Fe, iron; GPx, glutathione peroxidase; GSH, glutathione; GST, glutathione S-transferase; HO-1, heme-oxygenase-1; M, male; MDA, malond-
ialdehyde; MPO, myeloperoxidase; PC, protein carbonyl; Se, selenium; SEPP, selenoprotein P; SIRT1, sirtuin 1; SIRT3, sirtuin 3; SIRT5, sirtuin 5; SOD, superoxide dismutase; TAC, total antioxidant capacity; TAS, total
antioxidant status; TBARS, thiobarbituric acid reactive substances; UA, uric acid; Vit A, vitamin A; Vit C, vitamin C; Vit E, vitamin E; Zn, zinc.
Critical Reviews in Food Science and Nutrition
13
 14
Table 3. Characteristics of the reviewed studies regarding the association of the vegetarian and vegan diet with genome damage biomarkers.

∼Age (years, Type of diets Duration of

No. of mean or Vegetarian vegetarian Biomarkers tested (in relation to
Reference & country subjects range) Sex (vegan) Omnivores diet vegetarians/vegans) Main findings/Comments
Wulf et al. 1986 84 65 (15–88) M&F Lacto-ovo – – Lymphocyte SCE ↓ SDA had lower SCE frequency compared to
(Denmark) control omnivores. SDA vegetarians had
the same SCE frequency as the SDA
G. GAJSKI ET AL.

omnivores.
Fenech and Rinaldi 264 20–89 M&F Vegetarian Meat 5 days/week 3 y (min) Lymphocytes. Male: 20–40 y MNi The data from this study do not support
1995 (Australia) ↑, 41–60 y MNi ↓, 60+ y Ø. the hypothesis that vegetarians have a
Females: MNi Ø lower genetic damage rate than
non-vegetarians.
Verhagen et al. 1996 42 45 M&F Uncooked vegan – 5y Lymphocyte TL Ø, TM Ø, MNi Ø, There were no differences between the
(Finland) and induced MNi Ø groups.
Davies et al. 1998 39 58 & 54 F Vegetarian Meat eaters – Lymphocyte MNed cells Ø The consumption of meat elevated
(Canada) micronuclei frequency, although not
significantly.
Dhawan, Mathur, and 62 23–57 M&F Vegetarian – – Lymphocyte TL ↓, TI ↓, TM ↓ Differences in the extent of DNA damage
Seth 2001 (India) in the normal Indian population
depending on their eating and smoking
habits as well as age.
Bajpayee et al. 2002 230 25 M&F Vegetarian Occasional – Lymphocyte TL Ø, TI Ø, OTM Ø No differences between vegetarians and
(India) non-vegetarian (occasional) non-vegetarians.
Kažimírová et al. 2004 48 41 (20–69) M&F Lacto-ovo & Traditional mixed 11 & 8 y Lymphocyte CA Ø, MNi Ø, DNA Vegetarian diet can lead to a slight
(Slovakia) lacto strand breaks Ø, H2O2 challenge decrease in oxidative DNA damage in
Ø. SB + FPG sites ↓. Oxidized lymphocytes, but other markers of
pyrimidines genetic stability are not affected.
L-vegetarians < LO- vegetarians.
Krajčovičová-Kudláčková 48 41 (20–69) M&F Lacto-ovo Traditional mixed 10 y Lymphocyte DNA damage AU Ø, Vegetarian diet can lead to a decrease in
& Dušinská 2004 oxidized purines ↓ and oxidized purines in lymphocytes’ DNA
(Slovakia) pyrimidines Ø what is related to certain antioxidants.
Kažimírová et al. 2006 48 41 M&F Lacto-ovo & Traditional mixed 11 & 8 y Lymphocyte MN frequency Ø The micronucleus frequency did not differ
(Slovakia) lacto between vegetarians and
non-vegetarians, but was affected by
age, sex and total antioxidant capacity
of plasma.
Krajčovičová-Kudláčková 161 (20–30 & M&F Semi-vegetarians Traditional mixed 10 & 11 y Lymphocytes. Young: DNA damage Increase of oxidative damage in aging may
et al. 2008 60–70) (poll <2 (AU) Ø, oxidized purines Ø and be prevented by vegetarian nutrition.
(Slovakia) month) pyrimidines Ø. Elderly: DNA
damage (AU) ↓, oxidized
purines ↓ and pyrimidines ↓
Thaler et al. 2009 120 20–85+ M&F Lacto-ovo & Young and elderly 5 y (min) Buccal cells genome wide No difference in genome wide DNA
(Austria) pesco omnivores methylation Ø, telomerase methylation or telomerase activity in
activity Ø age matched vegetarians vs omnivores.
Such changes are age dependent.
Gadgil et al. 2014 49 27 & 28 F Lacto & vegan Egg/chicken/mutton/ – Plasma 5-methyl-20-deoxycytidine No differences in global DNA methylation
(India) fish 2 days/week Ø between the groups.
Tarallo et al. 2014 24 39 (21–60) M&F Vegetarian & – > 0 y (avr 3 Plasma miRNA-92a Different dietary habits influenced
(Italy) vegan y) Vegan > Vegetarian > Omnivores; individual miRNAs from the
Stool miRNA-92a concomitant analyses of plasma and
Vegan > Vegetarian and stool samples of healthy subjects.
Omnivores; Plasma and stool
miR-16 Ø, miR-21 Ø, miR-34 Ø,
miR-106a Ø, miR-146 Ø, and
miR-222 Ø
Kotova et al. 2015 53 21–37 M&F Vegetarian Traditional mixed 1 y (min) Transferrin positive reticulocytes Exclusion of meat and fish intake and the
(Sweden) MNi ↓ inclusion of protein-rich vegetarian
foods as an alternative, may be
beneficial in terms of reduced
cytogenetic damage.
Gajski et al. 2018 80 32 (19–59) M&F Lacto-ovo & Mixed diet 9 y (3–20) Blood cells TL ↑, TI ↑, TM ↑, MNi Lower nutritional status of some
(Croatia) vegan (Mediterranean) ↑, NPBs Ø, NBs Ø, telomere micronutrients accompanied with a
length (T/S ratio) Ø lower antioxidant defense system and
higher genome damage frequency in
vegetarians.
Cinegaglia et al. 2019 80 46 M&F Lacto-ovo, lacto At least 5 meat 4 y (min) Leukocyte telomere length (T/L No differences in LTL between the groups.
(Brazil) & vegan servings per week ratio) Ø LTL can be a marker of subclinical
atherosclerosis in the omnivorous
group.
Liu et al. 2020 96 61 M&F Semi-vegetarian, Meat more than 1 Range Plasma miRNA-3661 ↑, plasma Vegetarian diet has beneficial deregulation
(California, USA) vegetarian & time/week average 19 miRNA-204-3p ↑, plasma of miRNAs in terms of age-related
vegan to 24 y miRNA-320c ↑, plasma diseases
miRNA-29a-3p ↑, plasma
miRNA-320b ↑, plasma
miRNA-132-5p ↓
Miles et al. 2020 204 67 & 66 M&F Vegan Meat more than 1 – Modest ↓ leukocyte DNA Average CpG methylation across an entire
(California, USA) time/week methylation. gene showed 14 hypomethylated and 4
hypermethylated genes. Methylation of
genetic/intergenetic regions analysis
showed 21 differently methylated
genes. There is indication that more
differences exist.
Filippov et al. 2020 147 68, 70, 64 M&F Vegan & pesco Meat more than 1 – Ø methylation profiles from blood In relaxed statistical model ↑ methylation
(California, USA) vegetarian time/week cells DNA of PRR7, MRPL19, and SLC38A6 genes
in vegans. Hippo signaling and
Glutamatergic Synapse pathways were
enriched in omnivores and pesco
vegetarians.
↑ significant increase; ↓, significant decrease; Ø, no effect; AU, arbitrary units; CA, chromosome aberrations; F, female; FPG, formamidopyrimidine DNA glycosylase; M, male; MNi, micronuclei; MRLP19, mitochondrial
ribosomal protein L19; NBs, nuclear buds; NPBs, nucleoplasmic bridges; OTM, Olive tail moment; SB, DNA strand breaks; PRR7, proline rich 7; SCE, sister chromatid exchange; SLC38A6, solute carrier family 38 member
6; TI, comet assay’s tail intensity; TL, comet assay’s tail length; TM, comet assay’s tail moment.
Critical Reviews in Food Science and Nutrition
15
16 G. GAJSKI ET AL.
Figure 2. Geographical distribution of studies included in the review. Legend: The studies observed (a) inflammation biomarkers ( ), (b) oxidative stress bio-
markers ( ), (c) DNA damage biomarkers ( ), (d) biomarkers a + b ( ), (e) biomarkers a + c ( ), (f ) biomarkers b + c ( ), (g) biomarkers a + b + c ( ).
wall (Alizadeh Dehnavi et al. 2008), type 2 diabetes (Liu
et al. 2016) and different types of cancer (Il’yasova et al.
2005). A number of prospective epidemiological studies have
also shown that CRP is a strong and independent predictor
of cardiovascular risk (Ridker 2003; Ridker, Glynn, and
Hennekens 1998). Body mass index (BMI) and body fat,
suggested to be lower in vegetarians, were also shown to
correlate with CRP (Alizadeh Dehnavi et al. 2008).
In eight out of 26 studies, levels of CRP or hs-CRP
(which is more sensitive than the standard CRP test) were
lower in vegetarians as compared to omnivores, while only
one study showed an opposite pattern (Ganie et al. 2019).
No difference in the level of CRP and/or hs-CRP between
vegetarians and omnivores was detected in the remaining
studies.
In the American population, Fontana, Klein, and Holloszy
(2006, Fontana et al. 2005) found lower CRP values in
subjects on a raw vegan diet as well as low-calorie,
low-protein vegan diet as compared to a typical American
or Western diet, respectively. Jaceldo-Siegl et al. (2018)
found a significantly lower CRP level in vegetarians as
compared to omnivores, but only in those following a strict
vegetarian diet. In the other two studies carried out in the
American population (Haddad et al. 1999; Singh et al. 2019)
no difference in CRP levels was found between vegetarians
and omnivores. In a study by Singh et al. (2019), the inves-
tigated population had a predominantly Hispanic or Latino
background.
Lower hs-CRP in vegetarians (lacto-ovo and ovo) was
reported in a study in a Slovakian population by
Krajcovicova-Kudlackova and Blazicek (2005) showing a
decrease in hs-CRP with the duration of vegetarianism.
Consistently, the same research group confirmed decreased
CRP in participants who were on a lacto-ovo and semi
vegetarian diet (Krajcovicova-Kudlackova et al. 2011).
However, in two other studies on the Slovakian population,
no difference in hs-CRP between groups following a vege-
tarian and mixed diet was found (Sebeková et al. 2001;
Šebeková et al. 2006). Vučić Lovrenčić et al. (2020) com-
pared hs-CRP values of vegetarians with those of omnivores,
however no significant difference between these two groups
was found, which is in line with another study on the
Croatian population with the control group following a
mixed diet (Gajski, Gerić, Vučić Lovrenčić et al. 2018). No
difference in CRP levels regarding the vegetarian and omni-
vore diet was reported in studies carried out in the German
(Menzel, Biemann et al. 2020) and Australian (Grant et al.
2019) populations.
CRP levels in relation to the vegetarian diet have also
been investigated in several Asian populations. In the
Taiwanese population (Chen et al. 2008), a significantly
lower hs-CRP in vegetarians was reported as compared to
omnivores. However, no association between the vegetarian
diet and CRP was found in a large prospective cohort study
by Chuang et al. (2016) matched by age, sex, and study site,
although there was a tendency of lower CRP in vegetarians
in comparison with non-vegetarians. The absence of a sig-
nificant difference between vegetarians and omnivores in
the Taiwanese population was further reported in studies
by Su et al. (2011), Yang et al. (2020) and Yen et al. (2010).
In the Chinese population, a study by Szeto, Kwok, and
Benzie (2004) found significantly lower hs-CRP in vegetar-
ians while a more recent study found no effect of the veg-
etarian diet on CRP (Yang et al. 2011).
The effect of a vegetarian diet on CRP in the
South-American population was investigated in four studies.
In a cross-sectional study on the Brazilian population,
Acosta-Navarro et al. (2017) in a relatively small cohort of

male participants reported no difference in CRP between
vegetarians and omnivores, although the hs-CRP in vege-
tarians tended to lower values as compared to omnivores.
No effect of vegetarian diet on CRP was also reported in
a study by Cinegaglia et al. (2020). In contrast, another
study on the Brazilian population (Franco-de-Moraes et al.
2017) found lower CRP in vegetarians. The study conducted
in a Chilean population did not find a significant difference
in CRP between vegetarians and omnivores (Mezzano et al.
1999). Elevated hs-CRP levels in vegetarians were reported
only in one study, carried out in an Indian population
including apparently healthy vegetarian females compared
to BMI- and age-matched omnivores (Ganie et al. 2019).
Increased CRP values could, at least partially, be attributed
to dietary composition and/or the methods of food prepa-
ration, as the Indian vegetarian diet is rich in carbohydrates
and low in ω-3-fatty acids (Trichopoulou et al. 2014).
While most of the studies were carried out in adults,
two studies focused on the effect of a vegetarian diet in
children and teenagers. Ambroszkiewicz et al. (2017) found
that CRP was lower in Polish vegetarian prepubertal children
than in omnivores. Grant et al. (2019) in their investigation
of cardiovascular disease risk factors profile among Australian
teenagers found no difference in CRP levels between vege-
tarians and omnivores.
Three meta-analyses, which investigated relationships
between the vegetarian diet and levels of inflammatory
immune markers, showed that the vegetarian diet is asso-
ciated with lower levels of CRP (Craddock et al. 2019;
Haghighatdoost et al. 2017; Menzel, Jabakhanji et al. 2020).
In the most recent meta-analysis in apparently healthy
participants, lower CRP levels were found in both vegans
(3 studies) and vegetarians (14 studies) compared to omni-
vores (Menzel, Jabakhanji et al. 2020). Similar results of
lower CRP values in vegetarians compared to omnivores
(18 studies) were reported by Craddock et al. (2019),
although they included two studies with patients on dialysis
therapy in their analysis. In contrast, Haghighatdoost et al.
(2017) found no difference in CRP levels in subjects fol-
lowing a vegetarian diet compared to omnivores when
including subjects following a vegetarian diet for at least
6 months. However, subsequent subgroup analysis revealed
that lower CRP levels were observed in subjects following
a vegetarian diet for at least 2 years, indicating that there
is a minimum time from starting a vegetarian diet needed
for CRP to respond to the beneficial effects of a vegetarian
lifestyle.
Despite a substantial body of evidence that nutrition plays
an important role in innate, adaptive and cellular immunity,
there are very few studies on the effect of the vegetarian
diet on the expression of immunological markers (Table 1).
Most of the studies focused on the cytokines of the innate
immune system including the IL-1 cytokine family (Ganie
et al. 2019; Menzel, Biemann et al. 2020; Montalcini et al.
2015; Morgan-Bathke and Jensen 2019; Yu et al. 2014), IL-6
(Ganie et al. 2019; Jaceldo-Siegl et al. 2018; Montalcini et al.
2015; Morgan-Bathke and Jensen 2019; Pinto et al. 2017;
Singh et al. 2019; Yu et al. 2014) and TNF-a (Franco-de-
Moraes et al. 2017; Ganie et al. 2019; Montalcini et al. 2015;
Critical Reviews in Food Science and Nutrition 17
Morgan-Bathke and Jensen 2019; Yu et al. 2014), and only
a few studies have investigated biomarkers of adaptive
immunity (Menzel, Biemann et al. 2020; Su et al. 2006),
growth factors (Montalcini et al. 2015) and vascular cell
adhesion molecules (Menzel, Biemann et al. 2020; Su et al.
2006). None of the cytokine levels of the IL-1 family (IL-1a,
IL-1b, IL-1 RA, IL-18) were shown to be different between
vegetarians and omnivores (Ganie et al. 2019; Menzel,
Biemann et al. 2020; Montalcini et al. 2015; Yu et al. 2014).
There was also a lack of association between a vegetarian
diet and cytokine levels, namely IL-2, IL-4, IL-8, and IL-10
(Franco-de-Moraes et al. 2017; Ganie et al. 2019; Montalcini
et al. 2015; Yu et al. 2014). Reports comparing levels of the
most studied cytokine IL-6 among vegetarians and omni-
vores are inconclusive. Two of seven studies reported
increased values in vegetarians (Ganie et al. 2019; Yu et al.
2014), one study found decreased IL-6 values (Jaceldo-Siegl
et al. 2018), while in four studies no difference in IL-6
between vegetarians and omnivores was found. Lower IL-6
values reported in the study by Jaceldo-Siegl et al. (2018)
were explained by a mediating effect of lower BMI in veg-
etarians through an indirect pathway.
The proinflammatory cytokine TNF-α differed between
vegetarians and omnivores in only one of five published
studies and showed decreased expression in vegetarians
(Franco-de-Moraes et al. 2017). Among the studied param-
eters, vascular endothelial growth factor (VEGF), epidermal
growth factor (EGF) and insulin-like growth factor 1 (IGF-1)
(Fontana, Klein, and Holloszy 2006, Fontana et al. 2005;
Montalcini et al. 2015), known to moderate a variety of
immune events and secretion of cytokines, only IGF showed
a significant difference between vegetarians and omnivores
(being lower in vegetarians) (Fontana, Klein, and Holloszy
2006, Fontana et al. 2005). Haddad et al. (1999) and
Gorczyca, Prescha, and Szeremeta (2013) investigated the
levels of immunoglobulins IgG, IgM and IgA, but observed
no difference between vegetarians and omnivores. The lit-
erature on the impact of the vegetarian diet on T-cell medi-
ated immunity is scarce. Zhang et al. (2018) showed that a
long-term lacto-ovo-vegetarian diet was associated with a
lower diversity of T-cell repertoire and expression of IgE as
compared with subjects who consume an omnivorous diet.
Montalcini et al. (2015) investigated cytokines specific for
different subsets of T-cells such as IL-2, IL-4, and IL-10,
but found no difference between vegetarians and omnivores.
Surprisingly, only two studies investigated cell adhesion mol-
ecules, which are major regulators of leukocyte adhesion
and play an important role in inflammation (Kong et al.
2018). Levels of the intercellular adhesion molecule (ICAM)
did not show differences between vegetarians and omnivores
(Menzel, Biemann et al. 2020; Su et al. 2006), while levels
of vascular cell adhesion molecule (VCAM) were increased
in vegetarians (Su et al. 2006). Several studies focused on
inflammatory marker leptin which is mainly produced by
adipocytes and acts as a cytokine as well as a hormone
(Ambroszkiewicz et al. 2018, 2011; Menzel, Biemann et al.
2020). A consistent pattern was found, and the vegetarian
diet was associated with lower levels of leptin in all studies.
Inconsistent results have, however, been reported for
18 G. GAJSKI ET AL.
adiponectin, showing increased (Ambroszkiewicz et al. 2011)
or decreased (Ganie et al. 2019) levels in vegetarians, while
two studies found no difference between vegetarians and
omnivores (Ambroszkiewicz et al. 2018; Menzel, Biemann
et al. 2020). Evidence on the favorable adiponectin response
to a vegetarian diet in women, but not men, revealed an
interesting pattern of gender dimorphism regarding the met-
abolic effect of the vegetarian diet (Vučić Lovrenčić et al.
2020). The same sex-specific pattern was observed for adi-
ponectin in a recently published 4-week randomized con-
trolled trial by Lederer et al. (2020).
Due to its role in inflammation and immune response
(Zi and Xu 2018), the cysteine protease inhibitor cystatin
has been investigated in two studies (Sebeková et al. 2001;
Šebeková et al. 2006), however neither found a difference
between vegetarians and omnivores. Another inflammatory
marker, fibrinogen, has been investigated in four studies as
it has been shown to be associated with cardiovascular dis-
ease incidence (Pearson et al. 2003). In two studies, the
levels of fibrinogen were lower in vegetarians compared to
omnivores (Famodu et al. 1999; Mezzano et al. 1999), while
the other two studies found no difference between vegetar-
ians and omnivores (Pan et al. 1993; Suwannuruks
et al. 1990).
Taken together, the majority of studies found no differ-
ences in CRP (hs-CRP) levels between vegetarians/vegans
and omnivores. However, almost all of the studies that did
show a difference between these diet types, indicated a lower
CRP in vegetarians. This is consistent with recent
meta-analyses in three systematic reviews which suggest that
there is enough evidence of a beneficial effect of the veg-
etarian diet on keeping lower CRP levels, contributing to
lower cardiovascular risk. Regarding other inflammatory and
immunological markers, there is more evidence needed for
firm conclusions. Existing studies suffer from a small sample
size, differences in the definition of the vegetarian and con-
trol omnivore groups, as well as the duration of vegetari-
anism. Prospective studies in well-defined cohorts would
warrant better insight into the benefits of the vegetarian
diet regarding inflammatory processes.
Effects of vegetarian diets on oxidative status
Oxidative stress is defined as an imbalance between the
production of reactive species (ROS and RNS) and their
elimination by the cellular antioxidant defense system. The
consequences of uncontrolled oxidative stress can lead to
oxidative damage to biomolecules such as proteins, lipids
and DNA, which are critical to cellular structure and func-
tion. Luckily, powerful oxidative stress response mechanisms
have evolved to protect our organism from the damaging
effects of oxidative burden (Betteridge 2000; Ighodaro and
Akinloye 2018; Pruchniak, Aražna, and Demkow 2016;
Schmidt, Ghezzi, and Cuadrado 2021). However, uncon-
trolled oxidative stress has been implicated in a wide variety
of pathophysiology and age-related diseases such as cancer,
neurodegenerative disease, diabetes, cardiovascular condi-
tions, inflammatory disorders, asthma and male infertility
(Liguori et al. 2018; Pizzino et al. 2017). On the contrary,
ROS play an essential role in regulating various physiological
functions of living organisms (Brieger et al. 2012; Yang,
Chen, and Shi 2019). Therefore, understanding its role in
relation to dietary preferences was one of the goals of
this review.
Our search retrieved 59 studies investigating the impact
of the vegetarian diet on oxidative stress biomarkers yielding
different results (Table 2). It has been postulated that a
vegetarian diet may result in a higher intake of some vita-
mins and micronutrients that provide antioxidant defense,
while others might be deficient (Gajski, Gerić, Vučić
Lovrenčić et al. 2018; Kažimírová et al. 2006). Antioxidants
are substances that neutralize or remove free radicals by
donating an electron. The neutralizing effect of antioxidants
helps protect biological systems from oxidative stress (Lobo
et al. 2010). Examples of antioxidants include certain vita-
mins as well as micronutrients such as copper, zinc, sele-
nium and iron (Benzie and Choi 2014; Benzie and
Wachtel‐Galor 2009; Carlsen et al. 2010; Rahal et al. 2014).
Moreover, cells can also produce endogenous antioxidants
such as glutathione (GSH) (Duka et al. 2020; Jones et al.
2000; Pizzorno 2014). Some food nutrients, especially vita-
mins A (retinol), C (ascorbic acid), E (a group of eight
tocopherols and tocotrienols the most abundant of which
in human tissues is α‐tocopherol), and carotenoids, such as
β-carotene, lycopene, lutein, and zeaxanthin, are all
well-known antioxidants, as they can reduce oxidative stress,
which includes lipid peroxidation (LPO), protein carbonyla-
tion, and reduction of advanced glycated end-products
(Benzie and Wachtel‐Galor 2009; Eggersdorfer and Wyss
2018; Landete 2013; Rahal et al. 2014). As for vitamin
intake, the majority of the studies yielded contradictory
results, so we will highlight only those with the most
straightforward ones. Based on our literature review, we
found that different vegetarian diets could have a beneficial
effect in terms of higher vitamin intake (such as vitamins
A, C, and E) and their level in human body (Fenech and
Rinaldi 1995; Hänninen et al. 2000; Kažimírová et al. 2006;
Kováčiková et al. 1998; Krajcovicova-Kudlackova et al. 1994,
1995a, 1995b, 1995c, 1995d, 1996a, 1996b, 1997, 2007;
Krajcovicova-Kudlackova and Dusinská 2004; Lin et al. 2019;
Millet et al. 1989; Nagyová et al. 1998; Poornima et al. 2003;
Rauma et al. 1995; Somannavar and Kodliwadmath 2012;
Szeto, Kwok, and Benzie 2004; Verhagen et al. 1996) that
could also be sex-dependent (Kováčiková et al. 1998; Millet
et al. 1989). However, studies also suggest that there is no
difference in the level of vitamins (such as vitamins A and
E) between vegetarians and omnivores, and even that veg-
etarian vitamin pools (such as vitamins A and E) can be
lower (Elorinne et al. 2016; Fenech and Rinaldi 1995; Haldar
et al. 2007; Krajcovicova-Kudlackova et al. 2011, 2008; Li
et al. 2000; Malter, Schriever, and Eilber 1989; Miles et al.
2019; Nagyová et al. 1998; Šebeková et al. 2006; Szeto, Kwok,
and Benzie 2004; Tiahou et al. 2004; Weikert et al. 2020)
probably if their diet is not well balanced or due to the
poor bioavailability of these micronutrients from plant-based
foods (Platel and Srinivasan 2016). The same goes with

regard to carotenoids, whose concentration is generally
higher in vegetarians compared to omnivores (Hänninen
et al. 2000; Kažimírová et al. 2006; Krajcovicova-Kudlackova
et al. 1995b, 1995d, 1996a, 1996b, 1997, 2003, 2008, 2011;
Krajcovicova-Kudlackova and Dusinská 2004; Malter,
Schriever, and Eilber 1989; Markussen et al. 2015; Miles
et al. 2019; Rauma et al. 1995; Šebeková et al. 2006; Verhagen
et al. 1996) although some studies did not find significant
differences between dietary groups (Elorinne et al. 2016;
Haldar et al. 2007). The interpretation of antioxidant intake
as beneficiary should be taken with caution, since it was
shown that certain supplementations might even increase
cancer risks (Omenn et al. 1996).
Several micronutrients such as copper, zinc, selenium and
iron can be regarded as antioxidant micronutrients
(Ambroszkiewicz et al. 2017; Benzie and Wachtel‐Galor
2009; Craig 2010; Gibson 1994; Kadrabová et al. 1995) and
there are numerous studies that examined the status of such
antioxidant micronutrients in vegetarians compared to omni-
vores with slightly different outcomes. Vegetarians had a
significantly lower mean intake of selenium than meat eaters
and fish eaters, whose diets provided the higher amounts
of this micronutrient and this was shown in their status as
well, having lower blood/plasma selenium values (Akesson
and Ockerman 1985; Elorinne et al. 2016; Gajski, Gerić,
Vučić Lovrenčić et al. 2018; Haldar et al. 2007; Hoeflich
et al. 2010; Kadrabová et al. 1995; Kováčiková et al. 1998;
Sobiecki et al. 2016; Tiahou et al. 2004). However, there are
studies showing either the same or even higher levels of
selenium in vegetarians compared to omnivores (Debski
et al. 1989; Krajcovicova-Kudlackova et al. 1995b, 1996a,
1996b, 2003; Meißner 1997). As for levels of cooper, zinc
and iron, vegetarians and vegans tend to have a significantly
lower pool or comparable values of these particular antiox-
idant micronutrients compared to omnivore subjects
(Ambroszkiewicz et al. 2017; Gajski, Gerić, Vučić Lovrenčić
et al. 2018; Gorczyca, Prescha, Szeremeta, and Jankowski
et al. 2013; Haldar et al. 2007; Kadrabová et al. 1995;
Krajcovicova-Kudlackova, Spustová, and Pauková 2004;
Krajcovicova-Kudlackova et al. 1995b, 1996a, 2003; Weikert
et al. 2020). The lower level of such micronutrients com-
pared to omnivores may be the result of either their lower
bioavailability from this type of diet or the fact that some
of the accessed micronutrients are present in lower levels
in the vegetarian diet. Consequently, their lower status
caused by their inadequate intake may lower antioxidant
defense that in turn may contribute to a broad spectrum
of so-called redox diseases (Chiplonkar et al. 2004;
Cortese-Krott et al. 2017; Kadrabová et al. 1995). On the
other hand, lower iron stores have even been hypothesized
to reduce the risk of chronic diseases (Hunt 2003).
Cells produce GSH as an antioxidant to help resist oxi-
dative stress. GSH exists in a ratio of reduced (GSH) and
oxidized forms (GSSG) to maintain homeostasis. In normal
cells, more than 90% of GSH is in reduced form. An
increased GSSG-to-GSH ratio indicates oxidative stress
(Jones et al. 2000; Pizzorno 2014). Harmful hydrogen per-
oxide cellular levels are minimized by the enzyme
Critical Reviews in Food Science and Nutrition 19
glutathione peroxidase (GPx) using GSH as a reductant.
Furthermore, glutathione reductase (GR) helps maintain
GSH in reduced form, while glutathione S-transferase (GST)
conjugates reduced GSH to a variety of substrates for detox-
ification. Therefore, GSH, GPx, GR and GST can help eval-
uate the oxidative stress status of an organism and the
potential for downstream oxidative damage. Several studies
found significantly lower GSH concentrations in vegetarians
(Gajski, Gerić, Vučić Lovrenčić et al. 2018; Ingenbleek and
McCully 2012; Nagyova, Ginter, and Kovacikova 1995) indi-
cating a lower antioxidant defense in this group. This was
probably due to lower protein quality and quantity that the
vegetarian diet may provide, which leads to an insufficient
intake of sulphur-containing amino acids that may subse-
quently affect GSH values (Krajcovicova-Kudlackova et al.
1999). On the contrary, there are studies where GSH levels
were significantly higher in vegetarians compared to omni-
vores (Flagg et al. 1993) and those that found no difference
between the groups (Haldar et al. 2007; Manjari et al. 2001;
Nagyová et al. 1998; Poornima et al. 2003). Differences were
also found in the activities of GPx and GST, where several
studies showed either higher (Debski et al. 1989;
Krajcovicova-Kudlackova et al. 1996a; Somannavar and
Kodliwadmath 2012) or lower (Kadrabová et al. 1995;
Kováčiková et al. 1998; Nagyova, Ginter, and Kovacikova
1995; Tiahou et al. 2004) activities of measured enzymes or
no differences at all (Haldar et al. 2007; Hoeflich et al. 2010;
Kim, Cho, and Park 2012; Krajcovicova-Kudlackova et al.
1994, 1995c; Rauma et al. 1995; Song et al. 2014; Verhagen
et al. 1996). Vegetarians also showed significantly lower
activity of myeloperoxidase (MPO) (Navarro et al. 2016), a
heme peroxidase that catalysis the reaction between hydro-
gen peroxide and chloride ions to produce hypochlorous
acid as the primary oxidant (Frijhoff et al. 2015).
Concerning other antioxidant enzymes such as superoxide
dismutase (SOD) and catalase (CAT) contrasting findings
were also found between vegetarians and omnivores. The
major antioxidant enzymes, SODs, which includes copper-zinc
SOD, manganese SOD and extracellular SOD, all play critical
roles in scavenging superoxide radicals, the major and the
most toxic reactive species (Dikalov and Harrison 2014).
Decreased SOD activity results in elevated superoxide levels,
which in turn leads to decreased nitric oxide but increased
peroxynitrite concentrations. Increased SOD activity levels
are seen in Down’s Syndrome, while decreased activity is
seen in diabetes, rheumatoid arthritis, atherosclerosis, neu-
rodegenerative diseases, uremic anemia, certain cancers and
thyroid dysfunction (Fukai and Ushio-Fukai 2011; Liochev
and Fridovich 2007; McCord and Fridovich 1988; Younus
2018). CAT, another antioxidant enzyme, which is encoded
by a single gene and is highly conserved among species, is
one of the most efficient ways of removing cellular hydrogen
peroxide. Humans express CAT in all tissues, and a high
concentration of CAT can be found in the liver, kidneys as
well as erythrocytes. Deficiency or malfunction of CAT has
been postulated to be related to the pathogenesis of many
age-associated degenerative diseases like diabetes mellitus,
hypertension, anemia, vitiligo, cancer, neurodegenerative
20 G. GAJSKI ET AL.
diseases, bipolar disorder and schizophrenia (Deisseroth and
Dounce 1970; Glorieux and Calderon 2017; Nandi et al.
2019). Both SOD and CAT activities were generally increased
in vegetarians compared to omnivores
(Krajcovicova-Kudlackova et al. 1994, 1995a, 1995c; Manjari
et al. 2001; Rauma et al. 1995; Verhagen et al. 1996)
although there are studies indicating their lower activities
in vegetarian subjects or no differences between dietary
groups at all (Amirmozafari, Pourghafar, and Sariri 2013;
Boancă, Colosi, and Crăciun 2014; Haldar et al. 2007; Kim,
Cho, and Park 2012; Šebeková et al. 2006).
Several other biomarkers of oxidative stress were also
evaluated showing contradictory results. Plasma antioxidant
status (assessed with the FRAP assay) did not differ signifi-
cantly between vegetarians and omnivores (Haldar et al.
2007; Kažimírová et al. 2004, 2006; Šebeková et al. 2006;
Szeto, Kwok, and Benzie 2004; Vanacore et al. 2018), while
vegetarians had to some extent higher total antioxidant sta-
tus (Krajcovicova-Kudlackova et al. 1997; Nagyová et al.
1998) and lower values of reactive oxygen metabolites (Kim,
Cho, and Park 2012).
As a result of the imbalance between ROS production
and antioxidative defense, several products of oxidative stress
such as malondialdehyde (MDA), conjugated dienes (CD),
protein carbonyls’ (PC) and uric acid (UA), which can serve
as reliable biomarkers of oxidative insults, were also evalu-
ated in relation to dietary preferences. LPO is a well-known
mechanism of cellular injury in plants and animals and is
used as an indicator of oxidative stress in cells and tissues.
LPO products derived from polyunsaturated fatty acids
decompose to form a diverse mixture of compounds, includ-
ing MDA (Ayala, Muñoz, and Argüelles 2014; Esterbauer,
Schaur, and Zollner 1991). Generally, MDA and other reac-
tive aldehyde values were lower in vegetarians compared to
omnivores (Boancă, Colosi, and Crăciun 2014; Kováčiková
et al. 1998; Manjari et al. 2001; Nagyová et al. 1998;
Somannavar and Kodliwadmath 2012) although some studies
found opposite results or did not found significant difference
among the groups tested (Cariappa et al. 2005; Gajski, Gerić,
Vučić Lovrenčić et al. 2018; Nadimi et al. 2013; Nebl et al.
2019; Poornima et al. 2003; Šebeková et al. 2006; Szeto,
Kwok, and Benzie 2004; Tiahou et al. 2004; Vanacore et al.
2018). Furthermore, MDA inversely correlated with sirtuin
activities and was elevated in vegans after exercise, compared
to both omnivores and lacto-ovo vegetarians (Björn Potthast
et al. 2020). Similar results were found for CD as one of
the first peroxidation products of plasma fatty acids, where
generally lower values were observed in vegetarians
(Krajcovicova-Kudlackova, Spustová, and Pauková 2004;
Krajcovicova-Kudlackova et al. 1994, 1995a, 1995c, 1996a,
1997, 2011). As for PC, a biomarker of protein oxidation
(Augustyniak et al. 2015; Dalle-Donne et al. 2003), studies
found either no differences or lower levels in vegetarian
subjects (Krajcovicova-Kudlackova et al. 2008; Nadimi et al.
2013). Besides, circulating heme-oxygenase-1 (HO-1) level
was higher in healthy omnivores compared to vegetarians.
Since HO-1 is activated under oxidative stress, an increase
of circulating HO-1 in omnivores may indicate a pro-oxidative
status (Cinegaglia et al. 2020). Furthermore, there were gen-
erally no differences in UA level between vegetarians and
omnivores (Gajski, Gerić, Vučić Lovrenčić et al. 2018; Haldar
et al. 2007; Malter, Schriever, and Eilber 1989; Poornima
et al. 2003; Tiahou et al. 2004; Vučić Lovrenčić et al. 2020),
although some studies found decreased (Pan et al. 1993;
Szeto, Kwok, and Benzie 2004) as well as increased (Schmidt
et al. 2013) levels of UA in vegetarian subjects compared
to omnivores. UA is the final product of purine metabolism
in humans. Although its role in conditions associated with
oxidative stress is not entirely clear, evidence suggests that
increased serum levels of UA are a risk factor for cardio-
vascular disease where oxidative stress plays an important
pathophysiological role. Nevertheless, there is increasing
experimental and clinical evidence showing that UA has an
important role as an antioxidant (Glantzounis et al. 2005;
Pasalic, Marinkovic, and Feher-Turkovic 2012; Strazzullo
and Puig 2007).
The effect of oxidative stress was observed on the genome
level as well, where vegetarians exhibited lower oxidatively
damaged DNA compared to omnivores (Kažimírová et al.
2 0 0 4 ; Kr aj c ov i c ov a - Ku d l a c kov a e t a l . 2 0 0 8 ;
Krajcovicova-Kudlackova and Dusinská 2004) which will be
discussed in more details in the following chapter. Taken
together, most of the reviewed studies indicated favorable
effects of the vegetarian diet on oxidative stress parameters
compared to omnivores. Nevertheless, since some of the
studies suggested that a carnivorous diet rich in fruits and
vegetables is equally beneficial in terms of oxidative stress
it can be hypothesized that a well-balanced diet with the
consumption of both plant and animal derived food might
be as suitable as a plant-based one in terms of all of the
above-mentioned parameters.
Effect of vegetarian diets on genome integrity
The effects of the vegetarian diet on genome integrity were
reviewed as a possible link between inflammatory and oxi-
dative status and cancer occurrence; a growing global health
and socio-economic problem (Viegas et al. 2017). The
observed higher intake of several phytochemicals as well as
folate in the vegetarian diet are usually associated with
DNA-protective properties (Craig 2009; Crott et al. 2001;
Kimura et al. 2004). How is genome instability usually mea-
sured and how are those biomarkers related to health out-
comes? The frequency of micronuclei (MNi) is one of the
best validated and most commonly used biomarkers in
human biomonitoring studies evaluating DNA damage
(Gajski, Gerić, Oreščanin et al. 2018; Gajski et al. 2013;
Kopjar et al. 2010). The micronucleus reflects a piece of a
broken chromosome or an entire chromosome that has failed
to attach to a mitotic spindle (Fenech 2007; Garaj-Vrhovac,
Gajski, and Ravlić 2008; Guo et al. 2019). A higher MNi
frequency in a population indicates higher risk for all-type
cancer incidence (Bonassi et al. 2007). Within the analysis
of an MN sample, several other parameters can be traced.
Nucleoplasmic bridges (NPBs) reflect the existence of dicen-
tric chromosomes, fusion of telomere ends, or miss-repaired

DNA, while nuclear buds (NBUDs) represent over-amplified
DNA or DNA repair complexes extruded from the nucleus
(Fenech 2007; Kirsch-Volders et al., 2014; Thomas, Umegaki,
and Fenech 2003). Other commonly used biomarkers in
human biomonitoring are the comet assay’s tail length (TL),
tail intensity (TI), and tail moment (TM) if slides are ana-
lyzed using appropriate software, or arbitrary units (AU) if
slides are analyzed manually. The principle of the method
is to enable the release of DNA loops from gel-embedded
cells due to electric current in alkaline conditions. Single
and double strand breaks, alkali label sites, and strand breaks
associated with incomplete excision repair will facilitate the
relaxation of DNA, indicating higher DNA damage. During
the analysis, damaged cells appear as comets and the more
the DNA is damaged the higher the intensity of tail is
detected (Azqueta et al. 2020; Collins et al. 2014; Gajski,
Gerić, et al. 2020; Gajski, Langie, and Zhanataev 2020; Gerić
et al. 2018; Møller et al. 2021, 2020; Tice et al. 2000). TI
reflects the percentage of DNA in the tail and it is also
associated with increased risk of all-type cancer incidence
and may predict the risk of death (Bonassi et al. 2021; Milić
et al. 2021), while TL reflecting the comet tail’s length and
TM as a derivate of TL and TI have recently been discour-
aged from use. As carcinogenesis is a complex process, we
took into account the telomere shortening since critically
shortened telomeres lead to genomic instability and could
be modulated by a diet that might be sex-specific (Blasco
2005; Güneşliol et al. 2021; Öngel et al. 2020; Škrobot
Vidaček et al. 2018), as well as epigenetic changes respon-
sible for the structural adaptation of chromosomal regions
so as to register, signal or perpetuate altered activity states
(Bird 2007; Egger et al. 2004).
According to our search, 19 studies investigated the
impacts of the vegetarian diet on genome instability (Table
3). The results are not conclusive, though. Vegetarian
Seventh Day Adventists (SDA) were shown to have a lower
frequency of sister chromatid exchanges (SCEs) compared
to the general public. SCE is an exchange of genetic material
between two identical sister chromatids and has been used
as a mutagenic test in the past (Mourelatos 2016; Wilson
and Thompson 2007). However, within the group of SDA,
there were no differences between vegetarians and omni-
vores, indicating that total lifestyle might have been respon-
sible for the observed differences compared to omnivores
(Wulf et al. 1986). The SCE frequency serves as a good
biomarker for exposure to genotoxins, still due to an uncon-
firmed mechanism of biological significance and lack of
association with certain diseases, its application in human
biomonitoring is negligible (Bonassi et al. 2004; Wilson and
Thompson 2007). Epigenetic studies reveal that SDA vege-
tarians had a beneficial miRNA expression profile, showing
protective properties toward age-related diseases (Liu et al.
2020), as well as modestly hypomethylated leukocyte DNA
for genes responsible for metabolism, immune signaling,
protein degradation, lysosome regulation, RNA transport
and ribosome functions (Miles et al. 2020). Another study
also found modest changes in methylome, however the genes
related to ribosome structure, synapse-to-nucleus, and amino
acid solute carrier were highly methylated in SDA vegans
Critical Reviews in Food Science and Nutrition 21
(Filippov et al. 2020). The experience from the Slovak stud-
ies indicates that more abundant oxidative changes to DNA
bases might be observed in omnivores, particularly elderly
ones, while there were no differences in total DNA strand
breaks, chromosome aberrations, and MNi frequency, as
well as in the challenge assays between different dietary
groups (Kažimírová et al. 2004; Krajcovicova-Kudlackova
et al. 2008; Krajcovicova-Kudlackova and Dusinská 2004).
Vegetarians from the general Swedish population where a
lower frequency of MNi was observed in transferrin-positive
reticulocytes (Kotova et al. 2015) and in the general Indian
population where similar results were noticed (Dhawan,
Mathur, and Seth 2001), showed that vegetarians had lower
baseline cytogenetic status and less DNA strand breaks com-
pared to omnivores
The majority of the studies did not find any dietary
impacts on genome integrity. Interestingly, another report
from India, which covered a higher number of participants,
suggested no differences in DNA strand breaks between
vegetarians and omnivores (Bajpayee et al. 2002). No dif-
ferences in DNA strand breaks and lymphocyte MNi were
observed between groups practicing uncooked veganism and
matched omnivores (Verhagen et al. 1996). Among female
berry pickers of Asian descent, no differences in micronu-
cleated (MNed) cell frequency was found between meat
eaters and vegetarians (Davies et al. 1998). Leukocyte telo-
mere length (LTL), as a longevity biomarker, was not asso-
ciated to the vegetarian diet (Cinegaglia et al. 2019; Gajski,
Gerić, Vučić Lovrenčić et al. 2018), as well as telomerase
activity, genome wide and global DNA methylation in
plasma and buccal cells (Gadgil et al. 2014; Thaler et al.
2009). When assessing DNA damage at different age, Fenech
and Rinaldi (Fenech and Rinaldi 1995) found that young
(20–40 y) male vegetarians had a higher frequency of MNi,
middle-aged (41–60 y) male vegetarians had a lower fre-
quency of MNi, while elderly (60+ y) male vegetarians and
female vegetarians’ MN frequency at all ages did not differ
compared to omnivores.
Quite the opposite results were shown in certain other
studies. Higher DNA strand breaks and higher lymphocyte
MNi frequency was observed in a vegetarian group com-
pared to a matched group of omnivores, which was accom-
panied by several other blood/plasma biomarkers (Gajski,
Gerić, Vučić Lovrenčić et al. 2018). Negative epigenetic
trends were noted at the level of miRNA in both vegan and
vegetarian groups compared to omnivores. Groups avoiding
meat consumption showed higher plasma and stool
miRNA-92a concentration (Tarallo et al. 2014), associated
with tumor promotion, angiogenic functions, and metastatic
colorectal cancer mortality (de Miguel Pérez et al. 2020).
The miRNA-92a is also a part of the miRNA-17-92 cluster
linked to the pathogenesis of several malignancies (Di Leva
and Croce 2013; Tsuchida et al. 2011).
Taken together, most of the studies did not associate
particular dietary habits to DNA damage either “short-lived”
represented by repairable and repaired DNA (comet assay),
and “long-lived” represented by unrepaired, persistent breaks
(MNi) (Azqueta et al. 2014; Bolognesi et al. 1997; Collins
2004). Still, several conflicting studies suggest that carefully
22 G. GAJSKI ET AL.
designed studies with larger number of volunteers are
needed to reduce the variability of results and impacts of
possible confounders.
Discussion, future directions and conclusions
The scientific debate regarding vegetarian dietary health
benefits dates back as long as the late 19 th century
(Anonymous 1888a, 1888b; Green 1888) and it is still going
on in terms of whether the vegetarian as well as vegan diet
is healthier than a traditional one that includes food of
animal origin. It has been well-documented that long-term
avoidance of animal-originated food often results in a higher
intake of fiber, Mg, folate, vitamins C and E, and other
phytochemicals, while the intake of protein, vitamin B12 and
D, Ca, Fe, Zn, and ω-3 fatty acids are lower and potentially
leads to deficiencies. Although some plant-based foods are
quite abundant in some of these nutrients, food processing
and combinations of certain foods can decrease their bio-
availability (Agnoli et al. 2017; Craig 2009; Foster et al.
2013; Obersby et al. 2013; Platel and Srinivasan 2016).
Different dietary patterns impact human physiology, and
we will briefly give an overview of the processes on molec-
ular level influenced by an excess or lack of some nutrients.
The evidence on the effect of vegetarian diet on the inflam-
matory and immunological biomarkers is poor, which could
at least partly be explained by methodological constraints
such as small sample size, short duration of vegetarian diet,
and inconsistent definitions of the omnivorous diet. The
only inflammatory biomarker that seems to be associated
with the vegetarian diet was inflammatory mediator CRP,
which in several studies showed lower values in vegetarians
as compared to omnivores. There have been very few studies
on immunological markers and the results on the difference
between vegetarians and omnivores are inconclusive. In gen-
eral, a vegetarian diet results in a higher intake of some
vitamins and micronutrients used in antioxidant defense,
but it may also lead to a deficiency in other micronutrients
involved in DNA metabolism and stability (Claycombe and
Meydani 2001; Collins 2001; Field et al. 2018; Halliwell
2001; Hartwig 2001; Paoloni-Giacobino, Grimble, and
Pichard 2003). Dietary antioxidants have an important role
in cellular antioxidant defenses, enhancing DNA, protein
and lipid protection by modulating numerous signaling path-
ways and gene expression, protecting and repairing DNA
damage and increasing the free radical scavenging ability
that occurs during metabolic reactions (Khuda-Bukhsh, Das,
and Saha 2014; Nosrati, Bakovic, and Paliyath 2017; Varoni
et al. 2016). Besides, studies have shown that a human diet
with a high intake of fruits or vegetables rich in antioxidants
decreases the level of oxidative DNA damage (Duthie et al.
1996; Key, Appleby, and Rosell 2006; Pool-Zobel et al. 1997).
Since DNA damage itself is the initiating event in carcino-
genesis, this supports the idea that fruits and vegetables
might be beneficial in cancer protection by preventing free
radical attacks on macromolecules (Kažimírová et al. 2006;
Wasson, McKelvey-Martin, and Downes 2008). Furthermore,
antioxidants of plant origin such as vitamins C and E,
carotenoids and flavonoids are involved in protection from
diseases by decreasing the level of oxidative damage
(Smolková et al. 2004). Folate as an essential B vitamin
occurs naturally in a wide variety of plant foods and needs
to be obtained from dietary sources, as mammals are unable
to synthesize it de novo. Inadequate folate intake is associ-
ated with increased risk of neural tube defects, cardiovas-
cular disease, Down’s syndrome, Alzheimer’s disease as well
as various cancers (Beetstra et al. 2005). All of these findings
may indicate that vegetarianism increases antioxidant pro-
tection, implying a possible reduction in the risk of cardio-
vascular diseases and cancer (Kažimírová et al. 2004).
Moreover, vegetarian diets are associated with lower serum
concentrations of hs-CRP in long-term vegetarians
(Haghighatdoost et al. 2017) in addition to reduced body
weight and lower medical costs compared to non-vegetarian
diets (Berkow and Barnard 2006).
On the other hand, meats and dairy products are sources
of vitamin B12, which is required for cell division and eryth-
ropoiesis and its deficiency can cause anemia, lesions of the
nervous system and an increased level of homocysteine related
to increased risks for cardiovascular disease, atherosclerosis
and Alzheimer’s disease (Herrmann et al. 2001; Obersby et al.
2013). Vitamin D is another important vitamin that is present
mostly in food of animal origin (e.g., fatty fish and dairy
products) with a role in calcium metabolism (Holick 2007;
Keser et al. 2018; Pfotenhauer and Shubrook 2017). Besides
vitamins, in most vegetarians, iron, calcium and total protein
concentrations are lower, which may lead to genome instability
and oxidative stress (Ames 2001, 1999). Therefore, minimizing
or eliminating animal products from the diet decreases the
intake of some essential nutrients. Thus, the lack of balance
between the amount of “unhealthy” and “healthy” food also
leads to the accumulation of unrepaired damage, initiating
DNA instability and the possibility of inducing cancer devel-
opment (Kapiszewska 2006).
So, how is this molecular background reflected in epi-
demiological studies? The most straightforward effects of
long-term vegetarian diet are a reduction in BMI, as well
as reduction in cholesterol levels, and blood pressure.
Interestingly, the BMI reduction is applicable to western
societies, while no such effect was observed for the Indian
population (Appleby and Key 2016; Dinu et al. 2017; Key,
Appleby, and Rosell 2006). Lower cholesterol levels, blood
pressure and BMI in vegetarians reduce the risks for isch-
emic heart disease compared to non-vegetarians (Appleby
and Key 2016; Dinu et al. 2017). However, a balanced
plant-based diet that includes food of animal origin, where
∼20% of energy consumed from animal sources is replaced
by plant proteins, fat, polyunsaturated fatty acids, and whole
grain fatty acids and carbohydrates, also reduces coronary
hearth diseases risk and cardio-vascular diseases mortality
risk (Hemler and Hu 2019). A similar reduction of cardio-
vascular events was observed for long-term Mediterranean
diet adherence fortified by olive oil and nuts (Estruch et al.
2018). In general, high fruit, vegetable, and legume intake,
as well as individual types of fat were related to lower total
mortality, while processed carbohydrates, added sugars, and
 Critical Reviews in Food Science and Nutrition 23

processed grains have the opposite effects (Aune et al. 2017;
Dehghan et al. 2017; Miller et al. 2017; Ramsden and
Domenichiello 2017).
Another interesting point is the impact of the vegetarian
diet on cancer epidemiology. However, data are still incon-
clusive. While some studies report higher cancer rates in
vegetarians (Burkert et al. 2014), larger cohort studies
detected a slight reduction in overall cancer incidence for
vegetarians (Appleby and Key 2016; Dinu et al. 2017;
Segovia-Siapco and Sabaté 2019). Such a trend was observed
for stomach, lymphatic and hematopoietic cancers, while
reduced colorectal cancer rates were associated with intake
of fish and low meat consumption (Appleby and Key 2016;
Gilsing et al. 2016, 2015; Segovia-Siapco and Sabaté 2019).
The importance of a plant-based diet in reducing cancer
risks is probably associated with a high intake of fruit and
vegetables (Aune et al. 2017, 2018). On the other hand, red
and processed meat consumption and their impact on cancer
incidence is also still a matter of debate. Many studies have
found an association between meat consumption, DNA dam-
age, and different cancer incidences; however, due to the
research and publication bias, difficulties in assessing causal
relations and the contribution of meat compounds in car-
cinogenesis, high-quality research in order to give realistic
recommendations is needed (Lippi, Mattiuzzi, and Cervellin
2016; Pelland-St-Pierre et al. 2021). Lowering consumption
of meat tends to have a beneficial effect to human health,
however meat quality, as well as carbon footprint and envi-
ronment sustainability might also benefit from lowering
meat consumption (González et al. 2020).
The vegetarian diet is also associated with higher counts
of certain microbiota, which has been associated with high
fiber and polyphenol foods (Tomova et al. 2019), lower
urinary tract infections (Chen et al. 2020), lower rates of
diverticular diseases (Segovia-Siapco and Sabaté 2019), and
lower rate of type 2 diabetes (Appleby and Key 2016;
Segovia-Siapco and Sabaté 2019) as well as better periodontal
conditions (Staufenbiel et al. 2013). On the other hand, the
vegetarian diet reduces bone mineral density and increases
fracture risks mostly due to lower intake of proteins, Ca,
vitamin D and B12 (Appleby and Key 2016; Veronese and
Reginster 2019). The vegetarian diet is also associated with
a slightly higher prevalence of general psychopathology such
as depression, anxiety, and eating disorders, but the results
are still inconclusive and in need of more research or
meta-analyses (Askari et al. 2020; Baines, Powers, and Brown
2007; Beezhold et al. 2015, Beezhold, Johnston, and Daigle
2010; Burkert et al. 2014; Hibbeln et al. 2018; Jin et al.
2019; Lavallee et al. 2019; Matta et al. 2018; McComb and
Mills 2019; Michalak, Zhang, and Jacobi 2012; Paslakis
et al. 2020).
While exploring a complex outcome, such as disease
appearance or even mortality, it is usually difficult to dis-
entangle particular components impacting those outcomes.
To have more quality data and more accurate comparisons,
besides following the STrengthening the Reporting of
OBservational studies in Epidemiology (STROBE) checklist
(Gallo et al. 2012), we propose several recommendations
for further studies, summarized in Table 4.
• When performing a study to assess the effects of vegetar-
ian diet, the description of study groups is important to
eliminate volunteers consuming meat from the vegetar-
ian group. As previously discussed, a low meat, or fish
diet rich in vegetables and fruits showed certain health
benefits. Therefore we recommend a better description
of food consumption patterns and supplements intake,
using food frequency questionnaires and/or other tools
such as dietary indices (e.g., planet-based diet index,

Table 4. Recommendations for study design assessing the inflammatory, oxidative and DNA damage status and effects of vegetarian diets.
Issue Recommendation
Vegetarian group description Minimum criteria ruling out participants who consume meat, fish, and seafood (flexi-vegetarians,
semi-vegetarians, pesco-vegetarians, and pollo-vegetarians) from the vegetarian group
at least one year of adherence to vegetarian diet should be used as inclusion criteria for vegetarian group
reporting the average period and the range of vegetarian diet adherence
dietary supplementation reporting
reporting of the reason for turning vegetarian
Additional criteria at least two years of adherence to a vegetarian diet as the inclusion criteria for the
vegetarian group
subdividing vegetarians into lacto-, ovo-, lacto-ovo-vegetarians, or vegans
Recommended criteria the use of appropriate nutritional tools to assess food consumption pattern (food
frequency questionnaire, indices, etc.), energy and nutrient intake (food dairy, 24-hour recall, etc.)
Omnivore group description Minimum criteria describing the group as much as possible e.g., Western diet pattern, Mediterranean diet
pattern, carnivorous diet rich in fruits and vegetables, etc.
dietary supplementation reporting
Recommended criteria the use of an appropriate nutritional methodology to assess food consumption (food
frequency questionnaire, eating diaries, etc.), energy and nutrient intake
Groups matching Minimum criteria the compared groups should not differ statistically in volunteers’ number, age, and sex
other potential confounders should be addressed (e.g., number of cigarettes per day, residence, education level,
personal/house income, radiation exposure, medication use, family cancer history, medications, etc.)
Additional criteria the groups should be matched by volunteers’ number, age, and sex between the groups
Recommended criteria matching in more possible confounders
Lifestyle Minimum criteria the groups should report alcohol consumption and physical exercise, as well as meditation
Recommended criteria assessment of alcohol consumption by nutritional methodology (food frequency
questionnaire, food diaries, etc.) and physical activity/body fitness by appropriate quantitative methods
(questionnaires, simple physical tests, etc.)
Conflict of interest Recommended criteria the authors should declare their diet preferences, as well as funding by a certain
industry or membership in societies related to a specific human diet, animal welfare, or environment protection
24 G. GAJSKI ET AL.
Figure 3. The proportion of papers showing results in favor of vegetarian or
omnivore inflammatory, oxidative stress and DNA damage status according to
the studies selected in the review. Legend: vegetarian-beneficial papers ( ),
omnivores-beneficial papers ( ), no differences papers ( )
healthful and unhealthful plant-based diet index, and/
or pro-vegetarian diet index) that could better distin-
guish healthy from less-healthy plant-based diets (Kim,
Lee, et al. 2020; Satija et al. 2016, 2017; Thornton and
Villamor 2016). The length of practicing vegetarianism
is important to separate the short-term and long-term
effects of a diet. Motivations for becoming vegetarian
might be useful in interpreting results, particularly for
individuals driven by the health improvement effects
of a diet. Finally, the description of the non-vegetarian
group is important to minimize potential bias of a
highly imbalanced omnivorous diet.
• When comparing health effects of different groups where
a criterion such as age and sex might impact the results,
such data for two groups should be as similar as pos-
sible, preferably the same.
• Change in dietary habits usually leads to change in life-
style as well (e.g., physical activity, meditation, avoidance
of tobacco or alcohol products, better health awareness,
etc.). Description or measuring lifestyle increases the
quality of data interpretation.
• Since one’s diet is quite personal, and the debate regard-
ing health effects is still on, there is a possibility of
unintentional bias in data interpretation, so reporting of
dietary preferences as well as usual conflicts of interest
should be done.
To sum up, based on the reviewed literature, most of the
studies did not clearly associate particular dietary habits to
inflammatory biomarkers and genome damage, while most
did indicate the favorable effects of the vegetarian diet on
oxidative stress parameters compared to omnivores (Figure 3).
Nevertheless, since some of the studies suggested that an
omnivorous diet rich in fruits and vegetables is equally
beneficial in terms of oxidative stress, it can be hypothesized
that a well-balanced diet with the consumption of both
plant- and animal-derived food might be the most suitable.
There are also several alternatives how to balance our diet
in more environment-sustainable way and still improve per-
sonal health as well as to battle environmental degradation
to solve this diet–environment–health trilemma, such as the
Mediterranean or EAT-Lancet diet, or the use of low-food
chain animals (i.e., forage fish, bivalve mollusks, edible
insects) demanding less resources (i.e., greenhouse gas,
water, and carbon footprints) (Auestad and Fulgoni 2015;
Churchward-Venne et al. 2017; Clonan and Holdsworth
2012; D’Antonio, Serafini, and Battista 2021; Dernini et al.
2017; Kim, Santo, et al. 2020; Reisch, Eberle, and Lorek
2013; Tilman and Clark 2014; Willett et al. 2019, 1995).
Besides, despite the motivation to avoid consumption of
foods of animal origin, the diet should be carefully tailored
by a nutritionist and physician, thus avoiding health issues.
One of the key issues is also to consume less processed
food, both of animal or plant origin. We might say that the
future of the human diet is green, but it is more easily to
balance it with a pinch of quality meat.
Acknowledgements
This work was supported by affiliated institutions and by the European
Cooperation in Science and Technology (COST Actions CA15132 –
hCOMET, CA16112 – NutRedOx and CA16113 – CliniMARK). We
also thank Mr. Makso Herman for English language editing and
Flaticons for the icons used for graphical design.
Disclosure statement
GG, MG, IJ, IP, AMD, MVL, SK, and MB are omnivores, while VMA
was flexitarian for 30 years and has been an omnivore the last 3 years.
The authors also declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Funding
The author(s) reported there is no funding associated with the work
featured in this article.
ORCID
Goran Gajski http://orcid.org/0000-0002-1886-1453
Marko Gerić http://orcid.org/0000-0002-5886-4106
Ivone Jakaša http://orcid.org/0000-0002-7961-4069
Ines Peremin http://orcid.org/0000-0002-0517-4734
Ana-Marija Domijan http://orcid.org/0000-0001-5645-9732
Marijana Vučić Lovrenčić http://orcid.org/0000-0001-7365-0627
Sanja Kežić http://orcid.org/0000-0002-1063-4547
Martina Bituh http://0000-0002-3596-8383
Vanessa Moraes de Andrade http://orcid.
org/0000-0002-8474-4482

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