Trends in Food Science & Technology 109 (2021) 116–125

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Trends in Food Science & Technology

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

Oak trees (Quercus spp.) as a source of extracts with biological activities: A

narrative review
Diego Morales a, b, *
a
AZTI, Food Research Division, Astondo Bidea 609, 48160, Derio, Spain
b
Department of Production and Characterization of Novel Foods, Institute of Food Science Research, CIAL (UAM+CSIC), C/ Nicolas Cabrera 9, Campus de
Cantoblanco, Universidad Autónoma de Madrid, 28049, Madrid, Spain

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

Keywords: Background: Oak trees (Quercus spp.) constitute a relevant source of fractions and ingredients that have been
Oak tested regarding their biological activities. The high number of species, varieties and materials that can be found
Quercus through this genus has led to a great diversity of extracts that can be investigated regarding their potential to
Antioxidant
improve human health status.
Antitumoral
Immune-modulatory
Scope and approach: In this review, several biological activities were revised through the scientific literature
Antimicrobial (antioxidant, immune-modulatory, antiproliferative, hypoglycaemic, hypocholesterolemic, antihypertensive,
antimicrobial, etc.) paying attention to bioactive extracts obtained from different oak trees and parts, describing
the selected extraction technology, the utilised experimental model, the published results and their potential
impact on human health. Moreover, the related strengths and weaknesses were listed to elucidate the current
state of the related scientific evidence.
Key findings and conclusions: In vitro and in vivo analyses confirmed the potential of oak extracts particularly for
antioxidant, anti-inflammatory, antitumoral and antimicrobial effects. However, some points must be further
investigated: additional bioactive molecules and abilities testing, advanced extraction techniques evaluation and
more animal studies and clinical trials to validate the mentioned activities when the extracts are included in food
matrices.

1. Introduction
The use of natural sources of bioactive compounds have been utilised
since ancient times becoming a relevant complement of synthetic
pharmacological drugs.
An enormous diversity of plant species has been tested and subjected
to extraction procedures to obtain fractions with high levels of bioactive
molecules such as phenolic compounds (flavonoids, tannins), lipids
(triterpenoids, phytosterols), peptides, polysaccharides, saponins, alka­
loids, etc. (Essien, Young, & Baroutian, 2020; Morales, Miguel, &
Garces-Rimon, 2020; Patra, Das, Lee, Kang, & Shin, 2018). Among them,
species belonging to Quercus genus have been evaluated to recover ex­
tracts with biological activities such as antioxidant, anti­
tumoral/antiproliferative, anti-inflammatory/immune-modulatory,
antidiabetic/hypoglycaemic, hypocholesterolemic, antihypertensive,
antimicrobial, etc. These fractions might be used as medicinal nutra­
ceuticals or utilised in the design of novel functional foods.
Quercus trees (commonly named oaks) belong to one of the most
relevant genera in the Northern Hemisphere. Quercus genus has been
subjected to several re-classifications leading to the recognition of more
than 500 species distributed in different areas (Denk & Grimm, 2009)
(Table 1). Some of them are highly appreciated in Europe, Asia and
North America such as Quercus ilex, Quercus robur or Quercus arizonica.
The mainly used parts of these trees have been wood and bark as well
as acorns for human or animal feeding and leaves to prepare infusions
with medicinal or nutritional purposes (Costa, Lourenço, Oliveira, &
Pereira, 2019; Gamboa-Gomez et al., 2017; Vinha, Barreira, Costa, &
Oliveira, 2016). However, the search for bioactive fractions goes beyond
these widely utilised materials; therefore, other underexploited com­
ponents of the plants have been investigated. In this sense, Q. ilex roots
were macerated with methanol to obtain phenolic fractions that are able
to exert antioxidant and genoprotective effects (Meziti et al., 2019) and
Quercus infectoria galls (irregular plant protuberances provoked by some
insects) were extracted with different solvents to recover fractions with

* AZTI, Food Research Division, Astondo Bidea 609, 48160, Derio, Spain.
E-mail addresses: diegomoraleshdz@gmail.com, dmorales@azti.es.

https://doi.org/10.1016/j.tifs.2021.01.029
Received 20 November 2020; Received in revised form 15 December 2020; Accepted 9 January 2021
Available online 15 January 2021
0924-2244/© 2021 Elsevier Ltd. All rights reserved.
D. Morales Trends in Food Science & Technology 109 (2021) 116–125

Table 1 to recover not only phenolics but also bioactive lipids, polysaccharides
Geographical distribution and main parts of interest of the most studied Quercus and other molecules using supercritical fluids, pressurized liquids as
species. Used sources and databases: European Atlas of Forest Tree Species, In­ well as microwave- or pulsed electric fields-assisted extractions that
ternational Oak Society “Quercus Portal”, The Wood Database, USDA Plants have been widely utilised to obtain bioactive fractions from natural
Database. sources (Gil-Ramirez, Smiderle, Morales, Iacomini, & Soler-Rivas, 2019;
Quercus spp. Distribution Main parts of Patra et al., 2018; Tejedor-Calvo et al., 2020; Zia, Khan, Shabbir, Maan,
interest & Khan, 2020).
Q. arizonica USA, Mexico Acorns, leaves The Quercus spp. extracts and their isolated compounds were tested
Q. aucheri Greece, Turkey Seeds, galls regarding the previously mentioned biological activities. Together with
Q. brantii Iran, Iraq, Syria, Turkey Seeds, wood
in vitro methods that yielded promising results, animal models
Q. castaneifolia Azerbajan, Iran Acorns, leaves
Q. cerris Italy, Turkey, Balkans Acorns, leaves confirmed their potential and even several clinical trials were developed
Q. coccifera Spain, Northern Africa, Italy, Balkans, Acorns to evaluate wound-healing, antimicrobial and antitumoral effects of
Turkey Quercus materials-derived products (Afzali, Siahposh, Haghighi-Zadeh,
Q. convallata Mexico Leaves Farajzadeh, & Abbaspoor, 2020; Chokpaisarn, Chusri, & Vor­
Q. crassifolia Mexico, Guatemala Wood
avuthikunchai, 2020; Stauder et al., 2009; Zare et al., 2019). However,
Q. dilatata Afghanistan, Pakistan, Nepal Acorns
Q. durifolia Mexico Acorns, leaves the interactions between bioactive compounds and human gut micro­
Q. eduardii Mexico Leaves biome have not been deeply studied yet; only the effect of Quercus
Q. gilva Japan, Korea, China Leaves mongolica seeds consumption in rodents microbiome was assessed (Yi,
Q. grisea USA, Mexico Acorns, leaves
Guo, Wang, Xue, & Ju, 2020; Zhang, Bartlow, Wang, & Yi, 2018) and
Q. ilex Spain, Portugal Acorns
Q. incana USA Wood Quercus robur extracts were incubated with human faecal samples in an
Q. infectoria Middle East, Greece Galls ex vivo experiment (Piwowarski et al., 2014). This point must be
Q. laeta USA, Mexico Leaves considered in future research due to the impact of these interactions in
Q. mongolica Japan, Korea, China, Mongolia, Siberia Wood the biological activity and bioavailability of plant molecules (Vamanu &
Q. obtusata Mexico Leaves
Gatea, 2020).
Q. petraea Europe Wood, acorns
Q. phillyraeoides China, Japan Wood The scientific literature was reviewed in this article, paying special
Q. pyrenaica Spain, France, Portugal Wood, bark attention to the studied oaks species and raw materials, the reported
Q. resinosa Mexico Wood biological activities, the described experimental models, the extraction
Q. robur Europe, Caucasian region Acorns, bark
methods, the isolation and identification of active molecules (when it
Q. serrata China, Japan, Korea, Taiwan Acorns, galls
Q. sideroxyla Mexico Wood
was possible) and the potential effects on human health.
Q. suber Spain, Portugal Acorns, bark
2. Antioxidant activity

high levels of antioxidant and antitumoral tannins and other phenolic
compounds (Ahmad & Abdulah, 2018; Arina & Harisun, 2019; Cheng
et al., 2018; Kaur, Athar, & Alam, 2008; Yusof & Abdullah, 2020).
Additional extrinsic products of interest could be taken from oaks; for
instance, the honeydew honeys that are secreted by different insects that
feed on plant sap. These honeys showed high content of flavonoids and
other phenolics as well as in vitro antioxidant properties (Flores, Escur­
edo, & Seijo, 2015; Jara-Palacios et al., 2019; Kocygit et al., 2019).
Besides, Viscum album, a mistletoe species that normally grows on
Q. robur trees have been evaluated as a source of bioactive extracts such
as the fermented mistletoe extract named as ‘Iscador Qu/ISC-Qu’ that
was even tested as a supportive treatment for oncologic patients
(Stauder, Matthes, Friedel, & Bock, 2009).
As it can be noticed in Table 2, phenolic compounds (including fla­
vonoids and tannins) have been placed as the main research target and
just a few studies have gone beyond these molecules focusing on
bioactive lipids such as triterpenoids and jasmonates as well as poly­
saccharides or polymers such as suberin (Ahmadi, Sheikh-Zeinoddin,
Soleimanian-Zad, Alihosseini, & Yadav, 2019; Huang, Wang, Li, Wang,
& He, 2016; Krizkova et al., 1999; Mai, Wang, Wang, Xu, & He, 2020;
Wang et al., 2020). The phenolic compounds from oaks were extracted
with conventional techniques that involve utilising grinding and
solid-liquid extractions (SLE) of Quercus materials using solvents such as
water, ethanol and methanol (Amessis-Ouchemoukh et al., 2017; Anlas
et al., 2019; Skrypnik et al., 2019). In some cases, conventional SLEs
were replaced by Soxhlet extractions (Custodio et al., 2015; Gezici &
Sekeroglu, 2019; Kheirandish et al., 2016) and only Cetera, Russo,
Milella, and Todaro (2019) carried out ultrasound-assisted extractions
that were also applied for polysaccharide recovery by Tahmouzi (2014).
Same solvents and others with low polarity (hexane, chloroform) were
utilised to extract or fractionate triterpenoids and other lipids (Makh­
louf, Squeo, Barkat, Trani, & Caponio, 2018; Xu, Cao, Yue, Zhang, &
Zhao, 2018; Zehra et al., 2019). Nevertheless, this review evidenced the
lack of studies investigating the use of advanced extraction technologies
The obtention and evaluation of vegetal extracts, fractions and iso­
lated compounds have become in a relevant approach to obtain dietary
antioxidants that are potentially capable of avoiding the imbalance be­
tween free radicals production and antioxidant mechanisms, commonly
known as ‘oxidative stress’ (Morales, 2020; Wen, Zhang, Zhang, Duan, &
Ma, 2020). Different parts of oaks were investigated through several
works obtaining fractions that exerted in vitro or/and in vivo antioxidant
effects that were frequently correlated with high levels of phenolic
compounds. For instance, different parts of Quercus coccifera acorns
(shells, cups, inner parts) were extracted with ethanol and water and
they showed interesting in vitro antioxidant activities (Gezici & Seker­
oglu, 2019) as well as methanolic fractions that were obtained from stem
parts of these trees that showed high antioxidant capacity and high total
phenolics content (Anlas et al., 2019). Moreover, Pinto et al. (2019)
prepared a coffee-like beverage using Quercus cerris seeds kernels that
was capable of scavenging free radicals in vitro due to its content in
ellagic acid, gallotannins and elagitannins derivatives. The wood of this
tree was also subjected to a thermal treatment followed to maceration,
ultrasound-assisted or accelerated-solvent extraction using water
(Cetera et al., 2019). High contents of total flavonoids and total phenolic
compounds were recorded leading to potent in vitro antioxidant capacity
(Cetera et al., 2019), also noticed for extracts obtained from Q. robur
bark and Q. suber cork (Drozdz & Pyrzynska, 2019; Santos, Pinto, Sil­
vestre, & Neto, 2010; Skrypnik et al., 2019). Besides, Q. infectoria galls,
widely utilised because of their high tannin content, gave rise to the
obtention of antioxidant aqueous and ethanolic fractions and Q. ilex
roots were macerated with ethanol to generate an extract able to scav­
enge reactive oxygen species (ROS) but also to display a protective effect
against DNA oxidation induced by hydrogen peroxide and UV irradia­
tion and to avoid erythrocytes haemolysis caused by ROS (Meziti et al.,
2019).
Other promising in vitro results were reported for phenolics-rich
honeydews honeys obtained from Q. ilex and Quercus pyrenaica trees,
able to scavenge free radicals but also to inhibit lipid peroxidation

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D. Morales Trends in Food Science & Technology 109 (2021) 116–125

Table 2
Biological activities of oaks extracts evaluated in different experimental models.
Biological activity Quercus spp. Raw Bioactive compound Experimental model Extraction solvent/method Reference
material (s)

Antioxidant Q. brantii Acorns – Wistar rats Water Dogan et al. (2015)

Polysaccharides Radical scavenging Hot water (after grinding Ahmadi et al. (2019)
assays and deffating)
Leaves Ethanol and water/ Tahmouzi (2014)
Ultrasound-assisted
extraction
Q. castaneifolia – Radical scavenging 80% ethanol Paray et al. (2020)
assays, common carp
juveniles
Q. cerris Seeds Ellagic acid, Radical scavenging Hot water Pinto et al. (2019)
gallotannin/ assays
ellagitannin
derivatives
Wood Phenolic compounds 70% ethanol (after thermal Cetera et al. (2019)
(flavonoids) treatment, grinding and
deffating)/Ultrasound-
assisted extraction,
pressurized liquid
extraction
Q. cerris var cerris, Bark Phenolic compounds Methanol and water Sohretoglu et al. (2012)
Q. macranthera
subsp. syspirensis, Q.
aucheri
Q. coccifera Acorns Phenolic compounds 70% ethanol and water Gezici and Sekeroglu
(flavonoids) (2019)
Stems Phenolic compounds Methanol Anlas et al. (2019)
Q. convallata, Q. Leaves Radical scavenging Hot water Gamboa-Gomez et al.
arizonica assays, C57BL/6 mice (2017)
Q. dilatata Aerial parts Radical scavenging Ethanol + ethyl acetate; Kazmi et al. (2018)
assays, Sprague Dawley water + acetone
rats
Q. gilva Leaves Radical scavenging Methanol Indrianingsih et al. (2015)
Q. ilex Roots assays 80% methanol Meziti et al. (2019)
Acorns Ethanol Amessis-Ouchemoukh
et al. (2017)
Q. ilex, Q. pyrenaica Honeydew – Jara-Palacios et al. (2019)
honey
Q. ilex, Q. suber, Q. Acorns (oil) Phenolic compounds, Hexane/Soxhlet Makhlouf et al. (2018)
coccifera carotenoids,
tocopherols
Q. infectoria Fruit hulls Phenolic compounds Methanol/Soxhlet Kheirandish et al. (2016)
(flavonoids)
Galls Phenolic compounds Ethanol Kaur et al. (2008)
Tannins Hot water Arina and Harisun (2019)
Q. liaotungensis Acorns Galloyl triterpenes Radical scavenging and 70% ethanol + Resin Xu et al. (2018b)
ferrous ion chelating purification
assays
Q. petraea Bark Ellagitanins Wistar rats 70% ethanol (brandy) Panchal and Brown (2013)
Q. phyllyraeoides Leaves Tannins and Radical scavenging Methanol Indrianingsih and
β-sitosterol glucoside assays Tachibana (2016)
Q. pyrenaica Honeydew Phenolic compounds – Kocyigit et al. (2019)
honey Phenolic compounds Flores et al. (2015)
(flavonoids)
Q. resinosa Leaves – Hot water Gallegos-Infante et al.
(2013)
Phenolic compounds Rocha-Guzman et al.
(2009)
Q. resinosa, Q. Leaves THP-1 cells Vazquez-Cabral et al.
arizonica, Q. (2017)
convallata
Q. resinosa, Q. laeta, – Radical scavenging Water; hexane Sanchez-Burgos et al.
Q. grisea, Q. obtusata assays (2013)
Q. robur Acorns Phenolic compounds Pig fat Hot water Rakic, Povrenovic,
Tesevic, Simic, and Maletic
(2006)
Bark Radical scavenging 60% ethanol Drozdz and Pyrzynska
assays (2019)
Hot water Skrypkink et al. (2019)
Q. sideroxyla, Q. Leaves – Moreno-Jimenez et al.
durifolia, Q. eduardii (2015)
Q. suber Cork Methanol; 80% methanol Santos et al. (2010)
Leaves and Phenolic compounds Methanol; hexane/Soxhlet Custodio et al. (2015)
acorns
(continued on next page)

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D. Morales Trends in Food Science & Technology 109 (2021) 116–125

Table 2 (continued )
Biological activity Quercus spp. Raw Bioactive compound Experimental model Extraction solvent/method Reference
material (s)

Anti-inflammatory/ Q. castaneifolia Leaves – Common carp juveniles 80% ethanol Paray et al. (2020)
Immune-modulatory (plasma lysozyme
activity)
Q. coccifera Stems Phenolic compounds RAW 264.7 cells Methanol Anlas et al. (2019)
Q. ilex Acorns Xanthine oxidase Ethanol Amessis-Ouchemoukh
inhibition assay et al. (2017)
Q. infectoria Bark – Sprague Dawley rats Methanol Khouzami et al. (2009)
Nutgalls C75BL/6 mice 95% ethanol Chokpaisarn et al. (2020)
Q. petraea Bark Ellagitanins Wistar rats 70% ethanol (brandy) Panchal and Brown (2013)
Q. resinosa, Q. Leaves Phenolic compounds THP-1 cells Hot water Vazquez-Cabral et al.
arizonica, Q. (2017)
convallata
Q. robur Bark Tannins RBL-2H3 cells, HMC-1 Lorenz et al. (2016)
cells
Q. serrata Acorns Triterpenoids BV-2 microglia cells 95% ethanol Mai et al. (2020)
Q. serrata var Jasmonates 70% ethanol Wang et al. (2020)
brevipetiolata Triterpenes RAW 264.7 and MH7A Huang et al. (2016)
cells
Q. sideroxyla, Q. Leaves – HT-29 cells Hot water Moreno-Jimenez et al.
durifolia, Q. eduardii (2015)
Antitumoral/ Q. cerris var cerris, Bark Phenolic compounds Hep2 cells Methanol; water Sohretoglu et al. (2012)
antiproliferative Q. macranthera
subsp. syspirensis, Q.
aucheri
Q. coccifera Acorns – A549, MCF-7 and HeLa Ethanol Gezici (2019)
cells
Q. ilex Phenolic compounds U87 cells Amessis-Ouchemoukh
et al. (2017)
Q. incana Leaves Betulin NCI–H460 cells Methanol; chloroform Zehra et al. (2019)
Q. infectoria Acorn – HUVEC cells 50% ethanol Yarani, Mansouri,
shells Mohammadi-Motlagh,
Mahnam, & Aleagha
(2013)
Galls HeLa cells, Sprague Hot water Hazwani et al. (2018)
Dawley rats
Phenolic compounds DBTR-05MG cells Methanol; 70% methanol Cheng et al. (2018)
Phenolic compounds HeLa, MDA-MB-231 and Hexane; ethyl acetate; Yusof and Abdullah (2020)
(flavonoids, tannins), HepG2 cells methanol
saponins, terpenoids,
alkaloids, glycosides
Q. liaotungensis Acorns Galloyl triterpenes t-HSC/Cl-6 cells 70% Ethanol + Resin Xu et al. (2018b)
purification
Q. pyrenaica Honeydew Phenolic compounds AGS cells – Kocyigit et al. (2019)
honey
Q. resinosa Leaves HeLa cells Hot water Rocha-Guzman et al.
(2009)
Q. resinosa, Q. laeta, Leaves – Saccharomyces cerevisiae Water; hexane Sanchez-Burgos et al.
Q. grisea, Q. obtusata mutant strains (2013)
(antitopoisomerase
assay)
Q. robur – L1210 cells Methylene chloride; Goun et al. (2002)
ethanol
Mistletoe Clinical trial Fermentation Stauder et al. (2009)
Wood Triterpenoids PC3 and MCF-7 cells 70% methanol Perez et al. (2017)
Q. sideroxyla, Q. Leaves – Sprague-Dawley rats Hot water Moreno-Jimenez et al.
durifolia, Q. eduardii (2015)
Q. suber Cork HL-60 cells Dichloromethane/Soxhlet Bejarano et al. (2015)
DNA-protective Q. ilex Roots Phenolic compounds H202 photolysis 80% methanol Meziti et al. (2019)
Q. suber Cork – L929 fibroblasts 50% ethanol Araujo et al. (2015)
Suberin Euglena gracilis Alkaline methanolysis Krizkova et al. (1999)
Antidiabetic/ Q. brantii Acorns – Wistar rats Water Dogan et al. (2015)
Hypoglycaemic Q. coccifera Bark Chlorocatechin and α-glucosidase inhibition Methanol + Sari et al. (2019)
polydatin assay Chromatographic
purification
Q. convallata, Q. Leaves Phenolic compounds α-amylase, α-glycosidase Hot water Gamboa-Gomez et al.
arizonica and glucose diffusion (2017)
assays, C57BL/6 mice
Q. dilatata Fruits – Albino rats Methanol Shaheen et al. (2017)
Q. gilva Leaves Phenolic compounds α-glucosidase inhibition Indrianingsih et al. (2015)
assay
Q. mongolica Acorns Triterpenoids α-glucosidase, α-amylase 75% ethanol + Xu, Cao, Yue, Zhang, and
and protein-tyrosine Fractionation Zhao (2018)
phosphatase 1B
inhibition assays
Q. phyllyraeoides Leaves Methanol + Fractionation
(continued on next page)

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Table 2 (continued )
Biological activity Quercus spp. Raw Bioactive compound Experimental model Extraction solvent/method Reference
material (s)

Tannins and α-glucosidase inhibition Indrianingsih and

β-sitosterol glucoside assay Tachibana (2016)
Q. suber Leaves and Phenolic compounds AChE and BuChe Methanol; hexane Custodio et al. (2015)
acorns inhibition assays
Hypocholesterolemic Q. dilatata Fruits – Albino rats Methanol Shaheen et al. (2017)
Q. infectoria – HMGCoA reductase Gholamhoeisinian et al.
activity assay (2010)
Bark Sprague Dawley rats Khouzami et al. (2009)
Antihypertensive Q. infectoria Bark – ACE inhibition assay 80% methanol Sharifi et al. (2013)
Q. petraea Ellagitannins Wistar rats 70% ethanol (brandy) Panchal and Brown (2013)
Antimicrobial Q. brantii Fruits Tannins Clinical trial 80% ethanol Zare et al. (2019)
Galls – 70% methanol Afzali et al. (2020)
Leaves Polysaccharides Microbiological analyses Ethanol and water/ Tahmouzi (2014)
Ultrasound-assisted
extraction
Seed coats – Water Sharifi et al. (2019)
Q. castaneifolia Fruits Hot water; ethanol Bahador and Baserisalehi
(2011)
Leaves 80% ethanol Paray et al. (2020)
Q. coccifera Stems Phenolic compounds Methanol Anlas et al. (2019)
Q. crassifolia Bark Hot water Valencia-Aviles et al.
(2019)
Q. ilex – Methanol/Soxhlet + Berahou et al. (2007)
fractionation
Q. infectoria Methanol Khouzami et al. (2009)
Galls 70% ethanol Tayel et al. (2018)
Fruit hulls Phenolic compounds BALB/c macrophages (in Methanol/Soxhlet Kheirandish et al. (2016)
(flavonoids) vitro), BALB/c mice
Q. resinosa, Q. laeta, Leaves – Microbiological analyses Water; hexane Sanchez-Burgos et al.
Q. grisea, Q. obtusata (2013)
Q. suber – Phenolic compounds 70% acetone Akroum (2017)
Antithrombotic Q. robur – – Antithrombin bioassay Methylene chloride; Goun et al. (2002)
ethanol
Antihaemolytic Q. ilex Roots Phenolic compounds Mice erythrocytes 80% methanol Meziti et al. (2019)
Neuroprotective Q. suber Leaves and AChE and BuChe Methanol; hexane/Soxhlet Custodio et al. (2015)
acorns inhibition assays
Q. coccifera Acorns Phenolic compounds 70% ethanol and water Gezici and Sekeroglu
(flavonoids) (2019)
Anti-tyrosinase Q. coccifera Bark Chlorocatechin and Tyrosinase inhibition Methanol + Sari et al. (2019)
polydatin assay Chromatographic
purification
Anticalpain Q. ilex Acorns Phenolic compounds U87 cells Ethanol Amessis-Ouchemoukh
et al. (2017)
Gastroprotective Q. resinosa, Q. laeta, Leaves – KATO III cells Water; hexane Sanchez-Burgos et al.
Q. grisea, Q. obtusata (2013)
Anti-ulcerogenic Q. infectoria Bark – Sprague Dawley rats Methanol Khouzami et al. (2009)
Hepatoprotective Q. dilatata Aerial parts Phenolic compounds Ethanol + ethyl acetate; Kazmi et al. (2018)
water + acetone
Prebiotic Q. brantii Acorns Polysaccharides Lactobacillus plantarum Hot water (after grinding Ahmadi et al. (2019)
and deffating)
Wound healing Q. coccifera Stems Phenolic compounds 3T3 cells Methanol Anlas et al. (2019)
Q. infectoria Nutgalls – Clinical trial 95% ethanol Chokpaisarn et al. (2020)
Milk quality enhancer Q. brantii Acorns Tannins Kurdish goats – Alipanahi et al. (2019)
Q. infectoria Galls – Awassi ewes Barwary et al. (2019)
Microbiome- Q. mongolica Seeds Tannins Siberian chipmunks- Yi et al. (2020)
modulatory Zhang et al. (2020)
Q. robur Ellagitannins Human faeces Water Piwowarski et al. (2014)

(Flores et al., 2015; Jara-Palacios et al., 2019; Kocyigit et al., 2019).
Apart from phenolic compounds, other interesting molecules were
extracted from oaks products. In this sense, Soxhlet method using hex­
ane was applied to Q. ilex, Q. suber and Q. coccifera acorns to obtain oils
with high levels of tocopherols and carotenoids with potent in vitro
antioxidant abilities (Makhlouf et al., 2018), while Quercus liaotungensis
acorns were subjected to ethanol extraction and subsequent resin puri­
fication to recover fractions with high content of antioxidant galloyl
triterpenes (Gallegos-Infante et al., 2013). This antioxidant capacity was
also noticed for β-sitosterol-D-glucoside isolated from a methanolic
extract of Quercus phillyraeoides leaves (Indrianingsih & Tachibana,
2016) and for polysaccharide-rich fractions recovered after hot water
extraction and ethanol precipitation of defatted Q. brantii acorn flours
and ultrasound-assisted extraction of Q. brantii leaves (Ahmadi et al.,
2019; Tahmouzi, 2014).
The significant in vitro results were validated in many cases by a
significant quantity of in vivo studies. Q. brantii aqueous extracts were
lyophilized and included in the diet of diabetic Wistar rats causing a
reduction in the levels of biomarkers linked to diabetes and liver damage
(Dogan, Celik, & Kaya, 2015). Quercus dilatata aerial parts extracts were
obtained with mixed solvents (ethanol + ethyl acetate; water + acetone)
and they were able to restore levels of endogenous antioxidants (su­
peroxide dismutase, catalase, glutathione peroxidase) levels after hep­
atotoxicity induction in Sprague-Dawley rats (Kazmi et al., 2018).
Kheirandish et al. (2016) prepared infusions and fermented beverages
using Q. convallata and Q. arizonica leaves and observed marked anti­
oxidant effects in C57BL/6 mice. Panchal and Brown (2013) settled
brandy (70% ethanol) in contact with Quercus petraea bark at room

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D. Morales
temperature for 6 months. After dealcoholisation of the ellagitannin-rich
product, the authors observed a marked antioxidant response in Wistar
rats confirmed by a decrease in the oxidation stress markers. Further­
more, ethanolic extracts of Quercus castaneifolia leaves increased anti­
oxidant enzymes levels when they were dietary supplemented to
common carps (Paray, Hoseini, Hoseinifar, & Doan, 2020).
Despite the great number of articles that evidenced antioxidant ac­
tivities of oaks fractions using in vitro and animal models, no clinical
trials were found in the scientific literature.
3. Anti-inflammatory/immune-modulatory activity
During the last decades, vegetal edible materials have been tested as
bioactive food products able to inhibit inflammatory responses and to
exert additional immune-modulatory actions inducing changes in im­
mune system activity (Gandhi et al., 2020; Tasneem, Liu, Li, Choudhary,
& Wang, 2019). Among oaks immune-modulatory compounds, tannins
and other phenolic compounds have been evaluated in vitro and in vivo
but also some specific bioactive lipids.
Regarding this, phenolic-rich methanolic extracts obtained from
Q. coccifera stems were added to murine RAW 264.7 macrophages and
demonstrated to be capable of inhibiting lipopolysaccharide (LPS)-
induced secretion of TNF-α (Anlas et al., 2019). The same effect was
observed when infusions and kombucha beverages prepared with
different oaks leaves were applied to THP-1 cells, also decreasing IL-6
secretion (Vazquez-Cabral et al., 2017) while Quercus syderoxyla
leaves infusions downregulated NF-κβ, COX-2, COX-1 and IL-8 expres­
sion (Moreno-Jimenez et al., 2015). Moreover, tannins were extracted
with hot water from Q. robur bark and showed an inhibitory action
against basophils and mast cells degranulation decreasing the produc­
tion of IL-8, IL-6 and TNF-α production in RBL-243 rat basophils and
HMC-1 human mast cells (Lorenz et al., 2016). Another phenolic-rich
fraction obtained subjecting Q. ilex to ethanolic extraction displayed
the ability to inhibit xanthine oxidase activity that is involved in uric
acid production. This extract was even more effective than allopurinol, a
drug which is clinically used for gout treatment (Amessis-Ouchemoukh
et al., 2017).
Among the studied immune-modulatory lipids, triperpenes, tri­
terpenoids and jasmonates were extracted from Quercus serrata acorns
with ethanol:water mixtures. These extracts were supplemented to RAW
264.7, MH7A and BV-2 microglia cells and reduced LPS-induced NO and
pro-inflammatory cytokines production and blocked NF-κβ traslocation
(Huang et al., 2016; Mai et al., 2020; Wang et al., 2020).
In vitro studies were supported by animal experiments. For instance,
Paray et al. (2020) administered a Q. castaneifolia ethanolic extract to
common carps, noticing immune-system stimulatory responses since
plasma lysozyme activity was significantly increased. Q. infectoria was
also utilised in animal studies: bark was extracted with methanol and
this fraction reduced paw thickness caused by oedema in Sprague
Dawley rats that were used as models of acute and chronic inflammation
(induced by carrageenan and formalin, respectively) (Khouzami,
Mroueh, & Daher, 2009); nutgalls were extracted with 95% ethanol and
diabetic C75BL/6 mice fed the resulting fraction showing reduced
macrophage secretion of IL-1β and TNF-α (Chokpaisarn et al., 2017).
Moreover, the ellagitannin-rich Q. petraea product that was described in
the previous section, diminished plasma C-reactive protein levels, NF-κβ
expression, collagen deposition and infiltration of inflammatory cells in
Wistar rat hearts. These events seemed to be related with a decrease in
ventricular stiffness enhancing ventricular function (Panchal & Brown,
2013).
4. Antitumoral/antiproliferative activity
The search of novel and potent antitumoral agents is one of the main
goals of medical research since cancer is still the second leading cause of
death, only surpassed by cardiovascular diseases (Ahmadi &
121
Trends in Food Science & Technology 109 (2021) 116–125
Ebrahimzadeh, 2020). The existence of antiproliferative compounds in
different edible sources places food materials as an interesting target,
including vegetal matrices and particularly oaks products (Gezici, 2019;
Rayan, Raiyn, & Falah, 2017). Plenty of fractions obtained from
different oaks parts have demonstrated in vitro cytotoxicity against tu­
moral cell lines: Quercus macranthera subsp. syspirensis bark methanolic
and aqueous extracts against Hep2 cells (human larynx epidermoid
carcinoma) (Sohretoglu, Sabuncuoglu, & Harput, 2012); Q. coccifera
acorns ethanolic extracts against A549 (human lung carcinoma), MCF-7
(human breast cancer) and HeLa cells (human cervix carcinoma)
(Gezici, 2019); Q. ilex acorns ethanolic extract against U87 cells (human
glioblastoma) (Amessis-Ouchemoukh et al., 2017); Q. pyrenaica honey
honeydew (human gastric adenocarcinoma) (Kocyigit et al., 2019);
Q. resinosa leaves aqueous extracts against HeLa cells (Rocha-Guzman
et al., 2009).
In addition, Q. infectoria galls were used in different studies to obtain
antiproliferative extracts. Yusof and Abdullah (2020) recently extracted
flavonoids- and tannins-rich fractions using different solvents (hexane,
ethyl acetate, methanol) with cytotoxic activity against HeLa,
MDA-MB-231 (human breast cancer) and HepG2 cells (human liver
cancer). Other methanolic extracts with high levels of phenolic com­
pounds also exerted antiproliferative effects against DBTR-05MG cells
(human glioblastoma) (Cheng et al., 2018). Moreover, a vaginal cream
based on an aqueous extract showed cytotoxic effects against HeLa cells
and the authors also demonstrated that the formulation did not cause
adverse effects and it was not toxic when applied to Sprague Dawley
female rats (Hazwani, Hasmah, & Nizam, 2018).
Q. robur wood was utilised to obtain triterpenoids with high cyto­
toxicity against PCR3 (human prostate adenocarcinoma) and MCF-7
cells (Perez et al., 2017) and Q. suber cork extracts were also toxic to
HL-60 cells (human leukaemia) (Bejarano, Godoy-Cancho, Franco,
Martinez-Cañas, & Tormo, 2015).
Only one in vivo assay could be found in the scientific literature
regarding antitumoral activities of oaks extracts. Q. syderoxyla infusions
were administered to Sprague Dawley rats with induced colon cancer
and they were able to reduce tumor burden and multiplicity as well as
β-catenin level in adenocarcinomas (Moreno-Jimenez et al., 2015).
Besides, no material belonging to oaks have been tested in clinical
trials regarding anticancer capacity. However, as it was commented in
the Introduction section, Viscum album is a mistletoe species that grows
on Q. robur trees and it led to the generation of a fermented extract called
‘ISC-Qu’. This extract was utilised as a supportive treatment in a clinical
study carried out in 270 patients with pancreatic cancer and it succeed
in significantly prolonged overall survival (Stauder et al., 2009).
5. Antidiabetic/hypoglycaemic activity
Although the mainly used therapies for diabetic patients involved
drugs such as metformin or insulin, anti-hyperglycaemic dietary in­
gredients can become a relevant alternative for moderate clinical cases.
The use of functional foods might lead to avoid drug-related disadvan­
tages such as the doses administration and adverse effects (Harnedy
et al., 2018).
Quercus extracts have been tested in vitro and in vivo to prevent or
correct the complications related to diabetes. This is the case, for
instance, with methanolic extracts obtained from Quercus gilva leaves:
their high in vitro α-glucosidase inhibitory capacity correlated with high
phenolic compounds levels (Indrianingsih, Tachibana, Dewi, & Itoh,
2015). Also Quercus mongolica acorns were extracted with 75% ethanol,
later fractionated using different solvents (petroleum ether,
ethyl-acetate and n-butyl alcohol) and the resulting fractions and
bioactive triterpenoids were finally isolated exerting in vitro ability of
inhibiting α-glucosidase, α-amylase and protein-tyrosine phosphatase
1B activities (an enzyme that plays a key role in insulin signaling) (Xu,
Cao, et al., 2018). Q. phyllyraeoides leaves were extracted with methanol
and partitioned with different solvents (hexane, chloroform, ethyl
D. Morales
acetate and methanol) to finally isolate tannins and β-sitosterol gluco­
side that exerted α-glucosidase inhibitory in vitro (Indrianingsih &
Tachibana, 2016), as well as several compounds (cholorocatechin, pol­
ydatin) that were isolated from methanolic Q. coccifera bark extracts
(Sari, Barut, Ozel, Kuruuzum-Uz, & Sohretoglu, 2019). This activity was
also recorded for Q. convallata and Q. arizonica infusions and kombucha
beverages that even validated these antidiabetic effects in C57BL/6 mice
decreasing glucose levels and TyG index as hyperglycaemia and insulin
resistance markers (Gamboa-Gomez et al., 2017).
Other studies were carried out in animal models: diabetic Wistar rats
fed a lyophilized extract from Q. brantii acorns and they showed a
marked reduction in diabetes biomarkers and α-glucosidase activity
(Dogan et al., 2015); Q. dilatata fruits were extracted with methanol and
diabetic Albino rats were administered with the extracts leading to
reduced serum glucose levels (Shaheen, Khan, Ahmed, Mustaq, & Khan,
2017); methanol and hexane were used to extract Q. suber leaves and
acorns and the obtained fractions inhibited acetyl- and
butyl-cholinesterase activities that seemed to be linked to hyper­
glycaemia and related disorders (Custodio et al., 2015).
Up to now, no clinical trials regarding hypoglycaemic or antidiabetic
effects of oak extracts have been published.
6. Hypocholesterolemic/antihypertensive activity
Since cardiovascular diseases are the first cause of mortality world­
wide, the reduction of risk factors is a highly relevant goal. Among these
factors, high serum levels of total cholesterol and LDL-cholesterol as well
as elevated blood pressure are parameters that must be particularly
considered. Therefore, molecules with the ability of reducing cholesterol
absorption/biosynthesis or blood pressure levels have been investigated
through plenty of food matrices (Daskaya-Dikmen, Yucetepe,
Karbancioglu-Guler, Daskaya, & Ozcelik, 2017; Gil-Ramirez et al., 2017;
Morales et al., 2018 and 2019). Although these biological activities have
not been the most deeply studied in oaks, some studies were carried out
in this field.
For example, a Q. infectoria methanolic extract proved to be effective
as an in vitro inhibitor of the 3-hydroxy-3-methylglutaryl-coenzyme A
reductase (HMGCR), a crucial enzyme in cholesterol endogenous syn­
thesis (Gholamhoseinian, Shahouzehi, & Sharifi-Far, 2010). A similar
Q. infectoria fraction (obtained from bark) was not able to decrease total
and LDL-cholesterol levels in Sprague Dawley rats but it significantly
increased HDL-cholesterol values reducing the atherogenic index
(Khouzami et al., 2009). Moreover, diabetic albino rats fed the
Q. dilatata extract that was described in the previous section and it
succeed restoring normal lipemia by reducing total and LDL-cholesterol
as well as triglycerides levels while HDL-cholesterol levels were
increased (Shaheen, Khan, Ahmed, Mushtaq, & Khan, 2017).
Regarding antihypertensive activities, Q. infectoria bark extract
(obtained with 80% methanol) in vitro inhibited the angiotensin-
converting enzyme (ACE) action, a key enzyme in the renin-
angiotensin system that regulates blood pressure (Sharifi, Souri, Ziai,
Amin, & Amanlou, 2013). A decrease in blood pressure was confirmed in
vivo in Wistar rats that were supplemented with Q. petraea bark extracts
(Panchal & Brown, 2013).
No clinical trials can be found in the scientific literature reporting
hypocholesterolemic or hypotensive activities of oak extracts.
7. Antimicrobial activity
Oaks, as well as plenty of plants, constitute an interesting source of
natural antimicrobial compounds that can be utilised against pathogens
or non-desirable species in clinical or food preservation applications
(Sakkas & Papadopoulou, 2017; Sharifi, Azizi, Moradi-Choghakaboi,
Aghaeu, & Azizi, 2019; Tayel, El-Sefdy, Ibrahim, & Moussa, 2018).
The studies related with Quercus extracts have been mainly focused on
phenolics compounds, although other metabolites can be promising
122
Trends in Food Science & Technology 109 (2021) 116–125
candidates to be tested in the future.
Current evidence shows that different extracts are potent inhibitors
of microorganism growth in vitro. For instance, Q. castaneifolia extracts
exerted antimicrobial activity against Aeromonas hydrophila (leaves) or
Shigella dysenteriae, Escherichia coli, Salmonella typhimurium and Yersinia
enterocolitica (fruits) (Bahador & Baserisalehi, 2011; Paray et al., 2020).
Moreover, a polyphenolic fraction obtained from Q. crassifolia bark was
effective against Escherichia coli (Valencia-Aviles, Martinez-Flores, Gar­
cia-Perez, Melendez-Herrera, & Garcia-Perez, 2019) and an aqueous
extract recovered from Q. brantii seed coats displayed in vitro anti-­
Helycobacter pylori effects (Sharifi, Azizi, Moradi-Choghakabodi, Aghaei,
& Azizi, 2019). The latter oak variety was also subjected to
ultrasound-assisted extraction using ethanol and water and the obtained
product inhibited Salmonella typhi, Staphylococcus aureus, Candida albi­
cans and Penicillium citri growth (Tahmouzi, 2014). Other microorgan­
isms that were sensitive to oaks extracts were Proteus mirabilis and
Bacillius cereus, that were inhibited by phenolics-rich methanolic extract
from Q. coccifera stems (Anlas et al., 2019).
Q. infectoria parts were also studied regarding their antimicrobial
potential: bark methanolic extracts inhibited Citrobacter braakii and
Proteus mirabilis (Khouzami et al., 2009); galls hydroalcoholic extracts
decontaminated chicken eggs eliminating S. aureus, E. coli, Pseudomonas
aeruginosa, S. typhimurium and C. albicans (Tayel et al., 2018); fruit hulls
were subjected to Soxhlet extraction using methanol and the obtained
fraction exerted anti-Leishmania major activity in BALB/c mice (Kheir­
andish et al., 2016).
Furthermore, two clinical trials were published focused on the use of
vaginal creams against bacterial vaginosis based on Q. brantii extracts.
First, a tannin-rich ethanolic extract was recovered from fruits and used
in this cream combined with metronidazole tablets and this treatment
demonstrated to be more effective than the combination metronidazole
+ placebo (Zare et al., 2019). Later, a methanolic extract was obtained
from galls and the resulting cream led to similar results than a metro­
nidazole gel since both were able to eliminate symptoms in all patients
after one week (vaginal itching, burning sensation, bad odour, dysuria,
dyspareunia) (Afzali et al., 2020).
8. Other biological activities
Besides the previously described biological activities of Quercus
fractions, other not so deeply studied bioactivities were reported in
several works.
In this sense, a Q. robur methanolic extract showed promising in vitro
results inhibiting thrombin motivating further research related to
thrombosis treatment (Goun et al., 2002). Phenolics-rich fractions were
extracted from Q. ilex roots and they exerted an in vitro antihaemolytic
effect in mice erythrocytes (Meziti et al., 2019).
Moreover, Q. suber extracts with the ability to inhibit acetyl- and
butyl-cholinesterase activity were described in section 5 and this activity
was also reported to be linked to a neuroprotective effect since these
enzymes are involved in the hydrolysis of acetylcholine (Custodio et al.,
2015). This activity was also recorded for Q. coccifera acorns extracts
with high content of flavonoids (Gezici & Sekeroglu, 2019). The bark of
this tree was also utilised to isolate compounds such as chlorocatechin
and polydatin that exerted anti-tyrosinase activity that could be relevant
to reduce the risk of melanoma or Parkinson disease development (Sari
et al., 2019).
Furthermore, other claimed activities for oaks extracts were hep­
atoprotective, gastroprotective and anti-ulcerogenic (Kazmi et al., 2018;
Khouzami et al., 2009; Sanchez-Burgos et al., 2013). In fact,
wound-healing activity was observed in vitro (Anlas et al., 2019) and a
clinical trial was focused on the topical administration of a solution
based on Q. infectoria nutgalls extract for the treatment of chronic dia­
betic ulcers (Chokpaisarn et al., 2020). In addition, Q. brantii acorns
polysaccharides showed a prebiotic effect for Lactobacillus plantarum
(Ahmadi et al., 2019) and their tannins could even enhance goat milk
D. Morales
composition profile (Alipanahi et al., 2019). Significant interactions
were also noticed between Q. mongolica tannins and rodents gut
microbiome, being able to alter microbial populations in chipmunks (Yi
et al., 2020; Zhang et al., 2018). Moreover, ellagitannin-rich Q. robur
extracts were incubated with human faeces leading to the formation of
urolithins by tannin-degrading bacteria and the anti-inflammatory ef­
fects of these metabolites were confirmed in vitro applied to human
macrophages (Piwowarski et al., 2014).
9. Current limitations, conclusions and future perspectives
In this review, the scientific evidence around the biological activities
reported for Quercus spp. extracts was revised. The existence of prom­
ising results cannot be denied; however, both strengths and weaknesses
of its current state will be discussed in this section.
Regarding the nature of the bioactive compounds, a great effort was
carried out testing extracts with high contents of phenolic compounds
(particularly tannins and flavonoids). Nevertheless, although other
molecules such as polysaccharides or lipids (triterpenes, jasmonates,
etc.) were also evaluated by only a few works, further research targeting
other compounds such as bioactive peptides will be required.
The predominance of ‘phenolics-related studies’ led to a higher
number of projects focused on antioxidant capacity. Moreover, signifi­
cant research was carried out exploring antimicrobial, antiproliferative,
immune-modulatory and hypoglycaemic effects of oaks extracts. How­
ever, hypocholesterolemic and antihypertensive abilities, considered as
relevant capacities particularly to reduce cardiovascular disease risk,
have not so deeply analysed yet.
The potent bioactive effects that oak extracts demonstrated in vitro
constitute a powerful strength that was also confirmed utilising animal
models for most of the revised biological activities, although the number
of published in vivo studies is significantly lower than in vitro works.
Moreover, different oak-based products (from Q. brantii and
Q. infectoria) have been already tested in human trials. These products
were cream-like formulations or solutions that were topically applied
demonstrating effectiveness against vaginosis and exerting wound-
healing activities against diabetes ulcers (Afzali et al., 2020; Chokpai­
sarn et al., 2019; Zare et al., 2019). However, no food matrices including
oak ingredients (except for oak mistletoe) have been formulated or
clinically tested yet despite of the obtained results in vitro and in vivo.
The potential of these extracts motivates further research focused on
novel food design including the most active extracts. The development
of an oak-based alimentary product must include sensorial analysis and
consumers acceptance studies particularly if oak ingredients were not
obtained from usually consumed parts such as bark, roots or galls
instead of acorns or leaves.
The mentioned diversity of sources through Quercus genus also
represents an advantage: different species with varied characteristics as
well as different materials could be utilised.
Another gap that can be rapidly found in this field is the lack of
advanced extraction technologies in almost the totality of the works
since just two works applied pressurized liquids or ultrasound-assisted
extractions (Cetera et al., 2019; Tahmouzi, 2014). As it was com­
mented in the Introduction section, other techniques apart from pro­
cedures such as Soxhlet or other solid-liquid extractions are highly
recommended: supercritical fluids, microwave, pulsed electric
fields-assisted extractions, etc.
Moreover, the bioaccessibility and bioavailability of the bioactive
compounds, as well as their interactions with gut microbiome must be
investigated in future studies since these facts may be crucial for their
potential health benefits (Vamanu & Gatea, 2020).
In conclusion, further research is required including additional
compounds, bioactivities and extraction techniques evaluation as well
as more animal and clinical studies testing not only creams and solutions
but also food products incorporating oaks extracts and ingredients.
These advances might be useful to validate the promising results that
123
Trends in Food Science & Technology 109 (2021) 116–125
were included in this review.
Declaration of competing interest
None.
Acknowledgments
The author would like to thank EIT Food for the RIS Talent Fellow­
ship (EIT RIS Programmes).
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