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. 2021 Jun 18;12:695973. doi: 10.3389/fimmu.2021.695973

Table 1.

Extended summary of the current available literature on nutritional effects of Lamiaceae family and Allium sp. derived phytogenics upon fish mucosal immune response.

Phytogenic plant origin Supplemented form Inclusion dosage(s) tested Period of administration Main bioactive components (≤ 3) Fish species Performance Mucosal parameters evaluation Key benefits summary Reference
Lamiaceae family
Oregano (Origanum vulgare) Powder 0.5%,
1.0% and 2.0%
8 weeks N/I Zebrafish (Danio rerio) ↑ Final weight
↑ Weight gain
↓ FCR
↑ SGR
↑ Survival against A. hydrophila
↑ Skin mucus lysozyme activity
↑ Skin mucus alkaline phosphatase activity
↑ Skin mucus total Ig
↑ Skin mucus protease activity
↑ Skin mucus total protein
Beneficially affects the skin mucus immune parameters, growth performance and survival against pathogenic bacterial challenge Rashidian, Boldaji (56)
Oregano (Origanum vulgare) Powder 0.5% and
1.0%
15 and
30 days
N/I Gilthead seabream (Sparus aurata) No effect ↑ Skin mucus IgM
↑ Skin mucus bactericidal activity against P. damselae
Oregano improves humoral immunity and increases the antibacterial activity of skin mucus Beltrán, Gonzalez Silvera (57)
Oregano (Origanum vulgare) Ethanolic extract 0.2% and 0.5% 60 days + 7 days A. hydrophila challenge N/I Nile tilapia (Oreochromis niloticus) ↑ Final weight
↑ Weight gain
↓ FCR
↑ SGR
↑ Survival rate
↑ Survival against A. hydrophila
↑ Skin mucus total Ig Can effectively improve the fish growth, health, and immune status Mohammadi, Rafiee (58)
Oregano (Origanum vulgare) Powder 5.0, 10.0, 15.0 and 20.0 g kg-1 8 weeks Carvacrol and thymol
(Commercial product)
Common carp (Cyprinus carpio) ↑ Final weight
↑ Weight gain
↑ SGR
↑ Intestine villus height
↑ Intestine villus width
↑ Intestine crypt depth
Dose-dependent enhancement of intestinal morphometry, which subsequently lead to improvement of nutrients absorption Abdel-Latif, Abdel-Tawwab (59)
Oregano (Origanum vulgare) Essential oils 0.75, 1.5, 2.25 and 3.0 g kg-1 64 days Carvacrol, thymol and
p-cymene
Nile tilapia (Oreochromis niloticus) No effect ↑ Intestine villus height Increases intestinal villus size Heluy, Ramos (60)
Oregano (Origanum vulgare) Essential oils 0.5, 1.5 and 4.5 g kg-1 8 weeks + 7 days A. hydrophila challenge N/I Koi carp (Cyprinus carpio) ↑ Survival against A. hydrophila ↓ TNF-a and TGF-b gene expression in intestine
↑ Actinobacteria phylum, and Propionibacterium, Brevinema and Corynebacterium genera
↓ Bacteroidetes phylum and Vibrio genus
No effect on bacterial alpha diversity
Presents immunomodulatory effects and enhances disease resistance. Also beneficially alters the gut bacterial community composition of fish Zhang, Wang (61)
Oregano (Origanum vulgare) Powder 0.5% and 1.0% 30 days + cypermethrin exposure Carvacrol and thymol
(Commercial product)
Common carp (Cyprinus carpio) N/I ↓ Gill histopathologic lesions
↓ Gill proliferating cell nuclear antigen (PCNA) and caspase-3 immune positive cells
Protective roles against the adverse effects of cypermethrin, enhancing recovery from the exposure Khafaga, Naiel (62)
Oregano (Origanum vulgare) Essential oils 0.01%, 0.02%, 0.05% and 0.10% 24 days + 28 days I. salmonis and T. truttae challenge
(Total 52 days)
Carvacrol, p‐cymene and γ‐terpinene Chum salmon (Oncorhynchus keta) ↑ feed efficiency
I. salmonis infection
T. truttae infection
↓ cumulative mortality
Carvacrol content detected in the skin of fish fed the oregano supplemented diet Preventive effects against I. salmonis and T. truttae and suggests the possibility that its anti‐parasitic action is attributable to the bioactive component emergence through the skin Mizuno, Urawa (63)
Oregano (Origanum vulgare) Essential oils 0.5, 1.0, 1.5, 2.0
and 2.5 g kg-1
90 days N/I
(Commercial product)
Yellow
tail tetra (Astyanax altiparanae)
N/I ↑ Intestine villus length
↑ Intestine villus width
↑ Intestine absorption area
↑↓ Intestine goblet cells number
Promotes increased absorption surface area and modulates the number of goblet cells involved in protecting the intestinal mucosa Ferreira, Caldas (64)
Marjoram (Origanum majorana) Ethanolic extract 0.1, 0.2 and 0.4 g kg-1 60 days + 10 days A. hydrophila challenge N/I Common carp (Cyprinus carpio) ↑ Final weight
↑ Weight gain
↓ FCR
↑ SGR
↑ Survival against A. hydrophila
↑ Skin mucus alkaline phosphatase
↑ Skin mucus total Ig
↑ Skin mucus lysozyme activity
↑ Skin mucus alternative complement (ACH50) activity
Increase fish skin mucosal immunity and performance Yousefi, Ghafarifarsani (65)
Thyme (Thymus vulgaris) Essential oils 500 ppm 30 days + thiamethoxam exposure Thymol, p-cymene and γ-terpinene African catfish (Clarias garipenus) N/I ↓ Gill histopathologic lesions Mitigate the thiamethoxam induced toxicity El Euony, Elblehi (66)
Thyme (Thymus vulgaris) Aqueous extract 5.0, 10.0 and 20.0 g kg-1 2 weeks + oxytetracycline N/I Rainbow trout (Oncorhynchus mykiss) No effect ↑ Intestine antioxidant enzymes (SOD, CAT, GPx and GST) activity
↓ Intestine levels of the oxidative stress marker malondialdehyde
Mitigate adverse effects of oxytetracycline and improve the fish immune responses Hoseini and Yousefi (67)
Thyme (Thymus vulgaris) Essential oils 0.1%, 0.5%, and 1% 15 days Thymol, o-cymene and carvacrol Nile tilapia (Oreochromis niloticus) N/I No effect upon the population of beneficial Bacillus bacteria in the gut Stimulated the cellular components of the non-specific immune response without deleterious effects on the general health of the fish or the intestinal tract Valladão, Gallani (68)
Thyme (Thymus vulgaris) Essential oils 0.005, 0.010 and 0.02  g kg-1 5 weeks Thymol, p‐cymene and
linalool
Rainbow trout (Oncorhynchus mykiss) No effect No effect upon the allochthonous microbiota profile No toxic effects do not significantly alter the intestinal contents bacterial populations Navarrete, Toledo (69)
Spanish thyme (Thymus zygis subsp. gracilis) Essential oils 0.001, 0.002, 0.003 and 0.004 g kg-1 12 weeks Thymol, p‐cymene and carvacrol Gilthead seabream (Sparus aurata) No effect ↑ Anterior intestine lymphocyte aggregates in the lamina propria at low dose
↓ Anterior intestine lymphocyte aggregates in the lamina propria at high doses
Dose-dependent immuno‐modulatory effect upon the intestine Hernandez, Garcia (70)
Lemon balm (Melissa officinalis) Ethanolic extract 0.2% and 0.5% 60 days + 7 days A. hydrophila challenge N/I Nile tilapia (Oreochromis niloticus) ↑ Final weight
↑ Weight gain
↓ FCR
↑ SGR
↑ Survival rate
↑ Survival against A. hydrophila
↑ Skin mucus total Ig
↑ Skin mucus lysozyme activity
↑ Skin mucus protease activity
↑ Skin mucus alternative complement (ACH50) activity
Can effectively improve the fish growth, health, and immune status Mohammadi, Rafiee (58)
Peppermint (Mentha piperita) Powder 2.0, 3.0, and 4.0 g kg-1 8 weeks N/I Caspian roach (Rutilus caspicus) ↑ Final weight
↑ Weight gain
↓ FCR
↑ SGR
↓ Daily intake rate
↑ Secretion of skin mucosal protein pattern bands; higher lysozyme band intensity in particular
↑ Skin mucus lysozyme activity
↑ Skin mucus alkaline phosphatase activity
↑ Skin mucus soluble protein
Act as a growth promoter and immunostimulant Paknejad, Hosseini Shekarabi (71)
Peppermint (Mentha piperita) Essential oils 0.1 and 0.25 g kg-1 7, 14, 30 and 60 days Menthol, mentone and 1,8-cineole Nile tilapia (Oreochromis niloticus) N/I ↑ Intestine intraepithelial lymphocytes Show benefits in terms of intestinal health and on immune parameters Valladão, Gallani (72)
Peppermint (Mentha piperita) Ethanolic extract 1.0%, 2.0% and 3.0% 8 weeks N/I Rainbow trout (Onchorhynchus mykiss) ↑ Survival against A. hydrophila ↑ Skin mucus antibacterial activity against S. iniae, Y. ruckeri, A. hydrophila and L. garviea Triggers the immune system of rainbow trout against Y. ruckeri Adel, Pourgholam (73)
Peppermint (Mentha piperita) Ethanolic extract 1.0%, 2.0% and 3.0% 56 days N/I Caspian kutum roach (Rutilus frisii kutum) ↑ Weight gain
↓ FCR
↑ SGR
↑ Skin mucus antibacterial activity against S. iniae, Y. ruckeri, L. monocytogenes and E. coli
↑ Skin mucus protein level
↑ Skin mucus alkaline phosphatase activity
Increases the mucosal immune parameters and performance of fry in a dose dependent manner Adel, Amiri (74)
Peppermint (Mentha piperita) Ethanolic extract 1.0%, 2.0% and 3.0% 8 weeks N/I Caspian brown trout (Salmo trutta caspius) ↑ Weight gain
↓ FCR
↑ SGR
↑ Skin mucus protein level
↑ Skin mucus lysozyme activity
↑ Skin mucus alkaline phosphatase activity
Promote growth performance and have immunostimulant properties Adel, Safari (75)
Horsemint (Mentha longifolia) Ethanolic extract 2.0%, 4.0% and 6.0% 8 weeks + Y. ruckeri challenge N/I Caspian kutum roach (Rutilus frisii kutum) ↑ Weight gain
↓ FCR
↑ SGR
↑ Survival rate
↑ Skin mucus protein level
↑ Skin mucus lysozyme activity
↑ Skin mucus alkaline phosphatase activity
↑ Skin mucus protease activity
↑ Skin mucus esterase activity
↑ Skin mucus antibacterial activity against S. iniae, Y. ruckeri, A. hydrophila and L. garvieae
Improve growth performance and boost fish immune response in a dose‐related manner Gholamhosseini, Adel (76)
Horsemint (Mentha longifolia) Hydroalcoholic extract 0.1%, 0.2% and 0.3% 4 weeks + 10 days Y. ruckeri challenge N/I Rainbow trout (Onchorhynchus mykiss) ↑ Survival against Y. ruckeri ↑ Secretion of skin mucosal protein pattern bands; higher lysozyme band intensity in particular Dose‐related positive effect on immunogenicity and increased resistance to bacterial disease Heydari, Firouzbakhsh (77)
Thumbai (Leucas aspera) Powder 1.0, 2.0, 4.0 and
8.0 g kg-1
45 days + 15 days S. agalactiae challenge N/I Nile tilapia (Oreochromis niloticus) ↑ Weight gain
↓ FCR
↑ SGR
↑ Survival against S. agalactiae
↑ Skin mucus lysozyme activity
↑ Skin mucus peroxidase activity
Increase skin mucosal immune parameters, performance and survival against bacterial infection Kurian, Van Doan (78)
Shirazi thyme (Zataria multiflora) Hydroalcoholic extract 2.0 g kg-1 56 days Thymol and carvacrol? (N/I) Rainbow trout (Oncorhynchus mykiss) ↑ Survival rate ↑ Skin mucus bactericidal activity against A. hydrophila
↑ Skin mucus lysozyme activity
Increase skin mucosal immunity Mirghaed, Hoseini (79)
Shirazi thyme (Zataria multiflora)
+ Rosemary (Rosmarinus officinalis)
Powder (1:1)
+ aflatoxin B1
40 g kg-1
(20 g kg-1 each)
12 weeks N/I Common carp (Cyprinus carpio) No effect No effect Do not prevent intestinal tissue lesions induced by aflatoxin B1 Tasa, Imani (80)
Rosemary (Rosmarinus officinalis) Aqueous extract 10, 20, 40, 80 and 100 ml/100 g 20 days 1,8-Cineole Common carp (Cyprinus carpio) N/I ↑ Skin mucus level of 1,8-Cineole dose-dependent
No effect upon intestine histopathology
High volume of extracts might promote hepatic toxicity Zoral, Ishikawa (81)
Rosemary (Rosmarinus officinalis) Powder 0.6, 1.2, 1.8 and 2.4 g kg-1 4 and 12 weeks Carnosic acid and carnosol (1:1) Gilthead seabream (Sparus aurata) No effect No effect The histological examination of the intestine showed no aspects that might pose problems for absorption, or any immune system disorder associated with the intestine Hernandez, Garcia Garcia (82)
Oliveria (Oliveria decumbens) Essential oils and/or hydroethanolic extract 0.01%, 0.1% and 1.0% 60 days + 14 days S. iniae challenge γ-terpinene,
carvacrol and thymol
Nile tilapia (Oreochromis niloticus) ↑↓ Survival against S. iniae No effect Increase fish survival 14 days after challenge with S. iniae Vazirzadeh, Jalali (83)
Clove basil (Ocimum gratissimum) Ethanolic extract 5.0, 10.0, and 15.0 g kg-1 12 weeks + 14 days L. monocytogenes challenge N/I African catfish (Clarias gariepinus) ↑ final weight
↑ weight gain
↑ SGR
↑ Feed intake
↑ survival against L. monocytogenes
↑ intestine villus length
↑ intestine villus width
↑ intestine absorption area
Improve the fish performance, health, and immune response Abdel-Tawwab, Adeshina (84)
Clove basil (Ocimum gratissimum) Essential oils 0.5%, 1.0% and 1.5% 55 days + 10 days S. agalactiae challenge 1,8-cineole, eugenol and β-selinene Nile tilapia (Oreochromis niloticus) N/I ↑ Intestine villus height
↑ Intestine goblet cells number
↓ Gill epithelial detachment in the secondary lamellae
↓ Gill congestion at the base of the secondary lamellae
Ameliorate tissue damages, even in situations of infection Brum, Cardoso (85)
American basil (Ocimum americanum) Essential oils 0.25, 0.5, 1.0 and 2.0 g kg-1 7 weeks Linalool, eugenol and 1,8-cineole Red drum (Sciaenops ocellatus) No effect ↑ Stomach lysozyme activity
No effect upon the intestinal microbial community
Different supplementation levels do not influence growth performance and intestinal microbial community; however, show effects on immunological responses Sutili, Velasquez (86)
Savory (Satureja khuzestanica) Powder 1% 45 days N/I Common carp (Cyprinus carpio) N/I ↑ Intestinal lactic acid bacteria Improves intestinal health Mousavi, Mohammadiazarm (87)
Allium sp.
Garlic
(Allium sativum)
Aqueous extract 0.10, 0.15, and 0.20 ml kg-1 80 days N/I Guppy (Poecilia reticulata) No effect ↑ Skin mucus lysozyme
↑ Skin mucus alternative complement
↑ Skin mucus total Ig
↑ Skin mucus alkaline phosphatase
Administration of 0.15 mL of garlic extract per kg feed is suggested to obtain optimal skin mucus immunity Motlag, Safari (88)
Garlic
(Allium sativum)
Oil 50 µl kg-1 28 days + exposure to silver nanoparticles N/I Rohu (Labeo rohita) N/I ↓ Gill oxidative stress enzymes activity
↓ Gill histopathologic lesions
Amelioration of silver nanoparticles-induced oxidative stress and histoprotective effects Khan, Qureshi (89)
Garlic
(Allium sativum)
Powder 0.5 g and 1.0 g/100 g 2 months +
2 weeks S. iniae challenge
N/I Nile tilapia (Oreochromis niloticus) ↑ Survival against S. iniae ↑ Anterior intestine transcriptional levels of interleukin genes (IL‐10 and IL‐17F)
↑ OTU counts for the phylum of Proteobacteria and Tenericutes
Could be effective in the prevention of S. iniae infection in fish Foysal, Alam (90)
Garlic
(Allium sativum)
Powder 5.0%, 10.0% and 20.0% 14 or 28 or 32 days + C. irritans challenge Allicin (1.25 mg/g) Guppy (Poecilia reticulata) ↓↑ Gills and caudal fin C. irritans infection N/I No clear preventative effect against C. irritans Kim, Fridman (91)
Garlic
(Allium sativum)
Powder 1.0%, 1.5% and 2.0% 120 days Rainbow trout (Oncorhynchus mykiss) ↑ Weight gain
↑ SGR
↓ Bacterial diversity and richness
Deefgea, Mycoplasma, Exiguobacterium and Clostridium genera
↓↑ Aeromonas genus
Beneficial in terms of promoting growth and inducing changes in the intestinal microbiota in a dose-dependent manner Büyükdeveci, Balcázar (92)
Garlic
(Allium sativum)
Powder 5.0, 10.0 and 15.0 g kg-1 diet 8 weeks N/I Caspian roach (Rutilus rutilus) ↑ Weight
↑ Growth rate
↑ Skin mucus antibacterial activity against S. faecium, M. luteus, S. marcescens and E. coli
↑ Skin mucus protein level
↑ Skin mucus alkaline phosphatase activity
Beneficially affects the skin mucus immune parameters and growth performance Ghehdarijani, Hajimoradloo (93)
Garlic
(Allium sativum)
Lyophilized 2.0% 21 days + cadmium exposure N/I Prussian carp (Carassius gibelio) N/I ↓ Gill histopathologic lesions Shows chelating and antioxidant
potential
Nicula, Dumitrescu (94)
Onion (Allium cepa) Powder 1% 45 days N/I Common carp (Cyprinus carpio) N/I ↑ Intestinal lactic acid bacteria Improves intestinal health Mousavi, Mohammadiazarm (87)
Onion (Allium cepa) Ethanolic extract 0.5%, 1.0%, 1.5% and
2.0%
12 weeks N/I African sharptooth catfish (Clarias gariepinus) No effect ↓ Intestine villus length
↑ Intestine villus width
↑ Intestine absorption area
↑ Intestine cryptal depth
Increase the digestive and absorptive capacity of the intestine Bello, Emikpe (95)
Mongolian Wild Onion (Allium mongolicum) Ethanolic extract 0.04 g kg-1 4 weeks + chromium (Cr) exposure Flavonoids ≥90% (HPLC) Grass carp (Ctenopharyngodon idella) N/I ↓ Intestine and gill Cr accumulation
↓ Gill malondialdehyde content
↓ Gill protein carbonyl content
↑ Intestine lysozyme activity
↑ Intestine complement 3 levels
↑ Intestine and gill tight junction proteins gene expression
↑↓ Intestine and gill NF-κB signaling pathway gene expression
Decrease in Cr-accumulation, oxidative stress, immunosuppression and inflammatory response following Cr exposure Zhao, Yuan (96)
Single bioactive compounds
Thyme (Thymus vulgaris)* Ethanolic solution 0.15, 0.3, 0.45, 0.6, 0.75 g kg-1 56 days + 14 days A. veronii challenge Thymol (commercial product) Snakehead fish (Channa argus) ↑ Final weight
↑ Weight gain
↑ SGR
↑ Protein efficiency ratio
↓ FCR
↑ Survival to A. veronii
↑ Intestine SOD, CAT, GSH-Px activities
↓ Intestine malondialdehyde content
↑ IL-10 and TGF-b gene expression in intestine
↓ HSP70, TNF-a, IL-1b and IL-8 gene expression in intestine
Adequate dietary supplementation can effectively enhance the growth, antioxidant status, immune response and disease resistance Kong, Li (97)
Thyme (Thymus vulgaris)* N/I 0.1, 0.2 and 0.3 g kg-1 60 days + 8 days A. hydrophila challenge Thymol (commercial product) Grass carp (Ctenopharyngodon idella) ↑ Final weight
↑ Survival against A. hydrophila
↑ Gill enzymes of the phosphotransfer network: cytosolic and mitochondrial creatine kinases, adenylate kinase activities and ATP levels in infected fish
↓ Gill ROS levels in infected fish
Favors weight gain and fish longevity. Prevents A. hydrophila induced branchial bioenergetics. High concentrations deserve attention because of side-effects Morselli, Baldissera (98)
Thyme (Thymus vulgaris)* Crystals 0.5 g kg-1 70 days Thymol (99% purity; commercial product) Nile tilapia (Oreochromis niloticus) ↑ SGR
↑ Protein efficiency ratio
↓ Intestine total aerobic and anaerobic counts Improve some performance parameters and negatively modulates intestinal microbial communities. Demonstrates a notable synergistic interaction with chitosan nanoparticle with beneficial effects El-Naby, Al-Sagheer (99)
Thyme (Thymus vulgaris)* N/I 1.0 g kg-1 56 days Thymol (commercial product) Rainbow trout (Oncorhynchus mykiss) ↓ FCR ↓ Intestine culturable anaerobe bacteria Modulated intestinal microbial communities disfavoring total anaerobes Giannenas, Triantafillou (100)
Oregano (Origanum vulgare)* N/I 1.0 g kg-1 56 days Carvacrol (commercial product) Rainbow trout (Oncorhynchus mykiss) ↓ FCR ↓ Intestine culturable anaerobe bacteria Modulated intestinal microbial communities disfavoring total anaerobes Giannenas, Triantafillou (100)
Chinese skullcap (Scutellaria baicalensis)* Powder 0.4, 0.8 and 1.6 g kg-1 60 days + H2O2 challenge Baicalin (80% purity, commercial product) Nile tilapia (Oreochromis niloticus) ↓ FCR ↑ Gill glutathione level
↑ Gill total antioxidant capacity
Improves feed efficiency, enhance antioxidative ability and alleviate oxidative stress Jia, Du (101)
Garlic
(Allium sativum)*
Liquid 0.005%, 0.01% and 0.02% 30 days Allicin (98% purity, commercial product) Large yellow croaker (Larimichthys crocea) ↑ Final weight
↑ Final length
↑ SGR
↑ survival rate
↑ Intestine total antioxidant capacity
↑ Intestine antioxidant enzymes (CAT, NO and NOS) activity
↓ Intestine transcriptional levels of pro-inflammatory genes
Improve the survival and growth of large yellow croaker larvae probably by promoting intestinal development, alleviating inflammation and enhancing appetite Huang, Yao (102)
Phytogenics combinations
Phytogenics combination Essential oils 200 ppm 70 days + 15 days N. girellae challenge Garlic and Lamiaceae-plants oils (N/I; commercial additive) Greater amberjack (Seriola dumerili) No effect ↑ Skin mucus lysozyme activity 15 days post N. girellae challenge
↑ Piscidin gene expression in skin pre-challenge
↑ Proinflammatory cytokines (tnf-α and il1-β), AMPs (hep and cath), immunoglobulin (IgT), complement protein (c3) T-cells marker (cd8) and mucin (muc-2) gene expression in skin post-challenge
Casp3 gene expression in skin post-challenge
Facilitates the immunological response of skin once the parasite is fixed, generating a hostile microenvironment in skin and lowering the parasite load Fernández-Montero, Torrecillas (16)
Phytogenics combination Essential oils 0.3 g kg-1 8 weeks + 2 weeks hypoxia challenge
(Total 10 weeks)
Cinnamaldehyde, thymol and carvacrol Nile tilapia (Oreochromis niloticus) ↓ Hepatosomatic index ↑ Intestine villi density
↓ Intestine malondialdehyde content
Positive effects of digestion and antioxidative capacity Ning, Zhang (103)
Phytogenics combination Microencapsulated essential oils 0.5% 65 days Garlic essential oil (N/I), carvacrol and thymol
(Commercial additive)
Gilthead seabream (Sparus aurata) No effect ↑ Skin mucus inhibitory activity against V. anguillarum and P. anguilliseptica
↓ Cortisol in skin mucus
↑ Regulation of genes associated to the secretory pathway in skin
↑ Regulation of genes associated to non-specific immune response in skin
↑ Regulation of genes coding for oxidative stress enzymes in skin
Beneficially affects the skin and mucus immune and stress parameters, suggesting the stimulation and recruitment of phagocytic cells and a reduction in the fish allostatic load Firmino, Fernández-Alacid (14)
Phytogenics combination Microencapsulated essential oils 0.5% 65 days Garlic essential oil (N/I), carvacrol and thymol
(Commercial additive)
Gilthead seabream (Sparus aurata) No effect ↑ Regulation of genes related to processes of proteolysis and inflammatory modulation, immunity,
transport and secretion, response to cyclic compounds, symbiosis, and RNA metabolism in the mid-anterior intestine
No effect upon alpha diversity of bacterial communities in the anterior and posterior intestinal tract sections
↓ Spirochaetes phylum in the posterior intestine
Photobacterium and Corynebacterium genera in the anterior intestine
Comamonas in the anterior intestine, and Paracoccus, Prevotella and Rothia genera in the posterior intestine
↓ Bacterial sequences related to carbohydrate and drug metabolisms, and membrane transport
↑ Bacterial sequences related to glutathione and lipid metabolisms, naphthalene degradation and sulphur relay system
Evidence of host-microbial co-metabolism
The activation of leukocytes and crosstalk between gut and microbiota are suggested to regulate the inflammatory response induced by the additive Firmino, Vallejos-Vidal (12)
Phytogenics combination Microencapsulated essential oils 0.5% 65 days + 39 days S. chrysophrii challenge (total 104 days) Garlic essential oil (N/I), carvacrol and thymol
(Commercial additive)
Gilthead seabream (Sparus aurata) S.
chrysophrii total parasitation
↑ Regulation of genes related pro-inflammatory immune response arbitrated by degranulating acidophilic
granulocytes, sustained by antioxidant and anti-inflammatory responses in gills
↑ Carboxylate glycoproteins containing sialic acid in mucous and
epithelial gill’s cells
Promotes gill mucosal immunity and reduces gill ectoparasite incidence Firmino, Vallejos-Vidal (15)
Phytogenics combination Essential oils 0.02% 9 weeks + 1 week stress and V. anguillarum challenge Garlic and Lamiaceae-plants oils (N/I; commercial additive) European sea bass (Dicentrarchus labrax) ↑ Survival against Vibrio anguillarum when stress-challenged ↑ Skin mucus lysozyme activity when stress and bacterial challenge Attenuate the fish physiological response to stress increasing resistance to Vibrio anguillarum infection Serradell, Torrecillas (104)
Phytogenics combination Essential oils 0.02% 63 days Garlic and Lamiaceae-plants oils (N/I; commercial additive) European sea bass (Dicentrarchus labrax) No effect ↓ Shannon alpha diversity of mucosa-associated microbiota
↑Clostridiales order in intestinal content
↓ Coliforms and Vibrionales allochthonous microbiota
Reduction of orders containing potentially pathogenic species for fish, and enrichment of gut microbiota composition with butyrate producer taxa Rimoldi, Torrecillas (105)
Phytogenics combination Essential oils 200 ppm 63 days + 7 days stress and V. anguillarum challenge Garlic and Lamiaceae-plants oils (N/I; commercial additive) European sea bass (Dicentrarchus labrax) No effect ↓ Posterior intestine fold area covered by goblet cells
↓ Posterior intestine goblet cells area
↑ Intestine mucus coverage post-challenge
Protective effect focused mainly on the preileorectal valve region Torrecillas, Terova (13)
Phytogenics combination N/I extract 6.0 g kg-1 30 days + 10 days crowding stress (40 days total) Saint John’s wort (Hypericum perforatum, Hypericacea), lemon balm (Melissa officinalis, Lamiaceae) and rosemary (Rosmarinus officinalis, Lamiaceae) mixed at a ratio 3:2:1 Atlantic salmon (Salmo salar) No effect ↓ Gut lipid peroxidation Improves the gut antioxidant status Reyes-Cerpa, Vallejos-Vidal (106)
Phytogenics combination Powder 1% 45 days Savory (Satureja khuzestanica, Lamiaceae) 0.5% and Onion (Allium cepa, Alliaceae) 0.5% Common carp (Cyprinus carpio) N/I ↑ Intestinal lactic acid bacteria Improves intestinal health Mousavi, Mohammadiazarm (87)
Phytogenics combination Essential oils 0.06, 0.2, 0.4 and 0.8 g kg-1 6 weeks Thymol and carvacrol (1:1; commercial additive) Hybrid tilapia (O. niloticus ♀ × O. aureus ♂) N/I ↑ Posterior intestine villus height
↑ Posterior intestine goblet cell count per villus
↑ Anterior intestine intraepithelial leucocytes
↓ Distal intestine intraepithelial leucocytes
↑ OTUs, and PD whole tree and Chao1 diversity indexes
Thermi phylum and Bacteroides, Candidatus Cardinium, and Leptospirillum genera
Affect the immunity primarily through a direct effect on host tissue but also has an indirect effect mediated by microbial changes Ran, Hu (107)
Phytogenics combination Essential oils 100 ppm 9 weeks 25% thymol and 25% carvacrol (commercial additive) Gilthead seabream (Sparus aurata) ↓ FGR ↑ Intestine mucosal foldings
↑ Intestine enterocytes
↑ Intestine goblet cells
↓ Expression of genes related to cell differentiation and proliferation, intestinal architecture and permeability, immunosurveillance, such as cytokines, in the intestine
Induce an anti-inflammatory and anti-proliferative transcriptomic profile with probable improvement in the absorptive capacity of the intestine Perez-Sanchez, Benedito-Palos (108)

SGR, Specific Growth Rate.

FCR, Feed Conversion Ratio.

FGR, Feed Gain Ratio.

OTU, Operational Taxonomic Unit.

*Indicate the putative plant species with high content of the referred bioactive compound. N/I, not identified or not assessed.

Studies reporting the application of plant extracts or related compounds as bath treatments or evaluating bactericidal or antiparasitic effects in vitro were excluded from the selection. The table omits systemic immunity-related results, digestive enzymes or other complementary analysis performed within each study. Blends with other components besides the selected group of plants, terpenes or organosulfur compounds were excluded as well.