Citation: Huang, S.; Lin, L.; Wang, S.;
Ding, W.; Zhang, C.; Shaukat, A.; Xu,
B.; Yue, K.; Zhang, C.; Liu, F. Total
Flavonoids of Rhizoma Drynariae
Mitigates Aflatoxin B1-Induced Liver
Toxicity in Chickens via Microbiota-
Gut-Liver Axis Interaction
Mechanisms. Antioxidants 2023, 12,
819. https://doi.org/10.3390/
antiox12040819
Academic Editors: Fengna Li, Jie Yin,
Dan Wang and Sung Woo Kim
Received: 12 February 2023
Revised: 21 March 2023
Accepted: 24 March 2023
Published: 28 March 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
antioxidants
Article
Total Flavonoids of Rhizoma Drynariae Mitigates Aflatoxin
B1-Induced Liver Toxicity in Chickens via Microbiota-Gut-Liver
Axis Interaction Mechanisms
Shucheng Huang
1
, Luxi Lin
1
, Shiqiong Wang
2
, Wenli Ding
1
, Chaodong Zhang
1
, Aftab Shaukat
3
,
Bowen Xu
1
, Ke Yue
1
, Cai Zhang
4
and Fang Liu
1,
*
1
College of Veterinary Medicine, Henan Agricultural University, Zhengzhou 450046, China;
huang.sc@henau.edu.cn (S.H.); linmo0726@163.com (L.L.)
2
College of Food Science and Technology, Henan Agricultural University, Zhengzhou 450002, China
3
National Center for International Research on Animal Genetics, Breeding and Reproduction (NCIRAGBR),
Huazhong Agricultural University, Wuhan 430070, China
4
Laboratory of Environment and Livestock Products, Henan University of Science and Technology,
Luoyang 471023, China
* Correspondence: liufang.vet@henau.edu.cn; Tel.: +86-139-0383-1328
Abstract: Aflatoxin B1 (AFB1) is a common mycotoxin that widely occurs in feed and has severe
hepatotoxic effects both in humans and animals. Total flavonoids of Rhizoma Drynaria (TFRD), a
traditional Chinese medicinal herb, have multiple biological activities and potential hepatoprotective
activity. This study investigated the protective effects and potential mechanisms of TFRD against
AFB1-induced liver injury. The results revealed that supplementation with TFRD markedly lessened
broiler intestinal permeability by increasing the expression of intestinal tight junction proteins, as
well as correcting the changes in gut microbiota and liver damage induced by AFB1. Metabolomics
analysis revealed that the alterations in plasma metabolites, especially taurolithocholic acid, were
significantly improved by TFRD treatment in AFB1-exposed chickens. In addition, these metabolites
were closely associated with [Ruminococcus], ACC, and GPX1, indicating that AFB1 may cause
liver injury by inducing bile acid metabolism involving the microbiota–gut–liver axis. We further
found that TFRD treatment markedly suppressed oxidative stress and hepatic lipid deposition,
increased plasma glutathione (GSH) concentrations, and reversed hepatic ferroptosis gene expression.
Collectively, these findings indicate that ferroptosis might contribute to the hepatotoxicity of AFB1-
exposed chickens through the microbiota–gut–liver axis interaction mechanisms; furthermore, TFRD
was confirmed as an herbal extract that could potentially antagonize mycotoxins detrimental effects.
Keywords: aflatoxin B1; antioxidation; bile acid; Chinese medicinal herb; ferroptosis; gut microbiota
1. Introduction
Aflatoxin B1 (AFB1) is the largest proportion of known aflatoxins, with the highest
degree of mycotoxin contamination [1]. The International Agency for Research on Cancer
(IARC) evaluated AFB1 as one of the most toxic chemical carcinogens known to disrupt
the normal function of multiple organs and tissues [2,3]. It has been reported that poultry
might be most susceptible to AFB1 toxicity [4]. AFB1 exposure significantly impaired
poultry production performance and reproductive performance, such as meat quality,
feed intake, body weight, and egg production, which caused great economic losses to the
poultry breeding industry [4,5]. Recently, AFB1 was reported to cause liver oxidative stress,
inflammation, ferroptosis, and eventually liver cancer [6,7]. Thus, it is important to find
some green harmless substances that can prevent or treat AFB1 poisoning.
The gastrointestinal tract is the first place that encounters mycotoxins, as the absorp-
tion rate of aflatoxin is approximately 90% in poultry [8]. The intestinal mucosal barrier, a
semipermeable structure, consists of biological, physical, chemical, and immune barriers
Antioxidants 2023, 12, 819. https://doi.org/10.3390/antiox12040819 https://www.mdpi.com/journal/antioxidants