Food Chemistry 364 (2021) 130434

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Food Chemistry
journal homepage: www.elsevier.com/locate/foodchem

Polysaccharide from Artocarpus heterophyllus Lam. (jackfruit) pulp

modulates gut microbiota composition and improves short-chain fatty
acids production
Kexue Zhu a, e, Haofei Fan b, Shunjiang Zeng a, d, Shaoping Nie c, Yanjun Zhang a, Lehe Tan a, *,
Chuan Li d, Fei Xu a, Qibing Liu b, *, Gang Wu a
a
Spice and Beverage Research Institute, Chinese Academy of Tropical Agricultural Sciences, Wanning, Hainan 571533, China
b
Department of Pharmacology, School of Basic Medicine and Life Science, Hainan Medical University, Haikou, Hainan 571199, China
c
State Key Laboratory of Food Science and Technology, Nanchang University, Nanchang 330047, China
d
College of Food Science and Technology, Hainan University, Haikou, Hainan 570228, China
e
Key Laboratory of Processing Suitability and Quality Control of the Special Tropical Crops of Hainan Province, Wanning, Hainan 571533, China

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

Keywords: This study aimed to investigate the effects of polysaccharide from Artocarpus heterophyllus Lam. pulp (JFP-Ps) on
Artocarpus heterophyllus Lam. pulp gut microbiota composition and short-chain fatty acids production in mice. The microbial communities of V3 and
Polysaccharide V4 region 16S rRNA gene was amplified by PCR, then sequenced on an Illumina MiSeq PE250 platform and
Gut microbiota
analyzed by multivariate statistical methods. The concentrations of short-chain fatty acids (SCFAs) were
Short-chain fatty acids
measured using gas chromatography (GC) equipped with a flame ionization detector (FID). The results showed
that JFP-Ps significantly affected the levels of intestinal bacteria, including Bacteroidetes, Firmicutes, Proteo­
bacteria, Cyanobacteria, Actinobacteria, Tenericutes, Deferribacteres and TM7. The concentrations of acetic acid,
propionic acid, n-butyric acid and total SCFAs in mouse feces were significantly increased by treatment with JFP-
Ps for 2 weeks. These results indicate that JFP-Ps is beneficial to the gut health and can be developed as a
functional ingredient in relation to gut health.

1. Introduction polysaccharides and dietary fiber influences the intestinal microbiota,

mainly by decreasing the ratio of Firmicutes/Bacteroidetes (Fang, Hu,
The gastrointestinal tract hosts tens of trillions of microorganisms, Nie & Nie, 2019; Trompette et al., 2014).
known as gut microbiota or gut microbiome (Wang, Yao, Lv, Ling & Li, Artocarpus heterophyllus Lam. (known as jackfruit), of the Moraceae
2017). It is widely acknowledged that gut microbiota plays pivotal roles family, is one of the most important tropical fruit trees. Previous studies
in maintaining host health and/or inducing a wide variety of diseases. have shown that Artocarpus heterophyllus Lam. pulp was rich in carbo­
For example, gut microbiota can help harvesting energy for the host hydrate, crude fiber, protein, minerals and phytonutrients, which can be
(Patrice, 2009). In contrast, gut microbiota dysbiosis may be associated exploited as new sources for some important nutrients (Shafiq, Meh­
with many diseases, including inflammatory bowel disease (Kamada, mood, Yasmin, Khan, Khan & Ali, 2017; Swami, Thakor, Haldankar &
Seo, Chen & Núñez, 2013), colorectal cancer (Uronis, Mühlbauer, Her­ Kalse, 2012; Zhu et al., 2019). Some recent studies have focused on the
farth, Rubinas, Jones & Jobin, 2009), as well as extra-intestinal diseases biological effects of bioactive compounds, e.g. polysaccharide from
like obesity, diabetes, atherosclerosis and liver disease (Sekirov, Russell, Artocarpus heterophyllus Lam. (jackfruit). Tan, Li, Lai & Zhang (2013)
Antunes & Finlay, 2010). Diet has been widely considered as the pri­ found that polysaccharides isolated from jackfruit exhibited an immune-
mary food factor affecting the human gut microbiota composition and stimulating activity in a mouse model. Our research group also purified
functionality. For instance, a diet rich in non-starch indigestible a water-soluble polysaccharide from Artocarpus heterophyllus Lam. pulp,

* Corresponding authors at: Spice and Beverage Research Institute, Chinese Academy of Tropical Agricultural Sciences, Wanning, Hainan 571533, China (L. Tan).
Department of Pharmacology, School of Basic Medicine and Life Science, Hainan Medical University, No. 3, Xueyuan Road, Haikou 571199, Hainan Province, China
(Q. Liu).
E-mail addresses: tlh3687@163.com (L. Tan), Qibing.liu@hainmc.edu.cn (Q. Liu).

https://doi.org/10.1016/j.foodchem.2021.130434
Received 13 March 2021; Received in revised form 6 May 2021; Accepted 18 June 2021
Available online 23 June 2021
0308-8146/© 2021 Elsevier Ltd. All rights reserved.
K. Zhu et al. Food Chemistry 364 (2021) 130434

named JFP-Ps, which consisted of rhamnose, arabinose, galactose, 2.2. Animal experiment
glucose, xylose and galacturonic acid, with an average molecular weight
of 1668 kDa (Zhu et al., 2017). Moreover, JFP-Ps can be degraded under Twenty-four male Kunming mice (body weight (BW) 20 ± 2 g) were
gastrointestinal digestion process in vitro, and the intestinal digested purchased from Hunan SJA Laboratory Animal Co., Ltd (SCXK (xiang)
products exhibited strong antioxidant activity in vitro (Zhu et al., 2019). 2016-0002, Changsha city, Hunan Province, China). The animals were
However, to our knowledge, little is known about the functional kept in a 12-h light/12-h dark cycle room under controlled conditions of
interactions between JFP-Ps and gut microbiota, and its relevance to gut relative humidity (60 ± 10%) and temperature (23 ± 2 ◦ C), with ad
health. Thus, this study aimed to investigate the effect of JFP-Ps on gut libitum access to food and water. After acclimatization for 7 days, these
microbiota and short-chain fatty acids (SCFAs) production. mice were then randomly divided into 4 groups: (1) Control group,
orally administered once daily with distilled water, (2) JFP-Ps (50 mg/
2. Materials and methods kg BW) group, orally administered once daily with 50 mg/kg BW JFP-Ps,
(3) JFP-Ps (100 mg/kg BW) group, orally administered once daily with
2.1. Materials and reagents 100 mg/kg BW JFP-Ps, (4) JFP-Ps (200 mg/kg BW) group, orally
administered once daily with 200 mg/kg BW JFP-Ps. All animal care and
Artocarpus heterophyllus Lam. (Jackfruit) fruit was obtained from procedures were strictly conducted according to the protocols approved
Wanning city, Hainan Province, China. JFP-Ps was purified from Arto­ by the Animal Care and Use Committee of Hainan Medical College.
carpus heterophyllus Lam. (Jackfruit) pulp according to our previous
report (Zhu et al., 2017). 2.3. Sample collection and preparation
TIANamp Stool DNA Kit was acquired from Tiangen Biotech Co., Ltd.
(Beijing, China). Agarose and quant-iT PicoGreen dsDNA assay kit were After experimental treatment for 2 weeks, fresh fecal samples were
purchased from Invitrogen (Carlsbad, CA, USA). Q5 DNA polymerase collected and cooled in liquid nitrogen immediately, then stored at
(M0491L) was obtained from New England Biolabs (Beverly, MA, USA). − 80 ◦ C for gut microbiota and SCFAs analysis.
Acetic acid, propionic acid, i-butyric acid, n-butyric acid, i-valeric acid
and n-valeric acid were from Sigma (St. Louis, MO, USA). All other
chemicals and solvents used were of analytical grade.

Fig 1. Purity and quality of isolated DNA from fecal samples. (A) DNA quality (A260/A280 nm ratios), (B) 0.8% agarose gel electrophoresis of bacterial DNA, (C) 2%
agarose gel electrophoresis of bacterial 16S rRNA gene using the 338F-806R primer, (D) The numbers and length distribution based on DNA sequence data.

2
K. Zhu et al. Food Chemistry 364 (2021) 130434

Fig 1. (continued).

2.4. DNA extraction reverse primer, 2 μL DNA template, 0.25 μL Q5 DNA polymerase, and
8.75 μL ddH2O. The PCR cycling procedures were performed as follows:
The genomic DNA was extracted using a TIANamp stool DNA kit an initial denaturation at 98 ◦ C for 2 min, 30 cycles of 98 ◦ C denatur­
following the manufacturer’s instructions. The purity of isolated DNA ation for 15 s, 55 ◦ C annealing for 30 s, 72 ◦ C extension for 30 s, then
was assessed by comparing the absorbance ratio 260/280 using a final extension at 72 ◦ C for 5 min. The amplification products were
Nanodrop spectrophotometer (NC2000; Thermo Scientific, New York, assayed using 2% agarose gel electrophoresis on the DYY-6C electro­
USA). The quality of DNA was confirmed using electrophoresis with phoresis analyzer.
0.8% agarose gel, and the bands were observed using the DYY-6C The PCR products were purified using an Axygen gel extraction kit
electrophoresis analyzer (Beijing Liuyi Instrument Factory, China). (Axygen, Union City, CA, USA) and estimated according to a Quant-iT
PicoGreen dsDNA assay kit (Invitrogen, Carlsbad, California, USA)
2.5. 16S rRNA gene amplification and sequencing protocols on an FLx800 microplate reader (BioTek, New Jersey, USA).
The DNA sequencing library was constructed using TruSeq Nano DNA
The microbial communities of V3 and V4 region 16S rRNA gene was LT Library Prep Kit (Illumina, San Diego, USA). All the barcoded V3 and
amplified by PCR using a forward primer 338F: 5′ -ACTCCTACGG­ V4 PCR products were sequenced on an Illumina MiSeq PE250 platform
GAGGCAGCAG-3′ , and a reverse primer 806R: 5′ -GGAC­ (Illumina, San Diego, CA, USA).
TACHVGGGTWTCTAAT-3′ (Zheng et al., 2018). PCR was carried out in
25 μL PCR mixture containing 5 μL 5 × reaction buffer, 5 μL 5 × GC
buffer, 2 μL 2.5 mM dNTP, 1 μL 10 μM forward primer, 1 μL 10 μM

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Fig 2. (A) Venn diagrams of OTUs sharing > 97%

nucleotide sequence identity. There were 5067 spe­
cies in the control group, 5376 species in JFP-Ps (50
mg/kg BW) group, 4737 species in JFP-Ps (100 mg/kg
BW) group, and 5818 in JFP-Ps (200 mg/kg BW)
group. The total richness of all the samples was 9577.
(B) The bacterial taxonomic diversities classified in
phylum, class, order, family and genus formats. (C)
3D-PcoA plot with Unweighted UniFrac. (D) 2D plot
of NMDS score based on unweighted UniFrac
distances.

2.6. Bioinformatics analysis of gut microbiota profiles further compared using metastats.

The high-quality sequence data were selected and clustered into

operational taxonomic units (OTUs) at 97% similarity using the UCLUST 2.7. Determination of SCFAs production
function in QIIME. Then bacterial α-diversity (including the Simpson
index, Chao1 estimator, ACE estimator and Shannon diversity index) 10% (w/v) fecal homogenate were prepared by dissolving fecal
was determined using the observed number of OTUs. The β-diversity samples in ultrapure water and mixed on a vortex mixer, and then
analysis was examined based on Unweighted UniFrac principal co­ centrifuged at 4800 × g for 20 min at 4 ◦ C. The supernatant was
ordinates analysis (PCoA), Unweighted uniFrac nonmetric multidi­ collected and analyzed by GC.
mensional scaling (NMDS) and Unweighted pair-group method with GC analysis was performed on an Agilent 7890A GC system equipped
arithmetic means (UPGMA) in QIIME and R software. The potential with a flame ionization detector (FID) (Agilent Technologies Inc., USA)
microbial communities of different regions/species among groups were (Hu, Nie & Xie, 2013). A GC column (HP-FFAP, 30 m × 0.32 mm ID,
0.25 μm film thickness, J & W Scientific, Agilent Technologies Inc., USA)

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Fig 2. (continued).

was used for measuring SCFAs. The initial oven temperature was 80 ◦ C 2.8. Statistical analysis
for 0.5 min, and then was raised to 150 ◦ C by 4 ◦ C /min, subsequently
raised to 230 ◦ C at a rate of 20 ◦ C/min, and finally maintained at 230 ◦ C Results were presented as means ± standard error of the mean
for 10 min. As the carrier gas, nitrogen was applied at a flow rate of 19.0 (SEM). The statistical analysis was carried out using SPSS Statistics
mL/min. The temperatures of the FID and injection port were 240 ◦ C. 17.00 software (IBM Analytics, USA). Comparisons between groups
The flow rates of hydrogen and synthetic air were 30 and 300 mL/min, were analyzed using one-way analysis of variance (ANOVA), followed
respectively. The injected sample volume for GC analysis was 0.3 μL, and by the multiple comparisons test of Dunnett. p < 0.05 was considered to
the running time for each analysis was 32 min. The results were be statistically significant.
analyzed with an Agilent ChemStation plus software.

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Table 1 As shown in Fig. 1C, partial sequences of about 450 base pairs of V3
Effects of JFP-Ps on α-diversity index of gut microbiota. and V4 region 16S rRNA gene was obtained by the PCR amplification.
Control JFP-Ps(50 JFP-Ps(100 JFP-Ps(200 Fig. 1D confirmed that the sequences of DNA samples had an average
group mg/kg BW) mg/kg BW) mg/kg BW) length of 420–450 base pairs.
Simpson 0.977 ± 0.987 ± 0.007 0.981 ± 0.006 0.984 ± 0.009
0.017 3.2. OTU numbers
Chao1 1617.988 ± 1722.600 ± 1763.930 ± 2304.515 ±
74.046 90.991 170.064 89.167**
ACE 1670.238 ± 1762.816 ± 1795.346 ± 2348.116 ± A total of 9577 OTUs were detected at the level of 97% similarity,
70.501 105.534 175.471 98.464** 5067, 5376, 4737 and 5818 OTUs were identified in the control group,
Shannon 8.182 ± 8.540 ± 0.259 7.937 ± 0.379 8.577 ± 0.336 JFP-Ps (50 mg/kg BW) group, JFP-Ps (100 mg/kg BW) group and JFP-Ps
0.618
(200 mg/kg BW) group, respectively. The results showed that JFP-Ps
a
Results were presented as means ± SEM. stimulated the fecal gut microbiota composition. In addition, 1817
b *
P < 0.05, **P < 0.01 compared with the control group. OTUs were shared by JFP-Ps (50 mg/kg BW) group, JFP-Ps (100 mg/kg
BW) group and JFP-Ps (200 mg/kg BW) group, while 1296 OTUs were
3. Results shared by all the four groups. The unique OTUs of the control and JFP-Ps
groups were 711, 635, 850 and 787, respectively (Fig. 2A). According to
3.1. Purity and quality of isolated DNA the number of OTUs, bacterial taxonomic diversities were classified in
phylum, class, order, family, genus and species (Fig. 2B). The results
The purity of isolated DNA from fecal samples was confirmed by the showed that when compared to the control group, JFP-Ps treatment
absorbance ratio at 260/280 nm. As shown in Fig. 1A, the value of A260/ increased the OTUs numbers of fecal bacteria.
280 ratios were at 1.86 ± 0.07, 1.87 ± 0.06, 1.91 ± 0.05 and 1.88 ± 0.04
for the control group, JFP-Ps (50 mg/kg BW) group, JFP-Ps (100 mg/kg 3.3. α-Diversity
BW) group and JFP-Ps (200 mg/kg BW) group, respectively. These re­
sults indicate that the isolated DNA fractions were pure enough for α-Diversity of gut microbiota in the control group, JFP-Ps (50 mg/kg
further analysis. Moreover, the quality of the obtained DNA was assessed BW) group, JFP-Ps (100 mg/kg BW) group and JFP-Ps (200 mg/kg BW)
by 0.8% agarose gel electrophoresis. There were clear high molecular- group were described in Table 1. Compared to the control group, after
weight DNA bands without smears (Fig. 1B), indicating high quality treating with JFP-Ps for 2 weeks, the values of Chao1 and ACE were
DNA samples with good integrity status. found to have significantly higher levels in a dose-dependent manner.

Fig 3. (A) UPGMA cluster results of unweighted UniFrac distances based on microbial 16S rRNA sequences from the V3 and V4 region in fecal samples. (B) Effects of
JFP-Ps on the relative abundance of gut microbiota at the phylum level. (C) Effects of JFP-Ps on the taxa abundance of fecal microbiota in the two major gut bacterial
phyla (Bacteroidetes and Firmicutes) was performed using Metastats.

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Fig 3. (continued).

No significant differences have been found in the values of Simpson and were significant differences in bacterial communities after treating with
Shannon among JFP-Ps-treated groups and the control group. The re­ different doses of JFP-Ps (Fig. 2D). In dendrogram from UPGMA cluster
sults indicate a higher variation in intestinal bacteria after treating with analysis based on the unweighted UniFrac method (Fig. 3a), the species
JFP-Ps. composition of gut microbiota changed after treating with JFP-Ps for 2
weeks, but the relative abundance of each member fluctuated.
3.4. β-Diversity
3.5. Effect of JFP-Ps on microbiota taxonomic composition
The phylogenetic differences and similarities of the overall microbial
diversity among different groups were assessed by the unweighted The relative abundance of each phylum was calculated, according to
UniFrac PCoA at the genus level, which showed clear separation of the taxonomic analyses. Bacteroidetes, Firmicutes, Proteobacteria,
bacterial composition in the control group, JFP-Ps (50 mg/kg BW) Cyanobacteria, Actinobacteria, Tenericutes, Deferribacteres and TM7
group, JFP-Ps (100 mg/kg BW) group and JFP-Ps (200 mg/kg BW) group were the predominant bacterial phyla in fecal microbiota. A decrease of
(Fig. 2C). Unweighted uniFrac NMDS results showed a clear clustering of Bacteroidetes was observed in the JFP-Ps supplemented groups
the microbial communities for different groups, indicating that there compared with the control group. But the abundance of Firmicutes,

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Table 2
The concentration of acetic, propionic, i-butyric, n-butyric, i-valeric acid, n-
valeric and total SCFAs in different groups.
Control JFP-Ps(50 JFP-Ps(100 JFP-Ps(200
group mg/kg BW) mg/kg BW) mg/kg BW)

Acetic acid 5.51 ± 6.27 ± 0.40 6.57 ± 0.35* 7.22 ± 0.57**

(mmol/L) 0.94
Propionic acid 1.32 ± 1.41 ± 0.12 1.47 ± 0.16 1.76 ± 0.13*
(mmol/L) 0.29
i-Butyric acid 0.06 ± 0.08 ± 0.01 0.09 ± 0.01* 0.12 ± 0.01*
(mmol/L) 0.02
n-Butyric acid 1.07 ± 1.40 ± 0.10* 1.68 ± 0.14** 1.62 ± 0.07**
(mmol/L) 0.13
i-Valeric acid 0.10 ± 0.12 ± 0.01 0.14 ± 0.02* 0.18 ± 0.02**
(mmol/L) 0.03
n-Valeric acid 0.06 ± 0.07 ± 0.02 0.09 ± 0.03 0.11 ± 0.01*
(mmol/L) 0.01
Total SCFAs 8.13 ± 9.34 ± 0.41 10.05 ± 0.36* 11.02 ± 0.75*
(mmol/L) 1.15
a
Results were presented as means ± SEM.
b
*P < 0.05, **P < 0.01 compared with the control group.

Proteobacteria and Cyanobacteria tended to increase after supplemen­

tation with JFP-Ps (Fig. 3A, Fig. 3B). Taxonomic-based analysis sug­
gested that Bacteroidetes and Firmicutes were dominant in mouse gut
microbiota, and JFP-Ps modulated the composition of gut microbiota in
mice (Fig. 3C).

3.6. Effect of JFP-Ps on SCFAs production

As shown in Table 2, the concentrations of acetic acid, propionic

acid, i-butyric acid, n-butyric acid, i-valeric acid and n-valeric acid in
JFP-Ps-treated groups increased when compared to the control group (p
< 0.05). The most abundant SCFAs were acetic, propionic, and butyric
acids. It was also found that groups of JFP-Ps exhibited significantly Fig 4. Effect of JFP-Ps on gut microbiota and the SCFAs produced by microbial
fermentation.
higher production of total SCFAs as compared to the control group,
especially in JFP-Ps (100 mg/kg BW) and JFP-Ps (200 mg/kg BW)
groups (p < 0.05). The results revealed that JFP-Ps can be fermented to played an important role in improving gut microbiota composition,
SCFAs by intestinal SCFA-producing bacteria. which was in accordance with the results of previous studies.
Bacteroidetes and Firmicutes were the two most predominant phyla
in the gut, which accounted for almost 90% of total bacteria (Tang et al.,
4. Discussion
2018; Wang, Chen, et al., 2018; Wang, Zhang, et al., 2018). Bacter­
oidetes, a Gram-negative bacterial phylum, plays an important role in
Gut microbiota is considered as an essential part in maintaining
the carbohydrates metabolism. Foley, Cockburn & Koropatkin (2016)
human health, which provides protection against the pathogenesis of
confirmed that mammalian gut Bacteroidetes possess analogous Sus-like
several diseases (Xie et al., 2019). Accumulating evidence suggests that
systems that target numerous diverse glycans. The Firmicutes is a
natural products and their bioactive components have attracted wide
diverse group of gram-positive bacteria, considered as the main
attention due to their diverse bioactivities. Among them, poly­
butyrate-producer in the gut and a primary degrader of indigestible
saccharides have emerged as an important class of bioactive natural
polysaccharides (Ganesan, Chung, Vanamala & Xu, 2018; Graf et al.,
products. It was found that indigestible polysaccharides could improve
2015). The present work found that Bacteroidetes and Firmicutes were
the glycometabolism and prevent diseases by altering the composition of
two dominant bacterial phyla in the gut, and can be modulated by JFP-
gut microbiota (Fang et al., 2019). Our gut microbiota data profiles of
Ps.
the fecal samples were identified with bioinformatics analysis, and the
Polysaccharides widely existed in animals, plants and microorgan­
results showed that JFP-Ps administration increased the diversity of
isms (Bagchi & Kumar, 2016). Recent researches have indicated that
microbial community, suggesting that JFP-Ps could regulate the
polysaccharides are fermented and utilized by gut microbiota, and may
microbiota composition.
play a crucial role in host health (Wang, Chen, et al., 2018; Wang,
The abundance of bacterial species in the gut microbiota was
Zhang, et al., 2018). The degradation of polysaccharides through
assigned by OTUs. Tang et al. (2018) found that polysaccharides from
carbohydrate-active enzymes (CAZymes) encoded by gut microbiota is
purple sweet potato raised the OTU numbers and α-diversity index in
the first step in the production of SCFAs, and then the products are
normal mice. Consistent with these observations, JFP-Ps increased the
further fermented into SCFAs, including acetate, propionate, butyrate
OTUs numbers of fecal bacteria in mice. Meanwhile, Turnbaugh,
and valerate acid by SCFA-producing bacteria (Fang et al., 2019). Our
Ridaura, Faith, Rey, Knight & Gordon (2009) reported that the distal gut
results showed that JFP-Ps up-regulated the levels of acetic acid, pro­
microbiota of mice were composed of different bacterial phyla,
pionic acid and butyric acid compared to the control group (Table 2).
including Firmicutes, Bacteroidetes, Actinobacteria, Proteobacteria,
These results were in agreement with previous studies (Hu, Nie, Min &
Verrucomicrobia, Cyanobacteria, TM7, Fusobacteria and Spirochaeates.
Xie, 2012; Zhu et al., 2018).
Our results showed that the bacterial phyla of gut microbiota in mice
SCFAs are the important metabolites which are produced by gut
were composed of Bacteroidetes, Firmicutes, Proteobacteria, Cyano­
bacteria from undigested carbohydrates (Tan, McKenzie, Potamitis,
bacteria, Actinobacteria, Tenericutes, Deferribacteres and TM7. JFP-Ps

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K. Zhu et al. Food Chemistry 364 (2021) 130434

Thorburn, Mackay & Macia, 2014; Zhu, Nie, Tan, Li, Gong & Xie, 2016). Hu, J., Nie, S., & Xie, M. (2013). High pressure homogenization increases antioxidant
capacity and short-chain fatty acid yield of polysaccharide from seeds of Plantago
Among them, acetic acid is found to be the main SCFA and can be
asiatica L. Food Chemistry, 138(4), 2338–2345. https://doi.org/10.1016/j.
absorbed by the colonic epithelium, and then served as a source of en­ foodchem.2012.12.016.
ergy for brain, heart and peripheral tissues (Xu et al., 2019). Propionic Kamada, N., Seo, S., Chen, G. Y., & Núñez, G. (2013). Role of the gut microbiota in
acid is the second abundant SCFA in the colonic fermentation of indi­ immunity and inflammatory disease. Nature Reviews Immunology, 13(5), 321–335.
https://doi.org/10.1038/nri3430.
gestible carbohydrates in the gut and may have various beneficial effects Liu, Y., Gao, Y., Ma, F., Sun, M., Mu, G., & Tuo, Y. (2020). The ameliorative effect of
(Al-Lahham et al., 2012). For example, propionic acid was reported to Lactobacillus plantarum Y44 oral administration on inflammation and lipid
reduce hepatic and serum cholesterol content in rats (Liu, Gao, Ma, Sun, metabolism in obese mice fed with a high fat diet. Food & Function, 11(6),
5024–5039. https://doi.org/10.1039/D0FO00439A.
Mu & Tuo, 2020). Butyrate, an important source of energy for the in­ Patrice, D. C. A. N. (2009). The Role of the Gut Microbiota in Energy Metabolism and
testinal mucosa, was considered as a preventive factor against the Metabolic Disease. Current Pharmaceutical Design, 15(13), 1546–1558. https://doi.
development of colonic diseases (Finnie, Dwarakanath, Taylor & Rho­ org/10.2174/138161209788168164.
Sekirov, I., Russell, S. L., Antunes, L. C. M., & Finlay, B. B. (2010). Gut microbiota in
des, 1995). Our results showed that JFP-Ps was fermented into different health and disease. Physiological Reviews, 90(3), 859–904. https://doi.org/10.1152/
SCFAs, suggesting that the intake of JFP-Ps may be beneficial for the physrev.00045.2009.
colon health. Shafiq, M., Mehmood, S., Yasmin, A., Khan, S. J., Khan, N. H., & Ali, S. (2017).
Evaluation of phytochemical, nutritional and antioxidant activity of indigenously
In conclusion, Bacteroidetes, Firmicutes, Proteobacteria, Cyanobac­ grown jackfruit (Artocarpus heterophyllus Lam.). Journal of Scientific Research, 9(1),
teria, Actinobacteria, Tenericutes, Deferribacteres and TM7 were the 135–143. https://doi.org/10.3329/jsr.v1i1.29665.
major bacterial phyla of gut microbiota in mice. The diversity and Swami, S. B., Thakor, N. J., Haldankar, P. M., & Kalse, S. B. (2012). Jackfruit and its
many functional components as related to human health: A review. Comprehensive
composition of gut microbiota were significantly altered by treatment
Reviews in Food Science and Food Safety, 11(6), 565–576. https://doi.org/10.1111/
with JFP-Ps. Moreover, the intake of JFP-Ps could modulate SCFA- j.1541-4337.2012.00210.x.
producing bacteria, and then improved the levels of SCFAs (including Tan, J., McKenzie, C., Potamitis, M., Thorburn, A. N., Mackay, C. R., & Macia, L. (2014).
acetic acid, propionic acid, i-butyric acid, n-butyric acid, i-valeric acid The role of short-chain fatty acids in health and disease. Advances in Immunology,
121, 91–119. https://doi.org/10.1016/B978-0-12-800100-4.00003-9.
and n-valeric acid). This study has provided new insights into the po­ Tan, Y., Li, H., Lai, W., & Zhang, J. (2013). Crude dietary polysaccharide fraction isolated
tential application of JFP-Ps to be developed as a functional ingredient from jackfruit enhances immune system activity in mice. Journal of Medicinal Food,
in relation to gut health (Fig. 4). The potential physiological effects of 16(7), 663–668. https://doi.org/10.1089/jmf.2012.2565.
Tang, C., Sun, J., Zhou, B.o., Jin, C., Liu, J., Kan, J., … Zhang, N. (2018). Effects of
gut microbiota fermentation of JFP-Ps on colon health will be further polysaccharides from purple sweet potatoes on immune response and gut microbiota
investigated in our follow-up work. composition in normal and cyclophosphamide treated mice. Food & Function, 9(2),
937–950. https://doi.org/10.1039/C7FO01302G.
Trompette, A., Gollwitzer, E. S., Yadava, K., Sichelstiel, A. K., Sprenger, N., Ngom-
Declaration of Competing Interest Bru, C., … Marsland, B. J. (2014). Gut microbiota metabolism of dietary fiber
influences allergic airway disease and hematopoiesis. Nature Medicine, 20(2),
The authors declare that they have no known competing financial 159–166. https://doi.org/10.1038/nm.3444.
Turnbaugh, P. J., Ridaura, V. K., Faith, J. J., Rey, F. E., Knight, R., & Gordon, J. I. (2009).
interests or personal relationships that could have appeared to influence The effect of diet on the human gut microbiome: a metagenomic analysis in
the work reported in this paper. humanized gnotobiotic mice. Science Translational Medicine, 1(6), 6ra14. https://
doi.org/10.1126/scitranslmed.3000322.
Uronis, J. M., Mühlbauer, M., Herfarth, H. H., Rubinas, T. C., Jones, G. S., Jobin, C., &
Acknowledgement
Bereswill, S. (2009). Modulation of the intestinal microbiota alters colitis-associated
colorectal cancer susceptibility. PLoS One, 4(6), e6026. https://doi.org/10.1371/
This work is financially supported by the National Natural Science journal.pone.0006026.
Foundation of China (No. 31901649), the Key Research and Develop­ Wang, B., Yao, M., Lv, L., Ling, Z., & Li, L. (2017). The human microbiota in health and
disease. Engineering, 3(1), 71–82. https://doi.org/10.1016/J.ENG.2017.01.008.
ment Project of Hainan Province (No. ZDYF2020218) and the Central Wang, H., Zhang, X., Wang, S., Li, H., Lu, Z., Shi, J., & Xu, Z. (2018). Mannan-
Public-interest Scientific Institution Basal Research Fund for Chinese oligosaccharide modulates the obesity and gut microbiota in high-fat diet-fed mice.
Academy of Tropical Agricultural Sciences (No. 1630142020007). Food & Function, 9(7), 3916–3929. https://doi.org/10.1039/C8FO00209F.
Wang, M., Chen, Y., Wang, Y., Li, Y., Zheng, H., Ma, F., … Liao, Q. (2018). The effect of
probiotics and polysaccharides on the gut microbiota composition and function of
References weaned rats. Food & Function, 9(3), 1864–1877. https://doi.org/10.1039/
C7FO01507K.
Al-Lahham, S. A., Roelofsen, H., Rezaee, F., Weening, D., Hoek, A., Vonk, R., & Xie, S.-Z., Liu, B., Ye, H.-Y., Li, Q.-M., Pan, L.-H., Zha, X.-Q., … Luo, J.-P. (2019).
Venema, K. (2012). Propionic acid affects immune status and metabolism in adipose Dendrobium huoshanense polysaccharide regionally regulates intestinal mucosal
tissue from overweight subjects. European Journal of Clinical Investigation, 42(4), barrier function and intestinal microbiota in mice. Carbohydrate Polymers, 206,
357–364. https://doi.org/10.1111/j.1365-2362.2011.02590.x. 149–162. https://doi.org/10.1016/j.carbpol.2018.11.002.
Bagchi, S., & Kumar, K. J. (2016). Studies on water soluble polysaccharides from Xu, S., Aweya, J. J., Li, N., Deng, R., Chen, W., Tang, J., & Cheong, K. (2019). Microbial
Pithecellobium dulce (Roxb.) Benth. seeds. Carbohydrate Polymers, 138, 215–221. catabolism of Porphyra haitanensis polysaccharides by human gut microbiota. Food
https://doi.org/10.1016/j.carbpol.2015.11.018. Chemistry, 289, 177–186. https://doi.org/10.1016/j.foodchem.2019.03.050.
Fang, Q., Hu, J., Nie, Q., & Nie, S. (2019). Effects of polysaccharides on glycometabolism Zheng, J., Yuan, X., Cheng, G., Jiao, S., Feng, C., Zhao, X., … Liu, H. (2018). Chitosan
based on gut microbiota alteration. Trends in Food Science & Technology, 92, 65–70. oligosaccharides improve the disturbance in glucose metabolism and reverse the
https://doi.org/10.1016/j.tifs.2019.08.015. dysbiosis of gut microbiota in diabetic mice. Carbohydrate Polymers, 190, 77–86.
Finnie, I. A., Dwarakanath, A. D., Taylor, B. A., & Rhodes, J. M. (1995). Colonic mucin https://doi.org/10.1016/j.carbpol.2018.02.058.
synthesis is increased by sodium butyrate. Gut, 36(1), 93–99. https://doi.org/ Zhu, K., Nie, S., Tan, L., Li, C., Gong, D., & Xie, M. (2016). A polysaccharide from
10.1136/gut.36.1.93. Ganoderma atrum improves liver function in type 2 diabetic rats via antioxidant
Foley, M. H., Cockburn, D. W., & Koropatkin, N. M. (2016). The Sus operon: A model action and short-chain fatty acids excretion. Journal of Agricultural and Food
system for starch uptake by the human gut Bacteroidetes. Cellular and Molecular Life Chemistry, 64(9), 1938–1944. https://doi.org/10.1021/acs.jafc.5b06103.
Sciences, 73(14), 2603–2617. https://doi.org/10.1007/s00018-016-2242-x. Zhu, K., Yao, S., Zhang, Y., Liu, Q., Xu, F., Wu, G., … Tan, L. (2019). Effects of in vitro
Ganesan, K., Chung, S. K., Vanamala, J., & Xu, B. (2018). Causal relationship between saliva, gastric and intestinal digestion on the chemical properties, antioxidant
diet-induced gut microbiota changes and diabetes: A novel strategy to transplant activity of polysaccharide from Artocarpus heterophyllus Lam. (jackfruit) pulp. Food
Faecalibacterium prausnitzii in preventing diabetes. International Journal of Molecular Hydrocolloids, 87, 952–959. https://doi.org/10.1016/j.foodhyd.2018.09.014.
Sciences, 19(12), 3720. https://doi.org/10.3390/ijms19123720. Zhu, K., Zhang, Y., Nie, S., Xu, F., He, S., Gong, D., … Tan, L. (2017). Physicochemical
Graf, D., Di Cagno, R., Fåk, F., Flint, H. J., Nyman, M., Saarela, M., & Watzl, B. (2015). properties and in vitro antioxidant activities of polysaccharide from Artocarpus
Contribution of diet to the composition of the human gut microbiota. Microbial heterophyllus Lam. pulp. Carbohydrate Polymers, 155, 354–361. https://doi.org/
Ecology in Health and Disease, 26, 26164. https://doi.org/10.3402/mehd.v26.26164. 10.1016/j.carbpol.2016.08.074.
Hu, J., Nie, S., Min, F., & Xie, M. (2012). Polysaccharide from seeds of Plantago asiatica L. Zhu, Z., Zhu, B., Sun, Y., Ai, C., Wu, S., Wang, L., … Liu, X. (2018). Sulfated
increases short-chain fatty acid production and fecal moisture along with lowering polysaccharide from sea cucumber modulates the gut microbiota and its metabolites
pH in mouse colon. Journal of Agricultural and Food Chemistry, 60(46), 11525–11532. in normal mice. International Journal of Biological Macromolecules, 120, 502–512.
https://doi.org/10.1021/jf302169u. https://doi.org/10.1016/j.ijbiomac.2018.08.098.