Journal of Functional Foods 71 (2020) 104015

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Journal of Functional Foods

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

A systematic review on the composition, storage, processing of bamboo T

shoots: Focusing the nutritional and functional benefits
⁎
Yulin Wanga, Jia Chena, Damao Wanga, Fayin Yea, Yonglin Hea, Zicong Hua, Guohua Zhaoa,b,
a
College of Food Science, Southwest University, Chongqing 400715, People’s Republic of China
b
Chongqing Engineering Research Center of Regional Foods, Chongqing 400715, People’s Republic of China

A R T I C LE I N FO A B S T R A C T

Keywords: Bamboo shoot, the young culms of bamboo plants, has been utilized as a food item in Asian countries. Bamboo
Bamboo shoot shoots are rich in protein, fiber, vitamins and minerals as well as plenty of nutraceuticals including phenols and
Biological activities phytosterols. Bamboo shoots were historically used in medications and recently identified with the diverse
Storage health benefits such as hypolipidemic, prebiotic and anti-diabetic activities. However, the rapid postharvest
Processing
lignification deteriorated the nutritional quality and palatability of fresh bamboo shoots, and the presence of
Cyanogenic glucoside
cyanogenic glycosides made them inapplicable being eaten raw. To these ends, physical and chemical regimes
were invented for their postharvest storage and various processing methods were adopted to eliminate the toxins
and achieve a year-round supply. With their consumption spreading over the continents, bamboo shoots become
more important in human diet. The sustainable development, food safety control and the clarification of their
potential health effects are suggested in future consideration.

1. Introduction
As a giant perennial arborescent grass, bamboo belongs to the fa-
mily Poaceae and subfamily Bambuseae. It is native to China and widely
distributed across continents, especially in tropical, subtropical and
temperate regions with a mostly mesic to wet season (Chauhan, Unni,
Chitravathi, & Pakalapati, 2016; Singhal, Bal, Satya, Sudhakar, & Naik,
2013). To date, more than 1642 species in 75 genera of bamboo have
been evidenced to be distributed throughout the world, in which over
500 species in 39 genera grow in China (Choudhury, Sahu, & Sharma,
2012; Goyal & Brahma, 2014; Goyal, Ghosh, Dubey, & Sen, 2012). As
estimated, the global area of bamboo forest is up to 31.5 M ha, ac-
counting for approximately 1% of the total forest area. Over the last
30 years, the global forest area declines continuously, while the
bamboo forest area shows an average annual growth rate of 3% (FAO,
2010). In terms of the bamboo forest area, India and China are the first
two countries with the largest total areas of 9.57 M ha and 6.01 M ha,
respectively. Bamboo is regarded as the fastest-growing plant on the
planet. The fasting growing and quick maturation, as well as the short
production cycle, high biomass productivity and wide adaptability
make bamboo valuable in rapid forest establishment, sustainable con-
struction, environment-friendly furniture making and ecological food-
stuff producing (Goyal & Sen, 2016; He et al., 2014). Notably, it has
been proved that moderate harvesting could benefit the productivity of
bamboo. Thus, bamboo is regarded as one of the most culturally, eco-
logically and economically important plant or grass especially for the
livelihoods of some rural populations in Asia, Latin America and Africa.
In gross, approximate 2.5 billion people worldwide utilize various
bamboo-related products. The annually bamboo economic sector worth
in China was estimated up to USD 11.8 billion (Buckingham et al.,
2011).
Regarding the food applications, the leaves and shoots of edible
bamboo have been under consideration for many years. Published ar-
ticles had announced the anti-oxidant, anti-microbial, anti-in-
flammatory, anti-helminthic, anti-diabetic and anti-ulcer activities of
various extracts and components from bamboo leaves. This was mainly
ascribed to the phenolic components in bamboo leave such as orientin,
homoorientin, isoorientin, vitexin, homovitexin, and tricin and phe-
nolic acids (Nirmala, Bisht, Bajwa, & Santosh, 2018). Since 2004, the
bamboo leave extract has been authorized by The Chinese Ministry of
Health as a novel food antioxidant with a maximum addition of 0.5 g/
kg in edible oils, meat products, aquatic products, cereal products,
puffed food fruit and vegetable juices and tea beverage. Besides its wide
use as animal fodders, the leaves of bamboo have been used up to
thousands of years as wrapping materials for Chinese rice-pudding and
Japanese sushi. In China, the bamboo leaves were also made into a tea
form for drink.
In contrast to bamboo leaves, the bamboo shoot phytologically

⁎
Corresponding author at: College of Food Science, Southwest University, Tiansheng Road 2, Chongqing 400715, People’s Republic of China.
E-mail address: zhaogh@swu.edu.cn (G. Zhao).

https://doi.org/10.1016/j.jff.2020.104015
Received 6 January 2020; Received in revised form 6 May 2020; Accepted 10 May 2020
1756-4646/ © 2020 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/BY-NC-ND/4.0/).
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 1
Species of edible bamboo shoots in different countries.a.
Family Species (Countryb)

Acidosasa edulis (-)

Arundinaria alpine (Ethiopia); aristat (I); hirsute (I)
Bambusa arundianacea (I); arnhemica (A); balcooa (A, B, I, In); bambos (I, T, Uganda); edulis (C, T); glaucescens (I); kingiana (I); longispiculata (I); multiples (I, C); nana (I);
nutans (I); oldhamii (A, C, Japan, T); pallida (C, I, T); beecheyana (-); polymorpha (B, I, Myanmar, PR, T, Uganda); rigida (-); textiles (C); tulda (B, I, In,
Myanmar, Nepal, T, Vietnam); vulgaris (I, In)
Cephalostachyum capitatum (I); fuchsianum (I); pergracile (I)
Chimonobambusa callosa (I); hookerian (I); quadrangularis (-)
Dendrocalamus asper (A, B, C, I, In, Japan, Malaysia, Myanmar, Nepal, Philippines, T, Vietnam)
barbatus (Uganda); brandisii (A, C, I, Uganda); calostachyus (I); flagellifer (I); giganteus (B, Bhutan, C, Ghana, I, In, Kenyan, Myanmar, Nepal, Philippines, T,
Vietnam); hamiltonii
(Bhutan, I, Nepal); hookeri (Bhutan, I, Nepal); latifera (C, I); latiflorus (A, C, T); longispathus (I, In); membranaceus (I, PR); merrillianus (T); sikkimensis (Bhutan,
I, Nepal); stenoaurita (-); strictus (I, Myanmar, Nepal, T, Uganda); tulda (I); vario-striata (-)
Gigantochloa ablociliata (I); atter (A); apus (I, In, Malaysia, Myanmar, T); levis (C, In, Kalimantan, Malaysia, Philippines, PR, Vietnam); nigrociliata (In); pseudoarundinacea
(C, I, In, Malaysia); rostrate (I)
Guadua augustifolia (PR)
Melocanna baccifera (B, C, I, In, Myanmar, PR, Vietnam); bambusoides (US)
Oxytenanthera abyssinica (Ethiopia, Uganda); albociliata (I)
Phyllostachys bambusoides (I, Japan, US); dulcis (US); edulis (Japan, US); heterocyclavar (A, Korea); iridescens (C); makinoi (C); mitis (Japan); nigra (Korea); nuda (C, US);
praecox (C); prominens(-); pubescens (A, C, Japan, Korea, US); sulphurea (-); vivax (-); viridis (C, US);
Pleioblastus amarus (C)
Pseudoxytenanthera albociliata (T); polymorphum (I)
Qiongzhuea Tumidinada (-)
Sasa kurilensis (Japan)
Schizostachyum beddomei (I); capitatum (I); dullooa (I); funbomii (-);
Sinarundinaria elegans (I)
Sinocalamus oldhami (PR)
Teinostachyum wightii (I)
Thyrsostachys oliveri (I, Uganda); siamensis (B, C, I, In, Myanmar, T, Vietnam)
Yushania alpine (Kenyan)

a
Data in the table were cited from INBAR and FAO.
b
Refers to no country specified; A, B, C, I, In, PR, T and US are Australia, Bangladesh, China, India, Indonesia, Puerto Rico, Thailand and United States,
respectviely.

Table 2 refers to the aerial bud or meristematic tissue of bamboo plants. It is

World bamboo planting and import and export of bamboo shoots.a. often known as the young, juvenile, tender, immature or expanding
Country Bamboo forest No. of species Export Import
culms/stalks emerging from the nodes of the pseudo rhizomes of
(km2) (exclusive) products products bamboo plants. Bamboo shoot is the neonatal culm just sprouting from
(×103 USD) (×103 USD) the earth in full-diameter, in which the nodes and internodes were
densely-packaged. Fresh bamboo shoots own a crisp and crunchy tex-
Australia – 3 (2) 0 2,779
Bangladesh 830 18 (0) – –
ture as well as a sweet taste, imparting a unique flavor. However, not all
Bhutan – 21 (1) – – bamboo shoots are edible and the main edible species are summarized
Cambodia 290 4 (0) – – in Table 1. Although bamboo was called in ancient years as “the poor’s
Canada – – 0 2,791 timber”, the bamboo shoot was one of the highest palatable dishes in
China 67,000 637 (5 8 6) 284,633 142
delicacies recently honored as “the food of rich man”. Traditionally,
Ethiopia 14,744 2 (0) – –
EU countries – – 16,610 46,846 bamboo shoot has been treated as a regular cuisine in some Asia
Ghana 300,000 1 (0) – – countries. Nowadays, the consumption of bamboo shoots quickly
India 11,136,110 102 (40) 0 114 spreads all continents due to the worldwide food trade, popularizing
Indonesia 21,000 56 (29) – – Chinese restaurants and multi-region culture fusion. To date, the annual
Japan 1540 84 (75) 1,989 130,210
worldwide consumption of bamboo shoot has been beyond 2 million
Kenya 1332 1 (0) – –
Korea 60 6 (2) 0 4,384 tons. The worldwide production, trade and consumption of bamboo
Laos 16,000 13 (4) – – shoot are summarized in Table 2. In view of the increasing popular-
Madagascar 112 33 (32) – – ization of bamboo shoot in worldwide diets, this article aims to sum-
Malaysia 6770 50 (26) 211 0
marize the nutritional quality, health benefits, processing regimes and
Myanmar 8500 75 (30) 77 0
Nepal 629 25 (6) – – safety concerns of bamboo shoot.
Pakistan 200 3 (0) – –
Philippines 1720 26 (14) 8 0
Russia – 1 (0) 0 913 2. Nutrients and nutraceuticals in bamboo shoots
Sri Lanka 30 11 (6) – –
Sudan – 2 (0) 0 1187
Thailand 2610 36 (4) 12,724 0 Being a traditional forest vegetable with a long history of con-
The USA – 1(1) 278 28,843 sumption, bamboo shoots are not only tasteful and delicious but also
Uganda 5.4 3 (0) – – rich in nutrients (Nirmala, Bisht, & Haorongbam, 2011). It contains
Vietnam 15,325 69 (38) 405 0
high amounts of proteins, carbohydrates, vitamins and minerals but has
a
Data in the table were cited from INBAR.
low levels of fat and cholesterols, making it the most desired health
food among patients with lifestyle-related disorders. The nutrient
compositions of bamboo shoots are summarized in Table 3.

2
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 3
Proximate composition (g/100 g) of bamboo shoots.
Plant species Forma Moisture Protein FAAb TDFb Starch Sugar Fat Ash Reference

Bambusa bamboos F 89.93 3.57 3.98 3.54 0.25 5.42 0.50 1.38 Nirmala et al., 2007
Bambusa balcooa F 91.65 2.74 -e – – 3.90 – – Bhatt & Singh, 2005
Bambusa tulda F 83.60 3.69 3.65 3.97 0.59 6.92 0.48 0.85 Nirmala et al., 2007
Bambusa tulda F 88.17 2.88 2.99 15.47 0.48 4.43 0.40 0.89 Premlata, Saini, Nirmala, & Bisht, 2015
Bambusa tulda F – 3.69 3.65 3.97 – 6.92 0.48 Nirmala et al., 2011
Bambusa kingiana F 90.00 3.72 3.18 3.90 0.34 5.50 0.41 1.38 Nirmala et al., 2007
Bambusa nutans F 92.00 2.84 3.89 2.28 0.21 5.47 0.40 0.68 Nirmala et al., 2007
Bambusa nutans F 91.26 3.52 2.21 9.89 1.37 2.76 0.31 0.82 Premlata et al., 2015
Bambusa pallida F 92.20 2.31 – – – 3.83 – – Bhatt & Singh, 2005
Bambusa polymorpha F 90.26 3.64 3.42 3.82 0.38 5.44 0.46 0.76 Nirmala et al., 2007
Bambusa polymorpha F 91.65 2.1 – – – 4.86 – – Bhatt & Singh, 2005
Bambusa vulgaris F 90.60 3.64 3.57 4.24 0.27 6.51 0.50 1.01 Nirmala et al., 2007
Bambusa vulgaris F 91.40 2.33 – 2.60 – 2.43 0.14 1.15 Karanja, 2017
Dendrocalamus brandisii F 92.57 1.54 – 0.66 – – 0.28 0.89 Chen, Ma, Ding, Luo, & Liu, 2018
Dendrocalamus giganteus F 90.70 3.11 3.96 2.65 0.51 5.10 0.39 0.89 Nirmala et al., 2007
Dendrocalamus giganteus F 89.50 2.63 3.59 6.68 0.56 4.66 0.32 0.93 Premlata et al., 2015
Dendrocalamus giganteus F 91.19 2.59 – – – 4.78 – – Bhatt & Singh, 2005
Dendrocalamus giganteus F 90.7 3.86 3.11 4.80 0.51 5.10 0.39 0.89 Nirmala et al., 2008
Dendrocalamus giganteus F 91.20 2.30 – 1.60 – 3.60 0.17 1.17 Karanja, 2017
Dendrocalamus hamiltonii F 92.51 3.72 3.18 3.90 0.47 5.50 0.41 0.86 Nirmala et al., 2007
Dendrocalamus hamiltonii F 83.60 2.93 4.03 6.14 0.40 3.44 0.37 0.91 Premlata et al., 2015
Dendrocalamus hamiltonii F 92.37 2.60 – – – 4.00 – – Bhatt & Singh, 2005
Dendrocalamus hamiltonii F – 3.72 3.18 3.90 – 5.50 0.41 – Nirmala et al., 2011
Dendrocalamus membranaceus F 89.30 3.38 3.46 2.91 0.23 5.40 0.43 0.63 Nirmala et al., 2007
Dendrocalamus sikkimensis F 91.24 1.88 1.86 5.20 1.31 2.99 0.60 0.76 Premlata et al., 2015
Dendrocalamus strictus F 90.10 2.60 3.07 2.26 0.31 6.17 0.33 0.71 Nirmala et al., 2007
Dendrocalamus strictus F 85.98 1.98 – – – 9.94 – – Bhatt & Singh, 2005
Dendrocalamus yunnanicus F 90.44 1.61 – 0.67 – – 0.29 0.46 Chen et al., 2018
Dendrocalamopsis oldhami F 91.75 2.23 1.76 0.77 – – 0.48 0.86 Zhu & Luo, 2012
Gigantochloa albociliata F 89.23 3.05 3.52 4.15 0.32 4.59 0.51 0.73 Nirmala et al., 2007
Gigantochloa rostrate F 90.56 3.56 3.17 4.10 0.22 4.32 0.56 0.68 Nirmala et al., 2007
Melocanna bambusoides F 91.22 3.29 – – – 3.93 – – Bhatt & Singh, 2005
Phyllostachys praecox F 91.25 3.25 – 1.37 1.75 – 0.21 0.85 Zhang, Ji, Hu, Chen, & Ye, 2011
Bambusa bamboos F10 90.80 2.32↓c 2.52↓c 9.64↑c 0.20↓c 2.57↓c 9.64↑c 1.22↓c Nirmala et al., 2007
Bambusa tulda F10 89.33↑c 2.49↓c 2.21↓c 10.69↑c 0.35↓c 2.58↓c 10.69↑c 0.74 ↓c Nirmala et al., 2007
Dendrocalamus asper F10 90.20 2.17↓c 2.08↓c 10.85↑c 0.26↓c 4.46↓c 10.85↑c 0.94 Nirmala et al., 2007
Dendrocalamus giganteus F10 91.08 2.60↓c 2.23↓c 13.84↑c 0.46↓c 5.02 13.84↑c 0.74↓c Nirmala et al., 2007
Dendrocalamus hamiltonii F10 92.60 2.56↓c 2.11↓c 8.20↑c 0.41↓c 2.98↓c 8.20↑c 0.84 Nirmala et al., 2007
Pleioblastus amarus FH – 2.42 0.40 12.18d (C + L) 0.42 0.86 1.34 – Guo, Yang, Lin, Chen, & Yang, 2019
Pleioblastus amarus FM – 2.34 0.35 8.04d (C + L) 0.60 1.00 1.92 – Guo et al., 2019
Pleioblastus amarus FL – 2.70 0.40 10.67d (C + L) 0.56 1.00 1.53 – Guo et al., 2019
Bambusa balcooa D 86.30 3.30 – 26.40 – 5.20 1.00 3.10 Bhatt & Singh, 2005
Bambusa tulda D 92.80 3.40 – 24.60 – 4.70 0.90 3.00 Bhatt & Singh, 2005
Bambusa nutans D 94.70 3.30 – 28.50 – 4.90 1.00 2.70 Bhatt & Singh, 2005
Dendrocalamus giganteus D 92.00 2.80 – 27.60 – 4.90 0.80 2.20 Bhatt & Singh, 2005
Dendrocalamus hamiltonii D 93.60 3.70 – 25.40 – 4.80 0.90 2.80 Bhatt & Singh, 2005
Dendrocalamus hookerii D 93.50 3.40 – 34.70 – 4.50 1.00 3.20 Bhatt & Singh, 2005
Dendrocalamus latiflours D – 24.50 – 19.30 – 28.50 3.80 – Lu, 2007
Dendrocalamus longispathus D 91.70 2.60 – 26.70 – 4.70 0.60 3.40 Bhatt & Singh, 2005
Dendrocalamus sikkimensis D 92.40 3.10 – 23.50 – 4.70 – 2.60 Bhatt & Singh, 2005
Melocanna baccifera D 93.00 2.40 – 35.50 – 4.50 – 2.30 Bhatt & Singh, 2005
Phyllostachys bambusoides D 85.60 3.70 – 23.10 – 4.80 – 2.10 Bhatt & Singh, 2005
Phyllostachys bambusoides D – 28.15 21.57 – – – – – Xu et al., 2005
Phyllostachys heteroclada D – 20.10 12.90 – – – – – Xu et al., 2005
Phyllostachys iridenscens D – 22.14 18.75 – – – – – Xu et al., 2005
Phyllostachys pubescens D – 19.01 12.70 – – – – – Xu et al., 2005
Phyllostachys pubescens D 84.24 1.59 – 3.37 – 22.50 1.77 10.35 Lv, 2013
Phyllostachys praecox D – 30.66 19.60 – – – – Xu et al., 2005
Phyllostachys praecox D 90.62 2.85 – 1.27 – 19.20 1.51 5.93 Lv, 2013
Phyllostachys praecox D – 28.80 – 8.20 – 30.40 2.70 Lu, 2007
Phyllostachys tianmuensis D 91.36 2.41 – 1.19 – 15.90 1.85 9.76 Lv, 2013
Pleioblastus sulphurea D – 20.37 8.21 – – – – – Xu et al., 2005
Pleioblastus amarus D – 29.60 – 9.40 – 29.6 3 – Lu, 2007
Teinostachyum wighni D 78.60 3.60 – 23.7 – 4.90 – 3.70 Bhatt & Singh, 2005
Phyllostachys pubescens DT – 19.50 – 21.30 – 37.90 2.80 – Lu, 2007
Phyllostachys pubescens DB – 27.80 – 9.40 – 31.30 3.20 – Lu, 2007
Yushania alpine DT 92.20 33.40 – 23.90 – 23.60 2.00 17.10 Karanja et al., 2015
Yushania alpine DB 92.40 33.00 – 30.70 – 17.30 2.00 17.00 Karanja et al., 2015

a
F, F10 and D refer to the fresh, 10-day-old fresh and dry bamboo shoots, respectively. FH, FM and FA indicate the altitude of bamboo shoot growth, which are
high altitude, medium altitude and low altitude respectively. DT and DB are the tip and basal portions of dry bamboo shoots.
b
FAA and TDF are free amino acid and total dietary fiber, respectively.
c
↓ or ↑ followed the data of 10-day-old fresh bamboo shoots attested that these values are lower or higher than the corresponding values of the newly-emerging
fresh bamboo shoots.
d
C and L are cellulose and lignin, respectively.

3
Y. Wang, et al.
e
– means not determined.
2.1. Proteins, amino acids and peptides
Bamboo shoot is a potential source for proteins for human beings
with protein levels ranging from 1.49 g/100 g to 4.04 g/100 g and
21.1 g/100 g to 25.8 g/100 g on wet and dry weight basis, respectively
(Table 3). However, it must be noted that the protein levels reported by
Bhatt and Singh (2005) (2.6–3.7 g/100 g db) and Lv (2013)
(1.59–3.78 g/100 g db) were somewhat abnormal and much lower than
those reported by others (19.01–33.4 g/100 g db) on dry basis. Ob-
viously, the protein level in bamboo shoot is highly depended on the
bamboo species and the maturity. Within the available data, the highest
protein level on wet basis was found with the fresh bamboo shoots from
D. giganteus (3.86 g/100 g wb) and the one on dry basis was from Y.
alpine (33.4 g/100 g db) (Karanja et al., 2015; Nirmala, David, &
Sharma, 2007). In most cases, the protein level decreases with the age
of bamboo shoot, but a protein summit was also evidenced with some
species. For example, the shoot from B. tulda showed an increase in
protein content (0.51 → 1.29 g/100 g) in the first 14 days after the
emerging from the ground and followed a decrease (1.29 → 0.89 g/
100 g) until 20 days (Pandey & Ojha, 2013). Normally, the nutrients
were not evenly distributed in fresh bamboo shoots and a vertical dis-
tribution difference was often found from the tip to the basal. As re-
ported by Lin, Chen, Zhang, and Brooks (2018), the basal portion
(19.5 g/100 g) presented a lower protein level than the middle and tip
portions (27.8 g/100 g). Interestingly, it had been concluded that the
predominant proteins in bamboo shoots present a low molecular weight
(20.10–15.50 kDa). They were identified as the histone-like related
proteins, which mainly construct the nucleosome core and play an
important role in defense, stress and development (Waikhom,
Bengyella, Roy, & Talukdar, 2015). In addition, an antifungal protein,
designated dendrocin, was found in the shoots of D. latiflora Munro. It
has a molecular weight of 20 kDa and displayed strong inhibitory ca-
pacities on the mycelial growth of Botrytis cincerea, Fusarium oxysporum,
and Mycosphaerella arachidicola, indicated by the lower IC50 values of
1.8 μM, 1.4 μM and 5.1 μM, respectively. Structurally, the first fourteen
N-terminal amino acid residues of dendrocin are highly homological to
thaumatin-like proteins. Unlike thaumatin-like proteins, dendrocin was
devoid of ribonuclease and hemagglutinating activities (Wang & Ng,
2003).
As for the composition of bamboo shoot proteins, seventeen amino
acids were founded as arginine (Arg), serine (Ser), aspartic acid (Asp),
glutamic acid (Glu), glycine (Gly), threonine (Thr), alanine (Ala), pro-
line (Pro), methionine (Met), histidine (His), valine (Val), cysteine
(Cys), phenylalanine (Phe), leucine (Leu), isoleucine (Ile), lysine (Lys)
and tyrosine (Tyr). All eight or at least seven essential amino acids (Lys,
Ser, Met, His, Ile, Leu, Phe) for human beings are available in bamboo
shoots. In contrast to other vegetables, bamboo shoots presented ex-
tremely higher values in free amino acid content up to 4.03 g/100 g wb
in the fresh shoots from D. hamiltonii. More importantly, for free amino
acids of bamboo shoots, the fraction of overall essential amino acids
accounted for more than 30% in most cases and even up to 49.01%,
which was much higher than that of other common vegetables and
comparable to that of fish (44.80–45.92%) (Xu, Cao, Song, & Fang,
2005). On one hand, Tyr, the trace amino acid in common vegetables,
was evidenced as the most abundant amino acid in bamboo shoots, such
as these from D. hamiltonii (163.8 mg/100 g wb), P. pubescens. pubescens
(635–1100 mg/100 g wb; 21.4 g/kg db), P. sulphurea (10.5 g/kg db)
and P. phypramiens (28.3 g/kg db). On the other hand, Lys was con-
cluded as the first limiting amino acid of bamboo shoots in most cases.
As estimated by Xu et al. (2005), the amino acid score of six common
bamboo shoots located in the range of 26.5–90.3%. Besides their nu-
tritional implications, free amino acids are the key taste narrators of
bamboo shoots. Regarding their tastes, three groups of amino acids
were established, namely sweet amino acids (Gly, Ala, Pro, Ser), bitter
4
Journal of Functional Foods 71 (2020) 104015
amino acids (Val, Ile, Leu, Tyr, Phe, Try) and umami amino acids (Asp
and Glu). The results reported by Gao et al. (2019) concluded that the
bitterness of some bamboo shoots is largely due to L-phenylalanine,
uridine, L-ornithine, L-tryptophan, and adenine, with L-phenylalanine
being the greatest contributor. However, the results of Zhang, Ding,
Cui, Bai, and Cai (2017) uncovered that the bitter amino acids were not
the key determinators of the bitter and astringent taste of some bamboo
shoots, which was originated from the soluble tannin. The strong,
moderate and mild bitter sensations were precepted as the soluble
tannin in the range of > 3.0 mg/g, 3.0–1.2 mg/g and < 1.2 mg/g,
respectively.
As for the information on the peptides from bamboo shoots, only
two articles were available. It was founded that the water extract (the
mixture of boiled water, filled liquid and squeezed juice) from the
shoots of P. pubescens displayed angiotensin converting enzyme in-
hibitory (IC50 588.29 μg/mL) and antioxidant abilities (Liu, Liu, Lu,
Chen, & Zhang, 2013). A dipeptide, Asp-Tyr, was identified as the key
compound responsible for the angiotensin-converting enzyme in-
hibitory activity of bamboo shoots with an IC50 value as lower as
32.31 μg/mL. Fujimura, Ideguchi, Minami, Watanabe, and Tadera
(2005) isolated two antimicrobial peptides, designated Pp-AMP1 and
Pp-AMP2, from P. pubescens by using chitin affinity chromatography,
which has a high degree of homology with mistletoe toxins. In addition,
they displayed the common features of plant defensins and highly basic
with isoelectric points of 11.5 and 11.6, respectively. Structurally, Pp-
AMP1 presented a molecular mass of 4694 Da and composed of 44
amino acids with four cysteine residues and two disulfide bonds. Pp-
AMP2 presented a molecular mass of 4916 Da and composed of 45
amino acids with six cysteine residues and three disulfide bonds. Both
Pp-AMP1 and Pp-AMP2 demonstrated strong inhibitory activities
against the bacteria of Erwinia carotovora, Agrobacterium radiobacter,
Agrobacterium rhizogenes, Clavibacter michiganensis, Curtobacterium
flacccumfaciens, with IC50 values varied from 13 μg/mL to 25 μg/mL.
They inhibited more efficiently against the fungi of Fusarium oxysporum
and Geotrichum candidum at a rather low IC50 value of 2 μg/mL.
2.2. Carbohydrates
In view of total carbohydrates, polysaccharides, oligosaccharides
and monosaccharides were found in bamboo shoots. Generally, the
polysaccharides in bamboo shoots mainly include cellulose, hemi-
cellulose and starch as well as some other minor complex poly-
saccharides such as glycoproteins (Tanabe, Furuta, Maeda, & Kimura,
2017). Specifically, the hemicellulosic polysaccharides from bamboo
shoots mainly consisted of xyloglucan, β-D-glucan, arabinogalactan,
arabinoxylan and glucomannan. The major oligosaccharides in bamboo
shoots were identified as sucrose, arabinoxylan trisaccharide and tet-
rasaccharide and xyloglucan disaccharide. These non-sucrose oligo-
saccharides predominantly presented in feruloylated form with a minor
fraction in p-coumaroylated form (Edashige & Ishii, 1998; Ishii & Hiroi,
1990a, 1990b). In this sense, bamboo shoot is a good source of anti-
oxidant dietary fiber. Regarding the monosaccharides, fructose and
glucose were frequently found in bamboo shoots (Karanja, 2017). For
the distribution of fructose, glucose and sucrose in shoot body, different
results were reported. The study by Kozukue, Kozukue, and Kurosaki
(1983) revealed that, for the fresh shoots from P. pubestens, the levels of
fructose (218 mg/100 g wb), glucose (216 mg/100 g wb) and sucrose
(211 mg/100 g wb) were comparable in the tip quarter portion, while
fructose (597 mg/100 g wb) and glucose (489 mg/100 g wb) were three
times more than sucrose (151 mg/100 g wb) in the bottom quarter
portion. However, for the fresh shoots from Y. alpine, fructose (2.19 g/
100 g db) was much higher than glucose (0.65 g/100 g db) and sucrose
(0.68 g/100 g db) in the upper portion and only small differences were
observed the levels of fructose (0.62 g/100 g db), glucose (0.77 g/100 g
Y. Wang, et al.
db) and sucrose (0.58 g/100 g db) in the lower portion (Karanja, 2017).
However, it must be noted that the term “carbohydrate” used in
most research articles did not mean the sum of polysaccharides, oli-
gosaccharides and monosaccharides, but it varied article to article and
often pointed to the soluble carbohydrates, digestible carbohydrates or
reducing sugars as reflected by its measuring regimes. The carbohy-
drate content of bamboo shoots ranged from 2.0 g/100 g to 9.94 g/
100 g in common species of freshly emerged juvenile bamboo shoots
(Singhal et al., 2013). The carbohydrate level of bamboo shoots is
species specific and determined as 1.90 g/100 g wb, 5.10 g/100 g wb
and 9.94 g/100 g wb for Y. alpine, D. giganteus and D. strictus, respec-
tively (Bhatt & Singh, 2005; Karanja, 2017; Nirmala, Sharma, & David,
2008).
As reported by Felisberto, Miyake, Beraldo, and Clerici (2017),
starch is an energy source for the growth of bamboo shoots. As showed
in Table 3, starch is widely distributed in various bamboo shoots but its
level varies species to species. In fresh form, the shoot from P. praecox
gave the highest starch content up to 1.75 g/100 g wb, while the one
from the B. nutans was observed with the lowest starch content as
0.21 g/100 g wb. In dry form, the starch levels in the shoots of D. asper,
B. tuldoides and B. vulgaris were determined as 8.17–10.99, 1.75–2.65
and 15.30–16.89 g/100 g db. The starches isolated from the shoots of B.
tuldoides displayed similar apparent amylose to cereal starches, which
varied from 19.26 g/100 g db to 33.35 g/100 g db in bottom, middle
and top portions. The starch granules are in polyhedral shape and small
size (average 4.64 μm), which displayed an A-type crystalline pattern
with relative crystallinity around 25% (Felisberto et al., 2017). Similar
results were obtained for the starch from the roots of D. asper
(Felisberto, Beraldo, & Clerici, 2017).
Actually, bamboo shoot is a rich source of dietary fiber. The dietary
fiber in bamboo shoots is mostly insoluble, which accounts for 60–90%
of their total carbohydrate (Felisberto et al., 2017; Karnjanapratum
et al., 2019). On the dry mass basis, the total dietary fiber in bamboo
shoots varied from 19.3 g/100 g to 35.5 g/100 g. Among all bamboo
shoots, the one from M. baccifera demonstrated the highest values in
dietary fiber of 35.5 g/100 g db (Satya, Singhal, Bal, & Sudhakar,
2011). NDF (Nutrient dietary fiber), ADF (Acid detergent fiber), lignin,
hemicellulose and cellulose were identified as the major components of
the dietary fiber in bamboo shoots and their levels in the fresh shoots of
D. giaganteus were 2.645 g/100 g, 2.105 g/100 g, 0.56 g/100 g,
0.495 g/100 g, and 1.589 g/100 g wb, respectively (Nirmala et al.,
2008). It was well known that the dietary fiber in fresh bamboo shoots
sharply increased upon its growth and storage. A 10-day storage re-
sulted in double increase in dietary fiber content of fresh bamboo
shoots (Nirmala et al., 2008). In addition, the dietary fiber content also
varied in the different parts of bamboo shoots. For example, the dietary
fiber in the upper and the lower portions of the shoots from Y. alpina
was 23.9 g/100 g db and 30.7 g/100 g db, respectively (Karanja, 2017).
However, the results of Singhal et al. (2013) showed that the top and
basal portions of the shoots from D. giganteus were comparable and
determined as 0.96 g/100 g wb and 0.97 g/100 g wb, respectively.
More importantly, the level of dietary fiber and its composition is the
most critical determinants for the mouthfeel and taste of bamboo
shoots. The older bamboo shoots always presented higher total dietary
fiber, higher insoluble dietary fiber, higher lignin and lower eating
quality.
2.3. Fat
In general, bamboo shoots are characterized of a substantially low-
fat content ranging from 0.3 g/100 g to 3.97 g/100 g wb and thus were
considered a health food item for the overweight, diabetes, and hy-
perlipidemia (Karanja, 2017; Satya et al., 2011). More importantly,
bamboo shoots are rich in non-polar lipids, glycolipids and phospholi-
pids with a ratio of 17:27:56 (Singhal et al., 2013). Triglycerides are
predominant in non-polar lipid fraction, while the glycolipid fraction is
5
Journal of Functional Foods 71 (2020) 104015
mainly composed of phosphatidylcholine and phosphatidylethanola-
mine. There are nine components in bamboo shoot phospholipids, the
major being phosphatidylcholine and phosphatidylethanolamine. The
fatty acids in bamboo shoot oil mainly predominated by linoleic acid
(28.14 g/100 g), linolenic acid (17.57 g/100 g), palmitic acid (7.35 g/
100 g), oleic acid (6.42 g/100 g) and stearic acid (2.46 g/100 g). The
overall contents of unsaturated fatty acid and saturated fatty acid were
53.97 g/100 g and 16.07 g/100 g, respectively (Lu et al., 2010). It must
be noted that the fat content and composition in bamboo shoot were
age-dependent and unevenly distributed in tip, middle and basal parts.
In contrast to newly-emerged bamboo shoots, sharp increases in total
fat content were observed upon a ten-day growth for the shoots from B.
bamboos (3.53 → 9.64 g/100 g), B. tulda (3.97 → 10.69 g/100 g), D.
hamiltonii (3.90 → 8.20 g/100 g), D. giganteus (2.64 → 13.84 g/100 g)
and D. asper (3.54 → 10.85 g/100 g). However, in a single bamboo
shoot of P. pubescens, the total fat sharply decreased from the tip
(800 mg/100 g) to the basal (379 mg/100 g) (Kozukue & Kozukue,
1981).
2.4. Minerals
The available results have revealed that bamboo shoots are good
sources of both macro elements and microelements. Potassium (K),
phosphorus (P), sodium (Na), calcium (Ca) and magnesium (Mg) are
the major components for the macro elements, while the micro ele-
ments mainly included cobalt (Co), copper (Cu), nickel (Ni), manganese
(Mn), selenium (Se), iron (Fe) and zinc (Zn) (Christian et al., 2015;
Nirmala et al., 2011). In most study cases, K was determined as the
richest macro element in various bamboo shoots and followed by P and
Mg. All of these macro elements presented very large variations among
bamboo species. The highest and lowest K contents were evidenced as
19.95 mg/100 g wb (F. yunnanensis) to 576 mg/100 g wb (B. bambos),
respectively. The content of P and Mg were located in the ranges of
3.48–62.9 mg/100 g wb and 0.73–19.0 mg/100 g wb, respectively.
Occasionally, Na and Ca were identified as the third richest macro
element in B. blumeana and D. hamiltonii, respectively (Table 4). How-
ever, the results reported by Bhatt and Singh (2005) were distinctive, in
which Ca (1.2–1.9 g/100 g db) and P (0.5–1.0 g/100 g db) were de-
termined as the first and second richest macro element in eleven
bamboo shoots, respectively. It must be noted that Na was not de-
termined in this study. As for microelements, the situation became more
complex. In twenty-five fresh bamboo shoots, the probabilities for Fe,
Zn, Mn, Cu and Se being the richest microelement were 14/25, 4/25, 5/
25, 1/25 and 0/25, respectively. The ranges of content of Fe, Zn, Mn,
Cu and Se were 0.10–3.37 mg/100 g wb, 0.03–1.41 mg/100 g wb,
0.02–2.48 mg/100 g wb, 0.02–14 mg/100 g wb and 0.3–1.0 μg/100 g
wb, respectively. The results of Nirmala et al. (2008) showed that the
levels of mineral elements decreased with the age of bamboo shoots.
The abundance of minerals elements in the tip part of the shoot from Y.
alpine was significantly higher than those in the basal part (Karanja
et al., 2015). However, the tip part of the shoot from P. edulis was found
with lower element levels than the whole shoot. Obviously, bamboo
shoots are good sources of minerals, especially demonstrated by their
higher levels of micro minerals than common vegetables such as spi-
nach, potatoes, tomatoes and peppers (Huang & Yang, 2006). However,
the levels of the minerals in bamboo shoots were highly depended on
the plant species and growing age and were uneven across the shoot
body.
2.5. Vitamins
Regarding vitamins, most of investigations have focused on vitamin
C (ascorbic acid) and vitamin E (tocopherol). Vitamin C and vitamin E
are closely bound to the antioxidant capacity in vivo, more importantly,
vitamin E works synergistically with vitamin C in enhancing the im-
mune functions of the body (Nirmala et al., 2018). In contrast, vitamin
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 4
Composition of minerals and vitamins (mg/100 g) in bamboo shoots.
Plant species Forma K P Ca Na Mg Fe Zn Cu Mn Sec VC VE Reference

Bambusa balcooa F d
- – – – – – – – – – 2.21 0.62 Nirmala et al. (2014)
Bambusa bambos F 576 30.12 0.36 10.06 5.38 2.99 0.57 0.28 0.47 0.3 1.90 0.61 Nirmala et al., 2007
Bambusa blumeana F 253 44.95 21.75 0.3 13.78 0.58 0.54 0.12 0.50 – – – Niu, Tang, Zhang, Zhao, &
Wang, 2017
Bambusa nutans F – – – – – – – – – – 1.19 0.47 Nirmala et al. (2014)
Bambusa tulda F 408 19.31 4.06 12.96 8.68 3.02 0.72 0.44 0.7 0.4 1.42 0.61 Nirmala et al., 2007
Bambusa tulda F 338 – – 21.11 – – – – – – – – Premlata et al., 2015
Bambusa ventricosa F 41.40 47.90 16.60 1.04 15.00 0.57 0.53 0.12 0.23 – – – Niu et al., 2017
Bambusa vulgaris F – – – – – – – – – – 4.8 0.52 Nirmala et al., 2011
Chimonobambusa F 96.50 61.20 19 – 16.40 0.92 0.79 14 1.01 – – – Wang et al., 2017
yunnanensis
Dendrocalamus asper F 464 40.95 5.51 10.14 10.14 3.37 0.85 0.32 0.41 1 3.30 0.91 Nirmala et al., 2007
Dendrocalamus fugongensis F 275.7 42.2 8.60 – 13.3 0.45 0.48 0.09 0.59 – – – Wang et al., 2017
Dendrocalamus giganteus F 288 15.90 6.80 8.22 10.09 2.43 1.09 0.56 0.34 0.5 3.28 0.69 Nirmala et al., 2007
Dendrocalamus giganteus F 103.3 38.6 6.30 – 12.3 0.96 0.53 0.13 0.28 – – – Wang et al., 2017
Dendrocalamus hamiltonii F 416 28.12 3.00 9.32 6.09 2.69 0.70 0.29 0.16 0.8 2.45 0.71 Nirmala et al., 2007
Dendrocalamus hamiltonii F 304 38.20 22.40 – 16.10 0.37 0.67 0.11 1.98 – – – Wang et al., 2017
Dendrocalamus hamiltonii F 293 40.80 16.40 – 19.80 0.47 0.66 0.12 1.09 – – – Wang et al., 2017
Dendrocalamus hamiltonii F – – 2.16 – – 1.19 – – – – – – Premlata et al., 2015
Dendrocalamus membranceus F 114.30 49.20 12.70 – 12.70 0.52 0.59 0.13 0.3 – – – Wang et al., 2017
Dendrocalamus membranceus F – – – – – – – – – – 1.58 0.65 Nirmala et al. (2014)
Dendrocalamus semiscandens F 215.40 44.50 7.30 – 12.40 0.48 0.55 0.10 0.72 – – – Wang et al., 2017
Dendrocalamopsis oldhami F – 43.45 8.63 – – 0.86 – – – – 7.30 0.22 Zhu & Luo, 2012
Fargesia yunnanensis F 19.95 3.48 1.02 0.33 1.03 0.17 0.66 0.14 0.44 – – – Niu et al., 2017
Phyllostachys heterocycla F 28.82 46.38 0.36 0.07 0.73 0.10 0.03 0.02 0.02 – – – Niu et al., 2017
Phyllostachys iridescens F 27.33 51.99 0.34 0.03 1.10 0.65 0.03 0.03 0.02 – – – Niu et al., 2017
Schizostach yumfunghomii F 217.30 39.40 8.60 – 13.60 0.78 1.03 0.17 0.78 – – – Wang et al., 2017
Schizostach F 162.90 62.90 9.50 – 19.00 0.95 1.41 0.17 2.48 – – – Wang et al., 2017
yumpingbianeusus
Bambusa bambos F10 566 29.00↓b 0.30↓b 3.60↓b 5.20 1.31↓b 0.49↓b 0.20↓b 0.24↓b 0.3 1.31↓b 0.25↓b Nirmala et al., 2007
Bambusa tulda F10 398 18.70 2.42↓b 3.60↓b 7.50↓b 1.58↓b 0.54↓b 0.20↓b 0.12↓b 0.3↓b 1.0↓b 0.24↓b Nirmala et al., 2007
Dendrocalamus asper F10 460 29.08↓b 1.68↓b 4.42↓b 8.20↓b 2.52↓b 0.83 0.14↓b 0.16↓b 1.0 2.12↓b 0.42↓b Nirmala et al., 2007
Dendrocalamus giganteus F10 275 15.01↓b 1.47↓b 3.64↓b 9.57↓b 1.23↓b 0.56↓b 0.12↓b 0.16↓ 0.4↓b 2.15↓b 0.24↓b Nirmala et al., 2007
Dendrocalamus hamiltonii F10 210↓b 27.56 2.05↓b 3.64↓b 5.97↓b 0.76↓b 0.69 0.28 0.15 0.8 1.79↓b 0.31↓b Nirmala et al., 2007
Bambusa balcooa D 30 900 1400 – 40 – – – – – 27.74 – Bhatt & Singh, 2005
Bambusa nutans D 30 900 1500 – 40 – – – – – 100.00 – Bhatt & Singh, 2005
Bambusa tulda D 20 700 1300 – 40 – – – – – 105.56 – Bhatt & Singh, 2005
Chimonobambusa D 290 – 25.49 0.21 12.98 0.93 0.76 0.18 0.97 1.80 28.15 Gou et al., 2010
hejiangensis
Chimonobambusa utilis D 290 – 13.88 0.13 10.24 0.60 0.79 0.18 0.59 1.90 32.15 Gou et al., 2010
Chimonobambusa angustifolia D 250 – 18.24 0.25 11.97 1.12 0.73 0.23 0.76 2.00 24.56 Gou et al., 2010
Dendrocalamus giganteus D 30 700 1200 50 – – – – – 85.00 – Bhatt & Singh, 2005
Dendrocalamus hamiltonii D 20 600 1500 – 40 – – – – – 201.56 – Bhatt & Singh, 2005
Dendrocalamus hookerii D 20 1000 1600 – 40 – – – – – 152.31 – Bhatt & Singh, 2005
Dendrocalamus longispathus D 20 600 1300 – 40 – – – – – 63.86 – Bhatt & Singh, 2005
Dendrocalamus sikkimensis D 20 700 1900 – 40 – – – – – 39.47 – Bhatt & Singh, 2005
Melocana baccifera D 20 800 1400 – 40 – – – – – 108.57 – Bhatt & Singh, 2005
Phyllostachys bambusoides D 20 700 1300 – 40 – – – – – 42.36 – Bhatt & Singh, 2005
Phyllostachys edulis D 3503 – 286.01 2.58 132.78 9.25 5.98 0.69 33.51 – – – Huang et al., 2014
Phyllostachys nidularia D 530.54 – 23.62 2.12 9.84 0.62 0.54 0.12 0.18 – 4.23 – Chen, Gou, Sun, & Wang, 2017
Pleioblastus amarus D 110 – 5.01 0.23 10.65 2.50 0.54 0.08 0.21 240 6.00 0.05 Dang et al., 2017
Pleioblastus maculatus D 457.30 – 7.73 – 4.49 6.75 1.49 0.22 0.07 1.66 8.89 – Wang, Ma, & Gou, 2014
Teinostachyum wightii D 20 500 1300 – 40 – – – – – 30.84 – Bhatt & Singh, 2005
Phyllostachys edulis DT 2678 – 82.99 2.23 73.19 12.92 4.39 0.50 35.11 – – – Huang et al., 2014
Yushania alpina DT 3590 763 267 – 430 2.62 5.21 1.23 2.98 – 7.82 – Karanja et al., 2015
Yushania alpina DB 2760 481 106 – 127 2.22 3.81 0.37 2.03 – 5.12 – Karanja et al., 2015
Chimonobambusa D1020 220 – 15.66 0.22 11.32 0.57 0.78 0.17 0.76 1.60 28.69 – Gou et al., 2010
hejiangensis
Chimonobambusa D873 290 – 21.45 0.23 11.95 1.04 0.72 0.15 1.46 1.60 28.69 – Gou et al., 2010
hejiangensis
Chimonobambusa D780 270 – 19.56 0.24 11.69 0.50 0.76 0.22 0.75 2.20 25.12 – Gou et al., 2010
hejiangensis

a
F and F10 refer to the fresh and 10-day-old bamboo shoots and their values were expressed on wet basis. D, DT and DB are whole, tip and basal portions of
bamboo shoots and their values were expressed on dry basis. D1020, D873 and D780 indicate the altitude of bamboo shoot growth at 1020 m, 873 m and 780 m and
their values were expressed on dry basis.
b
↓ or ↑ followed the data of 10-day-old fresh bamboo shoots attested that these values are lower or higher than the corresponding values of the newly-emerging
fresh bamboo shoots.
c
Se in μg/100 g.
d
– means not determined.

6
Y. Wang, et al.
C is more abundant than vitamin E in fresh bamboo shoots, which was
also the fact in other common vegetables. On a wet basis, the levels of
vitamin C and vitamin E were in the ranges of 1.0–7.3 mg/100 g wb and
0.22–0.91 mg/100 g wb, respectively. On a dry basis, the level of vi-
tamin C ranged from 4.23 mg/100 g db to 201.56 mg/100 g db. Re-
grettably, the level of vitamin E was scarcely reported on a dry basis. In
addition, fresh bamboo shoots are a reasonable source of B-group vi-
tamins and β-carotene for certain locations. For example, in the shoots
of P. amarus, although vitamin A was not found, vitamin B1, vitamin
B2, vitamin B6 and β-carotene were determined at 0.09 mg/100 g db,
0.04 mg/100 g db, 0.38 mg/100 g db and 0.65 μg/100 g db, respec-
tively (Dang, Chen, & Yin, 2017). In the shoots from D. oldhami, the
levels of vitamin B1, vitamin B2 and vitamin B3 were determined as
1.7 mg/kg wb, 0.7 mg/kg wb and 3.5 mg/kg wb, respectively (Zhu &
Luo, 2012). Obviously, the giant variations were evidenced in the levels
of vitamins in bamboo shoots both on the wet and dry basis (Table 4).
The level of vitamins in various shoots is a multi-factor-depended event,
such as the species of the bamboo plant and the age of the shoot. As for
vitamin C in newly-emerged shoots on the wet basis, the highest value
was found with D. oldhami (7.3 mg/100 g wb) and the lowest value was
tested with B. nutans (1.19 mg/100 g wb) (Nirmala et al., 2008). The
study by Nirmala et al. (2007) uncovered that both the levels of vitamin
C and vitamin E sharply decreased with the age of the shoots from B.
bambos, B. tulda, D. asper, D. giganteus and D. hamiltonii. In depleting
percentage, vitamin E (59.70%, 60.46%, 53.85%, 65.51%, 56.56%)
decreased much rapidly than vitamin C (30.89%, 29.58%, 33.75%,
34.45%, 26.94%). In addition, bamboo shoots from different portions
(tip and basal) and altitude also had different degrees of influence on
vitamin C content. Similar to the effects of bamboo age, the tip portion
of the shoot from Y. alpina presented a higher level in vitamin C
(7.82 mg/100 g db) than the basal portion (5.12 mg/100 g db) (Karanja
et al., 2015). However, for the shoot from Y. alpine, the β-carotene in
the tip portion (9.9 μg/g db) was much lower than that in the basal
portion (15.7 μg/g db). Regarding the growing altitude, the previous
research revealed that vitamin C (25.12–28.69 mg/100 g db) in the
shoots from C. hejiangensis was independent on the altitude
(780–1020 m) (Gou, Ding, Yang, Li, & Jiang, 2010).
2.6. Phytosterols
Phytosterols, namely plant sterols, are a family of lipophilic sterols
naturally occurred in plant kingdom. More than 200 different plant
sterols have been identified in nature. Structurally, they are 28- or 29-
carbon alcohols derived from 5α-cholesten-3β-ol and functionally re-
sembled cholesterol in vertebrates. Phytosterols, a sub-group of tri-
terpenes, share a tetracyclic cyclopenta (a) phenanthrene structure with
a side chain at C17 and two methyl moieties C18 and C19 positions.
Nowadays, phytosterols were considered as valuable dietary supple-
ments due to their diverse health benefits such as serum cholesterol-
lowering, anti-ulcer, anti-cancer, anti-inflammatory and im-
munomodulation (Ogbe, Ochalefu, Mafulul, & Olaniru, 2015). As the
secondary metabolites, phytosterols are widespread in plants and
bamboo shoot was proved as a good source of phytosterols. However, to
the best of our knowledge, up to seventeen phytosterols were detected
in bamboo shoots (Lachance & He, 1998), although only part of them
was identified. Generally, six phytosterols, including β-sitosterol, cam-
pesterol, stigmasterol, cholesterol, ergosterol and stigmastanol, were
often confirmed in bamboo shoots (Fig. 1). On a dry basis, the level of
total phytosterol in bamboo shoots ranged from 66.60 mg/100 g db to
242.77 mg/100 g db (Table 5), which indicated the potential of bamboo
shoots in supplying functional phytosterols for human beings. As the
cases of other common vegetables, β-sitosterol (79.98–83.34%) was the
most abundant phytosterol in bamboo shoots followed by campesterol
(10.11–12.36%) and stigmasterol (4.84–5.74%). The levels of phytos-
terols in bamboo shoots depended on the plant species, parts and har-
vest seasons. The results of Lu, Ren, Zhang, and Gong (2009) showed
7
Journal of Functional Foods 71 (2020) 104015
the total phytosterol was in the plant species order of P. pubescens
(279.6 mg/100 g db) > P. amarus (221.7 mg/100 g db) > P. praecox
(165.4 mg/100 g db) > D. latiflorus (112.4 mg/100 g db) and the
phytosterol composition was independent of the plant species. As for
the distribution of phytosterols in bamboo shoots, the shell was found
with higher total phytosterol than the shoot and in the bamboo shoots,
the total phytosterol increased from the basal to the tip. For the shoots
from P. pubescens, the one harvested in summer presented a lower total
phytosterol (195.3 mg/100 g db) than the others harvested in spring
(279.6 mg/100 g db) and winter (226.2 mg/100 g db). Due to their
lipophilic nature, the phytosterols were enriched in extracted bamboo
shoot oil. In the shoot oil obtained by supercritical CO2 extraction, the
total phytosterol was 28.7 g/100 g with β-sitosterol as the predominant
(86%, w/w).
2.7. Phenols
As known to all, the phenolic compounds in plant foods demonstrate
diverse benefits in maintaining the health of human beings especially
due to their excellent anti-oxidant activities. Bamboo shoot was one of
the plant foods rich in phenols. Generally, gallic acid was applied as the
reference in the measurement of the total phenol in bamboo shoots and
the results were often expressed as milligrams of a gallic acid equivalent
(GAE). As showed in Table 5, the total phenol varied largely in different
bamboo shoots. On a wet basis, the lowest value was found with B.
pallida (79.85 mg GAE/100 g wb) and the highest value was concluded
for P. violascens (2541.02 mg GAE/100 g wb). In composition, the
phenols in bamboo shoots were mainly composed of phenolic acids and
flavonoids. On a dry basis, the total phenol and total flavonoid in P.
pubescens were determined as 5.48 mg GAE/g db and 1.3 mg QE/g db,
respectively. For phenolic acids, protocatechuic acid, p-hydroxybenzoic
acid, catechin, caffeic acid, chlorogenic acid, syringic acid, p-coumaric
acid, ferulic acid, gallic acid, vanillic acid were reported in bamboo
shoots (Pandey & Ojha, 2013; Park & Jhon, 2010) (Fig. 1). Among
them, protocatechuic acid, p-hydroxybenzoic acid and syringic acid
were the most abundant compounds (Nirmala et al., 2011). However, it
must be noted that, besides the plant species, the level of total phenol
and its composition is bamboo shoot age dependent, but the de-
pendences are compound specific. With the growth of the shoots from
D. asper, D. strictus and B. tulda, caffeic acid in all shoots increased.
Gallic acid remained constant in D. asper, gradually increased in D.
strictus and first increased then decreased in B. tulda. Vanillic acid
displayed an increasing trend in D. asper and D. strictus but showed an
opposite trend in B. tulda. Regrettably, the flavonoid monomers in
bamboo shoots were not elaborately characterized. In contrast, flavo-
noids such as orientin, isoorientin, isovitexin, vitexin and tricin were
identified in bamboo shoot and leaves (Liu, Zhang, Liu, & Duan, 2019)
(Fig. 1). Flavonoids mainly existed in the insoluble form of free agly-
cone or flavonoid ligand in different bamboo tissues including shoots,
sheaths and leaves (Su et al., 2019). As a by-product of bamboo shoot
processing, the bamboo shell was also plenty of phenols with the
highest total phenolic content recorded as 85.3 mg GAE/g db. In detail,
fifteen phenolic acids and seven flavonoids were identified in the
bamboo shell of P. pracecox. The phenolic acids included 3-O-caffeo-
lyshikimic acid, chlorogenic acid, p-coumaric acid, 3-p-coumar-
oylquinic acid, 5-p-coumaroylquinic acid, cryptochlorogenic acid, 1,3-
dicaffeoyl quinic acid, 3,5-dicaffeoyl quinic acid, ferulic acid, 3-O-fer-
uloylquinic acid, 5-O-feruloylquinic acid, gallic acid, p-hydroxybenzoic
acid, neochlorogenic acid and syringic acid. The seven flavonoids were
apigenin 6,8-di-C-α-L-arabinopyranoside, 6-C-β-D-glucopyranosyl-8-C-
α-L-arabinopyranosylchrysin, kaempferide 3-O-α-L-rhamnopyranosyl
(1,6)-β-D-glucopyranoside, narcissin, rutin, schaftoside and 5,7,4′-tri-
hydroxy-3′,5′-dimethoxyflavone. Quantitatively, p-coumaric acid
(11.9 mg/100 g db), chlorogenic acid (8.7 mg/100 g db), rutin (3.9 mg/
100 g db) and ferulic acid (1.7 mg/100 g db) were the most abundant
compounds.
Y. Wang, et al.
Fig. 1. Chemical structure of phytosterols (a. β-sitosterol; b. cholesterol; c. ergosterol; d. stigmasterol; e. campesterol; f. stigmastanol), phenolic acids (g. catechin; h.
caffeic acid; i. protocatechuic acid; j. p-coumaric acid; k. p-hydroxybenzoic acid; l. ferulic acid; m. syringic acid; n. chlorogenic acid; o. gallic acid; p. vanillic acid) and
flavonoids (q. isovitexin; r. isoorientin; s. tricin; t. orientin; u. vitexin) in bamboo shoots.
3. Health benefits of bamboo shoots
3.1. Traditional therapeutic effects
Historically, bamboo shoot was treated as a traditional medicine up
to 2500 years especially in China and this was recorded in the famous
Chinese book of Compedium of Materica Medica (Sarkar et al., 2020).
Nowadays, the application of bamboo shoots in various medications
still prevails in some countries to improve digestion, alleviate hy-
pertension, prevent cancer and cardiovascular disease (Table 6). In
South-Asian countries, bamboo shoot has been used to treat paralysis,
sweat and relieve hypertension (Basumatary et al., 2017). In Myanmar,
some female Burmese migrant workers use bamboo shoots with tradi-
tional Chinese medicine to abort the fetuses (Singhal et al., 2013). In
India, fermented bamboo shoot was used together with the dried and
powdered meat of bamboo rat and crushed berries to treat dysentery
and diarrhea. The mixture of fermented bamboo shoots, the crushed
leaves of Allium porrum Linn and chilli was proved with an ability to
cure influenza. Bamboo shoot soup was effective to treat gastric ulcers
(Singhal et al., 2013). For Indo-Persian and Tibetan medications, the
silicious concretions in bamboo shoots were considered a good tonic for
respiratory disorders. In China, bamboo shoot juice benefits the diges-
tion of protein and a poultice of bamboo shoots was useful in cleaning
wounds and healing infections (Devi, Chakma, & Yenkokpam, 2017;
Panee, 2015). In Java, bamboo shoot juice had been used to treat
jaundice (Kumar, Kumari, Devi, Choudhary, & Sangeetha, 2017). In
Korea, bamboo salt was applied to treat internal maladies. Frankly,
although bamboo shoots were widely applied in traditional medica-
tions, their working mechanisms were not well established.
8
Journal of Functional Foods 71 (2020) 104015
3.2. Modern bioactive activities
Beyond the traditional nutrients, a large number of phytochemicals
were founded in various plant foods and they were founded to provide
extra protections for human body. These phytochemicals were often
called bioactive substances or functional components. As mentioned
early, bamboo shoot is also a reservoir of bioactive substances, such as
phytosterols, polyphenols, polysaccharides and dietary fibers. In view
of these key components, bamboo shoots were evidenced with anti-
oxidant, hypolipidemic, prebiotic, anti-diabetic, anti-obesity, anti-in-
flammatory and anti-hypertensive activities and anti-microbial activ-
ities (Table 7). The anti-oxidant activity of bamboo shoots mainly de-
rived from its ascorbic acid and phenolic compounds, which exhibits
strong prevention against lipid oxidation, excellent scavenging capa-
cities to various radicals and comparable reducing abilities to oxidative
compounds, such as Fe3+. Overall, the indigestible carbohydrates in-
cluding dietary fiber and non-starch polysaccharides were the key de-
terminants of the hypolipidemic, prebiotic, anti-diabetic and anti-obe-
sity activities of bamboo shoots. In achieving the hypolipidemic
activity, the bamboo shoot fiber reduced the serum or plasma TG, TC,
HDL-C, LDL-C, non-esterified fatty acid and resulted in a decrease in
atherogenic index. Interestingly, bamboo shoot oil displayed strong
hypolipidemic activity in terms of lowering serum TC, TG and athero-
genic index, which were realized by decreasing hepatic lipase and in-
creasing fecal cholesterol excretion. The hypolipidemic activity of
bamboo shoot oil was ascribed to its high phytosterol content. The
prebiotic activity of bamboo shoots was generated by the fibers or
polysaccharide components, who improved the structure of gut flora by
enhancing the proliferation of beneficial microorganisms such as B.
adolescentis, B. infantis, B. bifidum, L. acidophilus as well as increased the
production of short-chain fatty acids. Upon the ingestion of bamboo
shoot fiber, the sugar metabolism was improved substantially in diverse
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 5
The levels of total phytosterols, total phenols and cyanogen in various bamboo shoots.a.
Plant species Content Plant species Content Plant species Content

b
Phytosterols (mg/100 g db)
Bambusa balcooa 165.5, 190.00 Bambusa khasiana 242.77 Bambusa kingiana 150.27
Bambusa nutans 66.60, 164.20 Bambusa tulda 130.00, 144.73 Cephalostachyum pergracile 180.03
Dendrocalamus brandisii 232.13 Dendrocalamus flagellifer 139.37 Dendrocalamus giganteus 136.23, 150.00, 175.57
Dendrocalamus hamiltonii 190.00, 198.27, 233.20 Dendrocalamus latiflorus 112.40, 146.33 Dendrocalamus sericeus 202.00
Dendrocalamus strictus 180.63 Phyllostachys praecox 165.40 Phyllostachys pubescens 279.60
Pleioblastus amarus 221.70

Phenols (mg GAE/100 g wb)c

Bambusa balcooa 191.37, 101.65 Bambusa nutans 489.83, 275.36 Bambusa pallida 79.85
Bambusa tulda 443.97, 80.54 Dendrocalamus asper 580.00 Dendrocalamus latiflorus 612.24
Dendrocalamus giganteus 336.56, 347.27, 616.50, 222.40 Dendrocalamus membranaceus 302.73 Dendrocalamus strictus 271.23, 630.00
Dendrocalamus hamiltonii 586.36, 505.93, 88.23, 264.83 Dendrocalamus sikkimensis 450.29 Phyllostachys aureosulcata 2643.90
Phyllostachys aureosulcata 2417.48 Phyllostachys bissetii 2387.62 Phyllostachys flexuosa 2612.10
Phyllostachys humilis 1966.36 Phyllostachys iridescens 2435.92 Phyllostachys nigra 2356.34
Phyllostachys mannii 2164.82 Phyllostachys sulphurea 1940.28 Phyllostachys vivax 1652.44
Phyllostachys violascens 2541.02 Phyllostachys viridiglaucescens 2407.68 Phyllostachys pubescens 5480.00d

Cyanogens (mg/kg wb)

Bambusa bambos 110.00, 678.39 Bambusa balcooa 1108.32 Bambusa tulda 160.00, 1412.4
Bambusa jaintia 434.02 Bambusa mizoranmeana 670.03 Bambusa nutans 1709.66
Bambusa tulda 1412.40 Chimonobambusa callosa 31.68 Dendrocalamus asper 160.00, 431.00
Dendrocalamus calostachys 636.77 Dendrocalamus flagellifer 1893.67 Dendrocalamus giganteus 988.17
Dendrocalamus hamiltonii 733.92, 338.00 Dendrocalamus hookerii 1315.49 Dendrocalamus longispathus 1951.49
Dendrocalamus manipureanus 1347.98 Dendrocalamus membranaceus 514.80 Dendrocalamus sikkimensis 778.27
Dendrocalamus strictus 180.00, 1717.85, 763.00 Melocanna baccifera 285.12, 526.00 Phyllostachys mannii 36.32
Phyllostachys aurea 494.00 Phyllostachys bambusoides 618.00 Phyllostachys pubescens 366.00
Thyrsostachys oliveri 1097.71

a
Data in the table were reported from the references of Badwaik, Gautam, and Deka (2015), Bhardwaj, Sharma, Bishist, Navale, and Kaushal (2019), Ingudam and
Sarangthem (2016), Nemenyi et al. (2015), Nirmala, Bisht, Bajwa, and Santosh (2018), Nirmala, Bisht, and Laishram (2014), Pandey and Ojha (2014), Pokhariya,
Tangariya, Sahoo, Awasthi, and Pandey (2018), Bajwa et al. (2015) and Rawat et al. (2015).
b
wb: wet basis; db: dry basis.
c
GAE means gallic acid equivalent.
d
Data were expressed on the dry basis.

Table 6
Traditional therapeutic effects of bamboo shoots.
Country Application in traditional medications

China Treatment of respiratory (bamboo manna); infectious diseases (bambusa textilis Mc Clure); Cleaning wounds and ulcers infected by maggots (bamboo
shoot decoction); Treatment of gastric ulcer (bamboo soup); Co-protein digestion (bamboo pressed juice); Delayed delivery (bamboo shoots mixed with
palm-jaggery); Treatment of cough and heart disease (bamboo juice beers); Prevention and treatment of cardiovascular disease and cancer (Choudhury
et al., 2012; Karanja, 2017; Panee, 2015)
India Treatment of asthma and gastric ulcer (bamboo soup); Treatment of chickenpox and other skin diseases (bamboo sap); Control of hypertension and
cardiovascular disease (Choudhury, Sahu, & Sharma, 2012; Nirmala, Bisht, & Laishram, 2014; Nongdam &Tikendra, 2014)
Indonesia Treatment of jaundice (bamboo sap) (Choudhury et al., 2012)
Korea Treatment of internal disease (bamboo salt tablets); As based cosmetics (bamboo bath salts) (Choudhury et al., 2012)
Myanmar Treatment of gynecological diseases (bamboo decoction) (Nongdam & Tikendra, 2014)
South-Asian countries Treatment of paralysis and sweating; Control hypertension (Basumatary et al., 2017)
– As a biological preservative (bamboo pastes or liquids, as well as tablets) (Akinlabi, Anane-Fenin, & Akwada, 2017)
– Treatment of cardiovascular disease; Treatments intestinal diseases (Basumatary et al., 2017)
– Treatment of cold, resolves phlegm, fever, or loss of consciousness associated with phlegm-heat (bamboo sap) (Sangeetha, Diea, Chaitra, Malvi, &
Shinomol, 2015)
– Treatment of diarrhea and dysentery; Treatment of fungal infections (bamboo paste); Treatment of pigs against Ascaris suum (with garlic); As
antimicrobial, astringent and eardrops (bamboo juice) (Singhal et al., 2013)

aspects such as the decreases in fasting glucose, fasting insulin, glucose
stimulated insulin, insulin resistance, glucose and insulin tolerance.
Apart from bamboo shoot oil, fiber and polysaccharides, bamboo shoot
alkaloids or peptides produced anti-inflammatory and anti-hyperten-
sive. In summary, bamboo shoot is turly a functional food material.
4. Storage of fresh bamboo shoots
Freshly-harvested bamboo shoots are often consumed as a vegetable
in the kitchen. However, the tissue of fresh bamboo shoots undergoes a
rapid lignification during the postharvest storage at room temperature,
which consequently leads to a sharp hardening process and thus the
edibility of fresh bamboo shoots heavily damaged (Li, Suo et al., 2019;
Li, Limwachiranon et al., 2019; Zheng, Li, Xu, & Zheng, 2019). At the
same time, browning or yellowing occurred with the fresh bamboo
shoots, especially at the harvest cut surface. In considering the fact that
bamboo shoots are mostly produced in remote mountainous areas, the
transportation up to several or tens of hours is required. In this scenario,
both the nutritional and sensory quality of fresh bamboo shoots sharply
decreased if no preventive measurement is applied. To this end, dif-
ferent methods were adopted to retard the lignification process of fresh
bamboo shoots. Commonly, both physical and chemical regimes were
invented for the preservation of fresh bamboo shoots.

9
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 7
Modern biological activities of bamboo shoots.
Substrate Activity narrators (reference)a

Antioxidant activity
Shoot juice lipid oxidation↓, shelf life of pork nuggets↑ (21 → 35 days at 4 °C) (Thomas, Jebin, Saha, & Sarma, 2016)
Phenolic extract scavenging DPPH and ABTS radicals, reducing Fe3+ (Milani et al., 2020)
Phenolic extract scavenging DPPH radical (IC50 0.4–5.3 mg/mL) (Park & Jhon, 2010)
Phenolic extract scavenging DPPH radical (IC50 137–260 μg/mL) (Wróblewska, Baby, Guaratini, & Moreno, 2019)

Hypolipidemic activity
Shoot oil serum TC↓, TG↓, hepatic lipase↓, atherogenic index↓, fecal cholesterol excretion↑ (Lu et al., 2010)
Shoot fiber serum TC↓, TG↓, LDL-C↓, atherogenic index↓ (Park & Jhon, 2009)
Shoot fiber plasma TG↓, TC↓, HDL-C↓, LDL-C↓, non-esterified fatty acid↓ (Li et al., 2018)
Shoot fiber plasma TG→, TC↓, HDL-C↓, LDL-C↓ (Li, Guo, Ji, & Zhang, 2016)

Prebiotic activity
Polysaccharides B. adolescentis↑, B. bifidum↑, organic acids↑ (He et al., 2016)
Polysaccharides B. adolescentis↑, B. infantis↑, B. bifidum↑, L. acidophilus↑, short chain fatty acids↑ (Chen et al., 2019)
Shoot fiber Bacteroidetes↑, Verrucomicrobia↓, Akkermansia↓, Bateroides↑, Bilophila↑, Prevotella↑ (Li et al., 2016)

Antidiabetic activity
Shoot fiber plasma fasting glucose↓, fasting insulin↓, glucose stimulated insulin↓, insulin resistance↓, glucose tolerance↓, insulin tolerance↓ (Li et al., 2018)
Shoot fiber plasma fasting glucose↓, fasting insulin↓, insulin resistance↓, glucose tolerance↓, leptin↓ (Li et al., 2016)

Antiobesitive activity
Shoot fiber body weight gain↓, body fat mass↓, adipose tissue (Li et al., 2016)

Anti-inflammatory activities
Shoot oil lesion severity↓, absolute prostate weight↓, prostate index↓, total acid phosphatase↓, prostatic acid phosphatase↓, white blood cell count↓, density of lecithin
corpuscles↑ (Lu et al., 2011)
Alkaloids nitric oxide murine macrophages↓, nitric oxidesynthase↓, cyclooxygenase-2↓, interleukin 1β↓, tumor necrosis factor α↓, phosphorylation protein kinase↓ (Ren
et al., 2019)

Antihypertensive activity
Peptide systolic blood pressure↓ (Liu et al., 2013)
Phenolic extract angiotensin converting enzyme activity↓ (Park & Jhon, 2010)

Antimicrobial activity
Shoot juice total viable count in nuggets upon the storage at 4 °C for 35 days↓ (Thomas et al., 2016)

Antianaemic activity
Shoot extract hemoglobin↑, red blood cells↑, hematocrit↑ (Chen, Shi, & Yang, 2008)

a
DPPH and ABTS refer to 1,1-diphenyl-2-picrylhydrazyl and 2, 2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic acid). TG, TC, HDL-C, LDL-C are triglyceride, total
cholesterol, high density lipoprotein cholesterol and low density lipoprotein cholesterol, respectively. IC50 is concentration of antioxidant required to quench 50% of
the initial radicals. ↓ and ↑ interpret the decreasing and increasing changes in specific parameter, respectively.

4.1. Physical regimes
In the group of physical regimes, heat treatment, ultraviolet-C ex-
posure (254 nm), gamma irradiation, hypobaric atmosphere and high
hydrostatic pressure were often applied. A heat treatment at 45 °C for
5 h was beneficial to the storage of fresh shoots from Phyllostachys
praecox at 20 °C by inhibiting disease incidence and respiration, re-
tarding ethylene production and delaying tissue lignification (Luo, Wu,
Xie, & Chen, 2012). For the fresh shoots from Zizania latifolia, a UV-C
treatment at 4.24 kJ/m2 and 20 °C for 2 d significantly retarded the
developments of greenness and toughness and increased the total
phenol concentration (Wen et al., 2019). Similarly, for the fresh shoots
from Phyllostachys prominens stored at 6 °C, the flesh firmness, re-
spiratory rate, weight loss, wound browning, disease incidence, cellu-
lose and lignin synthesis were substantially inhibited by a pre-storage
UV-C exposure at 4.0 kJ/m2 (Zheng & Zheng, 2018). In addition, the
ultraviolet-C exposure could alleviate the chilling injury of fresh
bamboo shoots of P. praecox at 1 °C (Zeng, Jiang, Wang, & Luo, 2015).
As for gamma radiation, a pre-treatment at a dose of 3 kGy was
selected for the storage of fresh shoots from Arundinaria oleosa at 4 °C
which retarded the increase of lignin and cellulose and prevented the
lignification and browning processes (Wang et al., 2019). For the fresh
shoots of P. praecox stored at 2 °C, a gamma radiation pre-treatment at
0.5 kGy decreased ethylene production, lignin accumulation (by
12.5%) and decay rate (by 71%) (Zeng, Luo, Xie, & Feng, 2015).
Hypobaric storage is to store the agriculture produces in an atmo-
sphere with a lower pressure. It has been successfully applied in in-
hibiting ripening and senescence and extending shelf-life of various
fruits and vegetables. The hypobaric storage at 50 kPa was proved ef-
fective in inhibiting the firmness development as well as the lignin and
cellulose accumulations in the fresh shoots from Phyllostachys violascens
at 2 °C (Chen et al., 2013). High hydrostatic pressure treatment is a
popular nonthermal technology often applied in food preservation. The
results of Miao, Wang, Zhang, and Jiang (2011) revealed that the high
hydrostatic pressure treatment at 600 MPa at 25 °C significantly re-
tarded the firmness development and lignin and cellulose accumula-
tions in the fresh shoots from Zizaniz caduciflora Turez stored at 4 °C.
4.2. Chemical regimes
In retarding the senescence, ripening, hardening and lignification of
fresh bamboo shoots, chemicals including melatonin, diphenyliodo-
nium iodide, oxalic acid, salicylic acid and nitric oxide (sodium ni-
troprusside) were applied. Usually, these chemicals were used in a form
of an aqueous solution to dip or soak the bamboo shoots with or
without sheath. With a pretreatment being dipped in 1.0 mM mela-
tonin, the shoots from Phyllostachys edulis stored at 4 °C showed an
effectively retarded shoot lignification with significant reductions in
hardening, yellowing and the biosynthesis of lignin and cellulose (Li,
Suo et al., 2019). Upon a pretreatment in 5 mM diphenyliodonium
iodide, the shoots from Phyllostachys praecox stored at 20 °C displayed
lower values in firmness (by 10.88%) and lignin content (by 19.23%)
(Li, Limwachiranon et al., 2019). Prior to the storage of the shoots from
P. prominens at 6 °C, the treatment with 10 mM oxalic acid for 10 min
decreased their respiration rates, disease incidence, weight loss, enzy-
matic browning and lignification process (Zheng et al., 2019). For the

10
Y. Wang, et al.
Fig. 2. Processing workflows of bamboo shoot products via drying, fermentation and canning.
shoots from P. violascens, the dipping in 0.5 mM sodium nitroprusside (a
nitric oxide donor) successfully prevented the firmness increase, lignin
and cellulose accumulations and external browning during the storages
at 20 °C or 10 °C (Yang, Wu, & Cheng, 2011; Yang, Zhou, Wu, & Cheng,
2010). Interestingly, the treatment with 1.0 mM salicylic acid could
suppress the chilling injury and flesh browning of the shoots from P.
praecox at 1 °C (Luo et al., 2012). In view of the mechanisms underlying
these chemicals, they were interpreted by the following ways: 1) bal-
ancing the formation and removal of active oxygen species, which are
crucial for flesh browning and lignification; 2) inhibiting the activities
of lignin and cellulose biosynthesis-related enzymes, such as poly-
phenol oxidase, peroxidase and phenylalamine ammonia lyase; 3) al-
tering the substance metabolism, such as soluble sugars and total
phenols.
5. Processing of bamboo shoots
As mentioned above, the fresh bamboo shoots lignified quickly upon
the harvesting and even lost their edible quality (Li, Limwachiranon
et al., 2019; Wang et al., 2019). The measures adopted for the post-
harvest storage could extend the shelf-life of fresh bamboo shoots, but it
cannot guarantee a year-round supply of bamboo shoots. To this end,
the fresh bamboo shoots were processed in different ways including
fermentation, roasting, boiling, blanching, canning and pickling.
Among them, canning, drying and fermentation are the most popular
methods (Fig. 2).
Canning is a common way to preserve vegetables usually by thermal
inactivating micro-organisms in hermetically packaged products. In the
present cases, the bamboo shoots with large body were often sliced
before canning and the ones with small body were processed without
cutting. To lower the sterilization temperature, protect the color and
guarantee the food safety in shelf-life, a dilute citric acid solution
(about 0.15%, w/w) containing preservatives (potassium sorbate and
sodium benzoate) was filled in the package. Under this circumstance, a
pasteurization in boiling water for 15 min could result in a shelf-life at
room temperature over 12 months.
Drying is effective in reducing water activity, inactivating enzymes
and inhibiting microbial growth and thus could be preserve the agri-
culture produces. Usually, the freshly-harvested bamboo shoots have a
moisture content over 90 g/100 g. As completely dried, the moisture
content of bamboo shoots decreased to an extent lower than 10 g/
100 g. In drying of bamboo shoots, solar drying, hot air drying, mi-
crowave drying and combined drying were applied (Sagar & Kumar,
11
Journal of Functional Foods 71 (2020) 104015
2010). In ancient years, the solar drying was widely used directly on the
fresh bamboo shoots. As a result, the dried bamboo shoots always gave
a dark appearance due to the enzymatic and non-enzymatic browning
during drying (Zheng, Zhang, Song, Lin, & Kan, 2014). In modern food
industry, the fresh bamboo shoots were firstly blanched either in
boiling water or dilute acid solution (Nimasow, Rawat, & Nimasow,
2010). Consequently, dried bamboo shoots with a yellowish appearance
were obtained. It is widely recognized that the freezing treatment
sharply damaged the texture of fresh bamboo shoots and resulted in
mesh-shaped spongy structure, which presented a terrible water
holding capacity and rather bad palatability. However, in terms of
nutrient retention and rehydration rate, freezing drying was often re-
commended by researchers (Singhal, Rudra, Singh, Satya, & Naik,
2018).
Fermentation, especially the lactic acid fermentation, is often
adopted to preserve various vegetables. Consequently, the fresh vege-
tables were converted to tasteful pickles and often used as table foods,
such as Chinese sauerkraut and Szechwan-style pickled vegetables. In
ancient years, the spontaneous fermentation under anaerobic condi-
tions was employed, while, in modern food industry, the preparatory
culture was used alternatively. Fermented bamboo shoots with different
processing methods and flavors were traditionally fabricated in
Table 8
Consumption of fermented bamboo shoots around the world.a.
Region Fermented product
Arunachal Jiang-sun, Ushoi
Pradesh
China Ulanzi
Indonesia Gulei rebung, Sayur ladeh lun-pia
India Amil, Edung, Eeku, Eepe, Ekung, Eup Hendua, Godhak, Hiring,
Hikhu, Hithyi, Ikung, Iromba, Khorisa-tenga, Kardi, Khorisa,
Kupe, Lung-siej, Lungseij, Midukey, Rawtuai rep, Soibum,
Soidon, Soijim, Syrwa, Tama, Ushoi
Kalahndi Handua
Manipur Soijim, Soidon, Soibum
Meghalaya Iromba
Nepal and Bhutan Alui-tama, Mesu, Tchang,
Philippines Dinengdeng na Labong, Ginataang Labong, Labong,
Thailand Naw-mai-dong, Nor-mai-dorng
a
Data in the table were cited from the references of Choudhury, Sahu, &
Sharma, 2011; Choudhury et al., 2012; Ingudam, Sarangthem, Nirmala, & Bisht,
2015; Nongdam & Tikendra, 2014; Kumar et al., 2017; Roy, Roy, & Rai, 2017;
Singhal et al., 2013 and Sonar et al., 2015.
Y. Wang, et al. Journal of Functional Foods 71 (2020) 104015

Table 9
Changes of phenol, phytosterol, cyanogen and other nutrients in bamboo shoots upon various processing methods.
Componenta Changes (phenol, phytosterol, vitamin C and E in mg/100 g; cyanogen and other nutrients in g/100 g)b

Soaking (water) (Nirmala, Bisht, & Laishram, 2014; Pokhariya, Tangariya, Sahoo, Awasthi, & Pandey, 2018; Bajwa et al., 2015; Rawat et al., 2015)
Phenol (wb) B. balcooa: 191.37 → 53.07 (↓77.27%); B. tulda: 443.97 → 58.17 (↓86.90%); D. giganteus: 347.27 → 187.97 (↓45.87%); D. hamiltonii: 505.93 → 56.74 (↓88.79)
Phytosterol (db) B. nutans: 164.20 → 97.24 (↓40.78%); D. giganteus: 136.23 → 89.56 (↓34.26%); D. hamiltonii: 198.27 → 108 (↓45.53%); D. latiflorus: 146.33 → 76.00 (↓
48.06%)
Cyanogen (wb) D. giganteus 0.099 → 0.040 (12 h ↓59.60%) → 0.018 (24 h ↓81.82%); D. hamiltonii 0.073 → 0.037 (12 h↓49.32%) → 0.011(24 h ↓84.93%); D. strictus
0.076 → 0.052 (5 h ↓31.58%) → 0.042 (10 h ↓44.73%) → 0.027 (15 h ↓64.47%) → 0.024 (20 h ↓68.42%)

Boiling (in water) (Nirmala, Bisht, & Laishram, 2014; Pandey & Ojha, 2014; Pokhariya, Tangariya, Sahoo, Awasthi, & Pandey, 2018; Zhang, Ji, Hu, Chen, & Ye, 2011; Bajwa et al., 2015;
Rawat et al., 2015)
Phenol (wb) B. balcooa: 191.37 → 56.83 (↓70.30%); B. tulda: 443.97 → 120.77 (↓73.00%); B. nutans: 489.83 → 180.21 (↓40.52%); D. giganteus: 347.27 → 182.80 (↓
47.36%); D. hamiltonii: 586.36 → 354.11 (↓39.61%); D. latiflorus: 612.24 → 482.43 (↓21.20%)
Cyanogen (wb) B. bambos 0.111 → 0.009 (10 min ↓91.89%) → 0.000 (25 min ↓100%); D. strictus 0.076 → 0.028 (10 min ↓67.161%) → 0.000 (25 min ↓100%); D. giganteus
0.099 → 0.023 (10 min ↓76.77%) → 0.014 (20 min ↓85.86%); D. hamiltonii 0.073 → 0.024 (10 min ↓67.12%) → 0.001 (20 min ↓98.63%)
Others (wb) P. praecox: moisture 91.25 → 91.76 (↑0.56%), proteins 3.25 → 3.01 (↓7.38%), TDF 1.37 → 1.38 (↑0.73%), starch 1.75 → 1.72 (↓1.71%), sugar 1.32 → 1.13 (↓
14.39%), fat 0.21 → 0.16 (↓23.81%), ash 0.85 → 0.81 (↓4.71%)

Boiling (in brine) (Pandey & Ojha, 2014)

Cyanogen (wb) 1) in 1% brine, B. bambos 0.111 → 0.006 (10 min, ↓94.59%) → 0.001 (25 min, ↓99.10%); 2) in 5% brine, B. bambos 0.111 → 0.005 (10 min ↓95.50%) → 0.001
(25 min ↓99.10%); 3) in 10% brine, B. bambos 0.111 → 0.007 (10 min ↓93.70%) → 0.001 (25 min ↓99.10%)

Drying (Bajwa et al., 2015; Chauhan et al., 2016; Pokhariya et al., 2018))
Phenol (db) B. nutans: 489.83 → 1930.66 (↑294.15%); D. giganteus: 336.56 → 1927.29 (↑472.64%); D. hamiltonii: 586.36 → 1934.54 (↑229.92%); D. latiflorus:
612.24 → 1950.67 (↑211.26%)
Cyanogen (wb) D. strictus 0.076 → 0.023 (60 °C ↓69.74%) → 0.034 (80 °C ↓55.26%) → 0.069 (100 °C ↓9.21%)
Others (wb1 db2) B. heterostachya: moisture1 92.6 → 4.6 (↓95.03%), proteins2 27.8 → 21.6 (↓22.30%), TDF2 5.2 → 5.0 (↓3.85%), starch2 28.3 → 9.2 (↓67.49%), Vc2 2.1 → 0.2
(↓ 90.48)

Fermentation (Nirmala, Bisht, & Laishram, 2014; Nirmala, Sharma, & David, 2008; Pokhariya, Tangariya, Sahoo, Awasthi, & Pandey, 2018; Rawat et al., 2015)
Phenol (wb) B. balcooa: 191.37 → 298.53 (↑56.00%); B. tulda: 443.97 → 641.73 (↑156.66%); D. giganteus: 347.27 → 891.33 (↑156.66%); D. hamiltonii: 505.93 → 745.56 (↑
47.36%);
Cyanogen (wb) D. hamiltonii 0.073 → 0.041(↓43.84%); B. vulgaris 0.043 → 0.016 (↓62.79%)
Others (wb) D. giganteus: moisture 90.70 → 88.83 (↓2.06%), proteins 3.11 → 2.57 (↓17.31%), FAA 3.86 → 2.01 (↓48.06%), TDF 4.80 → 7.46 (↑55.29%), starch
0.51 → 0.46 (↓10.08%), sugar 5.10 → 1.50 (↓70.53%), fat 0.39 → 0.32 (↓18.60%), ash 0.89 → 0.78 (↓12.36%), VC 3.28 → 1.09 (↓66.77%), VE 0.69 → 0.21 (↓
69.57%)

Canning (Nirmala, Bisht, & Laishram, 2014; Nirmala, Sharma, & David, 2008)
Phenol (wb) B. balcooa: 191.37 → 21.37 (↓88.83%); B. tulda: 443.97 → 67.23 (↓84.85%); D. giganteus: 347.27 → 80.36 (↓76.86%); D. hamiltonii: 505.93 → 34.47 (↓
81.33%)
Others (wb) D. giganteus: moisture 90.70 → 95.16 (↑4.92%), proteins 3.11 → 1.93 (↓37.9%), FAA 3.86 → 1.98 (↓48.70%), TDF 4.80 → 5.05 (↑5.33%), starch 0.51 → 0.44
(↓12.45%), sugar 5.10 → 1.45 (↓71.59%), fat 0.39 → 0.25 (↓35.40%), ash 0.89 → 0.75 (↓15.73%), VC 3.28 → 1.80 (↓45.12%), VE 0.69 → 0.3 (↓56.38%)

Pickling (5% brine) (Bajwa et al., 2015)

Phenol (wb) B. nutans: 489.83 → 291.34 (↓40.52%); D. giganteus: 336.56 → 157.41 (↓53.23%); D. hamiltonii: 586.36 → 192.77 (↓67.12%); D. latiflorus: 612.24 → 383.27 (↓
37.40%);
Phytosterol (db) B. nutans: 164.20 → 171 (↑4.14%); D. giganteus: 163.23 → 139.33 (↓14.64%); D. hamiltonii: 198.27 → 322.24 (↑62.52%); D. latiflorus: 146.33 → 175.23 (↑
19.75%)

Pickling (10% brine) (Bajwa et al., 2015)

Phenol (wb) B. nutans: 489.83 → 89.53 (↓63.21%); D. giganteus: 336.56 → 124.46 (↓81.71%); D. hamiltonii: 586.36 → 98.68 (↓83.71%); D. latiflorus: 612.24 → 109.29 (↓
82.15%)
Phytosterol (db) B. nutans: 164.20 → 81.77(↓50.20%); D. giganteus: 163.23 → 67.66 (↓58.55%); D. hamiltonii: 198.27 → 79.33 (↓59.99%); D. latiflorus: 146.33 → 57.00 (↓
61.05%)

Steaming (Pokhariya et al., 2018; Zhang et al., 2011)

Cyanogen (wb) D. strictus 0.076 → 0.040 (5 min ↓47.37%) → 0.021 (25 min ↓72.37%)
Others (wb) P. praecox: moisture 91.25 → 91.65 (↑0.44%), proteins 3.25 → 3.21 (↓1.23%), TDF 1.37 → 1.37 (-), starch 1.75 → 1.75 (-), sugar 1.32 → 1.31 (↓0.76%), fat
0.21 → 0.20 (↓4.76%), ash 0.85 → 0.85 (-)

Stir-frying (Zhang et al., 2011)

Others (wb) P. praecox: moisture 91.25 → 90.31 (↓1.03%), proteins 3.25 → 3.20 (↓1.54%), TDF 1.37 → 1.37 (-), starch 1.75 → 1.74 (↓0.57%), sugar 1.32 → 1.09 (↓
17.42%), fat 0.21 → 1.32 (↑528.57%), ash 0.85 → 0.84 (↓1.18%)

a
wb and db refer to wet basis and dry basis, respectively.
b
↓ and ↑ interpret the decreasing and increasing changes for the specific component and the percentages followed with them were the amplitude of variation.

different areas. There were twenty-six fermented products from
bamboo shoots were historically recorded in India (Table 8). In parti-
cular, the bamboo shoots were fermented into a sweet wine (ulanzi) in
China (Choudhury et al., 2012).
Although the above-mentioned processing methods successfully
preserved the bamboo shoots and brough about a year-round supply, it
must be noted that they exerted giant impacts on the nutritional quality
of bamboo shoots (Table 9). In terms of total phenols, soaking in water,
boiling, canning and pickling caused a substantial loss, while significant
increases were observed with fermentation and drying. The increase
upon drying was ascribed to the moisture loss and that upon fermen-
tation should be owed to the bioconversion from non-phenol substances
(Nongdam & Tikendra, 2014). For total phytosterol, only data on
soaking and pickling were available. The soaking and the pickling in
10% w/w brine led to a significantly decrease in total phytosterol,
while an increase was observed with the pickling in 5% w/w brine
(Bajwa, Nirmala, Koul, & Bisht, 2015). The total dietary fiber increased
with fermentation, canning and boiling, decreased with drying and
remained unchanged with steaming and stir-frying. For the common
nutrients, the crude protein, ash, sugar and vitamins (VC and VE) in
bamboo shoots decreased in all processing regimes. Except the increase
with stir-frying, the fat content of bamboo shoots decreased with other
processing regimes.

12
Y. Wang, et al.
Fig. 3. Biosynthesis (A) and biotransformation (B) of cyanogen glycosides (Møller, 2010; Sang-A-Gad et al., 2011).
6. Safety concerns of bamboo shoots
Apart from their nutritional benefits, the fresh bamboo shoots also
contain certain anti-nutrients even toxins such as tannin, oxalate, heavy
metals and cyanogenic substances or cyanides. Among them, cyano-
genic substances should be addressed seriously due to their high toxi-
city. In fresh bamboo shoots, cyanogenic substances presented mainly
in the form of cyanogenic glycosides, such as p-hydroxylated mande-
lonitrile, named taxiphyllin. Cyanogenic glycosides are formed by the
condensation of a hydroxyl group of cyanohydrin derivatives with D-
glucose. The metabolic pathway of cyanogenic glycosides in bamboo
shoots is shown in Fig. 3 (Møller, 2010). The cyanide level in fresh
bamboo shoots is plant species and age dependent. To the best of our
knowledge, the highest level of cyanide was found with the fresh
bamboo shoots from D. longispathus (1951.49 mg/kg wb), while the
lowest one was the shoots from C. callosa (31.68 mg/kg wb) (Table 5).
Regarding the shoot age, the emerging shoots always showed the lower
cyanogen content in contrast to its older counterparts (Pandey & Ojha,
2013).
When the fresh shoot tissues were physically disrupted (cutting or
peeling), the cyanogenic glycoside undergoes a hydrolysis to produce
HCN gas in contacting with a glycosidase. The long-time diet exposure
of HCN may lead to damages in pancreatic tissue, the central nervous
system and the thyroid gland (Sarkar et al., 2020). The acute lethal dose
of cyanide for human beings is 0.5–3.5 mg/kg. Generally, edible plants
with a cyanogen level over 20 mg/100 g wb are considered potentially
dangerous for human beings (Kingsbury, 1964). This suggested that
fresh bamboo shoots are dangerous for human consumption. However,
no cyanide poisoning has been reported from consuming bamboo shoot
but the poisoning events were reported for the workers dealing with the
pickled bamboo shoots in a well (Sang-A-Gad, Guharat, & Wananukul,
2011). This was due to the fact that fresh bamboo shoots were definitely
consumed after certain processing, such as soaking, boiling, drying,
fermentation and steaming, which could effectively eliminate cyanogen
from bamboo shoots (Table 9).
In contrast, the boiling either in water or in brine was the most
efficient way to eliminate cyanogen from bamboo shoots. Over 99% of
cyanogen was eliminated upon boiling for 25 min (Pandey & Ojha,
2014). As for soaking, treatment for 12 h achieved a decrease in cya-
nogen approximately 50% and this value was elevated to
68.42–84.93% as the treatment was extended to 24 h (Pokhariya,
Tangariya, Sahoo, Awasthi, & Pandey, 2018; Rawat, Nirmala, & Bisht,
2015). Drying, fermentation and steaming were somewhat less efficient
and no more than one-third of cyanogen could be eliminated by these
methods (Pandey & Ojha, 2014; Pokhariya et al., 2018; Rawat et al.,
2015). It must be noted that the operating temperature is extremely
13
Journal of Functional Foods 71 (2020) 104015
important for drying regime in terms of cyanogen elimination. The
reduction of cyanogen decreased with the increase in drying tempera-
ture and the values for 60 °C, 80 °C and 100 °C were determined as
69.74%, 55.26% and 9.21%, respectively (Pokhariya et al., 2018). This
was interpreted by the fact that the endogenous glucosidase in bamboo
shoots is most efficient around 55 °C and it is certainly inactivated at
higher temperatures as 80 °C and 100 °C. Therefore, the fresh bamboo
shoots are not recommended to be eaten raw and the processing is
essential before their consumption.
7. Conclusions
Since time immemorial, the bamboo shoots serve as a sustainable
food source. This article gave a systematic review of bamboo shoots by
addressing their nutritional values, health effects, processing and safety
concerns. The available knowledge attested that the bamboo shoot, on
one hand, is one of the most valued vegetables and it certainly playing
an important role in preventing malnutrition and lowering the in-
cidence of some diseases, such as hyperlipidemia and diabetes mellitus.
On the other hand, the presence of cyanogen glycosides is the biggest
hazard in consuming bamboo shoots and the boiling in water or brine is
the most efficient way to delaminate it. Due to its high nutritional
quality, diverse health benefits and low pollution, bamboo shoots be-
come more popular in recent years. In developing bamboo shoots as a
staple food, the following issues should be in serious consideration.
First, the harvest of bamboo shoots and the protection of the forest
should be carefully balanced. Second, the detection and detoxification
of cyanogen glycosides, even the breeding of edible bamboo shoots with
lower level of cyanogen glycosides should be highlighted in view of the
food safety. Third, the mechanisms underlying the traditional using of
bamboo shoots in regional medications deserve further exploration.
Ethics statement
This work did not use any human or animal subjects.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgements
This research was funded by The National Key Research and
Development Plan of China (2016YFD0400204-2) and National Natural
Y. Wang, et al.
Science Foundation of China (31771932).
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