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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6499 

 

Monopodial and Sympodial Bamboos Grown in Tropic 
and Sub-tropic Countries – A Review 
 
Norul Hisham Hamid,a,b Mohammad Jawaid,b,* Ummi Hani Abdullah,a,b and  

Taghrid S. Alomar c 

 
Bamboo belongs to the grass family and is an important non-timber forest 
product in tropic and sub-tropic countries. The global trade of bamboo 
products is worth billions of dollars and is mainly dominant with 
monopodial bamboo grown in sub-tropic countries such as China and 
Japan. Many researchers globally discuss that in addition to species and 
region, bamboo quality can differ based on its rhizome types because the 
physiology is different for both monopodial and sympodial bamboo. 
However, there is a massive competition within the yearly forest products 
due to the challenges posed by underground root system in agroforestry. 
This review studied the properties of bamboo with regards to their 
differences in terms of monopodial and sympodial types of rhizomes. It 
was found that most of the structural, chemical organic, and mechanical 
properties are higher in monopodial bamboo, but there is a greater fibre 
morphology and decay resistance in the sympodial bamboo.  

 

DOI: 10.15376/biores.18.3.Hamid 

 

Keywords: Bamboo; Rhizome; Growth; Anatomy; Physical; Mechanical properties; Chemical properties; 

Decay resistance 

 
Contact information: a: Faculty of Forestry and Environment Universiti Putra Malaysia 43400 UPM 

Serdang, Selangor, Malaysia; b: Institute of Tropical Forestry and Forest Products Universiti Putra 

Malaysia 43400 UPM Serdang, Selangor, Malaysia; c: Department of Chemistry, College of Science, 

Princess Nourah bint Abdulrahman University, Riyadh 11671, Saudi Arabia; *Corresponding author: 

h_noroul@upm.edu.my  

 

 
INTRODUCTION 
 

Bamboo is a monocotyledon plant and is classified as sub-family of Bambosoidea 

in the family of Graminae. Worldwide, bamboo consists of 119 genera and 1500 species 

under three main tribes, namely, Arundinarieae (temperate woody bamboo), Bambuseae 

(tropical woody bamboo), and Olyreae (herbaceous bamboo). They are distributed within 

the tropic and sub-tropic regions from sea level to the alpine, with the altitude or from 47 

S to 50 30’ N and latitude from sea level to 4300 m (Clark et al. 2015). Bamboo stands 

exist in all continents except for Europe and Antarctica (Ram et al. 2010; Hagarth and 

Belcher 2013; Mera and Xu 2014). 

Bamboo culms cover about 3.2% (37 million hectares) of the world forest area. 

Approximately 80% of the bamboo stand and species are distributed in the Asia and pacific 

regions (Mera and Xu 2014). China owns the largest bamboo forest area (601,000 km2) 

followed by India (108,630) and Myammar (8950 km2) (Fei et al. 2016). The deforestation 

that occurs in these countries is relieved by the emergence of bamboo forests and in turn 

this balances the ecosystem. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6500 

A bamboo forest is part of the forest ecosystem, and it acts as an important source 

of carbon and carbon sink (Li et al. 2003). Bamboo possesses a great advantage of reducing 

global warming by utilizing the carbon dioxide emission produced from modern vehicles, 

industries, and population growth. One hectare of bamboo culms is able to absorb more 

than twelve tons of carbon dioxide per year (Raka et al. 2011), while the dicotyledon trees 

only absorb slightly lower carbon dioxide, ranging from 1.1 to 9.5 tons per year (Chasan 

2019). This is due to the fact that bamboo achieves its maximum growth within a one year 

and matures within 3 to 4 years for sympodoal bamboo and 7 to 8 years for monopodial 

bamboo (Razak et al. 2010; Fangchun 2001a,b). Depending on the bamboo culm, the 

photosynthesis process that produces carbohydrates for the growth and maturation is also 

involved in nutrient uptake from soils.  

The bamboo culm grows toward maturation, and when it gets old, it loses its ability 

to uptake nutrients from soil. The amount of nutrients or inorganic contents in 

Gigantochloa scortechinii culm declines from a young age (6 months), toward its old age 

of 6.5 years (Norul Hisham et al. 2006). The oldest bamboo culm experiences an 

insufficiency in nutrients and will produce flowers and then die if it is not being utilised 

for any purpose. Therefore, mature bamboo culm needs to be harvested to allow a new 

shoot to emerge from the rhizome to produce a new culm. The age-maturation of bamboo 

culm is important to determine the harvesting cycle of each bamboo culm for production, 

propagation, and its overall sustainability for further utilization. The making of a specific 

bamboo product depends on the culm age for its maximum quality. Bamboo shoot as a 

food is best when it emerges less than 3 weeks from soils (Fangchun 2001b). The quality 

of bamboo higo products, such as barbeque and chop sticks, are maximum for the bamboo 

culm aged 1 to 2 years (Hamdan and Mohmod 1992). While the most suitable age for 

bamboo timber and flooring ranges from 3 to 4 years for sympodial bamboo to achieve its 

maximum strength (Sattar et al. 1994; Mohmod and Phang 2001; Banik 2015).  

A mature bamboo culm as a biological material for human products evolved from 

traditional to a modern application parallel to human civilization. Historical records show 

that bamboo was used as fire work and in rockets during the Chinese dynasties (Deluca 

2016). Indigenus people in Southeast Asia used bamboo as a rice cooker and in weapons 

in their daily life. Fei et al. (2016) mentioned that half of the world population utilizes 

bamboo products such as in housing, biocomposites, mats, chopsticks, charcoal, activated 

carbon, pulp, shoot, and other applications.  

Medicine is another category of a small amount of bamboo products. The 

Pleioblastus amarus leaves can remedy fever, fidgeting, and lung inflammation (Kiruba et 

al. 2007). The extracts of Sasa senanensis, Bambusa caulis, and Pseudosasa japonica have 

anti-cancer activity (Panee 2009; Seki et al. 2010; Kim et al. 2013). In addition, bamboo 

is reported to have great potential for soil erosion control, water conservation, land 

rehabilitation, and carbon sequestration (Zhou et al. 2005). 

The bamboo industry has evolved from being used as basic tools for domestic 

requirement to world commodities for international markets. The global bamboo market 

was about USD 68.8 billion in 2018 (Market Research Report 2019). The most popular 

product, woven bamboo, represents the largest proportion of global exports, estimated at 

USD 380 million in 2017 (Inbar 2020). China with the largest bamboo forest area is a main 

exporter for bamboo products. China’s bamboo industry success is not only related to its 

culture that has been ever expanding, but the proper selection of bamboo species for a 

specific product for optimum qualities is the main contribution for success. Norul Hisham 

et al. (2006) explained that there are thousands of bamboo species worldwide and their 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6501 

properties differ by species, age, location, and other external factors. According to the 

phenotype, bamboo can be divided into three types of rhizomes, namely monopodial 

(leptomorph), sympodial (pachymorph), and amphipodial. The different rhizome structure 

possibly influences its growth, development, and maturation (Fangchun 2001a; Fei et al. 

2016). 

Despite being in the same tribe, the growth and development phases in monopodial 

and sympodial bamboo culms may be different due to different rhizome structures. Zhao 

et al. (2014) made comparison between the micro Ribonuleic acid (mRNA) of monopodial 

bamboo (Phyllostachys pubescens) and sympodial bamboo (Dendrocalamus latiflorus). In 

their reports, the rhizome of monopodial bamboo can spread laterally while grown in soil, 

and can also be separated from the mother plant, while sympodial bamboo grows in clusters 

within a relatively small range. The result indicates that there are 19,295,759 and 

11,513,888 raw sequence reads, in which 92 and 69 conserve miRNAs, as well as 95 and 

62 novel miRNAs are identified in P. pubescens and D. latiflorus, respectively. The ratio 

of high conserved miRNA families in D. latiflorus is more than that in P. pubescens. In 

addition, a total of 49 and 106 potential targets are predicted in P. pubescens and D. 

latiflorus, respectively, in which several targets for novel miRNAs are transcription factors 

that play important roles in plant development. Experiments show that miR397, miR1432, 

and miR7748 are specifically conserved in the leaf sample of P. pubescens.  

Taken together, the comparison between P. pubescens and D. latiflorus indicate 

that monopodial and sympodial bamboo may share different miRNAs and target genes to 

have a better adaption for their development in different stages, and stress response in their 

diverse course of evolution. Therefore, monopodial bamboo requires more self-regulation 

to adapt to the environment than sympodial bamboo, which might be consistent with the 

generation of lower conserved miRNAs families. Fangchun (2001a) investigated the 

physical properties of 96 bamboo species and the mechanical properties of 65 bamboo 

species from both monopodial and sympodial bamboo. The moisture content, density, 

shrinkage, tensile, and compression properties vary by species, which originated from 

different rhizome type. The growth, development, and properties of some bamboo species 

either from monopodial and sympodial rhizomes may also be influenced by the site 

condition and climate of a specific area. 

The International Network for Bamboo and Rattan (INBAR) proposed priority 

species of bamboo and rattan that include 20 taxa (species and genera) of particular 

economic importance and another important 18 taxa (Rao et al. 1998). In this context, the 

priority is based on its specific uses, and the main criteria for the classification of species-

usage is general characteristics, such as diameter, wall thickness, internode length, and 

overall culm height. The priority species by the culm physical properties, P. pubescens, is 

categorized as medium to large monopodial bamboo with 10 to 20 m height, 18 to 20 cm 

diameter, internodes up to 45 cm long, and thick wall up to 2 cm (Rao et al. 1998). The P. 

pubescens is the most successful bamboo species in China for manufacturing glue-

laminated timber flooring. 

A similar characteristic in sympodial bamboo such as Dendrocalamus asper with 

up to 20 to 30 m height, internodes 20 to 45 cm long, diameter of 8 to 20 cm, and thick 

walls up to 2 cm, is that they are only used as furniture, musical instruments, chopsticks, 

household utensils, and handicrafts (Rao et al. 1998). The thick wall of D. asper does not 

give advantage for manufacturing glue-laminated board or flooring in Asian countries. The 

examples of end products from monopodial and sympodial bamboo species are shown in 

Table 1 and 2. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6502 

Table 1. Example of the Utilization of Monopodial and Sympodial Bamboo in 
China (Liu et al. 2018) 

Value Utilization Product Bamboo Species 

Economic Timber Bamboo flooring; 
Pulp and paper; 

Construction material; 
Bamboo chopsticks 

P. edulis (monopodial) 
D. giganteus (sympodial) 

Shoot Fresh shoot 
Drying shoot 

Canned shoots 
Flavored shoot 

P. edulis (monopodial) 
D. brandisii (sympodial) 
D. latiflorus (sympodial) 
P. praecox (monopodial) 

Skin/Bark Tables and chairs 
Basket 

Wall of house 

Neosinocalamus affinis 
B. textilis (sympodial) 
B. chungii (sympodial) 

Artistic  Musical instruments 
Bonsai 

Root carving 

Qiongzhuea tumidissinoda 
Chimonobambusa 

quadrangularis 
Pseudosasa amabilis 
P. nigra (monopodial) 

Ecological Water 
conservation 

Water conservation forest P. edulis (monopodial) 
D. giganteus (monopodial) 

Ecotourism Scenic spot P. aurea (monopodial) 
B. ventricosa (sympodial) 
Thyrsostachys siamensis 

 
Table 2. Example of the Utilization of Sympodial Bamboo in Malaysia 

Value Utilization Product Bamboo Species 

Economic Timber Structure 
Parquet 
Furniture 

Gigantochloa thoii 
G. scortechinii 

D. asper ; G. ligulata 
G. wrayi ; G. brang 

B. vulgaris ; B. blumeana 
B. heterostachya 

B. vulgaris cv. vittata 

 Shoot Fresh shoot 
Canned shoot 

B. vulgaris 
B. vulgaris var striata 

D. asper ; D. gigantues 
G. levis ; G. ligulata 

G. wrayi 
Schizostachyum brachycladum 

Trametes siamensis 

 Skin Basketry 
Blind 
Craft 

G. brang 
S. brachycladum 

S. grande 
S. zollingeri 
B. vulgaris 

B. blumeana 
B. heterostachya 

B. vulgaris cv vittata 

 Art and Craft Cooking vessel S. zollingeri 

 Utencil Chopstick 
Tooth pick 

Barbeque stick 

D. asper 
G. scortechinii 

Gigantochloa sp. 

 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6503 

Another example is the trial of glue laminated bamboo board and flooring from 

sympodial bamboo G. scortechinii in Peninsular Malaysia during 1998 to 2001. The G. 

scortechinii culm has an internode ranging 32 to 50 cm long, 11 to 18 cm in diameter, and 

6 to 9 mm in wall thickness (Norul Hisham et al. 2006), which is similar to P. pubescens 

grown in China. However, after two decades of trial, the quality appearance and marker 

acceptance of laminated bamboo flooring from G. scortechinii was not comparable to P. 

pubescens. In the period of 1998 to 2015, two laminated bamboo board factories in 

Malaysia were closed down due to lack of quality appearance for the local market. Another 

contributing factor is unsustainability of the material, as most of the stock is obtained in 

the natural forest and the bamboo plantation is not established at the beginning. This 

indicates that the quality of laminated bamboo flooring and other biocomposites is not only 

dependent on the basic and physical characteristics. Other properties, such as chemical, 

mechanical, machining, and appearance that vary between monopodial and sympodial 

bamboos, are a collective factor for optimum quality. In view of the above, this review 

aims to elaborate the properties characterization in monopodial and sympodial bamboos.  

 

  

RHIZOME  
 

The monopodial rhizome has a long and slender culm with a cylindrical or sub-

cylindrical form. Its diameter usually is less than that of the culm coming from it (Fig. 1).  

Its internode is longer, relatively uniform in length, rarely solid, typically hollow 

with interruptions at each node by a diaphragm; nodes in some genera usually elevated or 

inflated, in others no lateral buds in the dormant state are boat-shaped (McClure 1966). 

The monopodial bamboo is characterized as having strong frost resistance and is 

distributed in area of higher latitudes, such as Japan, Korea, Yellow River, and Yangtze 

Valley where there is a slight winter (Fangchun 2001a).  

The sympodial rhizome has 6 to 7 large lateral buds on either side of the thick 

rhizome proper, and the buds grow up to new bamboos with a short rhizome neck (Fig. 1). 

The rhizome internodes are broader, long, solid, and asymmetrical, while the nodes are not 

elevated. The underground rhizome consists typically of two parts: the rhizome proper and 

the rhizome neck. The neck is basal to the rhizome proper, generally shorter in length and 

obconical in shape. It connects the new rhizome to the mother rhizome. Rhizomes are 

usually more or less curve-shaped and rarely straight, with maximum thickness, typically 

somewhat greater than that of the culm (Liese and Kohl 2015). 

Being morphologically different, the monopodial bamboo rhizome can be extended 

horizontally under the soils depending on the species, and its length ranges from 50 to 70 

m for P. heteroxyxla, 90 to 250 m for P. viridis, and 200 to 350 m for P. niagra (Jinghua 

2000). Sympodial bamboo rhizome, such as B. tulda, can only be extended under the soils, 

up to 5.2 m (White and Childers 1945). Due to this capability, monopodial and sympodial 

bamboo rhizomes are commonly referred to as running and clumping bamboos, 

respectively. Amphipodial, which is a combination of monopodial and sympodial 

rhizomes, belongs to the genera including Bashania and Shibataea (Maoyi 2007). These 

genera originated from Japan. 

 

 

 

 



 

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Monopodial bamboo Sympodial bamboo 

  
Monopodial rhizome  Sympodial rhizome  

 

Fig. 1. Monopodial bamboo and rhizome and sympodial bamboo and rhizome (Redrawn with 
inspiration from Banik et al. 2015) 
 

 
SPROUT AND GROWTH 
 
Monopodial Bamboo 

The P. nigra var. henosis has the shortest sprouting phase (19 days), followed by 

P. nidularia (20 days), and P. makinio (25 days) as shown in Table 2. The P. heteroclade 

has the longest sprouting phase (45 days) among the Phyllostachys genera. The sprouting 

phase of P. pubescence grown in China (28 days) is quicker than those grown in the USA 

(44 days). This is reflected by the different site and climate between these countries. The 

growth phase is quickest for P. nigra (24 days) followed by P. pubescens (31 days) and P. 

makinoi (32 days) in China (Hwang and Ma 1994; Zhang et al. 1997; Li et al. 2005). 

 

Sympodial Bamboo 
The Fargesia spathecea has the earliest sprouting phase (59 days) amongst the 

sympodial bamboo, followed by F. robusta (80 days) and F. denudate (90 days). D. 

latiflorus has the longest sprouting phase (180 days) amongst the sympodial bamboo. The 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6505 

growth phase is the quickest for Fargesia robusta (70 days) followed by 

Dendrocalamopsis oldhani (80 days) and D. latiflorus (90 days) in China (Zhou 1999; Qin 

et al. 1993; Gao et al. 2000). 

 

Overall 
The sprouting and growth phases are significantly shorter in monopodial bamboo 

(32.7 and 33.2 days) compared to the sympodial bamboo (105.8 and 102.6 days), 

respectively, from further analysis of the statistical data (Table 3). The bamboo shoot 

elongation rate within 24 h depends on the genera, species and rhizome. P. reticulate 

records the fastest growth rate (maximum 120 cm/day) for monopodial bamboo (Ueda 

1960), and the rate is same as D. asper for sympodial bamboo (Subsansenee 1994). The 

growth rate of B. balcooa, D. gigantus, and B. vulgaris are recorded as 77 cm/day, 58 

cm/day, and 44 cm/day, respectively (Osmastos 1918; Banik 1993). 

 

Table 3. Sprouting and Growth Phases of Monopodial and Sympodial Bamboos 

Species Rhizome Location 
Sprout 
(days) 

Growth 
(days) 

Source 

Arundinaria amabilis Mono S.E China 38 44 Fangchun (2001a) 

A. fargesii Mono N. China 46 41 Fangchun (2001a) 

Brachyschyum densiflorum Mono S. China 28 27 Fangchun (2001a) 

Indocalamus barbatus Mono S. China 67 66 Fangchun (2001a) 

Indosasa crassiflora Mono S. China 40 32 Fangchun (2001a) 

P. august Mono S. China 22 26 Fangchun (2001a) 

P. bambusoides f. Tanakae Mono C. China 26 27 Fangchun (2001a) 

P. bambusoides Youngii Mono S. China 28 26 Fangchun (2001a) 

P. bissetii Mono S.W China 35 29 Fangchun (2001a) 

P. decora Mono E. China 22 24 Fangchun (2001a) 

P. glauca Mono E. China 29 31 Fangchun (2001a) 

P. glauca f. Yunzhu Mono C. China 30 28 Fangchun (2001a) 

P. parvifolio Mono E. China 32 30 Fangchun (2001a) 

P. praecox Mono S. China 29 28 Fangchun (2001a) 

P. meyeri Mono E. China 53 36 Fangchun (2001a) 

P. nigra Mono S. China 30 25 Fangchun (2001a) 

P. nigra var. henonis Mono S. China 26 27 Fangchun (2001a) 

P. nuda Mono S. China 28 33 Fangchun (2001a) 

P. pubescens Mono E. china 30 40 Fangchun (2001a) 

P. pubescens f. grammica Mono E. China 37 31 Fangchun (2001a) 

P. rubella Mono E. china 38 37 Fangchun (2001a) 

P. spextabilis Mono S. china 26 29 Fangchun (2001a) 

P. viridis f. houzeauana Mono S. china 29 28 Fangchun (2001a) 

P. vivax Mono S. China 28 28 Fangchun (2001a) 

Sinobambusa laeta Mono S.E. China 54 50 Fangchun (2001a) 

P. pubescens Mono S. China 31 NA Zheng et al. (1998) 

P. pubescens Mono E. China 28 33 Zhang et al. (1995) 
P. pubescens Mono USA 44 70 Lee and Addis (2001) 

P. nigra Mono E. China 27 24 Zhang et al. (1997) 

P. makinoi 
Mono 

E. China 25 32 
Huang and Ma 

(1994) 



 

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P. heteroclada Mono E. China 45 39 Jin et al. (1999) 

P. nidularia Mono W. China 20 45 Zhang et al. (1995) 

P. nigra var. henonis Mono C. China 19 34 Li et al. (2005) 

F. robusta Sym W. China 80 70 Qin et al. (1993) 
F. denudate Sym W. China 90 163 Wang et al. (1991) 

F. spathacea Sym C. China 59 110 Li (2003) 

Dendrocalamopsis oldhami Sym S. China 120 80 Gao et al. (2000) 

D. latiflorus Sym E. China 180 90 Zhou (1999) 

Mono – monopodial, Sym – sympodial, S.E – South east, N – North, S.W – South west, C- 
central, W – West, E- East, S – South 
 

Table 4. Statistical Analysis of Sprouting and Growth Phases for Monopodial and 
Sympodial Bamboos 

Phase Monopodial Sympodial DF F Significance 

Sprout (days) 32.7 
(10.77) 

105.8 
(49.94) 

1 16.3 0.000*** 

Growth (days) 33.2 
(8.94) 

102.6 
(36.86) 

1 20 0.000*** 

 *** Significant at P < 0.01. The value in the parenthesis is standard deviation. 

 
 
CULM DIAMETER AND HEIGHT 
 

Most bamboo culm achieves maximum height within one year, without showing 

any further growth for subsequent years (Liese 1985). The classification of culm diameter 

and height is useful in helping to identify the growth factors for individual species in 

different site location, topography, climate, and other conditions (Fangchun 2001a). It is 

also commonly used to classify bamboo according to its suitable usage (Benton 2015; Liese 

and Kohl 2015). Fangchun (2001a) classified the bamboo growth according to the average 

diameter namely: Class 1 (diameter more than 12 cm), Class 2 (10 to 12 cm), Class 3 (8 to 

10 cm), Class 4 (6 to 8 cm), and Class 5 (less than 6 cm). 

 

Monopodial Bamboo 
The P. nigra var. henonis grown in Central China achieves a maximum height of 

400 cm in only 34 days (Li et al. 2005). The culm diameter and height vary with genera 

and species in different site locations (Table 4). The P. makinoi grown in the Dasi site, 

West Taiwan records the highest diameter at breast height (DBH), height, and point density 

(5.9 cm, 11.1 m, and 18767 culm/ha) compared to Jhudong site, West Taiwan (4.7 cm, 10.7 

m, and 17567 culm/ha), respectively. The P. pubescence is grown in the Shi Zhua site, 

Taiwan with a lower temperature (11.5 C), and higher elevation that has a significantly 

higher DBH, height, and culm density (10.6 cm, 21.4 m, and 8344 culm/ha) compared to 

the Hui sun site (6.8 cm, 10.3 m, and 7933 culm/ha), which has temperature and elevation 

of 20.3 °C and 667 m, respectively (Wang and Chen 2015). This indicates that the P. 

pubescens prefers the mountain climate with elevation of 1000 to 1500 m. 

The majority of the monopodial bamboo (Table 5) species is classified as class 4 

and 5, with exception of P. pubescens (Class 2). Physically, P. pubescens is selected for 

manufacturing laminated flooring and other composite board in China. The quantity of 

bamboo strips is proportional to the culm diameter. The P. pubescens records the longest 

culm (21.4 m) for monopodial bamboo. Other monopodial bamboo species produce a culm 

length less than 12 m (Table 4).  



 

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Sympodial Bamboo 
Clump size also influences the culm DBH and height as in sympodial bamboo. 

Generally, the culm’s DBH and height increases with increasing clump size (Table 5). The 

small (4.5 m2), medium (7.13 m2), and large (9.3 m2) clump sizes of B. stenostachya 

produce the DBH (8.7 cm, 9.3 cm and 10.2 cm) and height (15.3 m, 17.4 m, and 20.3 m) 

respectively (Chen et al. 2012). The sympodial bamboos have all the classes as shown in 

Table 5. The species with diameter class 1 including D. asper, D. latiflorus, G. levis, 

Melocanna bambusoides, and Oxytenanthera abyssinica. Others, including B. vulgaris var. 

striata, B. stenostachya, Gigantochloa scortechinii, G. wrayi, and S. grande are classified 

as class 2, the same as P. pubescens. The D. latiflorus and T. oliveri are recorded as the 

longest culm (25 m) for sympodial bamboo. This is followed by B. vulgaris var. striata, D. 

asper, and M. bambusoides (23 m), and lastly, by B. stenostachya, G. ligulata, and S. 

funghomii (20 m).  

 

Overall 
The sympodial bamboo (9.3 cm and 17 m) has a significantly higher DBH and 

height than monopodial bamboo (5.6 cm and 10.8 m), as shown by the statistical analysis 

of the data (Table 5). This factor may be due to the fact that most sympodial bamboo are 

grown in tropical countries that are rich in sunlight and rain for photosynthesis. The starch 

stored in parenchyma is used for the culm growth. By contrast, most monopodial bamboo 

is grown in temperate countries with less sunlight, and the starch are stored in the 

parenchyma that are later used for their sustenance during winter and snow. 

 

 
Table 5. DBH and Maximum Height of Monopodial and Sympodial Bamboo 

Species Diameter Max. 
Height (m) 

Reference 
Max. DBH 

(cm) 
Class 

Monopodial 

P. pubescens 10.8 2 14.6 Inove et al. (2009) 

P. bambusoides 6.8 4 14.9 Inove et al. (2009) 
P. nigra var. henonis 2.3 5 5.9 Inove et al. (2009) 

P. nigra Munro 1.2 5 3.3 Inove et al. (2009) 

P. bambusoides f. Castillon 0.7 5 2.3 Inove et al. (2009) 

P. virida-glaucescens 3.9 5 10.1 Gratani et al. (2008) 

P. pubescens 10.7 3 14.3 Gratani et al. (2008) 
P. bambusoides 4.4 5 6.0 Gratani et al. (2008) 

P. pubescens 6.8 4 10.3 Wang et al. (2009) 

P. pubescens 10.6 2 21.4 Wang et al. (2009) 

P. makinoi 5.2 5 10.2 Wang and Shen (1987) 

P. makinoi 5.0 5 10.7 Wang and Shen (1987) 

P. makinoi 4.7 5 10.7 Wang and Shen (1987) 

P. makinoi 5.9 5 11.1 Wang and Shen (1987) 
P. makinoi 5.4 5 10.8 Wang and Shen (1987) 

P. makinoi 5.2 5 11.3 Wang and Shen (1987) 

Sympodial 

B. blumeana 9 3 13 Azmy and Razak (1991) 

B. heterostachys 5 5 18 Azmy and Razak (1991) 
B. vulgaris 9 3 18 Azmy and Razak (1991) 

B. vulgaris var. striata 10 2 23 Azmy and Razak (1991) 

B. stenostachya 8.7 4 15.3 Chen et al. (2012) 



 

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Table 6. Statistical Analysis of Basic Characteristics of Monopodial and 
Sympodial Bamboos 

Characteristic Monopodial Sympodial DF F Significance 

DBH (cm) 
5.6 

(3.1) 
9.3 

(3.7) 
9 1.93 0.09* 

Height (m) 
10.8 
(4.7) 

17.0 
(5.0) 

9 2.06 0.07* 

 *Significant at P < 0.1 

 

The application of NPK (the proportion of three plant nutrients in order: nitrogen 

(N), phosphorus (P) and potassium (K)) fertilizer of 0.75 to 0.93 kg/clump at 2 or 3 times 

per year could increase the bamboo production for Neosinocalamus affinis (Long and Jiang 

1996). The G. levis clump applied with 12 kg chicken manure gives the highest shoot 

sprouting. The fertilization using cow dung, chicken manure, and rusk husk ash at 0.5, 1, 

and 2 kg per clump during growth and development in G. leavis and D. asper were only 

important at the first year of plantation establishment (Fernandez et al. 2003). The average 

DBH of P. pubescens clump aged more than one year fertilized with N:P:K (244 kg N 

ha/year, 196 kg P/ha, 196 kg K ha/year) for 30 years is identical with the unfertilized clump 

(10.1 cm). This shows that the bamboo achieves its maximum growth in one year, as 

mentioned by Liese (1985) and Banik (2015). 

 

 
ANATOMY 
 
Microstructure 

In the transverse section, the microstructure of monopodial and sympodial bamboo 

is the same as they are both covered by epidermis, hypodermis, and cortex in the outermost 

zone. The epidermis consists of axially elongated cells, shorter cork, silica cells, and 

B. stenostachya 9.3 4 17.4 Chen et al. (2012) 

B. stenostachya 10.2 2 20.3 Chen et al. (2012) 

D. asper 13 1 23 Azmy and Razak (1991) 

D. latiflorus 20 1 25 Lu (2001) 
G. levis 13 1 10 Azmy and Razak (1991) 

G. ligulata 3.5 5 20 Azmy and Razak (1991) 

G. scortechinii 11 2 18 Azmy and Razak (1991) 

G. wrayi 10 2 12 Azmy and Razak (1991) 

S. brachycladum 7 4 21 Azmy and Razak (1991) 

S. grande 11 2 12 Azmy and Razak (1991) 
S. zollingeri 7 4 15 Azmy and Razak (1991) 

Guadua augustifolia 8.41 3 16.7 Riano et al. (2002) 

M. bambusoides 15 1 23 Liese (1985) 

Ochlandra travancorica 5 5 6 Liese (1985) 

Ox. abyssinica 15 1 15 Liese (1985) 

O. albociliata 3 5 10 Liese (1985) 
O. nigro-ciliata 10 3 15 Liese (1985) 

S. brachycladum 7 4 13 Liese (1985) 

S. funghomii 10 3 20 Inbar (2010) 

Teinostachym dulloa 10 3 23 Liese (1985) 

T. oliveri 8 4 25 Liese (1985) 
T. siamensis 6 4 13 Liese (1985) 



 

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stomata. The epidermis cells are always covered on the outside by a cutinized layer of 

cellulose and pectin with tangential lamellation. Its main function is for water blockage 

and tissue protection. The hypodermis is the next layer, and it consists of several layers of 

thick-walled scerenchymatous cells. This is followed by the pith ring or pith periphery, in 

which a non-vascular tissue is composed of layers of parenchyma cells, and often heavily 

thickened and lignified. They are commonly long in tangential direction, but small in radial 

and longitudinal directions. The ground tissue is next to the pith ring. The ground tissue 

contains parenchyma cells with embedded vascular bundle (Grosser and Liese 1971). 

 

Table 7. Vascular Bundle Type Classification by Grosser and Liese (1971, 1973)  

Type Characteristic Occurrence 

I 
Open-
type 

Consisting of one part (central 
vascular strand) with supporting 
tissue as sclerenchyma sheaths; 
intercellular space with tyloses. 
Type I is also called ‘open-type’ 

All species with leptomorph rhizomes 
(Arundinaria, PhylIostachys) 

II 
Tight-
waist 
type 

Consisting of one part (central 
vascular strand); supporting tissue 
as sclerenchyma sheaths; sheath 

at the intercellular space 
(protoxylem) strikingly larger than 

the two lateral ones and extends in 
a fan-like shape; intercellular 

space without tyloses. 
Type II is also called ‘tight-waist 

type’. 

In species with pachymorph rhizomes growing 
either in single culm formation (Melocanna) or in 
clumps (Cepha- lostachyum, Schizostachyum, 

and 
Teinostachyum). In Cephalostachyum, as only 

type throughout the culm; in Melocanna, 
Schizostachyum, and 

Teinostachyum in the base internodes often 
together with type III. 

III Consisting of two parts (central 
vascular strand and one fibre 
strand); fibre strand inside the 

central strand; sheath at the inner 
cellular space (protoxylem) 

generally smaller than the other 
ones. 

In dump-forming species with pachymorph 
rhizomes (Bambusa, Dendrocalamus, 

Gigantochloa, and Thyrsostachys); at the base 
internodes combined mostly with type IV, in the 

middle and upper parts as only type. In 
Melocanna, Schizostachyum, and 

Teinostachyum combined at the base 
internodes with type II. In some Oxytenanthera 

spp. as only type throughout the culm 

IV Consisting of three parts (central 
vascular strand and two fibre 

strands); fibre strands outside and 
inside the central strand. 

In clump-forming species with pachymorph 
rhizomes (Bambusa, Dendrocalamus, 

Gigantochloa, and Thyrsostachys); mostly at the 
base internodes, seldom at the middle part; 

always combined with type III 

 
The thin walls of parenchyma cells are connected to each other by numerous simple 

pits. The cell is thinner than the fibre wall, and they are located mainly on the longitudinal 

walls, while the horizontal walls are scarcely pitted. The parenchyma tissue is lignified 

during the sprouting of the culm and the cells may store a certain amount of starch. A small 

and elongated parenchyma cell, like a cubic cell, appear and are interspersed between the 

long cells. The cubic-like cell is characterized with a denser cytoplasm and thin walls, it is 

not lignified even in mature culm (Grosser and Liese 1971). 

 

Vascular Bundle 
The bamboo vascular bundle consists of two metaxylem vessels, phloem, and 

protoxylem attached to the fibre bundle. The size and shape vary with genera and species, 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6510 

as well as along the internode height and across the culm wall in transverse direction. The 

classification of vascular bundle was slightly different between European and Chinese 

researchers. Grosser and Liese (1971, 1973) on their detailed analysis of 52 species in 14 

genera classified the vascular bundle to four different classes as shown in Table 7. 

Types I, II, III, and IV of the vascular bundles obtained from sympodial bamboo 

are reported by researchers (Suzuki and Itoh 2001; Sharma et al. 2017; Nordahlia et al. 

2019). In contrast to the vascular bundle classified by Grosser and Liese (1971), Xin and 

Qion (1983) classified vascular bundles into four different classes using 10 genera and 45 

species belonging to monopodial bamboo natively from China (Table 8). 

 

Table 8. Classification of Vascular Bundle Type According to Xin and Qion 
(1983) 

Type Characteristic Occurrence 

I Vascular bundles are separated 
by parenchyma. No air canals 

in cortex. 

Bamboo with the following species: 
Phyllostachys arcana, P. aurea, P. 
aureosulcata, P. bambusoides, P. 

bambusoides var. tanakae, P. bambusoides 
var. castilloni, P. bambusoides var. castilloni-
inverssa, P. besseti, P. decora, P. dulcis, P. 
flexsuosa, P. glauca, P. glauca f. Yunzu, P. 
meyeri, P. nigra, P. nigra var. henonis, P. 

nuda, P. nuda f. Localis, P. viridis, P. 
platyglossa, P. praecox, P. Prapinqua, and P. 

pubescens. 

II Vascular bundles are isolated. 
Between them, there are fibre 

cell groups in rectangular, 
round, or variant forms, and 
they are never linked. These 

are mostly peripheral in 
position. No air canals in cortex 

Bamboo with the following species: 
Indocalamus longiauritus, I. victorialis, 

Pleioblastus amarus, PI. gramineus, PI. sp., 
Psudosasa amabilis, and Sinobambusa 

tootsik 
 

III Vascular bundles are linked, 
occasionally with very narrow 

gaps of parenchyma 
interrupting air canals in cortex 

Bamboo with the following species: 
Phyllostachys heteroclada and Ph. nidularia 

IV Vascular bundles are 
connected in the form of a ring 

enclosing stele. No air canals in 
cortex 

Bamboo with the following species: 
Arundinaria fargesii, Chimonobambusa utilis, 
Ch. quadrangular, Ch. purpurea, Qionzbuea 

lumidinoda, Sasa unbigena, and 
Sinarundinaria fangiana 

 

Vascular bundle classification made by Grosser and Liese (1971) is mostly used 

and referred to by many researchers worldwide. In an anatomical structure point of view, 

the sympodial bamboo has more fibre strands or island at the bottom (Type III), and both 

bottom and top (Type IV) of vascular bundle. This generally gives additional support and 

strength to the culm. The monopodial bamboo is structured by only a central vascular 

bundle (Type I) and surrounded with sclerenchyma sheaths. Therefore, the monopodial 

bamboo, such as P. pubescens, is more pliable and softer, which is advantageous during 

processing such as cutting, splitting, moulding, and sanding. The type of vascular bundle 

may be the same or differ within the individual culm of each species. In monopodial 

bamboo, the S. manii grown in India has vascular bundle Type I, as classified by Grosser 

and Liese (1971), along its culm height (Naithani et al. 2010). In contrast, Sharma et al. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6511 

(2017) found a Type III vascular bundle at the basal and middle portions, while the Type 

II of vascular bundle was at the top portion of S. manii. The P. pubescens, P. nigra, and P. 

bambusoides grown in South Korea have vascular bundle Type 1 with tylosoid in the 

intercellular space (Jeon et al. 2018).  

 
Monopodial Bamboo 

The vascular bundle dimension and shape vary with genera, species, site location, 

culm height, and diameter in transverse section (Mohamed et al. 2019). In monopodial 

bamboo (Table 9), the vascular bundle frequency differs across the transverse and 

longitudinal directions. The frequency of vascular bundle in 12 China bamboo species is 

biggest at the outer zone (14/mm2), followed by the middle zone (4.8/mm2), and smallest 

at the inner zone (2.9/mm2). The frequency is also increased with the internode height in 

P. pubescens (Fangchun 2001a). The frequency of vascular bundle is highest in order of 

Arundinaria japonica (7/mm2), Brachystachyum densiflorum (7/mm2), and Indocalamus 

mingoi (6.67/ mm2). 

 

Sympodial Bamboo 
In sympodial bamboo, the S. pergracile has Type II vascular bundle along the culm 

height (Sharma et al. 2017). The B. rigida grown in Sichuan, China has vascular bundle 

Type III at the middle zone of the transverse direction, while Type I and II are located at 

inner and outer zone, respectively (Huang et al. 2015). In the transverse direction, the D. 

brandisii grown in China has vascular bundle Type II, Type III, and Type V at the 

periphery, middle, and outer zones, respectively (Wang et al. 2016). The vascular bundle 

of bamboo grown in Malaysia is dominated with Type II, III, and IV. The species with 

Type II includes S. brachycladum and S. zollingeri. The Type III includes G. thoii, G. 

scortechinii, G. ligulata, G. wrayi, G. brang, S. grande, B. heterostachyum, and B. vulgaris 

cv. vittata. Type IV includes B. vulgaris, B. blumeana, and D. asper. In all cases, the 

vascular bundle Type IV is mostly classified as a thick-walled and large-diameter bamboo 

species (Nordahlia et al. 2019). This indicates that the vascular bundle type may be similar 

to or different from each of the individual culm of the same species. 

The vascular bundle dimension for radial/tangential (R/T) in sympodial bamboo, 

G. scortechinii increases from the inner zone to the outer zone, and it significantly differed 

at the middle portion with age. A smaller vascular bundle tends to be denser than a bigger 

one (Norul Hisham et al. 2006). The R/T ratio of vascular bundle increases from inner zone 

toward the outer zone in basal, middle, and top portion of B. rigida aged 1, 3, and 5 years. 

Overall, the R/T ratio of vascular bundle is not significantly different with culm height for 

all ages (Huang et al. 2015). The radial length and tangential diameter of vascular bundle 

in G. scortechinii is longer at the nodal portion compared to the internode portion (Table 

10). In the internode, the vascular bundle is longest in descending order of G. levis (1171.14 

µm), G. scortechinii (787.2 µm), and G. wrayi (754.1 µm). In the node, the vascular bundle 

length in descending order is G. levis (1193.2 µm), G. scortechinii (1078.2 µm), and G. 

wrayi (963.4 µm). In the internode, the vascular bundle width is highest in G. levis (798.3 

µm), followed by G. scortechini (544.6 µm), and G. wrayi (532.9 µm). In the node, the 

vascular bundle width in descending order is G. levis (720.4 µm), G. scortechinii (587.9 

µm), and G. wrayi (685.8 µm) (Mohd Tamizi et al. 2011). 

The metaxylem vessel diameter increases from the outer zone toward the inner zone 

in all age classes of G. scortechiii (0.5, 1.5. 3.5, 5.5, and 6.5 years) (Table 11). Vessel 

diameter is significantly bigger with age at the outer and inner zones, but this is not the 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6512 

same at the middle zone. The diameter gradually increased from the youngest age of 0.5 

years (0.51 µm) to a maximum diameter at the age of 5.5 years (0.62 µm). The vessel 

diameter in G. scortechinii is also smaller than P. pubescens, which has an average of 0.98 

µm (Fangchun 2000a). A smaller metaxylem vessel diameter is reported in D. brandisii 

(Wang et al. 2016), where it slightly increases with age from 139.4 µm (aged 1 year) to 

162.0 µm  (aged 3 years). The metaxylem vessel is slightly elliptical in B. rigida due to the 

radial length is longer than the tangential diameter (Table 9). The metaxylem vessel 

diameter is not significantly different with age but it tends to be slightly smaller from the 

basal portion toward the top. The diameter also significantly increases from outer part of 

the vessel towards the inner in all ages. The metaxylem diameter ranges from 112.3 m to 

127.4 m, 97.8 m to 127.5 m, and 104.6 m to 129.2 m for bamboo aged 1, 3, and 5 

years, respectively (Huang et al. 2015). Liu et al. (1998) mentioned that larger vessel 

diameter in laminated bamboo of D. latifflorus could probably cause its lower glue bond 

strength compared to P. edulis. 

 

Table 9. Vascular Bundle Frequency in Monopodial and Sympodial Bamboos 

Species Vascular Bundle Frequency (mm2) Reference 

Monopodial 

A. japonica 7.00 Fangchun (2001a) 

Brachystachyum densiflora 7.00 Fangchun (2001a) 

C. quadrangularis 5.00 Fangchun (2001a) 

I. migoi 6.67 Fangchun (2001a) 

P. bambusoides 3.67 Fangchun (2001a) 

P. glauca 5.67 Fangchun (2001a) 

P. pubescens 3.00 Fangchun (2001a) 

Pleioblatus amarus 4.67 Fangchun (2001a) 

S. manii 3.93 Fangchun (2001a) 

Sympodial 

S. pergracile 3.27 Fei et al. (2016) 

S. munroi 2.91 Fei et al. (2016) 

D. giganteus 1.13 Fei et al. (2016) 

B. sinospinosa 1.06 Fei et al. (2016) 

D. farinosus 3.08 Fei et al. (2016) 

B. rigida 1.69 Fei et al. (2016) 

B. pervariadilis 1.51 Fei et al. (2016) 

B. rigida (1 years) 4.98 Huang et al. (2015) 

B. rigida (3 years) 5.04 Huang et al. (2015) 

B. rigida (5 years) 5.04 Huang et al. (2015) 

G. brang 6.38 Mohd. Tamizi et al. (2011) 

G. levis 4.33 Mohd. Tamizi et al. (2011) 

G. scortechinii 7.73 Mohd. Tamizi et al. (2011) 

G. scortechinii (one month) 0.64 Mohamed et al. (2019) 

G. scortechinii (1 year) 0.64 Mohamed et al. (2019) 

G. scortechinii (2 years) 0.64 Mohamed et al. (2019) 

G. scortechinii (3 years) 0.64 Mohamed et al. (2019) 

G. wrayi 6.84 Mohd. Tamizi et al (2011) 

B. blumeana (1 year) 0.84 Mohmod et al. (1993) 

B. blumeana (2 years) 0.66 Mohmod et al. (1993) 

B. blumeana (3 years) 0.82 Mohmod et al. (1993) 

B. blumeana 2.89 Espiloy (1987) 

G. levis 1.56 Espiloy (1987) 

 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6513 

Table 10. Radial Length, Tangential Diameter, and R/T Ratio of Vascular Bundle 
in Monopodial and Sympodial Bamboos 

Species Vascular Bundle (µm) Reference 

 Radial 
(R) 

Tangential 
(T) 

R/T  

Monopodial 

A. japonica 0.40 0.36 1.11 Fangchun (2001a) 

Brachystachyum densiflora 0.34 0.31 1.10 Fangchun (2001a) 

C. quadrangularis 0.29 0.25 1.16 Fangchun (2001a) 

I. migoi 0.20 0.27 0.74 Fangchun (2001a) 

P. bambusoides 0.54 0.48 1.13 Fangchun (2001a) 

P. glauca 0.45 0.35 1.29 Fangchun (2001a) 

P. pubescens 0.50 0.49 1.02 Fangchun (2001a) 

Pleioblatus amarus 0.42 0.43 0.98 Fangchun (2001a) 

Sympodial 

G. scortechinii (0.5 year) - - 1.25  Norul Hisham et al. 
(2006) 

G. scortechinii (1.5 years) - - 1.28  Norul Hisham et al. 
(2006) 

G. scortechinii (3.5 years) - - 1.20  Norul Hisham et al. 
(2006) 

G. scortechinii (5.5 years) - - 1.20   Norul Hisham et al. 
(2006) 

G. scortechinii (6.5 years) - - 1.22   Norul Hisham et al. 
(2006) 

G. brang 0.79 0.51 1.55 Mohd Tamizi et al. (2011) 

G. levis 1.17 0.80 1.46 Mohd Tamizi et al. (2011) 

G. scortechinii 0.79 0.50 1.58 Mohd Tamizi et al. (2011) 

G. wrayi 0.75 0.53 1.42 Mohd Tamizi et al. (2011) 

D. giganteus 0.86 0.70 1.43 Fei et al. (2016) 

B. sinospinosa 0.71 0.74 0.96 Fei et al. (2016) 

D. farinosus 0.66 0.55 1.20 Fei et al. (2016) 

B. rigida 0.76 0.64 1.19 Fei et al. (2016) 

B. pervariadilis 0.79 0.70 1.13 Fei et al. (2016) 

B. rigida (1 year) - - 1.25 Huang et al. (2015) 

B. rigida (3 years) - - 1.18 Huang et al. (2015) 

B. rigida (5 years) - - 1.16 Huang et al. (2015) 

B. blumeana (1 year) - - 1.00 Mohmod et al. (1990) 

B. blumeana (2 years) - - 0.91 Mohmod et al. (1990) 

B. blumeana (3 years) - - 0.95 Mohmod et al. (1990) 

B. vulgaris (1 year) - - 1.38 Mohmod et al. (1990) 

B. vulgaris (2 years) - - 1.33 Mohmod et al. (1990) 

B. vulgaris (3 years) - - 1.33 Mohmod et al. (1990) 

G. scortechinii (1 year) - - 0.84 Mohmod et al. (1990) 

G. scortechinii (2 years) - - 1.36 Mohmod et al. (1990) 

G. scortechinii (3 years) - - 1.11 Mohmod et al. (1990) 

 
Overall Features 

The radial length and tangential diameter of the vascular bundle is significantly 

higher in monopodial bamboo (0.39 m and 0.37 m) than sympodial bamboo (0.28 m 

and 0.22 m), respectively, as shown in the statistical reanalysis results (Table 12). While 

the R/T ratio of vascular bundle is significantly higher in sympodial bamboo (1.23) than 

the monopodial bamboo (1.07), the metaxylem vessel diameter is significantly higher in 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6514 

sympodial (127.97 m) than the monopodial bamboo (72.38 m). In contrast, the vascular 

bundle frequency is significantly higher in monopodial (5.18/mm2) than the sympodial 

bamboo (2.80/mm2).  

 

Table 11. Metaxylem Vessel Diameter in Monopodial and Sympodial Bamboos 

Species Metaxylem Vessel 

Diameter (m) 

Reference 

Monopodial 

A. japonica 54.0 Fangchun (2001a) 

Brachystachyum densiflora 76.3 Fangchun (2001a) 

C. quadrangularis 57.3 Fangchun (2001a) 

I. migoi 45.3 Fangchun (2001a) 

P. bambusoides 109.7 Fangchun (2001a) 

P. glauca 78.3 Fangchun (2001a) 

P. pubescens 98.0 Fangchun (2001a) 

P. b.f zitchiku 58 Fangchun (2001a) 

P. congesta 65 Fangchun (2001a) 

P. nuda 68 Fangchun (2001a) 

Pleioblatus amarus 91.0 Fangchun (2001a) 

A. amabiris 89 Fangchun (2001a) 

Chimonobambusa marmorea 73 Fangchun (2001a) 

C. quadragularis 62 Fangchun (2001a) 

C. utilis 77 Fangchun (2001a) 

S. manii 56.2 Sharma et al. (2017) 

Sympodial 

B. rigida (1 year) 121.4 Huang et al. (2015) 

B. rigida (3 years) 113.2 Huang et al. (2015) 

B. rigida (5 years) 117.0 Huang et al. (2015) 

D. brandisii (1 year) 139.4 Wang et al. (2016) 

D. brandisii (2 years) 147.1 Wang et al. (2016) 

D. brandisii (3 years) 162.0 Wang et al. (2016) 

G. scortechinii (0.5 year) 51.0  Norul Hisham et al. (2006) 

G. scortechinii (1.5 years) 54.0  Norul Hisham et al. (2006) 

G. scortechinii (3.5 years) 57.0  Norul Hisham et al. (2006) 

G. scortechinii (5.5 years) 62.0  Norul Hisham et al. (2006) 

G. scortechinii (6.5 years) 50.0  Norul Hisham et al. (2006) 

Neosinocalamus affinis 191.5 Luo et al. (2019) 

B. intermedia 168.7 Luo et al. (2019) 

B. multiplex 170.4 Luo et al. (2019) 

B. rigida 185.1 Luo et al. (2019) 

B. blumeana 165.5 Espiloy (1987) 

G. levis 220.2 Espiloy (1987) 

 

  



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6515 

Table 12. Statistical Reanalysis of the Vascular Bundle Elements in Monopodial 
and Sympodial bamboos 

Characteristic Monopodial Sympodial DF F Significance 

Radial length 
0.39 

(0.11) 
0.28 

(0.40) 
1 0.08 0.44NS 

Tangential diameter 
0.37 

(0.09) 
0.22 

(0.25) 
1 0.14 0.20NS 

R/T 
1.07 

(0.16) 
1.23 

(0.19) 
1 0.16 0.04** 

Vascular bundle frequency 
5.18 

(0.69) 
2.80 

(0.43) 
1 8.47 0.000** 

Metaxylem vessel diameter 
72.38 

(17.69) 
127.97 
(49.92) 

1 14.6 0.001** 

DF– Degree of freedom, F– F ratio, NS is not significant at P > 0.1, * is significant at P < 0.1, ** is 
significant at P < 0.05. The value in parentheses is standard deviation. 
 

The proportion of metaxylem vessel diameter is significantly higher in sympodial 

(6.40%) than the monopodial bamboo (5.21%), but not for fibre and parenchyma 

proportions. The proportion of fibre and parenchyma are not significantly different for 

monopodial (40.92% and 45.56%) and sympodial (53.00% and 53.38%) bamboos, 

respectively. 
 

Fibre Morphology 
Monopodial bamboo 

In monopodial bamboo (Table 13), I. migoi and P. pubescens growing in China 

have the longest fibre (2250 m), while the Indocalamus tessellatus grown in China has 

the shortest fibre (1435 m). The Brachycladum densiflorus grown in China records the 

widest fibre (16.9 m), while the Arundinaria amabilis also grown in China has the thickest 

fibre wall, and the P. viridis grown in USA has the widest fibre lumen (5.63 m). The fibre 

of the three monopodial bamboos grown in Taiwan is the longest, when ranked in 

descending order of Phyllostachys bambusoides (2033 to 2239 m), P. nigra (1934 to 2199 

m), and P. pubescens (1375 to 1573 m). The outer zone has a longer fibre than the inner 

zone for all species (Jeon et al. 2018). 
 

Table 13. Fibre Morphology in Monopodial and Sympodial Bamboos 

Species Origin FL 

(m) 

FD 

(m) 

FLD 

(m) 

FWT 

(m) 

References 

Monopodial 

A. amabilis China 2338 15.3 3.81 5.23 Fangchun (2001a) 

A. japoniea China 1990 15.8 2.57 6.18 Fangchun (2001a) 

Brachystachyum densiflorum China 2175 16.9   Fangchun (2001a) 

C. quadrangularis China 1700 14.8 3.48 3.63 Fangchun (2001a) 

C. utilitis China 2230 11.9 4.67 2.41 Fangchun (2001a) 

I. migoi China 2250 14.0   Fangchun (2001a) 

I. tessellatus China 1435 13.5   Fangchun (2001a) 

P. congesta China 1784 12.7 2.82 4.02 Fangchun (2001a) 

P. flexuosa USA 1540 9.6 4.09 3.28 Fangchun (2001a) 

P. heterocycle Brasil 1690 8.7 3.17 3.94 Fangchun (2001a) 

P. pubescens China 2250 13.6 3.12 3.75 Fangchun (2001a) 

P. viridis China 1886 14.3 4.14 2.87 Fangchun (2001a) 

P. viridis USA 1690 11.4 5.63 2.10 Fangchun (2001a) 



 

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Pleioblastus amarus China 2129 14.4 2.49 5.78 Fangchun (2001a) 

P. pubescens Korea 1474    Jeon et al. (2018) 

P. nigra Korea 2066    Jeon et al. (2018) 

P. bambusoides Korea 2136    Jeon et al. (2018) 

Sympodial 

B. basihirsuta China 1667 14.4 2.35 2.14 Fangchun (2001a) 

B. boniopis China 1788 14.2 4.24 1.44 Fangchun (2001a) 

B. corginera China 2482 16.3 2.50 2.56 Fangchun (2001a) 

B. dessimulator China 1861 18.0 3.38 2.13 Fangchun (2001a) 

B. eutuldoides  China 1993 16.9 1.66 4.08 Fangchun (2001a) 

B. glaucescens China 2115 14.9 2.13 3.39 Fangchun (2001a) 

B. glaucescens  China 2079 15.9 2.49 2.85 Fangchun (2001a) 

B. gibboides China 2135 16.8 2.65 3.16 Fangchun (2001a) 

B. lapidea China 2390 10.8 5.59 2.05 Fangchun (2001a) 

B. lapidea China 2363 13.5 3.23 2.23 Fangchun (2001a) 

B. longiflora China 1806 16 2.05 3.61 Fangchun (2001a) 

B. multiplex China 2385 13.1 2.78 4.68 Fangchun (2001a) 

B. pervariabilis China 2036 14.1 3.71 2.83 Fangchun (2001a) 

B. rigida China 2230 13.7 5.77 1.22 Fangchun (2001a) 

B. sinospinosa China 2450 16.2 7.34 1.37 Fangchun (2001a) 

B. spinosa China 2270 14.7 5.28 1.94 Fangchun (2001a) 

B. textillis China 2480 14.9 3.37 4.63 Fangchun (2001a) 

B. textillis China 2236 14.4 2.68 4.10 Fangchun (2001a) 

B. textillis var. a China 1968 16.1 2.68 2.54 Fangchun (2001a) 

B. textillis var. g China 1842 14 4.08 1.42 Fangchun (2001a) 

D. gigantus Thai 2487 18.5 5.26 1.00 Fangchun (2001a) 

D. Oldhami China 2334 15.0 4.30 2.16 Fangchun (2001a) 

D. strictus China 2236 15.3 3.69 2.05 Fangchun (2001a) 

D. strictus China 2800 9.9 3.69 2.05 Fangchun (2001a) 

Dinochlua utilitis China 2340 16.3 5.48 1.26 Fangchun (2001a) 

Lignalia chungii China 2507 13.2 4.09 1.36 Fangchun (2001a) 

L. remotiflora China 2071 15.2 3.28 2.51 Fangchun (2001a) 

L. surectta China 2186 17 2.98 2.11 Fangchun (2001a) 

S. dumetorum China 2446 15 4.83 1.80 Fangchun (2001a) 

S. fungnomii China 2840 13.7 4.74 2.03 Fangchun (2001a) 

S. nalhannense China 2444 13.8 1.62 4.69 Fangchun (2001a) 

S. lima China 3190 15.2 5.04 2.05 Fangchun (2001a) 

S.pseudolima China 2135 17.8 3.22 2.43 Fangchun (2001a) 

Sinarundinaria chungii China 2260 11.7 1.59 4.53 Fangchun (2001a) 

Sinocalamus affinis China 2220 15 4.43 1.75 Fangchun (2001a) 

Sn. affinis China 2710 13.6 4.82 1.90 Fangchun (2001a) 

Sinocalamus pubescens China 1938 15.3 2.40 2.70 Fangchun (2001a) 

Sn. bicicatricatus China 2008 15 2.35 3.33 Fangchun (2001a) 

Sn. farinosus China 2670 16.9 3.28 5.03 Fangchun (2001a) 

Sn. latiflorus China 2880 14 4.74 2.62 Fangchun (2001a) 

Sn. latiflorus China 1830 14.5 3.78 3.01 Fangchun (2001a) 

Sn. minor China 2297 17.8 4.39 1.56 Fangchun (2001a) 

Sn. minor China 2920 10 3.28 3.00 Fangchun (2001a) 

Sn. oldhamii China 2480 13.8 4.46 2.69 Fangchun (2001a) 

Sn. stenoauritus China 1976 16.9 2.42 2.52 Fangchun (2001a) 

Sn. vario-striatus China 2198 16.5 3.2 1.94 Fangchun (2001a) 

Thamnocalamus siamensis China 2006 16.8 2.21 2.94 Fangchun (2001a) 

B. blumeana (1 year) Msia 1940 18 10 5 Mohmod et al. 
(1990) 

B. blumeana (2 years) Msia 1870 20 9 5 Mohmod et al. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6517 

(1990) 

B. blumeana (3 years) Msia 1950 20 9 5 Mohmod et al. 
(1990) 

B. vulgaris var. striata (1 
year) 

Msia 3340 17 2 7 Mohmod et al. 
(1990) 

B. vulgaris var. striata (2 
years) 

Msia 3300 17 3 7 Mohmod et al. 
(1990) 

B. vulgaris var. striata (3 
years) 

Msia 3760 17 2 6 Mohmod et al. 
(1990) 

G. scortechinii (1 years) Msia 3500 16 2 7 Mohmod et al. 
(1990) 

G. scortechinii (2 years) Msia 3800 17 2 7 Mohmod et al. 
(1990) 

G. scortechinii (3 years) Msia 4240 17 3 8 Mohmod et al. 
(1990) 

G. scortechinii (0.5 year) Msia 2230 26 10 8 Norul Hisham et al. 
(2006) 

G. scortechinii (1.5 years) Msia 2500 26 8 9 Norul Hisham et al. 
(2006) 

G. scortechinii (3.5 years) Msia 2500 26 8 9 Norul Hisham et al. 
(2006) 

G. scortechinii (5.5 years) Msia 2380 26 9 8 Norul Hisham et al. 
(2006) 

G. leavis Msia 2040 23.7 4.00 9.34 Razak et al. (2013) 

G. scortechinii Msia 1745 17.3 8.66 4.30 Razak et al. (2013) 

G. wrayi Msia 1799 17.9 3.83 7.02 Razak et al. (2013) 

G. brang Msia 1910 22.8 4.75 9.02 Mohd Tamizi et al. 
(2011) 

G. thoii Msia 4071 25.5 5.0 12.2 Nordahlia et al. 
(2019) 

G. ligulata Msia 3930 22.6 4.3 9.2 Nordahlia et al. 
(2019) 

G. wrayi Msia 2753 24 10.3 7.0 Nordahlia et al. 
(2019) 

G. brang Msia 3543 21.4 6 7.7 Nordahlia et al. 
(2019) 

S. brachycladum Msia 2840 22.2 6.4 7.9 Nordahlia et al. 
(2019) 

S. grande Msia 2451 15 3.1 6.1 Nordahlia et al. 
(2019) 

S. zollingeri Msia 2326 14.8 8.1 3.3 Nordahlia et al. 
(2019) 

B. vulgaris Msia 2494 14.1 3.5 7.1 Nordahlia et al. 
(2019) 

B. blumeana Msia 2905 18.9 7.6 5.7 Nordahlia et al. 
(2019) 

B. heterostachya Msia 3764 26.8 5.8 10.5 Nordahlia et al. 
(2019) 

B. vulgaris cv. Vitta Msia 3592 20.3 6.9 6.3 Nordahlia et al. 
(2019) 

D. asper Msia 2998 23.3 6.1 8.6 Nordahlia et al. 
(2019) 

FL: Fibre length; FD: fibre diameter; FLD: fibre lumen diameter; FWT: fibre wall thickness 

 

 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6518 

Sympodial bamboo 

In sympodial bamboo (Table 14), the longest and shortest fibres are recorded for 

G. scortecinii (4240 m) grown in Malaysia and B. basihirsuta (1667 m) grown in China, 

respectively. The fibre is widest in B. heterostachya (26.8 m) grown in Malaysia and 

narrowest in D. striticus (9.9 m) grown in China. The fibre wall is thickest in G. thoii 

(12.2 m) grown in Malaysia and thinnest in D. gigantus (1 m) grown in Thailand. The 

fibre lumen is largest in B. blumeana (10 m) grown in Malaysia and smallest in 

Sinarundina chungi (1.59 m) grown in China. The fibre wall diameter, thickness, and 

lumen diameter of B. blumena, B. vulgaris, and G. scortechinii are not significantly 

different with culm aged one to three years. The fibre length ranges from 1.89 m to 1.99 

m in B. blumeana, from 3.30 m to 3.76 m in B. vulgaris, and from 3.50 m to 4.24 m 

in G. scortechinii. The fibre diameter ranges from 0.018 m to 0.02 m in B. blumeana, 

0.017 m in B. vulgaris and G. scortechinii. The fibre wall thickness is 0.05 in B. 

blumeana, ranges from 0.06 to 0.07 in B. vulgaris, and 0.07 to 0.08 in G. scortechinii. The 

fibre lumen diameter ranges from 0.009 to 0.010 in B. blumana, from 0.002 to 0.003 in B. 

vulgaris, and G. scortechinii (Mohmod et al. 1990). 

In the G. scortechinii aged 0.5 to 6.5 years grown in the same clump, the youngest 

culm aged 0.5 years had the shortest fibre (2.23 m). Culm aged 1.5 years had the longest 

fibre, but the fibre tends to be shorter with ageing. The fibre diameter does not differ by 

age, and the mean is 26 μm. The widest fibre lumen diameter is recorded in the youngest 

culm aged 0.5 years (10 μm) and the diameter remains unchanged beyond age 1.5 years. 

The fibre wall is thinner at the early age of 0.5 years (8 μm) but thickens as much as 1 μm 

at age of 1.5 years (Norul Hisham et al. 2006). The fibre length is significantly higher for 

the internode than the node with an average of 2074.2 μm and 1672.6 μm in G. brang, G. 

levis, G. scortechinii, and G. wrayi. The fibre is longest in the middle section (2064.4 μm), 

followed by the inner (1861.4 μm) and outer (1698.5 μm) for the above three species. The 

fibre is wider in the node (22.0 μm) than the internode (18.2 μm). It is the widest in 

descending order of G. brang (22.8 μm), G. levis (22.7 μm), G. wrayi (17.9 μm), and G. 

scortechinii (17.3 μm), respectively. The fibre is also widest at the middle section (22.4 

μm), followed by the inner (19.6 μm) and outer sections (18.5 um) across the culm wall.  

The fibre lumen is widest in descending order of G. scortechinii (8.60 um), G. 

brang (4.75 um), G. levis (4.75 um), and G. wrayi (4.75 um). The lumen is also larger in 

node compared to the internode. The lumen diameter is largest at the inner section (5.96 

um) and smaller toward the outer section (5.44 um). The fibre wall is thickest in descending 

order of G. levis (9.34 m), G. brang (9.02 m), G. wrayi (7.02 m), and G. scortechinii 

(4.30 m). The node has a thicker fibre wall than the internode. The fibre wall is also 

thickest in the middle zone (8.43 m) followed by the outer (7.03 m) and inner (6.80 m) 

zones (Mohd Tamizi et al. 2011). The fibre length in Schizostachyum manii, Scizostachyum 

munroi, and Schizostachyum pergracile grown in India increased from the inner zone and 

reached a maximum at the middle zone but further decreased toward the inner zone in the 

transverse section of all culm height. The fibre characteristics, such as diameter, lumen 

diameter, and wall thickness decreased along the culm height (Sharma et al. 2017). 

Amongst the 4-year-old G. scortechinii, G. thoii, G. ligulate, G. wrayi, G. brang, S. 

brachycladum, S. grande, S. zollingeri, B. vulgaris, B. blumeana, B. heterostachya, B. 

vulgaris cv Vittata, and D. asper grown in Malaysia, G. thoii has the longest fibre (4070 

m) and the fibre length for all species are ranged from 2330 m to 4070 m (Norhadlia 

et al. 2019). 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6519 

Table 14. Statistical Reanalysis of the Fibre Morphology in Monopodial and 
Sympodial Bamboos 

Fibre Monopodial Sympodial DF F Significance 

Length 1929.7(273.6) 2494.8 (6.09) 1 9.08 0.00*** 

Diameter 12.95 (2.33) 17.06 (3.92) 1 11.34 0.00*** 
Lumen diameter 3.64 (0.96) 4.49 (2.25) 1 1.52 0.22NS 

Wall thickness 3.93 (1.32) 4.39 (2.72) 1 0.31 0.58NS 

DF– Degree of freedom, F– F ratio, NS is not significant at P > 0.1, *** is significant at P < 0.01. 
The value in parentheses is standard deviation 
 

The statistical reanalysis of the fibre morphology data shows that the fibre length 

and diameter are significantly longer and wider in sympodial bamboo (2494.8 m and 

17.06 m) than the monopodial bamboo (1929.7 m and 12.95 m), respectively. In 

contrast, fibre lumen diameter and wall thickness are not significantly different in 

monopodial (3.63 m and 3.93 m) and sympodial (4.49 m and 4.39 m) bamboos, 

respectively. 
 

 

PHYSICAL PROPERTIES 
 
Basic Density and Volumetric Swelling 

The density is closely related to the mechanical properties of bamboo and it differs 

by species, age, culm height, and portion. The density increases from the basal portion 

towards the top as well as from inner to the outer culm wall. The main reason for this trend 

is that the top and outer portion are heavily distributed by vascular bundles and a thinner 

vessel diameter.  

 

Monopodial Bamboo 
The density progressively increases with age as occurs in P. pubescens and S. affinis 

aged 1 (0.43 g/cm3 to 0.49 g/cm3) to 7 years (0.62 g/cm3 and 0.63 g/cm3), respectively, but 

it tends to decrease beyond 8 years (Table 16). The density is higher for bamboo grown in 

drought and low temperature areas, as it has compact tissue. In contrast, the bamboo density 

is lower in warm, moist climates and nourishing soil because its thick culm wall tissue is 

loose (Fangchun 2001a). 

The same trend occurs for N. affinis grown in China, for which the density 

significantly increases with age for 1 year (0.56 g/m3), 2 years (0.68 g/m3), and 3 years 

(0.77 g/m3). However, all bamboo age classes fertilized with potassium, calcium, and 

nitrogen show a lower density than the unfertilized bamboo especially with a higher dose 

of nitrogen (Xie et al. 2019). These results are in a good agreement with the findings of 

Yang et al. (2014), in which the long-term nitrogen fertilization significantly decreased the 

basic density of P. pubescens. The density increases from basal portion (0.60 g/m3) toward 

the middle (0.69 g/m3) and top (0.79 g/m3) portions of B. vulgaris aged 4 years grown in 

China (Huang et al. 2014). The volumetric swelling is highest and lowest in B. sinospinosa 

(32.8%) and P. glauca F. Youzhu (7.4%). 

 

Sympodial Bamboo 
The density decreases with age for 1-year (1.10 g/m3), 2-years (1.04 g/m3), and 3-

years-old (1.00 g/m3) wild B. blumeana culms. In contrast, the density increases with ages 

in 1-year (0.29 g/m3), 2-years (0.51 g/m3), and 3-years-old (0.54 g/m3) wild B. vulgaris 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6520 

var. striata. The same trend occurs in wild G. scortechinii, where the density increases with 

age in 1-year (0.47 g/m3), 2-years (0.53 g/m3), and 3-years-old (0.56 g/m3) culms. Overall, 

B. blumeana has the highest density follow by G. scortechinii and, lastly B. vulgaris (Abdul 

Latif et al. 1990). The density of wild G. scortechinii grown in the same clump also 

increased with age for 0.5 (0.53 g/m3), 1.5 (0.59 g/m3), 3.5 (0.61 g/m3), 5.5 (0.63 g/m3), 

and 6.5 years (0.68 g/m3). Along the internode height, the density trends are also slightly 

increased from the bottom portion toward the top portion along the sixth internode, for all 

age classes ( Norul Hisham et al. 2003; Norul Hisham et al. 2006).  

The densities of 4-year-old Malaysian bamboo, as reported by Nordahlia et al. 

(2019) in descending order, are G. thoii (0.75 g/m3), G. scortechinii (0.64 g/m3), G. wrayii 

(0.63 g/m3), S. grande (0.63 g/m3), B. vulgaris (0.61 g/m3), S. brachycldum (0.59 g/m3), D. 

asper (0.56 g/m3), B. vulgaris cv vittata (0.55 g/m3), G. brang (0.54 g/m3), B. heterostachya 

(0.53 g/m3), B. blumeana (0.48 g/m3), G. ligulata (0.44 g/m3), and S. zollingeri (0.36 g/m3). 

Within the culm region, the node density is not significantly different from internode 

density either at basal or top portions for D. asper grown in Thailand. The density at basal 

portion of culm without node (0.71 g/m3) is not significantly different from the one with 

node (0.69 g/m3). The density at the top portion of culm without node (0.90 g/m3) is not 

significantly different from the one with node (0.92 g/m3). The same trend is reported for 

P. bambisoides (monopodial type) by Tomak et al. (2012) as well as for Guadua 

angustifolia (sympodial type) by De Vos (2010). 

 

Overall Analysis 
The statistical reanalysis of the density in monopodial and sympodial bamboos 

shows that there is no significant different of density (0.58 g/cm3 and 0.59 g/cm3) for both 

types of bamboo. In contrast, the volumetric shrinkage is significantly higher for sympodial 

(16.92%) than the monopodial (12.63%) bamboos. 

 

Table 15. Basic Density and Shrinkage Properties of Monopodial and Sympodial 
Bamboo 

Species Basic Density 

(g/cm3) 
Shrinkage 

(Volume, %) 
Reference 

Monopodial 

Arundinaria 

A. amabilis 0.63 10.2 Fangchun (2001a) 

A. fargesii 0.54 15.2 Fangchun (2001a) 

A. japonica 0.63 - Fangchun (2001a) 

A. spongisa 0.50 22.4 Fangchun (2001a) 

Chimonobambusa 

C. marmorea 0.60 - Fangchun (2001a) 

C. quadrangularis 0.51 - Fangchun (2001a) 

C. utilis 0.58 - Fangchun (2001a) 

Indosasa 

I. longspicta 0.51 15.4 Fangchun (2001a) 

Phyllostachys 

P. angusta 0.49 - Fangchun (2001a) 

P. aurea 0.91 - Fangchun (2001a) 

P. spec bilis 0.51 - Fangchun (2001a) 

P. bambusoidea 0.72 10.4 Fangchun (2001a) 

P. bambusoidea  f. 
Tanaka 

0.51 13.6 Fangchun (2001a) 



 

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P. bambusoidea. f. 
Zitchiku 

0.53 10.3 Fangchun (2001a) 

P. congesta 0.65 - Fangchun (2001a) 

P. decora 0.55 - Fangchun (2001a) 

P. filifera 0.63 - Fangchun (2001a) 

P. flexuosa 0.57 - Fangchun (2001a) 

P. gluaca 0.68 10.4 Fangchun (2001a) 

P. glauca f. Youzhu 0.63 7.4 Fangchun (2001a) 

P. glauca f. 
variabilis 

0.66 9.2 Fangchun (2001a) 

P. decora 0.55 - Fangchun (2001a) 

P. heteroclada 0.52 - Fangchun (2001a) 

P. iridensclada 0.43 - Fangchun (2001a) 

P. kwangsinsis 0.59 - Fangchun (2001a) 

P. meyeri 0.43 12.4 Fangchun (2001a) 

P. nidularia 0.45 - Fangchun (2001a) 

P. nigra 0.72 - Fangchun (2001a) 

P. nigra var. henosis 0.44 - Fangchun (2001a) 

P. nuda 0.64 - Fangchun (2001a) 

P. parvifloria 0.59 - Fangchun (2001a) 

P. platyglossa 0.47 - Fangchun (2001a) 

P. praecox 0.66 - Fangchun (2001a) 

P. pubescens 0.66 14.7 Fangchun (2001a) 

P. viridi-
glaucessens 

0.50 - Fangchun (2001a) 

P. viridis 0.62 - Fangchun (2000) 

P. vivax 0.64 - Fangchun (2000) 

Pleioblastus 

P.amarus 0.57 - Fangchun (2001a) 

Sympodial 

Bambusa 

B. badihirsuta 0.67 18.5 Fangchun (2001a) 

B. breviflora 0.57 10.2 Fangchun (2001a) 

B. cornigera 0.60 15.1 Fangchun (2001a) 

B. dissemualator 0.60 - Fangchun (2001a) 

B. dissimilis 0.60 - Fangchun (2001a) 

B. dolichoclad 0.73 - Fangchun (2001a) 

B. eatuldoies 0.57 14.2 Fangchun (2001a) 

B. e. var. basistriata 0.77 17.2 Fangchun (2001a) 

B. flexuosa 0.45 13.8 Fangchun (2001a) 

B. gibba 0.57 18.1 Fangchun (2001a) 

B. gibboides 0.54 18.6 Fangchun (2001a) 

B. lapidea 0.50 18.7 Fangchun (2001a) 

B. longiflora 0.57 20.2 Fangchun (2001a) 

B. multiplex cv. 0.50 - Fangchun (2001a) 

B. multiplex f. lutea 0.55 18.4 Fangchun (2001a) 

B. pervarialis 0.58 22.0 Fangchun (2001a) 

B. p. var. viridi-st 0.52 15.9 Fangchun (2001a) 

B. rigida 0.55 16.0 Fangchun (2001a) 

B. rigida 0.69 14.2 Huang et al. (2014) 

B. rutita 0.61 11.1 Fangchun (2001a) 

B. sinospinosa 0.50 32.8 Fangchun (2001a) 

B. stemostachy 0.64 23.1 Fangchun (2001a) 

B. spimosa 0.36 - Fangchun (2001a) 

B. textilis 0.69 - Fangchun (2001a) 



 

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B. t. var. sracillis 0.58 - Fangchun (2001a) 

B. tulda 0.65 13.0 Fangchun (2001a) 

B. tubdoides 0.59 14.3 Fangchun (2001a) 

B. ventricosa 0.42 10.5 Fangchun (2001a) 

B. vulgaris 0.68 10.2 Fangchun (2001a) 

B. vulgaris 0.61 - Nordahlia et al. (2019) 

B. vulgaris cv vittata 0.55 - Nordahlia et al. (2019) 

B. vulgaris var. 
striata 

0.65 13.7 Fangchun (2001a) 

B. vulgaris var. striata 

1 year 0.30 - Mohmod et al. (1990) 

2 years 0.51 - Mohmod et al. (1990) 

3 years 0.54 - Mohmod et al. (1990) 

B. bambos 0.87 - Srivaro and Jakranod (2016) 

B. longispiculata 0.80 - Srivaro and Jakranod (2016) 

B. blumeana 0.77 - Srivaro and Jakranod (2016) 

B. blumeana 

1 year 1.03 - Mohmod et al. (1990) 

2 years 1.04 - Mohmod et al. (1990) 

3 years 1.00 - Mohmod et al. (1990) 

B. blumeana 0.48 - Nordahlia et al. (2019) 

B. heterostachya 0.53 - Nordahlia et al. (2019) 

Dendrocalamus 

D. beecheyanus 0.59 26.7 Fangchun (2001a) 

D. beecheyanus 
var. pubuscens 

0.72 9.6 Fangchun (2001a) 

D. bicicatriatus 0.53 16.0 Fangchun (2001a) 

D. gigantens 0.55 19.6 Fangchun (2001a) 

D. gromdis 0.48 21.8 Fangchun (2001a) 

D. hamilthonii 0.69 20.3 Fangchun (2001a) 

D. mino var. 
pubencen 

0.56 14.3 Fangchun (2001a) 

D. stenoauritus 0.74 14.0 Fangchun (2001a) 

D. stricticus 0.50 - Fangchun (2001a) 

D. validus 0.47 24.4 Fangchun (2001a) 

D. vario-striatus 0.75 11.8 Fangchun (2001a) 

Gigantochloa 

G. scortechinii 

1 year 0.47 - Mohmod et al. (1990) 

2 years 0.53 - Mohmod et al. (1990) 

3 years 0.56 - Mohmod et al. (1990) 

G. scortechinii 0.64 - Nordahlia et al. (2019) 

G. thoii 0.75 - Nordahlia et al. (2019) 

G. ligulata 0.44 - Nordahlia et al. (2019) 

G. wrayi 0.63 - Nordahlia et al. (2019) 

G. brang 0.54 - Nordahlia et al. (2019) 

G. brachycladum 0.59 - Nordahlia et al. (2019) 

G. atter 

0.3 year 0.35 - Marsoem et al. (2015) 

1.3 years 0.43 - Marsoem et al. (2015) 

3.3 years 0.59 - Marsoem et al. (2015) 

Lignania 

L. cerosissina 0.69 - Fangchun (2001a) 

L. chungii 0.42 - Fangchun (2001a) 

L. papilata 0.77 16.9 Fangchun (2001a) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6523 

Schizostachyum 

S. fungbomii 0.50 - Fangchun (2001a) 

S. hainanense 0.46 - Fangchun (2001a) 

S. pseudolima 0.48 - Fangchun (2001a) 

S. manii 0.50 - Nordahlia et al. (2019) 

S. munroi 0.57 - Nordahlia et al. (2019) 

S. pergracile 0.64 - Nordahlia et al. (2019) 

S. grande 0.63  Nordahlia et al. (2019) 

S. zollingeri 0.36 - Nordahlia et al. (2019) 

Sinobambusa 

S. laeta 0.45 - Fangchun (2001a) 

 

Table 16. Basic Density and Volumetric Shrinkage of Monopodial and Sympodial 
Bamboos 

Fibre Monopodial Sympodial DF F Significance 

Basic density 

(g/cm3) 
0.58 (0.10) 0.59 (0.14) 1 0.11 0.74NS 

Volumetric 
shrinkage (%) 

12.63 (3.99) 16.92 (5.10) 1 6.93 0.01*** 

 DF– Degree of freedom, F– F ratio, NS is not significant at P > 0.1, *** is significant at P < 0.0. 
The value in parentheses is standard deviation 

 
 

MECHANICAL PROPERTIES 
 

Bamboo has high mechanical and workability properties and is a preferable 

material for many uses in agricultural, industrial, building, and architecture sectors. The 

flexural ductility of bamboo (Ph. pubescens) is 3.06 times that of wood (Tectona grandis). 

The bending MOR of bamboo (P. pubescens) is 1.72 times that of wood (Tectona grandis), 

while its MOE is 0.84 higher than wood at similar density. High length-to-width ratio, 

density, and strength of bamboo fibers, as well as their parallel orientation all contributed 

to the excellent ductility and strength properties of bamboo (Chen et al. 2020). The 

mechanical properties differ with the bamboo culm position, portion, and section. 

Generally, it is higher at the upper position than the lower position, and higher at the outer 

section of the culm wall than the inner section. This gives rise to a denser and larger 

vascular bundle in the outer section than at the inner section (Fangchun 2001b). 

 

Monopodial Bamboo 
The effect of age on the mechanical properties of bamboo is significantly important 

for choosing an optimum harvesting age, the quality of end products, and service life ( 

Norul Hisham et al. 2006). The compression and tensile strengths of Ph. pubescence 

gradually increase from one to five years but are almost constant from six to eight years 

before slightly declining from nine to ten years. Therefore, it is assumed that the best 

harvesting age for P. pubescens ranges from six to eight years (Fangchun 2001b). The 

mechanical properties of bamboo are clearly influenced by the species seen as shown in 

Table 17. The highest compression, tensile, and modulus of rupture for monopodial 

bamboo grown in China are P. pubescens (86.5 MPa), P. decora (310.8 MPa) and P. glauca 

(213.4 MPa). 

  



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6524 

Sympodial Bamboo 
The highest compression, tensile, and modulus of rupture strenghts for sympodial 

bamboo species grown in China are B. rigida (79.8 MPa), D. latiflorus (199.1 MPa), and 

B. rigida (196.4 MPa). It is G. scortechinii (59.1 MPa), D. asper (222 MPa) and G. wrayi 

(201 MPa) for Malaysia bamboo respectively. The B. bambos has a highest tensile (260 

MPa) and modulus of rupture (225 MPa) for the Thailand bamboo (Table 17). 

In contrast, the lowest compression, tensile and modulus of rupture strengths for 

sympodial bamboo grown in China is recorded in D. beecheyanis and the values are 19.4 

MPa, 92.9 MPa and 40.5 MPa respectively. In Malaysia bamboo, the lowest is B. blumeana 

(27.1 MPa), D. asper (222 MPa) and B. heterostachya (120 MPa) respectively. The lowest 

tensile (110 MPa) and modulus of rupture (145 MPa) is recorded in B. longispiculata for 

Thailand bamboo. 

The shear, compression, and static bending strength of B. blumeana, B. vulgaris 

var. striata and G. scortechinii aged 1, 2 and 3 years are significantly increased with age 

(Mohmod et al. 1990). Nordahlia et al. (2019) also found that the MoR and MoE of 13 

Malaysian bamboos are increased with the culm height and the trend is the same for G. 

scortechinii (Shahril and Mansur 2009), D. latiflorus, D. meerrillanus, B. vulgaris 

(Leoncio 2017) and D. strictus (Bhonde et al. 2014). The increment of MoR and MoE 

along the culm height is accompanied by the higher number of vascular bundles (Nordahlia 

et al. 2019). 

Hamdan et al. (2009) reported that the MoR of bamboo without node is 

significantly higher at the top (258 MPa) than the basal (140 MPa) portions. The MoR at 

the top portion (147 MPa) is also significantly higher than at the basal portion (95 MPa) 

for the bamboo with node. A higher MoR at the internode portion is also contributed to its 

longitudinal straight cells as compared to the partially interrupt of radial cells at the node 

region. Bamboo with node fails quickly at the node region itself as compared to bamboo 

with internode. This trend is the same for sympodial bamboo, Guadua angustifolia (De 

Vos 2010) and Gigantochloa scortechinii. However, it is contradicted with monopodial 

bamboo, P. pubescens (Shoa et al. 2010; De Vos 2010), which the MoR is not significantly 

different in bamboo with and without the node. The MoR of bamboo with node is 32% 

(basal portion) and 43% (top portion) lower than bamboo without node in D. asper (Srivaro 

and Jakronod 2016), 20% lower than bamboo without node in G. angustifolia (De Vos 

2010) and 27% lower for bamboo without node in G. scortechinii (Hamdan et al. 2009).  

There is no significant difference of the MoE for D. asper with or without node 

(Srivaro and Jakronod 2016), as well as in P. pubescens and G. angustifolia (De Vos 2010); 

P. bambusoides (Lee et al. 1994; Tomak et al. 2012). The mechanical properties are higher 

at the top portion in the same or different species due to a higher volume fraction of 

vascular bundles and density (Amada et al. 1996; Malanit 2009; Dixon and Gibson 2014). 

The shear strength of D. asper aged 10 years without node are not significantly different 

for basal (12.3 MPa) and top (14.2 MPa) portions. The same trend occurs for D. asper with 

node for basal (11.9 MPa) and top (13.1 MPa) portions. The shear strength is higher at the 

top portion than the lower portion regardless of node or internode as the shear strength is 

mostly influenced by both density and culm height. This demonstrates that failure occurs 

by slipping of two shearing planes and a tearing failure of the soft parenchyma ground 

tissue as a result of the shear force (Srivaro and Jakronod 2016). 

In the same study, Srivaro and Jakronod (2016) reported that the tensile strength of 

D. asper without node is significantly higher at the top (299 MPa) than the basal (165 MPa) 

portions. The same trend for the D. asper with node for the top (145 MPa) and basal (73 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6525 

MPa) portions. A higher tensile strength at the internode than the node of the region for all 

culm heights is also obtained in P. bambusoides and P. pubescens (Lee et al. 1994; Shao 

et al. 2010). A higher tensile strength at the internode region is due to its longitudinal 

straight cell as compared to partly radial aligned cells at the node of the region, which is 

likely a loose compact structure (Wang et al. 2014). 

 
Overall 

The monopodial bamboo (207.18 MPa and 160.66 MPa) has a significantly higher 

tensile and modulus of rupture than the sympodial bamboo (122.27 and 72.63 MPa), 

respectively, as shown in the statistical reanalysis of the data. However, the compression 

strength is not significantly different from the rhizome as shown in Table 17. 

 

Table 17. The Compressive, Tensile and Modulus of Rupture for Monopodial and 
Sympodial Bamboos  

Species Origin Compressive 
(MPa) 

Tensile 
(MPa) 

MoR 
(MPa) 

References 

Monopodial 

Arundinaria 

A. amabilis China  82.5 - - Fangchun (2001a) 

A. amabalis China 80.8 280 - Fangchun (2001a) 

A. fergesia China  41.4 - - Fangchun (2001a) 

A. spongiosa China  64.5 169 - Fangchun (2001a) 

Indosasa 

I. crassiflora China  50.8 - - Fangchun (2001a) 

I. longispicata China  54.6 139.7 - Fangchun (2001a) 

Chimonobambusa 

C. quadragularis China 65.9 - - Fangchun (2001a) 

Phyllostachys 

P. angsiensis China  60.0 171.4 - Fangchun (2001a) 

P. angusta China  63.1 177.0 - Fangchun (2001a) 

P. aureosulcata China  72 252.4 - Fangchun (2001a) 

P. bambusoides China  64.3 239.8 - Fangchun (2001a) 

P. decora China  77.5 310.8 - Fangchun (2001a) 

P. frimbrigula China  52.4 189 - Fangchun (2001a) 

P. flexxuosa China  70.8 278.3 - Fangchun (2001a) 

P. glauca China  76.0 255.3 - Fangchun (2001a) 

P. glauca China  - 185.9 213.4 Fangchun (2001a) 

P. heteroclada China  65.4 250.2 - Fangchun (2001a) 

P. iridenscens China  57.6 185.6 - Fangchun (2001a) 

P. makinoi China  55.8 -  Fangchun (2001a) 

P. meyeri China  68.8 203.5 - Fangchun (2001a) 

P. nidularia China  57.9 176.2 - Fangchun (2001a) 

P. nigra China  40.7 - - Fangchun (2001a) 

P. nigra. var. 
henonis 

China  59.6 267.8 - Fangchun (2001a) 

P. nuda China  75.7 262.1 - Fangchun (2001a) 

P. platyglossa China  60.5 227.3 - Fangchun (2001a) 

P. praecox China  59.2 166.2 - Fangchun (2001a) 

P. pubescens China  71 198.4 - Fangchun (2001a) 

P. pubesces China  61.1 185.4 136.9 Fangchun (2001a) 

P. pubescens China  61.1 186 131.9 Fangchun (2001a) 

P. pubescens (1 China  49 135  Fangchun (2001b) 



 

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year) 

P. pubescens (2 
years) 

China  61 175  Fangchun (2001b) 

P. pubescens (3 
years) 

China  65 200  Fangchun (2001b) 

P. pubescens (4 
years) 

China  69 186  Fangchun (2001b) 

P. pubescens (5 
years) 

China  68 184  Fangchun (2001b) 

P. pubescens (6 
years) 

China  70 181  Fangchun (2001b) 

P. pubescens (7 
years) 

China  67 192  Fangchun (2001b) 

P. pubescens (8 
years) 

China  76 215  Fangchun (2001b) 

P. pubescens (9 
years) 

China  65 185  Fangchun (2001b) 

P. pubescens (10 
years) 

China  63 186  Fangchun (2001b) 

P. fimbriligula (1 
year) 

China  46 117  Fangchun (2001b) 

P. fimbriligula (2 
years) 

China  48 133  Fangchun (2001b) 

P. fimbriligula (3 
years) 

China  49 149  Fangchun (2001b) 

P. fimbriligula (4 
years) 

China  50 163  Fangchun (2001b) 

P. fimbriligula (5 
years) 

China  52 177  Fangchun (2001b) 

P. fimbriligula (6 
years) 

China  52 189  Fangchun (2001b) 

P. fimbriligula (7 
years) 

China  53 201  Fangchun (2001b) 

P. fimbriligula (8 
years) 

China  54 212  Fangchun (2001b) 

P. pubescens China 86.5   Li (2004) 

P. (Se.) 
pubescens 

China  65 189.5 127 Fangchun (2001a) 

P. (uns.) 
pubescens 

China  71.2 175.6 - Fangchun (2001a) 

P.viridi-
glaucescen 

China  67.2 230.7 - Fangchun (2001a) 

P. viridi China  48.1 158.2 - Fangchun (2001a) 

P. viridi China  - 289.1 194.1 Fangchun (2000b) 

P. vivax China  50.2 146.8 - Fangchun (2001a) 

Pleioblatus 

P. amara China 66 173.4 - Fangchun (2001a) 

Sympodial 

Bambusa 

B. breviflora China 58.0 - - Fangchun (2001a) 

B. corginera China 50.6 136 - Fangchun (2001a) 

B. lapidea China 44 - - Fangchun (2001a) 

B. flexuasa China 38.9 - - Fangchun (2001a) 

B. gibba China 31.3 - - Fangchun (2001a) 

B. pervariabilis China 34.8 - - Fangchun (2001a) 

B. pervariabilis China 48.5 191.2 - Fangchun (2001a) 



 

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B. rigida China 51 - - Fangchun (2001a) 

B. rigida China 79.8  196.4 Huang et al. (2014) 

B. rutila China 43.6 - - Fangchun (2001a) 

B. spp China 46 - - Fangchun (2001a) 

B. textilis China 58.7 - - Fangchun (2001a) 

B. textilis var. 
fusca 

China 36.7 123.4 63.3 Fangchun (2001a) 

B. textilis var. 
gracilis 

China 54.1 - - Fangchun (2001a) 

B. tulda China 43 - - Fangchun (2001a) 

B. tuldoides China 46.3 - - Fangchun (2001a) 

B. ventricosa China 33.7 - - Fangchun (2001a) 

B. ventricosa var. 
striata 

China 54.8 - - Fangchun (2001a) 

B. vulgaris Malaysia   172 Nordahlia et al. (2019) 

B. blumeana Malaysia   116 Nordahlia et al. (2019) 

B. blumeana Malaysia 27.1   Mohmod et al. (1990) 

B. blumena 
(without nodes) 

Thailand  170 128 Srivaro and Jakranod 
(2016) 

B. blumena (with 
nodes) 

Thailand  100 100 Srivaro and Jakranod 
(2016) 

B. heterostachya Malaysia   120 Nordahlia et al. (2019) 

B. vulgaris cv. 
vittata 

Malaysia   176 Nordahlia et al. (2019) 

B. vulgaris Malaysia  232.1  Awalludin et al. (2017) 

B. vulgaris Malaysia 28.2   Mohmod et al. (1990) 

B. longispiculata 
(without nodes) 

Thailand  180 175 Srivaro and Jakranod 
(2016) 

B. longispiculata 
(with nodes) 

Thailand  110 145 Srivaro and Jakranod 
(2016) 

B. bambos 
(without nodes) 

Thailand  260 225 Srivaro and Jakranod 
(2016) 

B. bambos (with 
nodes) 

Thailand  125 150 Srivaro and Jakranod 
(2016) 

Dendrocalamus 

D. beecheyanus China 19.4 92.2 40.5 Fangchun (2001a) 

D. gigantens China 41.6 161.7 - Fangchun (2001a) 

D. hamiltonii China 47.5 170.5 - Fangchun (2001a) 

D. latiflorus China 19.7 - - Fangchun (2001a) 

D. latiflorus China - 199.1 - Fangchun (2000b) 

D. asper Malaysia   150 Nordahlia et al. (2019) 

D. asper Malaysia  222  Awalluddin et al. 
(2017) 

Lingnania 

L. chungii China 27.3 - - Fangchun (2001a) 

L. chungii China 47.8 173.4 119.3 Fangchun (2001a) 

L. fimbrigulata China 55 - - Fangchun (2001a) 

L. spp. China 69.1 - - Fangchun (2001a) 

L. wenchouensis China 46.6 103 76.1 Fangchun (2001a) 

Schizostachyum 

S. funghoamii China 40.3 - - Fangchun (2001a) 

S. affinis China - 227.6 - Fangchun (2000b) 

S. brachycladum Malaysia   263 Nordahlia et al. (2019) 

S. grande Malaysia   184 Nordahlia et al. (2019) 

S. zollingeri Malaysia   143 Nordahlia et al. (2019) 



 

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Sinobambusa 

S. laeta China 40.9 - -  

Sinocalamus 

S. beecheyana China 38.9 - - Fangchun (2001a) 

S. beecheyana 
var. pubescens 

China 22.1 - - Fangchun (2001a) 

S. farinosus China 48.5 187.9 - Fangchun (2001a) 

S. latiflorus China 48.6 141.3 - Fangchun (2001a) 

S. latiflorus China 28.8 85.3 65.1 Fangchun (2001a) 

S. oldhamii China 39.3 156.3 71.5 Fangchun (2001a) 

S. oldhamii China 42 - - Fangchun (2001a) 

S. sffinis China 30.7 - - Fangchun (2001a) 

S. spp.  38 - - Fangchun (2001a) 

Gigantochloa 

G. thoii Malaysia   163 Nordahlia et al. (2019) 

G. scortechinii Malaysia   125 Nordahlia et al. (2019) 

G. scortechinii Malaysia 28.9   Mohmod et al. (1990) 

G. scortechinii 
(with node) 

Malaysia 39.6   Noor Zuraida et al. 
(2013) 

G. scortechinii 
(with internode) 

Malaysia 51.9   Noor Zuraida et al. 
(2013) 

G. ligulata Malaysia   180 Nordahlia et al. (2019) 

G. wrayi Malaysia   201 Nordahlia et al. (2019) 

G. brang Malaysia   159 Nordahlia et al. (2019) 

Guadua 

G. angustifolia Columbia 30.7   Camargo (2006) 

G. angustifolia (2 
years) 

Columbia 28.6  95.8 Correal and Camargo 
(2010) 

G. angustifolia (3 
years) 

Columbia 41.0  92.7 Correal and Camargo 
(2010) 

G. angustifolia (4 
years) 

Columbia 40.4  103.8 Correal and Camargo 
(2010) 

G. angustifolia (5 
years) 

Columbia 35.2  107 Correal and Camargo 
(2010) 

 

 

Table 18. Statistical Reanalysis of the Compressive, Tensile and Modulus of 
Rupture in Monopodial and Sympodial Bamboo 

Property 
Rhizome 

DF F Significance 
Monopodial Sympodial 

Compression 37.44 
(34.21) 

36.43 
(10.86) 

1 0.05 0.95NS 

Tensile 207.18 
(45.82) 

122.27 
(35.82) 

1 11.95 0.007*** 

MoR 160.66 
(40.08) 

72.63 
(25.95) 

1 19.42 0.002*** 

DF– is degree of freedom, F– is F ratio, NS is not significant at P > 0.1, *** is significant at P < 
0.0. The value in parentheses is the standard deviation. 

 

  



 

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CHEMICAL PROPERTIES 
 

Bamboo tissue consists of cellulose, hemicellulose, lignin, extractives, ash, silica 

(as a component of ash), starch, as well as various sugars akin to other monocotyledon and 

dicotyledon plants. The bamboo cellulose is composed of a longer chain of C12H10O15, and 

its molecular weight is approximately 1,500,000 g/mol. The cellulose in bamboo can be 

further separated into 70% to 80% alpha cellulose, 25% beta cellulose, and 1% to 5% 

gamma cellulose (Fangchun 2001a). The bamboo hemicellulose is mainly composed of 

pentosane with a small quantity of hexoan. About 90% of bamboo hemicellulose is made 

from xylan. The bamboo xylan is made up of D-glucuranate arabinoxylan, which 

comprises 4-oxygen-methyl-D-glucuranate, L-arabinose, and D-xylose. The composition 

of bamboo arabinoxylan is different from conifers and broad-leaved trees. The polymerized 

molecules of bamboo xylan are more than that of trees. The content of pentose in bamboo 

ranges from 19% to 23%, which is close to broad leaves and much higher than that of 

conifers, which is 10% to 15% (Maoyi et al. 2007). 

The structure of hemicellulose is determined by mainly arabinoxylans linked via 

(1-4)-ß-glycosidic bonds with branches of arabinose and 4-O-methyl-D-glucuronic acid 

(Lou et al. 2012). The bamboo lignin is a typical herbaceous lignin, consisting of three 

phenyl propane units, i.e., paradinum, guaiacyl, and mauve in the ratio of 5:34:11. The 

specific features of bamboo lignin lie in the presence of dehydrogenated polymerides and 

5% to 10% of acrylic ester. The lignin content of one-year-old bamboo ranges from 20% 

to 25%, approaching a broad-leaved wood and some grass (such as wheat straw 22%), and 

lower than that of conifers. The specific features of bamboo lignin lie in the existence of 

dehydrogenated polymerides and 5% to 10% of acrylic ester (Maoyi et al. 2007). 

Information on the content and distribution of lignin at each developmental stage is crucial 

for the exploitation of bamboo biomass (Chang and Holtzapple 2000; Shimokawa et al. 

2009). 

The lignin and the process of lignification in bamboo cells are used by researchers 

to study the earliest growth of bamboo cells toward its maturation. In bamboo, lignification 

in the culm proceeds acropetally, whereas lignification in each internode proceeds 

basipetally (Itoh 1990). Fibres and parenchyma cells of bamboo develop thick secondary 

walls that are composed of polylamellate structures containing broad and narrow lamellae 

(Parameswaran and Liese 1976; 1980). The deposition of lignin is much denser in the 

narrow lamella, and the distribution of the lignin-rich layers in bamboo fibres shows 

concentric rings in cross-sectioned walls (Parameswaran and Liese 1976). The pores of cell 

walls and cell corners are filled with lignin as seen in rapid-freezing and deep-etching 

(RFDE) electron microscopy (Nakashima et al. 1997; Fujino and Itoh 1998; Hafrén et al. 

1999). In contrast to the findings made by Itoh (1990), the lignification of bamboo cells in 

various age classes of bamboo P. pubescens still occurred after the first growing season.  

The protoxylem vessels are lignified in the early stage of vascular bundle 

differentiation. Upon completion, metaxylem vessel and fibre walls initiate lignification 

from the middle lamella and cell corners. Most of the parenchyma cell walls are lignified 

after the stem reaches its full height, while a few parenchyma cells remain non-lignified 

even in the mature culm. The cell walls of fibres and most parenchyma cells are further 

thickened during the stem growth to form polylamellate structure and the lignification 

process of these cells may last even up to 7 years. The fibre walls are rich in guaiacyl lignin 

in the early stage of lignification, and lignin rich in syringyl units are deposited in the later 

stage. Vessel walls mainly contained guaiacyl lignin, while both guaiacyl and syringyl 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6530 

lignin are present in the fibre and parenchyma cell walls (Lin et al. 2002). The unlignified 

primary wall (ULP) of P. pubescens is characterized by the narrow spacing between the 

cellulose microfibrils in fibres, but not in parenchyma cells. The unlignified secondary wall 

(ULS) largely consisted of dense cellulose microfibrils with narrow spacing or “slit-like” 

pores. 

The cell wall architecture of the delignified secondary wall (DLS) in fibres showed 

porosity similar to that of ULS. Pores in the middle lamella and secondary walls of ULS in 

fibres are reduced significantly or disappear immediately after lignification. However, the 

pores reappear following delignification. The deposition of lignin in ULS immediately 

proceeds in the pores during maturation to LS. The pore sizes of primary and secondary 

fibre walls are significantly smaller in bamboo than in either Eucalyptus or Pinus, 

suggesting a denser arrangement of cellulose microfibrils in bamboo fibre walls than in 

either tree species. The narrow spacing between cellulose microfibrils in bamboo fibres 

may be one of the reasons for the deposition of less lignin in bamboo than in tree species 

(Suzuki and Itoh 2001). 

In Sinobambusa tootsik (Tsuyama et al. 2017), the content of monolignol 

glucosides is maximum during the early stages of lignification, whereas the contents of 

monolignols peak at later stages of lignification. Elongation growth is ended by the culm 

lignin content, which is approximately half that of mature culms.  

 
Monopodial Bamboo 

In relation to species (Table 19), the -cellulose, hemicellulose, and lignin contents 

in monopodial bamboo P. heterocycla (Carr.) Mitford cv. Pubescens aged 4 years are 

42.7%, 25.52%, and 21.15%, respectively (Wang et al. 2021). The Melocacna baccifera 

contains 52.78% -cellulose, 21.1% hemicellulose, 25.2% lignin, 4.13% hot water soluble, 

3.24% hot water soluble, 2.45% ash, 19.5% NaOH, and 3.48% ethanol/toluene extractives 

(Tripathi et al. 2018). In relation to the culm age (Table 20), the holocellulose, lignin, 

ethanol/toluene extractive of monopodial bamboo P. edulis are not significantly different 

with age (1, 2, and 3 years) ranging from 65.97% to 67.24%, 30.48% to 32.09%, and 4.59% 

to 5.11%, respectively. The ash content is significantly lower in the oldest culm (0.89%), 

and it is not significantly different for culm aged 1 (1.83%) and 2 years (1.68%). In contrast, 

the silica content is significantly high for the oldest culm (0.30%), while it is not 

significantly different for culm aged 2 (0.28%) and 3 years (0.21%) (Ju et al. 2021). 

In relation to the culm portion (Table 20), the holocellulose, ethanol/toluene, ash, 

and silica contents in monopodial bamboo P. edulis aged 1, 2, and 3 years are not 

significantly different with the culm height ranging from 65.77% to 67.69%, 4.58% to 

4.85%, 1.36% to 1.72%, and 0.23 to 0.33%, respectively, with the exception of the lignin 

content. The lignin content is significantly highest at the middle portion of the culm 

(32.92%), while it is not significantly different at the basal (31.01%) and top portions 

(30.67%), regardless of age (Zhan et al. 2021).  

 

Sympodial Bamboo 
The chemical composition of cultivated sympodial bamboos G. brang, G. levis, G. 

scortechinii, and G. wrayi varies by species, portion (node or internode), and section (outer, 

middle, or inner) of the culm (Table 20). The -cellulose is highest in descending order of 

G. brang (51.58%), G. scortechinii (46.87%), G. wrayi (37.66%), and G. leavis (33.81%). 

Regardless of species, the -cellulose is not significantly different with node (42.74) and 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6531 

internode (42.22), but the outer culm layer has a significantly highest -cellulose (49.07%), 

followed by middle (41.28%), and inner (37.09%) layers. The holocellulose is highest in 

G. wrayi (84.53%) but it is not significantly different from G. levis (84.52%). The G. brang 

(79.70%) has a significantly higher holocellulose content than the G. scortechinii 

(74.63%). Overall, the node (81.66%) has a significantly higher holocellulose than the 

internode (80.03%) and it is significantly in descending order of outer (82.99%), middle 

(80.89%), and inner (78.65%) layers of the culm wall. 

The lignin is significantly highest in G. scortechinii followed by G. wrayi (37.66%), 

but it is not significantly different for G. brang (24.83%) and G. levis (26.50%). Overall, 

the lignin content is significantly higher in internode (32.295) than the node (24.76%) 

portions, and it is significantly highest at the outer (33.43%), followed by inner (30.03%) 

and middle (21.98%) culm wall layers. The ash content is significantly highest in G. 

scortechinii (2.84%) followed by G. levis (1.30%), G. brang (1.26%), and G. wrayi 

(0.88%). Overall, the ash is significantly higher at the node (1.6%) than the internode 

(1.54%) portions. Opposite of --cellulose and lignin, the ash is significantly highest at the 

inner (1.89%) culm wall, followed by the outer (1.52%) and lastly middle (1.28%) culm 

wall layers. The ethanol/toluene extractive is significantly highest in G. levis (9.23%), 

followed by G. wrayi (8.62%). The extractive content is not significantly different for G. 

brang (8.30%) and G. scortechinii (8.00%). The node (8.63%) has a significantly higher 

extraction content than the internode (8.46%), while it is significantly highest in the inner 

(13.42%) culm wall followed by the middle (7.21%) and lastly the outer (4.99%) layers of 

the culm wall (Razak et al. 2013). 

In seven sympodial bamboo species grown in India, B. nutan Dehradun has 51.6% 

cellulose, 26% lignin, 8.1% hot water soluble, 7.1% cold water soluble, 28.1% NaOH 

soluble, and 4.2% ethanol/toluene extractive contents. The B. tulda has 56.2% cellulose, 

24% lignin, 7.8% hot water soluble, 5.5% cold water soluble, 26.1% NaOH soluble, and 

3.2% ethanol/toluene extractive. The B. arundinacea Allahabad has 47.7% cellulose, 

26.5% lignin, 11.4% hot water soluble, 9.8% cold water soluble, 27.0% NaOH soluble, and 

3.5% ethanol/toluene extractives. The B. pallida IWST, Bangalore has 46.5% cellulose, 

20% lignin, 12.6% hot water soluble, 11.1% cold water soluble, 27% NaOH, and 5.2% 

ethanol/extractive contents. The B. bambos IWST, Bangalore has 50.5% cellulose, 21.5% 

lignin, 11.2% hot water soluble, 10% cold water soluble, 27.4% NaOH soluble, and 4.2% 

ethanol/toluene extractive.  

The D. strictus Alalhabad has 53.6% cellulose, 25% lignin, 8.4% hot water soluble, 

6.7% cold water soluble, 27.9% NaOH soluble, and 4.2% ethanol/toluene extractives. The 

D. strictus Teri has 53.4% cellulose, 27% lignin, 8.3% hot water soluble, 6.2% cold water 

soluble, 28.0% NaOH soluble, and 4.9% ethanol/toluene extractive contents (Kaur et al. 

2016ab). The chemical composition of sympodial bamboo B. garuchokua, B. pallida, and 

B. assamica aged 3 years grown in India behave differently (Brahma and Brahma 2018). 

The -cellulose is highest in B. pallida (38.29%), followed by B. garachokua (36.75%) 

and B. assamica (31.37%). Regardless of species, the -cellulose content is highest at the 

outer culm all section followed by the middle (37.31%) and inner (35.90%) sections. In 

contrast, the holocellulose is highest in ascending order of B. assamica (60.18%), B. pallida 

(65.92%), and B. garuchokua (68.86%).  

Overall, the holocellulose content is highest at the outer section (69.63%), followed 

by the middle (62.81%, and inner (62.51%) sections. The lignin content is highest in 

ascending order of B. assamica (18.29%), B. pallida (22.42%), and B. garuchokua 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6532 

(23.03%). Overall, the outer section has the highest lignin content (23.98%) and it is not 

much different with the inner (22.23%) and lastly the middle (17.78%) sections. The ash 

content is increased in ascending order of B. garachukua (0.99%), B. pallida (1.08%), and 

B. assamica (1.12%), while it is increased from the outer wall section (0.95%) toward the 

middle (1.04%) and inner (1.20%) sections. The hot water soluble is highest in B. 

garuchokua (5.90%) followed by B. pallida (5.70%) and lastly B. assamica (4.24%), while 

it gradually increases from outer section (4.43%) toward the middle (5.32%) and inner 

(6.09%) sections. The alcohol/toluene content is almost identical for B. garuchokua 

(4.12%) and B. pallida (4.37%), but it is lowest in B. assamica (3.44%). Across the culm 

wall, the extractive content is gradually decreased from the inner (4.55%) section toward 

the middle (4.02%) and outer sections (3.36%). 

In relation to the culm section (Table 19), the contents of holocellulose, -cellulose, 

lignin, ash, hot water soluble, cold water soluble, and 1% NaoH soluble are not 

significantly different with node or internode regions at the basal, middle, and top portions 

of sympodial bamboo D. asper aged 3 years ranging from 75.65% to 77.36%, from 67.07% 

to 69.64%, from 26.47% to 30.86%, from 0.92 to 2.29%, from 6.47 to 9.63%, from 4.23 to 

14.05%, and from 23.40 to 26.78% (Kamthai and Puthson 2005). Almost all major 

chemical constituents in sympodial bamboo G. scortechinii aged 0.5, 1.5, 3.5, 5.5, and 6.5 

years are relatively low at the youngest age of 0.5 years (Table 18). The starch content is 

low at the age of 1.5 years (0.6%) but increases to 3.5% at the age of 3.5 years and remain 

unchanged toward the later years. This is probably because no further increase of 

parenchyma length and lumen diameter occurred beyond the age of 3.5 years. The starch 

granule is situated or stored in vertically elongated cells of ground parenchyma (Liese and 

Weiner 1996).  

The authors reported that younger 1-year-old bamboo culms do not contain any 

starch during the growing phase, because all the nutrients must be utilized immediately for 

metabolic processes. However, Mohmod et al. (1992) reported minor trace of starch 

content (0.8%) at the basal portion of 1-year-old G. scortechnii. Lignin content drastically 

increased at the age of 1.5 years (14.5%) and then gradually increased thereafter. No 

specific trend for alcohol/toluene extractive was observed, but it was high at age of 0.5, 

3.5, and 6.5 years. The -cellulose content remained unchanged, but the holocellulose 

content slightly increased beyond 3.5 years. The -cellulose, holocellulose, hot water 

soluble, NaOH soluble, and ash contents in G. scortechinii are not significantly different 

with bamboo aged 1, 2, and 3 years (Table 20). The averages are 40.7%, 41.41%, and 

40.49% for -cellulose; 66.7%, 67.8%, and 67.9% for holocellulose; 6.3%, 5.9%, and 5.4% 

for hot water soluble; 19.6%, 19.2%, and 19.6% for NaOH soluble; and 11.0%, 11.1% and 

11.4% for ash contents, respectively (Mohmod et al. 1994).  

The lignin content is not significantly different in culm aged 1 (25.7%) and 2 years 

(24.9%), but significantly highest in the oldest culm (28.0%). The same trend occurs for 

cold water soluble and ethanol/toluene extractive. The average cold water soluble is 4.3%, 

4.4%, and 5.5% in culm aged 1, 2, and 3 years, while it is 3.2%, 3.2%, and 3.5% for the 

ethanol/toluene extractives. Zhang et al. (2015) examined the chemical composition of F. 

fungosa aged 1, 2, and 3 years. The holocellulose content is not significantly different, with 

age ranging from 69.86% to 70.77%. Regardless of age, the holocellulose content is also 

not significantly different at the basal (70.11%), middle (69.99%), and top (70.56%) 

portions of the culm. Same as holocellulose, the lignin is not significantly different with 

age ranging from 22.66% to 24.21%. The alcohol/toluene extractive is significantly highest 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6533 

for the oldest culm (4.14%) and it is not significantly different for culm aged of 1 year 

(3.01%) and 2 years (3.22%). The ash content is significantly highest at a youngest age of 

1 year (2.88%) but declines toward 2 (1.51%) and 3 years (1.40%). The silica content is 

significantly decreased in descending order of 1 year (0.48%), 2 years, (0.38%) and 3 years 

(1.40%). 

In sympodial bamboo of D. hamiltonii (Zhan et al. 2016), the holocellulose content 

is significantly increased at the youngest age of 1 year toward the older age of 2 (68.7%) 

and 3 (76.7%) years. In contrast, the lignin content is not significantly different with ages 

ranging from 21.4% to 23.6%, but it is significantly different with the culm portion. The 

ethanol/toluene extractive content is significantly increased at the youngest age of 1 year 

(0.9%) toward the older aged of 2 years (1.2%) and 3 years (1.9%). The ash content is not 

significantly different with age ranging from 2.0% to 2.9%. In contrast to F. fungosa (Zhan 

et al. 2015), the silica content is significantly increased with age from 1 year (0.3%), 2 

years (0.7%), and 3 years (2.1%) (see Fig. 2).  

 

  

  

Fig. 2. The effect of ageing on the chemical properties in monopodial and sympodial bamboos 

 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6534 

In relation to the culm portion (Table 19), the -cellulose, lignin, holocellulose, 

cold water soluble, and ash contents in G. scortechinii aged 1, 2, and 3 years are not 

significantly different with the culm height (basal, middle, and top portions); ranging from 

40.3% to 41.1%, 25.5% to 26.6%, 67.2% to 68.90%, 4.4% to 4.9%, and 1.10% to 1.13%, 

respectively. The hot water soluble is significantly increased at the basal (5.3%) toward the 

middle (6.0%) and top (6.5%) portions. A same trend occurs for NaoH soluble and 

ethanol/toluene extractives. The average NaoH soluble is 18.49% (basal), 19.8% (middle), 

and 20.1% (top), while it is 3.0% (basal), 3.3% (middle), and 3.6% (top) for the ethanol 

extractive contents (Mohmod et al. 1994). The holocellulose content in D. hamiltonii (Zhan 

et al. 2016) is not significantly different with portions ranging from 69.2% to 73.4%. The 

lignin content is significantly decreased in descending order of top (24.8%), middle 

(22.4%), and basal (21.60%) portions. Within the culm height, the extractive content is not 

significantly different with the culm portion ranging from 1.1% to 1.5%. The ash and silica 

contents are not significantly different with the culm portion ranging from 1.8% to 3.1% 

and 0.4% to 1.6% respectively. 

The lignin content of F. fungosa aged 1, 2, and 3 years is significantly highest at 

the top (24.66%) but not significantly different from the basal (23.76%) portions. The 

middle portion has the lowest lignin content (21.86%). The ash content is not significantly 

different from the culm portion, ranging from 1.90% to 1.96%. Within the culm wall 

section, the silica content is significantly decreased in descending order of top (0.43%), 

middle (0.39%), and basal (0.33%) portions (Zhan et al. 2015). The maximum cellulose, 

hemicellulose, lignin, ash, silica, hot water soluble, cold water soluble, NaOH soluble, and 

ethanol/toluene extractive contents in monopodial bamboo are recorded for P. heterocycle 

(75.30%), P. edulis (29.66%), Sasa albomarginate Makin (39.76%), S. albomarginate 

Makin (3.38%), P. sp (0.13%), P. heterocycla (15.94%), P. heteroclada (13.57%), and P. 

mayeri (35.31%). While the minimum contents in monopodial bamboo are obtained in P. 

bambusoides (12.02%), M. baccifera (15.03%), P. bissetii McClure (14.69%), C. utilis 

(0.74%), B. garuechokua (0.99%), S. affinis (0.17%), P. bisseti McClure (1.68%), P. 

pubescens Mazel (2.38%), P. bambusoides (14.60%), and P. pubescens (1.60%) (Table 

20). 

In contrast, the maximum cellulose, hemicellulose, lignin, ash, silica, hot water 

soluble, cold water soluble, NaOH soluble, and ethanol/toluene extractive contents in 

sympodial bamboo are recorded by D. striticus (68.00%), S. affinis (25.41%), D. striticus 

(32.20%), D. sp (4.39%), S. funghpmii McClure (3.76%), B. sinaspinosa (9.91%), B. 

sinaspinsa McClure (10.53), B. textilis (32.27), and B. sinaspinosa (28.02%); while, the 

minimum contents for sympodial bamboo are recorded in B. sinanpinosa (17.10%), D. 

striticus (15.05%), D. sp (17.62%), B. garuechokua (0.99%), S. affinis (0.17%), O. 

travancorica (3.13%), T. sp (1.61%), D. hamiltonii (13.81%), and D. hamiltonii (0.9%). 

 

Overall 
Regardless of the rhizome, the overall monopodial type of bamboo has a 

significantly higher cellulose, hemicellulose, lignin, hot water soluble, cold water soluble, 

and 1% NaOH soluble (47.63%, 20.81%, 25.57%, 6.97%, 6.89%, and 28.09%) contents 

compared to the sympodial type of bamboo (40.42%, 14.24%, 24.33%, 5.75%, 5.61%, and 

24.69%), respectively. In contrast, the sympodial type of bamboo has a significantly higher 

ash content (2.06%) compared to the monopodial type of bamboo (1.54%). Both the silica 

(0.27% and 0.92%) and ethanol/toluene extractive (4.72% and 7.20%) are not significantly 

different in monopodial and sympodial types of bamboo (Table 21).



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6535 

Table 19. Chemical Properties in Monopodial and Sympodial Bamboos 

Species 

Percentage 
References 

Origin Cel Hemi Lig Ash Sil Hw Cw NaOH Eth/Tl  

Monopodial 

Arundinaria 

A. fargesii China 45.10 21.8 25.26 1.54 - - - 24.35 - Fangchun (2001a) 

A. murielea Gamble China 61.06 - 21.18 2.92 - - - -  Fangchun (2001a) 

Chimonobambusa 

C. quadragularis China 50.53 18.47 20.78 2.97 0.59 - 4.18 - - Fangchun (2001a) 

C. utilis Keng F. China 55.07 22.25 26.08 0.74 - - - 29.14 - Fangchun (2001a) 

C. delicatus Hsu et China 48.34 17.58 20.01 2.68 0.51 - 6.61 - - Fangchun (2001a) 

Indocalamus 

I. farinosus Keng et K China 57.02 18.67 24.69 2.03 - - - 27.06 - Fangchun (2001a) 

Melocanna 

M. baccifera China 62.25 15.13 24.13 1.87 - 6.48 3.26 18.97 - Fangchun (2001a) 

M. baecifera China 75.30 15.03 23.20 1.74 - - - 17.13 - Fangchun (2001a) 

M. baccifera India 52.78 21.1 25.2 2.45  4.13 3.24 19.5 3.48 Tripathi et al. (2018) 

Phyllostachys 

P. bambusoides Seib China          Fangchun (2001a) 

P. bambusoides  China 55.7 - 15.84 - - - - 17.02 - Fangchun (2001a) 

P. bambusoides China 37.51 - 39.51 - - - - 14.60 - Fangchun (2001a) 

P. bambusoides (0.5 
year) 

China 48.92  24.51 2.22 - 5.93 4.62 27.60 1.81 Jiang (2002) 

P. bambusoides (1 year) China 56.74  29.93 1.25 - 8.97 10.49 29.93 7.34 Jiang (2002) 

P. bambusoides (3 
years) 

China 12.02 - 25.15 0.98 - 7.32 6.11 31.33 5.86 Jiang (2002) 

P. meyeri (0.5 year) China 49.97 - 23.58 1.68 - 5.15 3.60 27.27 1.81 Jiang (2002) 

P. meyeri (1 year) China 57.88 - 23.62 1.29 - 8.91 10.70 34.28 7.04 Jiang (2002) 

P. meyeri (3 years) China 39.95 - 23.35 1.85 - 12.7
1 

8.81 35.31 7.52 Jiang (2002) 

P. bissetii McClure China 52.14 19.26 24.02 0.89 - 1.68 - 31.05 - Fangchun (2001a) 

P. bissetii McClure China 55.53 20.11 25.36 1.07 - 8.50 - 27.35 - Fangchun (2001a) 

P. bissetii McClure China 55.38 19.88 25.60 1.41 - 8.03 - 26.91 - Fangchun (2001a) 

P. bissetii McClure China 69.25 25.31 14.69 2.78 - 8.58 - 30.21 - Fangchun (2001a) 



 

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P. glauca China - 22.64 33.46 1.43 - 5.24 - 28.96 - Fangchun (2001a) 

P. makinoi Hayata China 50.55 24.79 30.88 - - - - - - Fangchun (2001a) 

P. nidularia Munro China 54.83 21.39 25.58 0.87 - - - 26.69 - Fangchun (2001a) 

P. nigra var. Menoni China 51.34 19.76 33.45 1.38 - 5.84 - 33.07 - Fangchun (2001a) 

P. nigra (0.5 year) China 45.38 - 28.94 1.98 - 8.30 6.72 31.83 4.12 Jiang (2002) 

P. nigra (1 year) China 58.85 - 23.90 1.81 - 8.53 10.69 32.24 5.29 Jiang (2002) 

P. nigra (3 year) China 13.79 - 25 1.71 - 8.36 6.50 33.65 5.58 Jiang (2002) 

P. nigra Japan 42.3 - 23.80 2.0  - - - 3.4 Higuchi (1958) 

P. pubescens China - 21.12 30.67 1.10 - 5.96 - 30.98 - Fangchun (2001a) 

P. pubescens Mazel China 45.94 18.81 27.83 0.81 0.17 - 4.17 - - Fangchun (2001a) 

P. pubescens Mazel China 45.50 21.12 30.67 1.10 - 5.96 2.38 30.98 - Fangchun (2001a) 

P. sp. China 45.80 18.39 23.06 1.73 0.13 - 3.91 - - Fangchun (2001a) 

P. pubescens (0.5 year) China 61.97 - 26.36 1.77 - 3.26 5.41 27.34 1.60 Jiang (2002) 

P. pubescens (1 year) China 59.82 - 34.77 1.13 - 6.31 8.13 29.34 3.67 Jiang (2002) 

P. pubescens (3 years) China 60.55 - 26.20 0.69 - 5.11 7.10 26.91 3.88 Jiang (2002) 

P. pubescens 7 years) China 59.09 - 26.75 0.52 - 5.17 7.14 26.83 4.78 Jiang (2002) 

P. pubescens (1 year) U.S.A 47.11 - 21.78 1.90 - 5.56 - - 3.22 Li (2004) 

P. pubescens (3 years) U.S.A 46.63 - 23.62 1.36 - 6.89 - - 4.73 Li (2004) 

P. pubescens (5 years) U.S.A 47.21 - 22.97 1.30 - 5.31 - - 6.92 Li (2004) 

P. heteroclada (1 year) China 58.15 - 22.42 1.24 - 9.60 13.57 30.89 5.38 Jiang (2002) 

P. heteroclada (3 years) China 38.96 - 22.75 1.27  15.9
4 

9.68 34.84 9.11 Jiang (2002) 

P. heterocycla Japan 49.1 - 26.1 1.3 - - - - 4.6 Higuchi (1958) 

P. heterocycla China 42.7 25.52 21.15 - - - - - - Wang et al. (2020) 

P. praccox (0.5 year) China 42.23 - 26.74 3.24 - 8.57 6.72 33.36 2.25 Jiang (2002) 

P. praccox (1 year) China 56.03 - 21.68 1.96 - 7.68 11.21 32.84 3.80 Jiang (2002) 

P. praccox (3 years) China 40.81 - 25.65 2.26 - 9.09 7.18 23.26 5.64 Jiang (2002) 

P. reticulata Japan 25.30 - 25.3 1.9 - - - - 3.4 Higuchi (1955) 

P. edulis Taiwan 45.39 29.66 21.51 1.48 - - - - 6.39 Li et al. (2018) 

Sasa 

S. albomarginate Makin China - - 39.76 3.38 - - - - - Fangchun (2001a) 

Sympodial 

Bambusa 

B. arundinacea China 57.56 19.62 30.90 3.26 - 5.25 4.59 19.35 - Fangchun (2001a) 

B. breviflora Munro China - 20.80 24.20 - - - - - - Fangchun (2001a) 

B. nutans China - - 21.70 - - - - - - Fangchun (2001a) 



 

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B. pervariabilis McCl China 55.77 16.19 23.28 3.00 - 5.30 4.29 29.12 - Fangchun (2001a) 

B. pervariabilis McCl China 48.51 15.81 23.98 1.56 0.54 - 8.03 - - Fangchun (2001a) 

B. polymor pha China - 18.50 24.70 - - - - - - Fangchun (2001a) 

B. polymor pha Munro China 46.90 17.05 23.86 2.10 0.21 - 5.00 - - Fangchun (2001a) 

B. rigida Keng et Keng China 46.71 18.78 22.16 1.20 0.54 - 6.98 - - Fangchun (2001a) 

B. rigida Keng et Keng China 56.98 19.19 23.51 1.49 - - - - 25.44 Fangchun (2001a) 

B. sinaspinosa McClure China 47.50 16.14 24.35 3.72 2.26 - 10.53 - - Fangchun (2001a) 

B.sinaspinosa McClure China 55.46 20.30 23.17 2.86 - - - - 28.02 Fangchun (2001a) 

B. sinaspinosa (0.5 year) China 52.58  19.90 2.69  8.23 7.29 29.98 4.23 Jiang (2000) 

B. sinaspinosa (1 year) China 49.45  20.51 1.92  9.91 8.08 30.25 5.49 Jiang (2000) 

B. sinaspinosa (3 years) China 17.10  24.17 1.84  9.27 9.07 26.92 5.88 Jiang (2000) 

B. sterstachya Hacel China 52.20 19.61 30.88 2.45 - - - - - Fangchun (2001a) 

B. textilis McClure China 58.48 18.87 20.19 2.24 - 7.60 4.88 25.11 - Fangchun (2001a) 

B. textilis (0.5 year) China 51.96  18.67 2.39 2.39 8.03 6.64 32.27 - Jiang (2002) 

B. textilis (1 year) China 50.40  19.39 2.08 2.08 7.55 6.30 30.57  Jiang (2002) 

B. textilis (3 years) China 45.50  23.81 1.58 1.58 8.75 6.84 28.01  Jiang (2002) 

B. tulda China 64.36 18.42 24.16 2.02 - 4.97 2.64 21.89 - Fangchun (2001a) 

B. tulda China - 18.10 23.10 - - - - - - Fangchun (2001a) 

B. vulgaris China 41.00 21.00 28.10 1.70 0.71 - 4.60 21.50 - Fangchun (2001a) 

B. vulgaris China 43.00 22.50 25.80 - - - 3.70 20.20 - Fangchun (2001a) 

B. vulgaris China - 21.00 22.90 - - - - -  Fangchun (2001a) 

B. vulgaris China 43.80 20.60 27.90 1.88 0.57 - 3.00 -  Fangchun (2001a) 

B. sp China - 19.19 23.51 1.49 - 8.22 - 25.44  Fangchun (2001a) 

B. sp China - 18.17 22.71 1.14 - 8.45 - 25.27  Fangchun (2001a) 

B. sp China 61.79 21.48 21.99 1.67 - 6.88 2.93 18.39  Fangchun (2001a) 

B. sp China 45.03 17.64 23.04 2.42 1.19 - 14.34 -  Fangchun (2001a) 

B. sp China 47.62 15.68 26.41 1.30 0.25 - 6.53 - - Fangchun (2001a) 

B. garuchokua India 36.75  23.03 0.99  5.90   4.12 Brahma and Brahma 
(2018) 

B. pallida India 38.29  22.42 1.08  5.70   4.37 Brahma and Brahma 
(2018) 

B. assamica India 31.37  18.29 1.12  4.24   3.44 Brahma and Brahma 
(2018) 

Cepholostachyum 

C. fuchsiamum Gamb China 56.12 17.45 22.15 - - - 4.84 - - Fangchun (2001a) 



 

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C. pergracila Munro China 49.05 17.55 22.44 2.67 1.31 - 7.05 - - Fangchun (2001a) 

C. pergracile China - 18.40 24.90 - - - - - - Fangchun (2001a) 

Dendrocalamus 

D. giganteus China 51.49 17.83 24.44 2.16 0.96 - 5.65 - - Fangchun (2001a) 

D. giganteus China 39.40 18.40 25.30 2.87 0.37 - 5.10 24.40 - Fangchun (2001a) 

D. hamiltonii China 63.26 21.49 26.21 1.80 - 4.42 2.47 20.81 - Fangchun (2001a) 

D. hamiltonii China - 16.90 22.40 - - - - 13.81 - Fangchun (2001a) 

D. hamiltonii (1 year) China - - 23.5 2.3 0.3 - - - 0.9 Zhan et al. (2016) 

D. hamiltonii (2 years) China - - 23.6 2.0 0.7 - - - 1.2 Zhan et al. (2016) 

D. hamiltonii (3 years) China - - 22.9 2.9 2.1 - - - 1.9 Zhan et al. (2016) 

D. latiflorus Munro China 52.84 19.78 26.25 3.03 - - - 21.81 - Fangchun (2001a) 

D. longispathus China - 18.60 25.00 - - - - - - Fangchun (2001a) 

D. membranaceus 
Munro 

China 47.61 16.60 26.59 1.83 0.87 - 6.54 - - Fangchun (2001a) 

D. sericeus Munro China 50.34 16.27 23.50 1.94 0.77 - 5.63 - - Fangchun (2001a) 

D. sinicus Chia China 47.53 15.97 26.59 3.44 2.07 - 6.40 -  Fangchun (2001a) 

D. strictus China 68.00 19.56 32.20 2.10 - 5.93 4.20 15.00 - Fangchun (2001a) 

D. strictus China 66.40 15.06 27.87 2.32 - - - 30.61  Fangchun (2001a) 

D. strictus China - 23.20 26.00 - - - - - - Fangchun (2001a) 

D. sp China 53.46 16.13 19.61 4.39 1.11 - 6.48 -  Fangchun (2001a) 

D. sp China 44.01 19.79 23.54 3.63 0.55 - 8.74 -  Fangchun (2001a) 

D. sp China 50.07 13.57 24.80 2.14 0.61 - 5.79 - - Fangchun (2001a) 

D. sp China 50.25 16.56 18.67 2.55 0.17 - 5.58 -  Fangchun (2001a) 

Dinochloa 

D. sp China 54.04 17.99 17.62 2.89 1.13 - 5.52 - - Fangchun (2001a) 

Ochlandra 

O. travancorica China 61.76 17.84 26.91 2.60 - 3.13 3.59 19.98  Fangchun (2001a) 

Schizostachyum 

S. fumghpmii McClure China 52.49 16.76 23.90 5.73 3.76 - 3.44 -  Fangchun (2001a) 

S. affinis McClure China 63.98 - 22.08 1.85 0.17 - - 24.93 - Fangchun (2001a) 

S. affinis China - 25.41 31.28 1.20 - 9.78 - 31.24  Fangchun (2001a) 

S. affinis Keng China 59.06 18.88 28.96 1.62 - 7.10 - 26.20  Fangchun (2001a) 

S. affinis Keng China 60.68 18.15 24.27 2.47 - 7.75 - 25.31 - Fangchun (2001a) 

S. affinis Keng China 57.07 18.03 23.11 2.51 - 8.68 - 26.89  Fangchun (2001a) 

S. affinis Keng China 62.57 19.17 21.35 1.69 - 7.52 - 27.82  Fangchun (2001a) 

S. distegius Keng et K China 50.84 17.74 22.65 1.79 0.51 - 5.56 - - Fangchun (2001a) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6539 

S. distegius Keng et K China 59.07 19.00 23.31 1.13 - - - 27.71 - Fangchun (2001a) 

S. farinosus Keng China 51.47 16.11 24.22 2.78 1.84 - 3.39 - - Fangchun (2001a) 

S. farinosus Keng et K China 58.11 19.20 25.25 2.63 - - - 27.37  Fangchun (2001a) 

Thyrsostachys 

T. oliveri China - 18.50 20.90 - - - - - - Fangchun (2001a) 

T. oliveri Gamble China 51.16 16.31 23.92 2.49 0.63 - 4.44 - - Fangchun (2001a) 

T. siamensis Gamble China 48.83 15.72 21.19 3.80 0.98 - 9.33 - - Fangchun (2001a) 

T. sp. China 51.37 16.25 24.55 3.64 2.08 - 3.73 - - Fangchun (2001a) 

T. sp. China 66.72 17.41 27.09 1.25 - 3.39 1.61 17.11 - Fangchun (2001a) 

Fargesia 

F. fungosa (1 year) China - - 22.66 2.88 0.48 - - - 3.01 Zhan et al. (2015) 

F. fungosa (2 years) China - - 24.21 1.16 0.38 - - - 3.22 Zhan et al. (2015 

F. fungosa (3 years) China - - 24.04 1.40 0.30 - - - 4.14 Zhan et al. (2015 

Gigantochloa 

G. brang Malaysia 51.58 - 24.83 1.26 - - - - 8.30 Razak et al. (2013) 

G. levis Malaysia 33.81 - 26.50 1.30 - - - - 9.23 Razak et al. (2013) 

G. scortechinii Malaysia 46.87 - 32.55 2.84 - - - - 8.00 Razak et al. (2013) 

G. wrayi Malaysia 37.66 - 30.04 0.88 - - - - 8.62 Razak et al. (2013) 

G. scortechinii (0.5 year) Malaysia 64.6 - 23.4 1.9 0.6 5.8 - - - Norul Hisham et al. 
(2006) 

G. scortechinii (1.5 y) Malaysia 64.1 - 26.8 2.5 1.1 3.4 - - - Norul Hisham et al. 
(2006) 

G. scortechinii (3.5 y) Malaysia 64.6 - 27.8 2.8 1.7 5.3 - - - Norul Hisham et al. 
(2006) 

G. scortechinii (5.5 y) Malaysia 63.3 - 28.7 3.0 2.2 3.5 - - - Norul Hisham et al. 
(2006) 

G. scortechinii (6.5 y) Malaysia 64.4 - 29.0 3.5 2.0 5.6 - - - Norul Hisham et al. 
(2006) 

 

  



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6540 

Table 20. Statistical Reanalysis of the Chemical Properties in Monopodial and Sympodial Bamboos 

Chemical Rhizome DF F Significance 

Monopodial Sympodial 

Cellulose 47.63 
(15.20) 

40.42 
(20.38) 

1 3.73 0.06* 

Hemicellulose 20.81 
(3.41) 

14.24 
(7.92) 

1 14.29 0.00*** 

Lignin 25.57 
(4.83) 

24.33 
(3.10) 

1 3.29 0.07* 

Ash 1.54 
(0.80) 

2.06 
(1.06) 

1 8.89 0.00*** 

Silica 0.27 
(0.21) 

0.92 
(0.55) 

1 1.71 0.20NS 

Hot water 6.97 
(2.95) 

5.75 
(2.84) 

1 2.93 0.09* 

Cold water 6.89 
(2.94) 

5.61 
(2.52) 

1 3.52 0.07* 

NaoH soluble 28.09 
(5.30) 

24.69 
(4.91) 

1 7.25 0.01** 

Ethanol/toluene extractive 4.72 
(1.94) 

7.20 
(7.54) 

1 2.59 0.12NS 

NS –Not significant at P > 0.1, * - significant at P < 0.1, **- significant at P < 0.05, and ***- significant at P < 0.01



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6541 

SUSCEPTIBILITY AND RESISTANCE TO FUNGI 
 

Akin to other monocotyledon and dicotyledon plants, bamboo contains structural 

organic chemicals, inorganic chemicals, and extractives. It is easily decayed by micro-

organisms, such as mold, fungi, and insects, under appropriate conditions. Although it is 

well known that the bamboo has been used by human and animals, a few hundred years 

ago both in tropic and temperate climate, minimal information is known regarding its 

susceptibility to different fungi, mode of attack, resistance, and the classes. In the earliest 

research on susceptibility of bamboo to micro-organism, Liese (1985) reported that it is 

attacked especially by insects at ambient temperature; while the white, brown, and soft rot 

fungi are able to attack above the bamboo fibre saturation point. Liese (1985) and George 

(1985) both agree that the bamboo service life is estimated between 6 months to 3 years 

when in contact with soils. This finding is in a good agreement with Kaur et al. (2016b), 

which reveals that the untreated D. strictcus damages at 60% within 3 months and is 

completely destroyed within 6 months of exposure to termites.  

The soft rot decay on the different ages of Sinobambusa tootsik (Makino) is 

characterized by cavities only in the fibre cell walls, but parenchyma and vessel 

components are unattacked. The rate of decaying is influenced by the location of the fibre, 

culm age, and the degree of lignification of individual fibers (Murphy et al. 1991). In 

bamboo, the brown rot fungi consume the carbohydrate fraction of the walls and modifying 

lignin during the process. The white rot is able to consume both the carbohydrate and lignin 

fraction of the walls, while the soft rot consumes the carbohydrate fraction but probably 

not the lignin. Each fungal type gives its own microscopic characteristics at the cell wall 

levels after the decaying process. Brown rot gives indication of widespread amorphous 

degradation of the bulk cell wall, and white rot gives indication of localized degradation 

adjacent to the hyphae (bore holes and erosion troughs). The soft rot shows indication of 

discrete cavities within the secondary wall, erosion of the wall from the lumen, and 

possibly, a generalized dissolution of the wall (Sulaiman and Murphy 1994). 

The compound middle lamellae (CML) encompassing the cell corner regions are 

preferentially degraded in P. pubescens at an early stage of decaying process by white rot 

Lentinus edodes. The fibre secondary walls remain largely intact during this period. The 

preferential degradation of the CML compared to the fiber secondary walls strongly 

involves not only enzyme systems of the white rot fungus, but also a relationship to 

physicochemical properties of bamboo cell walls, particularly the influence of lignin 

composition and distribution (Kim et al. 2008). Same as the mode of white rot attacks, the 

CML in P. pubescens fibers are degraded at an early stage of decaying process by the G. 

trabum, which is confirmed by the distribution of H-unit lignin in the middle lamella. The 

absorbance bands assigned to lignin are decreased in the Fourier transform infrared spectra. 

The decay of bamboo fiber walls by G. trabeum is influenced by lignin distribution in the 

fiber walls. Polylaminate layers in bamboo fibers had an influence on cell wall degradation, 

with the narrow layers showing greater resistance than the broad layers (Cho et al. 2008). 

In G. scortechinii decayed by white rot (Coriolus versicolor) and brown rot 

(Coniophora puteana) fungi, the mode of hyphae attacks is penetration of the larger 

methaxylem vessel cell and further to the neighbor parenchyma and fibre cells through 

small pit membrane cells (Norul Hisham et al. 2012). Similarly, the hyphae of white rot P. 

chrysosporium and brown rot G. trabum attack the P. edulis through the parenchyma cells. 

Places near the inner skin are the most frequently attacked, and the vessels are the primary 

paths for the spread of mycelium. The bamboo crystalline structure decreases after being 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6542 

decayed by both fungi and the crystalline cellulose in bamboo is well deteriorated. The 

white rot strongly degrades the lignin component, then the hemicellulose and cellulose 

components. The brown-rot selectively degrades the hemicellulose fraction over cellulose 

and lignin. The decaying process is accompanied by oxidation and hydrolysis surface 

reactions, but the reaction rates behave differently for cellulose and lignin (Xu et al. 2013). 

In the decay resistance class of several bamboo species (Wei et al. 2013), the G. 

angustifolia is rather resistant to Trametes versicolor and same with D. asper against 

Chaetomium globosum. For brown-rot fungi, the Coniophora puteana and Gloeophyllum 

trabeum produce a low mass loss (maximum 2.9%). In contrast, the white-rot, T. versicolor 

yields the highest decay (max. 15.3%), while the Schizophyllum commune is considered as 

inactive (max. 3.2%). For soft-rot fungi, Ch. globosum gives a medium degradation (max. 

9.6%) and Paecilomyces variotii exhibits low degradation (max. 3.1%). The decay 

resistance depends on the bamboo species, type of fungi, the standard of testing, incubation 

period, and its environmental conditions. The deterioration is always expressed in 

percentage weight loss of oven-dried specimen before and after the exposure. For instance, 

the G. scortechinii exposed to brown rot Coniophora puteana showed 8.90% of weight 

loss after 8 weeks incubation period (Norul Hisham et al. 2012), while it was 18.70% 

weight loss after 52 weeks of incubation (Schmidt et al. 2013). The D. asper recorded 

15.3% weight loss after it was exposed to Schizosphyllum commune for 12 weeks (Suprapti 

2010), and it was only 4.3% weight loss after it was exposed to the same fungus for 52 

weeks (Schmidt et al. 2013). The P. pubescens showed only 5.3% weight loss after 52 

weeks was exposed to Gloeophyllum trabeum (Schmidt et al. 2011), and it recorded 

54.36% of weight loss after it was exposed for 8 weeks (Li et al. 2020). The weight loss of 

P. pubescens decayed by brown rot Coniophora puteana contact with soil was 6.3% 

compared to without soil contact, 25% (Schmidt et al. 2011) 

 

Overall 
Regardless of fungi (Table 22), the decay resistance is not significantly different 

for both monopodial (16.72%) and sympodial (14.22%) bamboo. The resistance is also not 

significantly different with fungi, either white, brown, or soft rot for both monopodial 

(19.06%, 15.92%, and 11.05%) and sympodial (14.03%, 12.88%, and 17.38%) bamboos, 

respectively. Generally, in all type of fungus, the monopodial bamboo is less resistant 

toward white, brown, and soft rot compared to the sympodial bamboo. 

 

Table 21. Weight Loss of Monopodial and Sympodial Bamboos Decayed by 
Fungi 

Species Week Fungi Strain % WL References 

Monopodial 

P. pubescens 52 WR Pleurotus ostreatus 
11 

21.0 Schmidt et al. 
(2011) 

P. pubescens 52 WR Schizophyllum 
commune 87 

5.2 Schmidt et al. 
(2011) 

P. pubescens 52 WR Schizophyllum 
commune 98 

4.4 Schmidt et al. 
(2011) 

P. pubescens 52 WR Trametes versicolor 
63 

47.8 Schmidt et al. 
(2011) 

P. pubescens (with soil 
contact) 

52 WR Schizophyllum 
commune 87 

6.3 Schmidt et al. 
(2011) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6543 

P. pubescens (without 
soil contact) 

52 WR Schizophyllum 
commune 87 

5.8 Schmidt et al. 
(2011) 

P. pubescens 8 WR Coriolus versicolor 60.48 Li et al. (2020) 

P. pubescens 52 BR Coniophora puteana 
167 

4.7 Schmidt et al. 
(2011) 

P. pubescens 52 BR Gloeophyllum 
trabeum 183 

5.3 Schmidt et al. 
(2011) 

P. pubescens (with soil 
contact) 

52 BR Coniophora Puteana 
167 

6.3 Schmidt et al. 
(2011) 

P. pubescens (without 
soil contact) 

52 BR Coniophora Puteana 
167 

25.0 Schmidt et al. 
(2011) 

P. pubescens 52 SR Chaetomium 
globosum 10 

38 Schmidt et al. 
(2011) 

P. pubescens 52 SR Paecilomyces 
variotii 13 

3.9 Schmidt et al. 
(2011) 

P. pubescens 8 BR Gloeophyllum 
trabeum 

54.36 Li et al. (2020) 

P. pubescens 52 BR Schizophyllum 
commune 87 

5.7 Schmidt et al. 
(2013) 

P. pubescens 52 BR Coniophora puteana 
167 

38.3 Schmidt et al. 
(2013) 

P. nigra 52 BR Coniophora puteana 
167 

32.6 Schmidt et al. 
(2013) 

P. nigra 52 WR Schizophyllum 
commune 87 

9.1 Schmidt et al. 
(2013) 

P. nigra (with soil 
contact) 

52 BR Coniophora puteana 
167 

15.5 Schmidt et al. 
(2011) 

P. nigra (without soil 
contact) 

52 BR Coniophora puteana 
167 

40.3 Schmidt et al. 
(2011) 

P. nigra (with soil 
contact) 

52 WR Schizophyllum 
commune 87 

16.4 Schmidt et al. 
(2011) 

P. nigra (without soil 
contact) 

52 WR Schizophyllum 
commune 87 

7.4 Schmidt et al. 
(2011) 

P.nigra Boryana (with 
soil contact) 

52 BR Coniophora puteana 
167 

35.3 Schmidt et al. 
(2011) 

P. nigra Boryana 
(without soil contact) 

52 BR Coniophora puteana 
167 

38.5 Schmidt et al. 
(2011) 

P. nigra Boryana (with 
soil contact) 

52 WR Schizophyllum 
commune 87 

19.7 Schmidt et al. 
(2011) 

P. nigra Boryana 
(without soil contact) 

52 WR Schizophyllum 
commune 87 

5.8 Schmidt et al. 
(2011) 

P. edulis 12 WR Phanerochaete 
chrysosporium 

40.0 Xu et al. (2013) 

P. edulis 12 BR Gloeophyllum 
trabeum 

34.6 Xu et al. (2013) 

Phyllostachys vivax 16 WR Trametes versicolor 19.0 Xu et al. (2013) 

Phyllostachys vivax 16 WR Xylaria polymorpha 17.5 Xu et al. (2013) 

Phyllostachys vivax 16 BR Coniophora puteana 9.2 Xu et al. (2013) 

Phyllostachys vivax 16 BR Oligoporus placenta 6.9 Xu et al. (2013) 

Phyllostachys vivax 16 SR Xylaria longipes 18.2 Xu et al. (2013) 

P. bambusoides (top 
with node) 

8 BR Coniophora puteana 6.9 Tomak et al. 
(2013) 

P. bambusoides (top 
with internode) 

8 BR Coniophora puteana 7.9 Tomak et al. 
(2013) 

P. bambusoides (middle 
with node) 

8 BR Coniophora puteana 6.4 Tomak et al. 
(2013) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6544 

P. bambusoides (middle 
with internode) 

8 BR Coniophora puteana 7.1 Tomak et al. 
(2013) 

P. bambusoides (basal 
with node) 

8 BR Coniophora puteana 4.1 Tomak et al. 
(2013) 

P. bambusoides (basal 
with internode) 

8 BR Coniophora puteana 6.1 Tomak et al. 
(2013) 

P. bambusoides (top 
with node) 

8 BR Poria placenta 6.5 Tomak et al. 
(2013) 

P. bambusoides (top 
with internode) 

8 BR Poria placenta 2.7 Tomak et al. 
(2013) 

P. bambusoides (middle 
with node) 

8 BR Poria placenta 3.4 Tomak et al. 
(2013) 

P. bambusoides (middle 
with internode) 

8 BR Poria placenta 0.5 Tomak et al. 
(2013) 

P. bambusoides (basal 
with node) 

8 BR Poria placenta 6.2 Tomak et al. 
(2013) 

P. bambusoides (basal 
with internode) 

8 BR Poria placenta 1.8 Tomak et al. 
(2013) 

A. amabilis 52 BR Coniophora puteana 
167 

38.6 Schmidt et al. 
(2013) 

A. amabilis 52 BR Schizophyllum 
commune 87 

10.8 Schmidt et al. 
(2013) 

Sympodial 

G. atroviolacea 52 WR Pleurotus ostreatus 
11 

10.6 Schmidt et al. 
(2011) 

G. atroviolacea 52 WR Schizophyllum 
commune 87 

6.7 Schmidt et al. 
(2011) 

G. atroviolacea 52 WR Schizophyllum 
commune 98 

5.6 Schmidt et al. 
(2011) 

G. atroviolacea 52 WR Trametes versicolor 
63 

51.6 Schmidt et al. 
(2011) 

G. atroviolacea 12 WR Coriolus versicolor 
1030 

7.7 Suprapti (2010) 

G. atroviolacea 12 WR Phanerochaete 
chrysosporium 

HHBI-320 

7.6 Suprapti (2010) 

G. atroviolacea 12 WR P. sordida HI IBI-
321 

5.4 Suprapti (2010) 

G. atroviolacea 12 WR Phlebia brevispora 
Mad. 

4.1 Suprapti (2010) 

G. atroviolacea 12 WR Postia placenta 
Mad- 696 

5.1 Suprapti (2010) 

G. atroviolacea 12 WR Pycnoporus 
sanguineus FIHBI-

324 

14.4 Suprapti (2010) 

G. atroviolacea 12 WR P. sanguineus 
HHBI-8149 

3.8 Suprapti (2010) 

G. atroviolacea 12 WR Schizophyllum 
commune FIHBI-204 

5.8 Suprapti (2010) 

G. atroviolacea 12 WR S. commune HHBI-
222 

2.8 Suprapti (2010) 

G. atroviolacea 52 BR Coniophora puteana 
167 

5.6 Schmidt et al. 
(2011) 

G. atroviolacea 52 BR Gloeophyllum 
trabeum 183 

5.7 Schmidt et al. 
(2011) 

G. atroviolacea 12 BR Dacryopinax 3.4 Suprapti (2010) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6545 

spathularia HHBI-
145 

G. atroviolacea 12 BR D. spathularia HHBI-
223 

7.7 Suprapti (2010) 

G. atroviolacea 12 BR Lentinus lepideus 
Mad-534 

3.9 Suprapti (2010) 

G. atroviolacea 12 BR Polyporus sp. HHBI-
209 

20.9 Suprapti (2010) 

G. atroviolacea 12 BR Tyromyces palustris 
FRI Japan-507 

21.0 Suprapti (2010) 

G. atroviolacea 12 SR Chaetomium 
globosum FRI Japan 

5-1 

4.6 Suprapti (2010) 

G. atroviolacea 52 SR Chaetomium 
globosum 10 

9.4 Schmidt et al. 
(2011) 

G. atroviolacea 52 SR Paecilomyces 
variotii 13 

3.6 Schmidt et al. 
(2011) 

B. maculate 52 WR Pleurotus ostreatus 
11 

28.2 Schmidt et al. 
(2011) 

B. maculate 52 WR Schizophyllum 
commune 87 

2.8 Schmidt et al. 
(2011) 

B. maculate 52 WR Schizophyllum 
commune 98 

1.8 Schmidt et al. 
(2011) 

B. maculate 52 WR Trametes versicolor 
63 

62.5 Schmidt et al. 
(2011) 

B. maculate 52 WR Schizophyllum 
commune 87 

5.1 Schmidt et al. 
(2013) 

B. maculate 52 BR Coniophora puteana 
167 

3.6 Schmidt et al. 
(2011) 

B. maculate 52 BR Gloeophyllum 
trabeum 183 

1.9 Schmidt et al. 
(2011) 

B. maculate 52 BR Coniophora puteana 
167 

20.4 Schmidt et al. 
(2013) 

B. maculate 52 SR Chaetomium 
globosum 10 

31.8 Schmidt et al. 
(2011) 

B. maculate 52 SR Paecilomyces 
variotii 13 

1.2 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 WR Schizophyllum 
commune 3 

7.4 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 WR Schizophyllum 
commune 3 

6.9 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 WR Schizophyllum 
commune 3 

5.0 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 WR Schizophyllum 
commune 3 

4.6 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 WR Schizophyllum 
commune 4 

9.1 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 WR Schizophyllum 
commune 4 

6.4 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 WR Schizophyllum 
commune 4 

5.6 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 WR Schizophyllum 
commune 4 

4.8 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 WR Trametes versicolor 
63 

28.3 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 WR Trametes versicolor 16.9 Schmidt et al. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6546 

63 (2011) 

M. bambusoides (2 
years) 

24 WR Trametes versicolor 
63 

12.5 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 WR Trametes versicolor 
63 

14.7 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 BR Coniophora puteana 
1 

11.2 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 BR Coniophora puteana 
1 

13.7 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 BR Coniophora puteana 
1 

8.4 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 BR Coniophora puteana 
1 

9.9 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 BR Oligoporus placenta 
120 

5.6 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 BR Oligoporus placenta 
120 

6.8 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 BR Oligoporus placenta 
120 

5.4 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 BR Oligoporus placenta 
120 

6.0 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 SR Chaetomium 
globosum 76 

52.7 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 SR Chaetomium 
globosum 76 

31.4 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 SR Chaetomium 
globosum 76 

32.3 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 SR Chaetomium 
globosum 76 

27.9 Schmidt et al. 
(2011) 

M. bambusoides (0.5 
year) 

24 SR Paecilomyces 
variotii 92 

19.7 Schmidt et al. 
(2011) 

M. bambusoides (1 year) 24 SR Paecilomyces 
variotii 92 

9.6 Schmidt et al. 
(2011) 

M. bambusoides (2 
years) 

24 SR Paecilomyces 
variotii 92 

8.2 Schmidt et al. 
(2011) 

M. bambusoides (3 
years) 

24 SR Paecilomyces 
variotii 92 

7.5 Schmidt et al. 
(2011) 

B. polymorpha (Top) 24 WR Schizophyllum 
commune 4 

5.9 Schmidt et al. 
(2011) 

B. polymorpha (Basal) 24 WR Schizophyllum 
commune 4 

4.1 Schmidt et al. 
(2011) 

B. polymorpha (Top) 24 BR Coniophora puteana 
1 

12.1 Schmidt et al. 
(2011) 

B. polymorpha (Basal) 24 BR Coniophora puteana 
1 

15.5 Schmidt et al. 
(2011) 

B. polymorpha (Top) 24 BR Oligoporus placenta 
120 

12.8 Schmidt et al. 
(2011) 

B. polymorpha (Basal) 24 BR Oligoporus placenta 
120 

5.7 Schmidt et al. 
(2011) 

B. polymorpha (Top) 24 SR Chaetomium 
globosum 76 

33.7 Schmidt et al. 
(2011) 

B. polymorpha (Basal) 24 SR Chaetomium 
globosum 76 

23.3 Schmidt et al. 
(2011) 

B. polymorpha (Top) 24 SR Paecilomyces 
variotii 92 

14.1 Schmidt et al. 
(2011) 

B. polymorpha (Basal) 24 SR Paecilomyces 9.4 Schmidt et al. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6547 

variotii 92 (2011) 

D. strictus (Top) 24 WR Schizophyllum 
commune 4 

2.7 Schmidt et al. 
(2011) 

D. strictus (Basal) 24 WR Schizophyllum 
commune 4 

4.7 Schmidt et al. 
(2011) 

D. strictus, Teri 12 WR Polyporus versicolor 54.1 Kaur et al. (2016b) 

D. strictus, Allahabad 12 WR Polyporus versicolor 54.3 Kaur et al. (2016b) 

D. strictus (Top) 24 BR Coniophora puteana 
1 

38.2 Schmidt et al. 
(2011) 

D. strictus (Basal) 24 BR Coniophora puteana 
1 

12.7 Schmidt et al. 
(2011) 

D. strictus (Top) 24 BR Oligoporus placenta 
120 

11.3 Schmidt et al. 
(2011) 

D. strictus (Basal) 24 BR Oligoporus placenta 
120 

9.8 Schmidt et al. 
(2011) 

D. strictus (Top) 24 SR Chaetomium 
globosum 76 

28.2 Schmidt et al. 
(2011) 

D. strictus (Basal) 24 SR Chaetomium 
globosum 76 

6.7 Schmidt et al. 
(2011) 

D. strictus (Top) 24 SR Paecilomyces 
variotii 92 

9.3 Schmidt et al. 
(2011) 

D. strictus (Basal) 24 SR Paecilomyces 
variotii 92 

2.1 Schmidt et al. 
(2011) 

O. nigro-ciliata (Top) 24 WR Schizophyllum 
commune 4 

2.1 Schmidt et al. 
(2011) 

O. nigro-ciliata (Basal) 24 WR Schizophyllum 
commune 4 

2.8 Schmidt et al. 
(2011) 

O. nigro-ciliata (Top) 24 BR Coniophora puteana 
1 

38.5 Schmidt et al. 
(2011) 

O. nigro-ciliata (Basal) 24 BR Coniophora puteana 
1 

4.0 Schmidt et al. 
(2011) 

O. nigro-ciliata (Top) 24 BR Oligoporus placenta 
120 

7.4 Schmidt et al. 
(2011) 

O. nigro-ciliata (Basal) 24 BR Oligoporus placenta 
120 

1.6 Schmidt et al. 
(2011) 

O. nigro-ciliata (Top) 24 SR Chaetomium 
globosum 76 

41.1 Schmidt et al. 
(2011) 

O. nigro-ciliata (Basal) 24 SR Chaetomium 
globosum 76 

21.1 Schmidt et al. 
(2011) 

O. nigro-ciliata (Top) 24 SR Paecilomyces 
variotii 92 

7.4 Schmidt et al. 
(2011) 

O. nigro-ciliata (Basal) 24 SR Paecilomyces 
variotii 92 

6.0 Schmidt et al. 
(2011) 

T. oliveri (Top) 24 WR Schizophyllum 
commune 4 

6.8 Schmidt et al. 
(2011) 

T. oliveri (Basal) 24 WR Schizophyllum 
commune 4 

3.2 Schmidt et al. 
(2011) 

T. oliveri (Top) 24 BR Coniophora puteana 
1 

18.5 Schmidt et al. 
(2011) 

T. oliveri (Basal) 24 BR Coniophora puteana 
1 

8.2 Schmidt et al. 
(2011) 

T. oliveri (Top) 24 BR Oligoporus placenta 
120 

6.4 Schmidt et al. 
(2011) 

T. oliveri (Basal) 24 BR Oligoporus placenta 
120 

2.3 Schmidt et al. 
(2011) 

T. oliveri (Top) 24 SR Chaetomium 47.2 Schmidt et al. 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6548 

globosum 76 (2011) 

T. oliveri (Basal) 24 SR Chaetomium 
globosum 76 

27.2 Schmidt et al. 
(2011) 

T. oliveri (Top) 24 SR Paecilomyces 
variotii 92 

7.7 Schmidt et al. 
(2011) 

T. oliveri (Basal) 24 SR Paecilomyces 
variotii 92 

5.2 Schmidt et al. 
(2011) 

B. vulgaris 12 WR Coriolus versicolor 
1030 

7.2 Suprapti (2010) 

B. vulgaris 12 WR Phanerochaete 
chrysosporium 

HHBI-320 

8.7 Suprapti (2010) 

B. vulgaris 12 WR P. sordida HI IBI-
321 

5.4 Suprapti (2010) 

B. vulgaris 12 WR Phlebia brevispora 
Mad. 

4.8 Suprapti (2010) 

B. vulgaris 12 WR Postia placenta 
Mad-696 

4.5 Suprapti (2010) 

B. vulgaris 12 WR Pycnoporus 
sanguineus FIHBI-

324 

22.5 Suprapti (2010) 

B. vulgaris 12 WR P. sanguineus 
HHBI-8149 

5.0 Suprapti (2010) 

B. vulgaris 12 WR Schizophyllum 
commune FIHBI-204 

8.8 Suprapti (2010) 

B. vulgaris 12 WR S. commune HHBI-
222 

4.6 Suprapti (2010) 

Bambusa vulgaris 
Schrad. 

3 WR Trametes versicolor 48.09 Poonia et al. 
(2021) 

B. vulgaris 12 BR Dacryopinax 
spathularia HHBI-

145 

5.2 Suprapti (2010) 

B. vulgaris 12 BR D. spathularia HHBI-
223 

4.7 Suprapti (2010) 

B. vulgaris 12 BR Lentinus lepideus 
Mad-534 

4.1 Suprapti (2010) 

B. vulgaris 12 BR Polyporus sp. HHBI-
209 

36.2 Suprapti (2010) 

B. vulgaris 12 BR Tyromyces palustris 
FRI Japan-507 

37.4 Suprapti (2010) 

Bambusa vulgaris 
Schrad. 

3 BR Rhodonia placenta 47.65 Poonia et al. 
(2021) 

B. vulgaris 12 SR Chaetomium 
globosum FRI Japan 

5-1 

7.0 Suprapti (2010) 

B. nutans, Dehradun 12 WR Polyporus versicolor 58.2 Kaur et al. (2016b) 

B. arundinacea, 
Allahabad 

12 WR Polyporus versicolor 55.7 Kaur et al. (2016b) 

B. tulda, Bihar 12 WR Polyporus versicolor 57.4 Kaur et al. (2016b) 

B. bambus, IWST, 
Bangalore 

12 WR Polyporus versicolor 56.9 Kaur et al. (2016b) 

B. pallida, IWST, 
Bangalore 

12 WR Polyporus versicolor 59.2 Kaur et al. (2016b) 

D. asper 12 WR Coriolus versicolor 
1030 

15.2 Suprapti (2010) 

D. asper 12 WR Phanerochaete 7.2 Suprapti (2010) 



 

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chrysosporium 
HHBI-320 

D. asper 12 WR P. sordida HI IBI-
321 

7.5 Suprapti (2010) 

D. asper 12 WR Phlebia brevispora 
Mad. 

11.1 Suprapti (2010) 

D. asper 12 WR Postia placenta 
Mad-696 

3.7 Suprapti (2010) 

D. asper 12 WR Pycnoporus 
sanguineus FIHBI-

324 

19.0 Suprapti (2010) 

D. asper 12 WR P. sanguineus 
HHBI-8149 

8.9 Suprapti (2010) 

D. asper 12 WR Schizophyllum 
commune FIHBI-204 

15.0 Suprapti (2010) 

D. asper 12 WR S. commune HHBI-
222 

4.6 Suprapti (2010) 

D. asper 52 WR Schizophyllum 
commune 87 

4.3 Schmidt et al. 
(2013) 

D. asper 12 BR Dacryopinax 
spathularia HHBI-

145 

6.7 Suprapti (2010) 

D. asper 12 BR D. spathularia HHBI-
223 

8.8 Suprapti (2010) 

D. asper 12 BR Lentinus lepideus 
Mad-534 

5.8 Suprapti (2010) 

D. asper 12 BR Polyporus sp. HHBI-
209 

21.0 Suprapti (2010) 

D. asper 52 BR Coniophora puteana 
167 

29.3 Schmidt et al. 
(2013) 

D. asper 12 BR Tyromyces palustris 
FRI Japan-507 

16.5 Suprapti (2010) 

D. asper 12 SR Chaetomium 
globosum FRI Japan 

5-1 

8.0 Suprapti (2010) 

D. gigantues 12 BR Gloeophyllum 
trabeum 

13.36 Brito et al. (2020) 

D. gigantues 12 BR Postia placenta 22.47 Brito et al. (2020) 

G. apus 12 WR Coriolus versicolor 
1030 

4.8 Suprapti (2010) 

G. apus 12 WR Phanerochaete 
chrysosporium 

HHBI-320 

6.5 Suprapti (2010) 

G. apus 12 WR P. sordida HI IBI-
321 

5.4 Suprapti (2010) 

G. apus 12 WR Phlebia brevispora 
Mad. 

3.8 Suprapti (2010) 

G. apus 12 WR Postia placenta 
Mad-696 

4.8 Suprapti (2010) 

G. apus 12 WR Pycnoporus 
sanguineus FIHBI-

324 

9.0 Suprapti (2010) 

G. apus 12 WR P. sanguineus 
HHBI-8149 

4.0 Suprapti (2010) 

G. apus 12 WR Schizophyllum 
commune FIHBI-204 

4.5 Suprapti (2010) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6550 

G. apus 12 WR S. commune HHBI-
222 

3.2 Suprapti (2010) 

G. apus 12 BR Dacryopinax 
spathularia HHBI-

145 

5.80 Suprapti (2010) 

G. apus 12 BR D. spathularia HHBI-
223 

5.9 Suprapti (2010) 

G. apus 12 BR Lentinus lepideus 
Mad-534 

4.3 Suprapti (2010) 

G. apus 12 BR Polyporus sp. HHBI-
209 

21.7 Suprapti (2010) 

G. apus 12 BR Tyromyces palustris 
FRI Japan-507 

23.8 Suprapti (2010) 

G. apus 12 SR Chaetomium 
globosum FRI Japan 

5-1 

7.5 Suprapti (2010) 

G. pseudoarundinacea 12 WR Coriolus versicolor 
1030 

20.7 Suprapti (2010) 

G. pseudoarundinacea 12 WR Phanerochaete 
chrysosporium 

HHBI-320 

16.6 Suprapti (2010) 

G. pseudoarundinacea 12 WR P. sordida HI IBI-
321 

5.3 Suprapti (2010) 

G. pseudoarundinacea 12 WR Phlebia brevispora 
Mad. 

10.3 Suprapti (2010) 

G. pseudoarundinacea 12 WR Postia placenta 
Mad-696 

13.0 Suprapti (2010) 

G. pseudoarundinacea 12 WR Pycnoporus 
sanguineus FIHBI-

324 

32.6 Suprapti (2010) 

G. pseudoarundinacea 12 WR P. sanguineus 
HHBI-8149 

8.7 Suprapti (2010) 

G. pseudoarundinacea 12 WR Schizophyllum 
commune FIHBI-204 

26.6 Suprapti (2010) 

G. pseudoarundinacea 12 WR S. commune HHBI-
222 

4.0 Suprapti (2010) 

G. pseudoarundinacea 12 BR Dacryopinax 
spathularia HHBI-

145 

4.1 Suprapti (2010) 

G. pseudoarundinacea 12 BR D. spathularia HHBI-
223 

3.3 Suprapti (2010) 

G. pseudoarundinacea 12 BR Lentinus lepideus 
Mad-534 

11.0 Suprapti (2010) 

G. pseudoarundinacea 12 BR Polyporus sp. HHBI-
209 

27.4 Suprapti (2010) 

G. pseudoarundinacea 12 BR Tyromyces palustris 
FRI Japan-507 

26.9 Suprapti (2010) 

G. pseudoarundinacea 12 SR Chaetomium 
globosum FRI Japan 

5-1 

9.3 Suprapti (2010) 

G. scortechinii (0.5 year) 8 WR Coriolus versicolor 9.90 Norul Hisham et al. 
(2012) 

G. scortechinii (3.5 
years) 

8 WR Coriolus versicolor 9.24 Norul Hisham et al. 
(2012) 

G. scortechinii (6.5 
years) 

8 WR Coriolus versicolor 5.30 Norul Hisham et al. 
(2012) 



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6551 

G. scortechinii (0.5 year) 8 BR Coniophora puteana 9.95 Norul Hisham et al. 
(2012) 

G. scortechinii (3.5 
years) 

8 BR Coniophora puteana 9.49 Norul Hisham et al. 
(2012) 

G. scortechinii (6.5 
years) 

8 BR Coniophora puteana 8.90 Norul Hisham et al. 
(2012) 

G. scortechinii 52 BR Coniophora puteana 
167 

18.7 Schmidt et al. 
(2013) 

G. scortechinii 52 BR Schizophyllum 
commune 87 

4.7 Schmidt et al. 
(2013) 

 

Table 22. Statistical Analysis of Weight Loss of Decayed Monopodial and 
Sympodial Bamboo  

Fungus Rhizome DF F Significance 

Monopodial Sympodial 

White rot 19.06X 

(17.21) 
14.03X 

(16.49) 
1 0.02 0.08NS 

Brown rot 15.92X 

(15.55) 
12.88X 

(10.57) 

Soft rot 11.05X 

(10.11) 
17.38X 

(13.98) 

Average 16.72 

(15.77) 
14.22 

(14.25) 

DF– Degree of freedom, F– F ratio, NS is not significant at P > 0.1, *** is significant at P < 0.0. 
The value in parentheses is standard deviation. 

 
 

CONCLUSIONS 
 
1. The bamboo properties are not only different with genera, species, and site location, 

but also with rhizome type. The monopodial bamboo has shorter sprouting time, growth 

phase, diameter breast height, and overall height than the sympodial bamboo. 

2. Anatomically, the monopodial bamboo contains a higher radial length and tangential 

diameter, but its radial length/tangential diameter is smaller than the sympodial 

bamboo. The vascular bundle frequency is higher in monopodial bamboo. 

3. The monopodial bamboo contains higher -cellulose, hemicellulose, lignin, hot water 

soluble, cold water soluble, and 1% NaOH soluble contents. Monopodial bamboo has 

a higher tensile and modulus of rupture but there is not much difference in the 

compression strength. 

4. The fibre length and diameter are longer and wider in sympodial bamboo, but the fibre 

lumen diameter and wall thickness are not different in either monopodial or sympodial 

bamboos. 

5. The proportion of metaxylem vessel diameter is higher in sympodial bamboo but the 

proportion of fibre and parenchyma are not much different for monopodial and 

sympodial bamboos. The volumetric shrinkage is higher in sympodial bamboo, but the 

density is not much different for both rhizome types. 

  



 

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Hamid et al. (2023). “Mono & sympodial bamboo,” BioResources 18(3), 6499-6560.  6552 

6. Sympodial bamboo has a significantly higher ash content, but both the silica and 

ethanol/toluene extractive contents are not much different from either monopodial or 

sympodial bamboo. Overall, the sympodial bamboo is more resistant toward white, 

brown, and soft rot compared to the monopodial bamboo. 

 

 

ACKNOWLEDGMENTS 
 

The authors are grateful for the support of the Ministry of Higher Education (Grant 

No: PRGS/1/2019/WAB07/UPM/02/1-5534900)  

 
 
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Article submitted: June 9, 2022; Peer review completed: November 30, 2022; Revisions 

received: April 18, 2023; Revisions accepted: April 24, 2023; Published: May 25, 2023. 

DOI: 10.15376/biores.18.3.Hamid