Anatomy and Properties of

Bamboo
W. Liese
Institute of Wood Biology and Wood Preservation of the Federal
Research Centre for Forestry and Forest Products,
Leuschnerstr, 91, 2050 Hamburg, Federal Republic of Germany

Abstract
The numerous alternatives in the use of bamboo depend on
the unique properties of its culm. In order to understand the
anatomical and chemical make-up and its ensuing
mechanical properties, an attempt has been made to
summarize the accessible. information.

Anatomy
Gross anatomy: The properties of the culm are determined
by its anatomical structure. The culm consists of internodes
and nodes. At the internodes, the cells are axially oriented,
whereas at the nodes, cells provide the transverse
interconnections. No radial cell elements, such as rays, exist
in the inter-nodes. Within the nodes an intensive branching
of the vessels occurs. These also bend radially inward and
provide
transverse
conduction
through
the
nodal
diaphragms, so that all parts of the culm are interwoven.
The outer part of the culm is formed by two epidermal cell
layers, the inner appearing thicker and highly lignified. The
surface of outermost cells are covered by a cutinized layer
with a wax coating. The inner parts of the culm consist of
numerous sclerenchyma cells. Any lateral movement of
liquids is therefore much hindered. Pathways for penetration
are thus only the cross ends of the culm and to a much
smaller extent the sheath scars around the nodes. The gross
anatomical structure of a transverse section of any culm
internode is determined by the shape, size, arrangement
and number of the vascular bundles. Thev are clearly
contrasted by the darker colored sclerenchymatous tissue

against the paren-chymatous ground tissue. At the

peripheral zone of the culm the vascular bundles are smaller
and more numerous,' in the inner parts larger and fewer
thin the culm wall the total number of vascular bundles
decreases from bottom towards the top, while their density
increases at the same time. The culm tissue is mostly
parenchyma and the vascular bundles which are com-posed
of vessels, sieve tubes with companion cells and fibres. The
total culm comprises about 50% parenchyma, 40% fibre,
and 10% conducting tissues (vessels and sieve tubes) with
some variation according to species. The percentage
distribution and orientation of cells show a definite pattern
within the culm, both horizontally and vertically. Parenchyma
and conducting cells are more frequent in the inner third of
the wall, whereas in the outer third the percentage of fibers
is distinctly higher. In the vertical direction the amount of
fibres increases from bottom to top and that of parenchyma
decreases The common practice of leaving the upper part of
a cut culm unused in the forest is therefore a waste with
regard to its higher fibre content.

Parenchyma:
The ground tissue consists of parenchyma cells, which are
mostly vertically elongated (l00 x 20 um) with short, cubelike ones interspersed in between. The former are
characterized by thicker walls with a polylamellate structure
they become lignified in the early stages of shoot growth.
The shorter cells have a denser cytoplasm, thinner walls and
retain their cyto-plasmic activity for a long time. The
function of these two different types of parenchyma cells is
still unknown. Of interest in the structure of parenchyma
walls is the occurrence of warts in many taxa like Bambusa,
Cephalostachyum, Dendrocalamus, Oxytenanthera, and
Thyrostachys, which have not been observed so far in the
parenchyma of hardwoods. Genuine warts have to be
carefully distinguished from cytoplasmic debris, which are
also frequent in parenchyma cells after the death of the
protoplast. Their distribution is variable from verydense to

sparse. Among the species examined the parenchyma cells

appear to .possess a even higher number and density of
warts than fibres and vessel members. Their size varies from
120 - 520 nm. The occurrence of warts in the lignified
parentchyma cells of bamboo is perhaps an expression of
the close association of lignin-like nature of warts, since
warts have not been observed in non-lignified cells
(Parameswara"and Liese, 1977).Vascular bundles: The
vascular bundle in the bamboo culm consists of the xylem
with one or two smaller protoxylem elements and two large
metaxylem vessels (40 - 120 u)and the phloem with thin
walled, unlignified sieve tubes connected to companion cells
The vessels possess large diameters in the inner parts of the
culm wall and become small towards outside. These water
conducting elements have to function throughout the
lifetime of a culm without the formation of any new tissue,
as in the case of hardwoods and softwoods with cambial
activity. In older culms, vessels and sieve tubes can become
partly impermeable due to depositions of gum-like
substances, thus losing their conductivity which may cause'
death of the aged culms. The one or two tracheary elements
of the protoxylem have mostly annular thickenings. They are
local areas of stasis accumulating wall material, which are
connected with each other by membranes in the early
stages of development. During extension growth of the cell,
they are disrupted. The walls of metaxylem vessels of
bamboo are characterized by a middle lamella and a primary
wall together with a well developed zonation of the
secondary wall into 51 and 52. Whereas the 51 possesses a
flat spiral arrangement of fibrils (90 - 95) the 52 zone shows
a slight deviation from the known fibril orientation in
tracheids. The fibrils are arranged at an angle of 30 - 90 to
the cell axis; also micro lamellae are present with fibrils
arranged in a fan-like fashion. This wall structure perhaps to
be considered as "normal", is modified in some taxa like
Oxy-tenathera abysinica and Me/ocanna bam-busoides to
such an extent that a polylamellae construction results,
resembling a paren-chyma wall with the herringbone pattern
of fibrillar arrangement whereby the number of layers are

mostly restricted to two to four (Parameswaran and Liese,

1980). Warts have been observed in the metaxylem of
vessels
of
Oxytenanthera
nigrociliata,
Me/o-canna
bambusoides, Gigantoch/oa alter.nigra, Schizostachyum
b/umei, and S.brachyc/adiUm. The pits of these vessels
towards the surrounding parenchyma of adjacent vessel
elements are slightly bordered. Their membrane consists of
fibrils with a net-like texture, resembling hardwood pits.

CHEMICAL PROPERTIES
Chemical constitution:
The main constituents of the bamboo culms are cellulose,
hemicellulose and lignin; minor constituents consist of
resins, tannins, waxes and inorganic salts. The composition
varies according to species, the conditions of growth, the
age of the bamboo and the part of the culm. Because the
bamboo culm tissue matures within a year when the soft
and fragile sprout becomes hard and strong, the proportion
of lignin and carbohydrates is changed during this period.
However, after the full maturation of the culm, the chemical
composition tends to remain rather constant. Approximate
chemical analysis for some bamboo species. Small
differences exist along a culm, The nodes contain less watersoluble extractives, pentosans, ash, and lignin but more
cellulose than the inter-nodes.The season influences the
amount of water-soluble materials, which are higher in the
dry season than in the rainy season. The starch content
reaches' its maximum in the driest months before the rainy
season and sprouting. The ash content (1 - 5%) ishigher in
the inner part than in the outer one. The silica content varies
on an average from 0:5 to 4%. increasing from bottom to
top. Most silica is deposited in the epidermis, the skin
zone", whereas the nodes contain little silica and the tissues
of the internodes almost none. Silica content affects the
pulping properties of bamboo. Cellulose and hemicellulose:
The cellulose in bamboo amounts - as holocellulose - to
more than 50 % of the chemical constituents. As in other
plants it consists of linear chains of 1, 4 bonded
hydroglucose units (C2H1206).The number of glucose units

in one molecular chain is referred to as the degree of

polymerization (OP). The OP for bamboo is considerably
higher than for dicotyledoneous woods. Cellulose is difficult
to isolate in pure form because it is closely associated with
the hemicelluloses and the lignin. More than 90% of the
bamboo hemicelluloses consist of axylan which seems to be
aI,4-linked linear polymer forming a 4-0-methyl-O-glucuronic
acid, L-arabinose, and O-xylose in a molar ratio of 1.0 : 1.3 :
25 respectively. It is in the main chain linear, but appears to
be different from the xylan found in the woods of
gymnosperms with regard to the degree of branching and
molecular properties. Further-more, the bamboo xylan
contains 6 - 7% of native acetyl groups, which is a feature
shared by hardwoods. With regard to the presence of
arabinose it is closer to softwoods. Thus, the bamboo xylan
is intermediate between hard-wood and softwood xylans.
These results indicate that the bamboo xylan has the unique
structure
of
Gramineae
(Higuchi,1980).Lignin:
After
cellulose, lignin represents the second most abundant
constituent in the bamboo and much interest has been
focused on its chemical nature and structure. Bamboo lignin
is a typical grass lignin, which is built up from the three
phenyl-propane units p-coumaryl, coniferyl, and sinapyl
alcohols interconnected through biosynthetic pathways.
Bamboo grows very rapidly and completes the height growth
within a few months reaching the full size. The growing
bamboo shows various lignifications stages from the bottom
to the top portions of the same culm (ltoh and Shimaji,
1981). The lignifications within every internode proceeds
downward from top to bottom, whereas transversely
proceeds from inside to outside. During the height growth
lignifications of epidermal cells and fibres precede that of
ground tissue parenchyma. Full lignifications of bamboo
culm is completed within one growing season, showing no
further ageing effects. No difference has been detected in
lignin
composition
between
vascular
bundles
and
parenchyma tissue (Higuchi et al., 1966). Bamboo has been
chosen as one of the suitable plants to study the
biosynthesis of lignin. Initially, these investigations were

almost exclusively based on feeding experiments with radioactive precursors and it has been known that lignin is
synthesized from glucose formed by photosynthesis via the
"Shikimic acid pathway" (Higuchi, 1969) . Several key
enzymes involved in the synthesis of shikimic acid were
isolated' from bamboo shoots (Fennel and Shako, 1984;
1985).

Physical and Mechanical Properties

Moisture content: The moisture content varies within one
culm and is influenced by its age, the season of felling and
the species. In the green stage greater differences exist
within one culm as well as in relation to age, season and
species. Young, one-year old shoots have a high relative
moisture content of about 120 - 130% both at bottom and
top. The nodes, however, show lower values than the
internodes. These differences can amount to 25 % of the
water content and are larger at the base than at the top. In
culms of 3 - 4 years the base has a higher moisture content
than the top, e.g. for Dendrocalamus strictus about 100%
and 60% relative moisture content respectively. The
moisture content across the culm wall is higher in the inner
part than in the outer part. The season has a great influence
on the water content of the culm, with a minimum atthe end
of the dry period, followed by a maximum in the rainy
season. During this period the stem can double its water
content. The variation due to the season is higher than the
differences between base and top as well as between
species. Among species the water content varies even in the
same locality. This is mainly due to the variation in the
amount of parenchyma cells, which corresponds to water
holding capacity (Liese and Grover,1961). The considerable
differences in the moisture content of freshly felled culms
have to be considered when determining the yield of
bamboo expressed by its fresh weight.Fibre saturation point
and shrink age The fibre saturation point is influenced by the
composition of the tissue and the amount of hygroscopic
extractives. Since fibres and parenchyma have apparently a

different fibre saturation point, their varying amount within a

culm leads to different values. The fibre saturation point
consequently differs within one culm and between species.
For Dendroca Jamus strictus the mean value was determined
to be about 20%, for Phyllostachys pubescens about 13%
(Ota, 1955).Unlike wood, bamboo begins to shrink right from
the beginning of seasoning. The shrinkage affects both the
thickness of the culm walls and the circumference.
Seasoning of mature bamboo from green condition to about
20% moisture content leads to a shrinkage of 4 to 14% in
the wall thickness and 3 to 12% in diameter. Bamboo tissue
shrinks mainly in the radial direction, and the minimum
deformation occurs in the axial direction. The tangential
shrinkage is higher in the outer parts of the wall than in the
inner parts. The shrinkage of the whole wall appears to be
governed by the shrinkage of the outermost portion, which
possesses also the highest specific gravity. Mature culms
shrink less than immature ones. Value of shrinkage from
freshly felled to the oven-dry state were determined for
Phyl-lostachys pubescens as follows: tangential: 8.2% for the
outer part of the wall and 4.1 % for the inner; radial: 6, 8%
for the outer part and 7.2% for the inner; longitudinal: 0.17%
for the outer part and 0.43% for the il1ner. Shrinkage starts
simultaneously with the decrease of moisture content but
does not continue regularly as water content diminishes
from 70 to 40%, shrinkage stops; below this range it can
again be initiated. Parenchyma tissue shrinks less in bamboo
than in timber, while vascular fibers shrink as much as in
timbers of the same specific gravity. When the moisture
content is low, swelling due to absorption of water is almost
equal to shrink-age. Moist heating leads to irreversible
swelling in all directions. The percentage of swelling
decreases with an increase of basic density (Kishen et al.,
1958; Sekhar and Rawat, 1964).

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