bamboo structures in colour
133
Fig. 133: Vietnamese ladies’ hats made from Neohouzeaua dulloa
128 the anatomy of bamboo culms
structuralChapter Five modifications +107
129 CHAPTER FIVE structural modifications
Growth Effects
DEVELOPMENT AND AGEING
The previous chapters have outlined the general structure of mature culms and its variations. Few investigations have dealt with structural development until maturation (Grosser 1971; Fujii 1985; Alvin and Murphy 1988). Recent observations have shown that some structural modifications occur after maturation also (Liese and Weiner 1996, 1997). The ageing of a bamboo culm influences certain properties and, consequently, its processing and utilization. This section describes the changes during culm elongation and maturation, and certain modifications that occur in mature culms.
The elongation of a culm results from the expansion of individual internodes, already present in the bud. Differentiation starts at the upper part of an internode by elongation of different cell types, and proceeds down to its base. The differentiation of an individual internode is completed in just a few days (Hsiung et al. 1980). Longitudinally, there is no major difference in the composition and structure of the tissue within an internode, except for the expansion of the cells, especially the fibres. The fibres undergo a significant elongation from only a few microns to about 2 mm. Within the internode, the fibres near the nodes are always shorter than in the middle portion (see page 61 & Fig. 82). Elongation, however, causes only a few anatomical changes along culm length. The narrowing of the culm wall in the upper part results in a reduction of its inner portion with less parenchyma, and the vascular bundles decrease in size but increase in number (see page 27). The upper culm part has a higher fibre content with a higher specific gravity.
The younger stage of a culm is marked by certain external characteristics that are useful indicators to avoid premature harvesting. These include the presence of culm sheaths, bud break, branching pattern, number of leaf scars, and colour changes of the stem ranging from fresh green to often yellow-grey (Banik 1993). A young hairy culm becomes glabrous at maturity.
Structural modifications during the maturation phase and the years following relate to fibres and parenchyma cells. The results obtained by Alvin and Murphy (1988), Majima
130 the anatomy of bamboo culms
et al. (1991), Liese and Weiner (1996, 1997) and Murphy and Alvin (1997 a,b) have demonstrated that the maturation process of fibres proceeds quite differently over the transverse section of a culm wall. It is influenced by the position of vascular bundles and by the position of the fibre within the vascular bundle. Murphy and Alvin (1997b) have charted the maturation in developing bamboo culms based on observations of some bamboo species. Fig. 99 summarizes the measurements by Liese and Weiner of fibre wall thickness across the culm wall. Generally, fibres at the outer culm wall have thicker and more lamellated walls than those in the inner part. Cell wall thickening of bundle sheath fibres starts from the inner vascular side and proceeds to the outer parenchyma side. Whereas at the outer culm part the fibres near the protoxylem have thicker walls than those near the phloem, the reverse is true near the lacuna with thicker cell walls in the fibre sheath between ground parenchyma and phloem. Fibre maturation can be a process prolonged over many growing seasons. Any investigation of changes
Fig. 99: Fibre wall thickness across the culm wall for the fibre sheath between ground parenchyma (GP) and phloem (PH), as well as between protoxylem (PX) and ground parenchyma (GP), 1 cortex, 9 = tear lacuna - Phyllostachys viridiglaucescens
FIBRE WALL THICKNESS (µm)
11 10 9 8 7 6 5 4 3 2 1 0 1 2 3456 7 8 910 FIBRE SHEATHS OVER CROSS CULM WALL
fibre sheath GP-PH fibre sheath PX-GP
131 structural modifications
related to ageing must also recognize the distinct pattern of differences in the structure of the fibre wall and the location in the culm wall.
Nomura (1993) observed the ageing of Phyllostachys heterocycla culms from seedling stage through the first five years and found an increase off about 6.7 times in the diameter of the vascular bundle fibre sheath, 4.4 times in the metaxylem vessels and about 1.9 times in fibre length.
Detailed measurements of changes with age were undertaken with culms of Phyllostachys viridiglaucescens aged up to 12 years by Liese and Weiner (1996, 1997). During the first month of the growing period, most fibres were still unlignified — e.g., at the 20th
Fig. 100: Fibre wall thickness over the transverse section at the 20th internode from 3-month, 1-year, 6-year and 11-year-old culms of Phyllostachys viridiglaucescens
FIBRE WALL THICKNESS (µm)
8
6
4
2
0 epidermis middle lacuna FIBRE SHEATHS OVER CROSS SECTION
1994 1993 1987 1983
132 the anatomy of bamboo culms
101
10µm
Fig. 101: Fibre cells with three lamellae and without starch in a 1-year-old culm of Phyllostachys viridiglaucescens
internode they had a very thin cell wall of only 1.5-1.7 µm, which increased to 2.3 µm in the fully elongated culm. At this stage, the fibre wall consisted of three lignified lamellae.
Fig 100 shows, at the 20th internode, the cell wall thickness of fibre sheaths in 3-month, 1-year, 6-year and 11-year-old culms across the culm wall, near the epidermis, in the middle and near the lacuna. The 11-year old culm has thicker fibre walls than the 7-year old one. Such increases have been measured both in lower and in upper internodes.
The increase in wall thickness is not caused by the thickening of the existing cell wall, but by the deposition of additional lamellae. Whereas the wall of a 1-year-old culm
133 102
10µm
Fig. 102: Fibre cells with eight lamellae in a 12-year-old culm of Phyllostachys viridiglaucescens
103
5µm
Fig. 103: Wall thickening owing to the development of septation in a fibre in Dendrocalamus latiflorus
134 the anatomy of bamboo culms
shows three lamellae at the base (2.6 µm), a culm of 12 years has about eight lamellae (about 8 µm) (Figs. 101, 102). The polylamellation of fibre walls is especially high in cells adjacent to the ground tissue parenchyma and somewhat less pronounced in fibres near the vascular tissue as also shown by Murphy and Alvin (1997a). The multi- layered texture of different lamellae has already been described earlier. The addition of new wall layers during ageing is also reflected in the development of septated fibres (Fig. 103).
These results demonstrate that cell wall thickening of fibres occurs not only during the maturation period but also in later years. Corresponding observations on developmental changes of fibre structures owing to ageing were also made on palms (Calamus axillaris, Rhapis excelsa) by Weiner et al. (1996).
In a similar way, thickening of parenchyma cell walls was noted in older culms (Alvin and Murphy 1988) up to 3 years. In Figs. 104 and 105, the parenchyma tissue of an 1- year-old and a 12-year-old culm is shown, with distinct thickened cell walls in the latter one. The polylamellate parenchyma wall of a 12-year-old culm can be seen in Fig. 106; the fine structural details of the fibrillar orientation of the parenchyma cell wall were shown in Fig. 60. The wall thickening of parenchyma cells requires their state as living cells, which is shown by the storage of starch as energy reserve also in older culms (Fig. 107).
A similar increase of wall thickness in fibre cells necessitates their living state at a higher age. Generally, fibres lose their protoplast soon after their wall differentiation as known from fibres in dicotyledonous woody tissue. In bamboo, however, fibres remain alive for a long period, as demonstrated by their capacity to store starch and to undergo septation (see page 70 & Fig. 107).
The wall thickening may explain the increase in density observed even in older culms. For validating such observations, the presence and variability of the starch content must be considered. The structural changes influence certain culm properties, like their resistance to splitting. Hence, in well-managed Chinese bamboo forests, only culms that are at least 5-6 years old are harvested.
135 structural modifications
104
250µm
Fig. 104: Ground parenchyma with vascular bundle in a 1-year-old culm of Phyllostachys viridiglaucescens
The lignification of Phyllostachys heterocycla during its growth from a sprout to a 14-year-old culm was studied by Itoh (1990), revealing full lignification of the component cells within one growing season. Changes in the chemical composition of the culm wall of P. pubescens up to 7 years have been investigated by Chen et al. (1985).
Older culms show symptoms of senescence which affect their functional efficiency, especially the conductivity of the metaxylem vessels for water and the sieve tubes for the assimilates (Grosser and Liese 1971). It is remarkable that these crucial pathways are formed within a few days during shoot differentiation and have to function for
136 the anatomy of bamboo culms
105
250µm
Fig. 105: Ground parenchyma with vascular bundle in a 12 - year - old culm of Phyllostachys viridiglaucescens
106
10µm
Fig. 106: Polylamellate parenchyma wall of a 12-year-old culm of Phyllostachys viridiglaucescens
137 structural modifications
107
50µm
Fig. 107: Starch granulae in the parenchyma of a 12-year-old culm of Phyllostachys viridiglaucescens
138 the anatomy of bamboo culms
108
30µm
Fig. 108: Metaxylem vessel filled with tyloses in a 12 - year - old culm of Phyllostachys viridiglaucescens
10 years or longer, with no chance of restoration as it is possible for trees using their secondary meristem. However, with increasing age, certain changes occur in the conductive tissue. Tyloses develop within the metaxylem vessels from the surrounding parenchyma cells and block water conduction (Fig. 108). Slime/gum-like substances contribute to the sealing effect. The sieve tubes are blocked by slime and tylosoids, whereas the companion cells become sclerified (Fig. 109). The combination of such age-related changes leads to breakdown of the transport systems, resulting in the dying of individual culms within a clump or grove.
139 109
50µm
Fig. 109: Ageing effects at the phloem, blockage of sieve tubes and sclerification of companion cells in Phyllostachys edulis
The flowering of culms is mostly followed by their widespread death. Dying culms become brittle and often bend down and break. Since this phenomenon is not associated with fungal degradation, it must be the result of some chemical/structural changes. Despite the importance of this event in terms of culm utilization, the process is still inadequately understood. A significant reduction in specific gravity and strength properties of flowered culms of Phyllostachys heterocycla var. pubescens has been observed by Kitamura (1975). It may be noted that complete exhaustion of starch precedes the flowering of culms.
SITE CONDITIONS AND FERTILIZATION
There are only a few observations on the possible influence of site conditions and fertilizers on the anatomical characters of a culm. Ueda (1960) reported that the quality of bamboo was influenced by environmental factors, especially soil conditions. It is reported that the Sundanese of West Java prefer the culms of Gigantochloa pseudoarundinacea harvested from slopes to those from valleys. A critical analysis by Soeprayito et al. (1990) revealed a higher specific gravity, and bending and tensile strengths in the former, but no significant differences in the anatomical parameters,
140 the anatomy of bamboo culms
such as fibre length, wall thickness and frequency of the fibrovascular bundles in relation to the habitat. According to Gnanaharan (1994), Dendrocalamus strictus from moist areas in Kerala (India) has longer internodes, larger diameter and poorer modulus of rupture (MOR) and modulus of elasticity (MOE) than culms from dry areas which are much stronger although culm length, internode length and diameter are lower. Possible changes in the anatomical structure are not mentioned. Abd. Latif (1995) compared the anatomical and technical properties of Bambusa vulgaris and Gigantochloa scortechinii using 1 920 culms from four sites. While all morphological characteristics (except culm wall thickness) differed significantly with site, the variations in anatomical properties were insignificant, except for a minor effect on fibre width and fibre lumen diameter, which were partly associated with culm wall diameter. He concluded from this extensive study that both bamboo species possess stable anatomical characteristics which do not change with topographical and environmental factors. Average product quality varied insignificantly among sample products from the four study sites.
Fertilization of bamboo stands is being increasingly practised, not only for shoot production but also for culm wood. As a consequence of fertilization, the number of shoots increases, and the yield, diameter and height of culms may increase. However, the anatomical composition remains apparently unchanged and hence, the main technical properties also. This is unlike wood, wherein an increased cambial growth owing to fertilization influences the properties of softwoods and ringporous hardwoods. Investigations by Azmy (1996) on the effects of various rates of felling intensity and fertilizer regimes on Gigantochloa scortechinii showed that felling intensity and fertilizer application have an impact on shoot production, although the timing of fertilizer application is significant. Studies by Lakshmana (1994) and Shanmughavel and Francis (1996) showed that fertilizer application resulted in higher culm production and larger culm size in Bambusa bambos.
Wound Effects
A bamboo plant may experience damage to its culm and rhizome which seriously affects its functioning. As said earlier, the vegetative propagation method that employs culm,
141 structural modifications
branch or rhizome cuttings leads to considerable injuries. Another wounding practise is the cutting of the upper part of a newly planted bamboo to reduce transpiration. In China, even fully grown culms are thus cut to prevent snow breakage. Mechanical damage resulting from harvesting cannot also be excluded. Although the culm wall appears rather resistant to physical damage owing to its hard, siliceous cortex, insects may penetrate the wall to breed in the lacuna (Singh 1990; Kovacs and Azare 1995). Rhizomes of monopodial bamboos are frequently cut to restrict their further expansion and this results in a large wound surface.
In each case of wounding, the water and sap transport system in the vascular bundles is interrupted. As a monocotyledon plant, bamboo lacks a secondary meristem and cannot develop new cellular pathways or a callus for wound closure to prevent further expansion of vessel embolism. Nevertheless, the damaging influence of the invading air needs to be effectively blocked to protect the exposed tissue. Even branches must have such defence mechanisms to seal off a dying part from the remaining culm. Whereas such defence reactions of dicotyledonous trees have been investigated intensively during recent years (Shigo 1984; Blanchette and Briggs 1992; Liese and Dujesiefken 1996; Schmitt and Liese 1995), information on the wound reactions of monocotyledons is meagre, especially for bamboo. Wound responses of the rhizome of banana were reported by Van der Molen (1977) and of the stem of the Royal palm (Roystonea regia) by Weiner and Liese (1995). However, the defence reactions of bamboo have not been investigated until very recently, although they appear to be of special significance — the culm growth is completed within only a few months, but the transport system has to function for a decade, and even longer, with no chance for a damage repair by new tissue formation.
Some recent studies have investigated, on a larger scale, the reactions to wounding of bamboo culms and rhizome (Ding et al. 1997b; Liese and Weiner 1997; Weiner and Liese 1997). Nearly 400 samples from 11 genera, both monopodial and sympodial, of different ages and climatic origins were wounded using a drill in various seasons. The tissue response was regularly monitored, macroscopically as well as using light and electron microscopy, up to one year. Despite many variations in the experimental design, no significant differences were found in the cellular reaction of different genera and species, but a general sequence of wound responses became obvious.
142 the anatomy of bamboo culms
Macroscopically, a dark narrow discoloration develops after some days around the injury on the outer side of the culm and more extensively on its inner side. This discoloration gradually extends axially on the epidermis as a small stripe, and on the inner side as a discoloration of the pith ring. In some species, the discoloration develops along the whole internode (Fig. 110 - see page 119). Microscopically, as initial cellular reaction, plugging of the sieve plates occurs near the wound surface (Fig. 111 - see page 119). Slime is then formed in the parenchyma cells of the vascular bundles and extruded through the pit membranes into the adjacent metaxylem vessels to fill their lumen (Figs. 112, 113 - see page 120). The metaxylem vessels of some species also develop
112
5µm
Fig. 112: Contact parenchyma secretes slime into adjoining vessel - Phyllostachys viridiglaucescens
143 structural modifications
tyloses (Fig. 114). The protoxylem tracheids, however, show tyloses in all species, and often in the unwounded controls also. The cell walls of the short parenchyma cells become lignified and the longitudinal cells show a brownish discoloration (Fig. 115 - see page 120). Starch often accumulates in the neighbouring parenchyma cells.
About two weeks after wounding, an additional wall layer forms in the vascular bundle parenchyma and the longitudinal parenchyma. The layer shows characteristic lamellation of suberin (Fig. 116). Later on, another layer develops upon this suberin lamella (Fig. 117). The cells still contain wall-attached cytoplasm, indicating their vitality (Fig. 118).
114
100µm
Fig. 114: Tyloses in metaxylem vessel - Phyllostachys edulis
144 the anatomy of bamboo culms
116
1µm
Fig. 116: Suberinization as additional wall layer in parenchyma cell - Sinarundinaria nitida
117
0.5µm
Fig. 117: Formation of additional wall lamellae on top of suberin layer in Phyllostachys viridiglaucescens
145 structural modifications
118
2µm
Fig. 118: Parenchyma cell with wall-attached cytoplasm and plastid - Sinarundinaria nitida
Phenolic compounds are present as a thin layer in the sieve tubes (Fig. 119 - see page 121). From the wound edge towards the inner tissue, a distinct axial gradient of wound reactions becomes obvious: up to 5 mm phenolics, up to 10 mm lignification of cell walls and about 20-25 mm plugs and slime in vessels and sieve tubes. Table 6 summarizes the wound reactions of the cell types.
As part of the study, similar wounds were inflicted also in the nodes. Although the anatomical arrangement of the cells in a node is quite different from that in an internode (see page 91), wounds induce similar cellular responses in the same time scale as in internodes. Wound reactions may not be confined to the wounded node and pass also through the nodes. For instance, a solid culm of Chusquea sp. exhibited a response spread along three internodes.
146 the anatomy of bamboo culms
Table 6: Wound reactions of the various cell types of a bamboo culm (Weiner and Liese 1997)
Cell type Reaction
Sieve tubes closing of sieve plates with “plugs” formation of slime lignification of sieve tubes formation of phenolics Metaxylem vessels formation of slime partial formation of tyloses formation of tyloses Protoxylem tracheids formation of slime formation of tyloses Vascular bundle parenchyma accumulation of starch formation of slime partial formation of phenolics formation of suberin lamella formation of additional wall layers Ground parenchyma - short cells lignification formation of phenolics - long cells accumulation of starch formation of phenolics formation of additional wall layers Fibres fibres with starch formation of septa formation of additional wall layers
The branches of a culm experience a natural wounding when they die sequentially from node to node and finally at their culm base (Fig. 120 - see page 121). Effective protection, especially of the vascular pathways, must develop inside a branch before the dead part breaks off. Observations on dying branches revealed cellular reactions similar to those
147 structural modifications
within the culm: formation of plugs, slime, tyloses, phenolics and lignification. To a distance of about 10-20 cells from the branch separation, the parenchyma cells showed additional lignified wall layers (Fig. 121 - see page122). This extra thickening of parenchyma walls extended over the whole cross section of the branch (Fig. 122 - see page 123). Thus, an efficient pre-sealing of the prospective break-off region was achieved.
The rhizome of monopodial species can also be severely wounded, when it is cut for offset propagation or for restricting the expansion of running bamboos. Ding et al. (1997b) investigated the wound reactions of Phyllostachys edulis. Although the rhizome has some structural differences compared with the culm (see page 101), the cellular wound response was quite similar in both. The phloem was blocked by phenolic substances, and the metaxylem vessels filled with slime (Fig. 123 - see page 124). The walls of sieve tubes and short parenchyma cells near the wound edge became lignified. Around the wounded tissue, a layer of longitudinal parenchyma cells developed additional wall lamellae and separate the uninjured tissue from the injured (Fig. 124 - see page 125). The fibres adjacent to the wounded parenchyma also showed additional wall lamellae. Such demarcation resembled zonation in dying branches, but it was not observed in wounded culms.
Further differences relate to the large intercellular spaces present in rhizomes alongside short parenchyma cells filled with a brown-black substance. Tyloses in metaxylem vessels were not observed in the rhizome, whereas they develop in the culm metaxylem as response to wounding. Wound reactions in rhizomes were always strongly limited to the injured tissue in the axial and lateral directions. Their higher parenchyma and higher moisture contents may be beneficial for strong wound reactions.
Summarizing the various cellular reactions in culms and rhizomes to wounding reveals a time sequence. First, plugs form to close the sieve tubes, and then a suberin layer and finally additional wall layers in parenchyma cells and fibres are formed. Fig. 125 illustrates the various components of the efficient defence system of bamboo culms against wounding.
Bamboo cannot form a barrier zone by developing a callus to close the wound, as can dicotyledons with their secondary meristem. Therefore, the term
148 the anatomy of bamboo culms
“compartmentalization” used by Shigo (1984) for restricting a wound in a tree appears not suitable for the defence system of bamboo. This has been shown also for the Royal palm (Roystonea regia) by Weiner and Liese (1995). Wounded bamboos show a lateral limitation of the wound response, but no defined axial blocking of the wound effects. Their wound reactions resemble a gradual “fading out” of the cellular response.
======➧ additional wall layer ➧ phenolic compounds »»»»»»»»»»»»»»»»»»»»»»»»» ➧ suberin layer
{{{{{{{{{{{{{{{{{{{{{{{{{{ ➧ tyloses
➧ slime ▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲▲ ➧ plugs ▲
1 3 7 10142128 56 360 days
Fig. 125: Sequence of wound reactions in a bamboo culm
149 150 the anatomy of bamboo culms
implicationsChapter Six of structure +132
151 CHAPTER SIX implications of structure
Taxonomy
The taxonomy of bamboos is still far from satisfactory. Often, herbarium specimens represent only flowering branches and are mostly inadequate to represent the species. Beginning with Munro’s monograph on world bamboo (1868) a number of classification systems have been proposed (Dransfield and Widjaja 1995). As flowering is rare, many attempts were made to use anatomical characters for taxonomy. Bamboo leaves were preferred for these studies, since they are easier to prepare and bear considerable diagnostic differences (Brandis 1907; Metcalfe 1956; Luxmi Chauhan et al. 1988; Agrawal and Luxmi Chauhan 1990; Ding and Zhao 1994).
Studies on the micromorphology of leaves show that the morphological features can be observed precisely and in more detail through a combination of light and electron microscopies (Ding 1998). The form and arrangement patterns of the prickles, as well as the presence or absence of macro hairs, are significant for specific identification, but less significant for systematic reconstruction of Phyllostachys.
Although the culm structure may appear rather uniform among taxa, the components show considerable differences of taxonomic value. Many investigations have revealed distinct anatomical differences among various genera, and even within a genus, related to specific structures. However, so far only a few attempts have been made to integrate characteristic anatomical differences into an overall compilation to assist taxonomy. The following section aims to present some results to underline the importance of comparative anatomical studies, and also to stimulate further research work.
Some recent studies were concerned with the overall differences between species, their distinction and taxonomic relations. These include:
Grosser (1971) developed an identification key for 25 species based on detailed anatomical studies of various cell structures.
Alam and Dransfield (1981) studied the taxonomic value of the leaf and culm anatomy of Melocalamus compactiflorus, confirming that Melocalamus is related to Melocanna and Dinochloa.
152 the anatomy of bamboo culms
Wu and Hsieh (1990) presented structural differences of nine introduced bamboos in Taiwan-China with an identification key and analysed (1991) the fine structural characteristics of Dendrocalamus latiflorus and Phyllostachys edulis.
Jiang (1992) proved the value of microstructural details for the characterization and identification of major Chinese bamboos, and Yao and Xu (1992) analysed 71 native Chinese species in this respect.
Sekar and Balasubramanian (1994b) investigated the culm anatomy of Guadua and its systematic position. They further showed (1994c) the unique structures of Melocanna with clear distinction from genera like Bambusa and Ochlandra, and analysed (1994d) the structural diversities in Bambusa vulgaris with taxonomic conclusions.
More specific investigations discussed below relate to individual structures of the culm.
OUTER AND INNER LAYERS
The structures of the outer and the inner culm layers are heterogenous with considerable taxonomic value. The cortex shows remarkable differences between species and genera (Ghosh and Negi 1960; Pattanath and Rao 1969; Wu and Hsieh 1991; Luxmi Chauhan et al. 1992). Rong (1958) described differences in the “rind” structure among six varieties of Phyllostachys heterocycla. Scanning electron microscopic studies by Bisen et al. (1988) on the culm and leaf epidermis of 37 Indian bamboos revealed the leaf epidermal features to be very useful in differentiating genera and species. Leaf epidermis does not show great variation with age and location in contrast to the less reliable culm epidermis, which exhibits variations with height associated with masking encrustations. Detailed studies by Sekar (1992) on the outer cortex and the inner lining cavity have led to a diagnostic key for 15 Indian bamboo species.
GROUND TISSUE
The parenchymatous ground tissue of bamboo species shows some variation in the percentage and pattern of its two cell types (see page 74). Their amounts depend much
153 implications of structure
on the thickness of the culm wall and the form and height of the culm itself. Taxonomically important differences between taxa are not known. Beside the changes related to the position towards fibre sheaths and fibre bundles and the metaxylem vessels, the tissue appears somewhat homogeneous. A closer look, however, reveals a varied appearance and detailed investigations on a possible structural pattern within an internode are still warranted. Imai et al. (1995) have identified for the parenchyma of three Phyllostachys species a barrel type (P. bambusoides), a thick-walled barrel type (P. pubescens) and cylindrical type cells (P. nigra var. henonis).
VASCULAR BUNDLE TYPES
The vascular bundle is the most varied structural component of a bamboo culm. Differences in vascular bundles have been analysed with regard their diagnostic value (Velasquez and Santos 1931; Lee and Chin 1960). Grosser and Liese (1973) discussed at length the designation by Holtum (1956) of bundle types into the natural systematic units based on the structure of the ovary and older classification systems, and revealed a close correlation. A number of further investigations described the bundle types and their variations with diagnostic and taxonomic significance, of which only a few are mentioned here (Grosser 1971; Grosser and Zamuco 1973; Wu and Wang 1976; Wen and Chou 1984; Wu and Leu 1987; Wu and Hsieh 1990, Qiao 1991; Muller 1996; Wei et al. 1998) (see page 26). Also, the detailed investigation by Sekar (1992) on 15 Indian bamboos showed considerable structural differences of diagnostic value. For their identification, a key based on the bundle pattern and the characters of cortex and lining cavity was developed. A preliminary study on bamboo taxonomy based on vegetative characters was presented by Keng and Wen (1991), wherein the vegetative characters were combined with the five types of vascular bundles, resulting in a new key for Chinese bamboo genera.
PHLOEM, PROTOXYLEM AND METAXYLEM
So far, no generic differences have been observed in the structure and arrangement of the phloem and the protoxylem. In certain species, the phloem field is reported to be
154 the anatomy of bamboo culms
larger, and its position at the inner culm side is often inverted. Further investigations have to clarify their taxonomic significance.
In contrast to this, the metaxylem has been found to be of interest because of its importance for water conduction as well as for the sap displacement treatment. Beside the considerable changes in size within a culm, distinct differences between species also exist (see page 60).
FIBRES
Fibres occur as vascular bundle sheaths of different shape and, in certain genera, as additional fibre bundles, thus having a considerable taxonomic value. The individual fibres are of different lengths and diameters, influenced by their position within the culm. Nevertheless, average fibre length and width vary significantly between species (see page 60). Sekar (1992) recognized four types in wall thickness — very thick, thick, thin and very thin — with a varied distribution among the taxa studied. Fibre septation could be of some diagnostic value — e.g., two species of Ochlandra could be separated based on the presence or absence of striated septa. According to Younus-uzzaman (1991), the sclerenchyma sheaths of Sinobambusa tootsik are considerably thicker than those of Arundinaria falconeri, which has smaller and more numerous fibre bundles. However, in general, fibre dimensions are not of much significance in classification. Parameswaran and Liese (1977b) found that the warty layer which covers the inner wall of fibres, vessels and parenchyma (see page 69) has no taxonomic value.
INCLUSIONS
The inclusions — both organic and inorganic — show a certain species relationship, but they are much influenced by site, age and season. The appearance of the epidermis with its apposition of cutin and wax can be species-dependant, but as a taxonomic parameter needs additional anatomical criteria.
The presence of starch not only varies with season, but also shows a relation to species (see page 84). Thus, Bambusa vulgaris is known to be rich in starch, whereas Gigantochloa atter has a lower content.
155 implications of structure
Also, the ash contents — especially the amount of silica — vary between species, influenced by age and site. Bambusa vulgaris has a low silica content while Schizostachyum has a higher content. Although these differences influence certain properties — such as the taste of shoots, and pulping and cutting properties — they have no taxonomic relevance.
NODAL STRUCTURES
Differences exist between the nodes of monopodial and sympodial bamboos and, within these groups, in the variation of fibre length between species. However, no major distinctions of taxonomic interest have been found so far.
SUBTERRANEAN STEM
The subterranean stem (rhizome in monopodial bamboos) reveals considerable structural differences, between and within the monopodial and the sympodial bamboos. The cortex of the former group reveals certain differences, related to the arrangement of the vascular bundles beneath and the occasional presence of air canals. As a result, four types have been identified that can be of value for the taxonomy of individual species within a genus (see page 106).
Similar differences have been recognized in the culm neck of the sympodial bamboos. Anatomic studies by Ding (1998) on 39 specific and 12 infra-specific taxa reveal that the air canal in the cortex of the rhizome is very characteristic and could be used as criterion for dividing the genus Phyllostachys into two sections — Sect. Phyllostachys and Sect. Heteroclada (see page 109). Biological Resistance
Bamboo, in general, is vulnerable to attack by fungi (brown rot, white rot, soft rot) and insects (beetles, termites) as it has a low resistance to such organisms. This, however, is influenced by structural characteristics only to a small extent. The culm tissue does not
156 the anatomy of bamboo culms
contain phytotoxic substances like the heartwood of many tree species. Nevertheless, certain differences in biological resistance seem to exist. It may be noted that most observations to date on the durability of bamboo are based on practical experiences, and only a few tests have been carried out using comparable samples in service.
Sometimes it is reported from local customs that the durability improves when the culms are cut before sunrise. This may refer to the influence of the starch/sugar content, which might be lowered during the night by respiration, leading to a slightly improved resistance against moulds and beetles. However, susceptibility to true wood destroying fungi and termites does not change. Harvesting during the descending moon phase is also considered to increase the durability, but intensive investigations have not shown any proof for this belief.
FUNGI
Fungi attack only bamboo tissues with sufficient moisture content, at least above fibre saturation point (20-22%); air-dry bamboo is protected against fungal degradation. Moisture content may be high in processed culms because they have been either insufficiently seasoned or improperly stored. Water uptake occurs easily through the cut ends with their wide metaxylem vessels and, to a much lesser extent, through the sheath scars at the nodes. Vessel blocking through slime and tyloses can retard moisture penetration, but cannot prevent it. Lateral uptake through the outer waxy epidermis is very little and slightly easier through the inner culm zone.
Regarding the natural resistance of the culm itself, it appears that the bottom part has a higher “durability” than the middle and top parts. The inner part of a culm is attacked faster than the outer one, which is attributed to the higher content of nutritious parenchyma in the inner part. The starch content of the parenchyma cells influences to a larger extent the susceptibility to attacks by fungi, especially blue stain fungi, and beetles. The variation in starch and its significance on beetle susceptibility have been described earlier. Laboratory experiments have revealed that bamboo gets deteriorated faster by white rot and soft rot than by brown rot. Immature culms are degraded faster than mature ones by all fungal types (Liese unpubl.). The degradation of bamboo tissue
157 implications of structure
by soft rot fungi and the influence of the cell wall structure on the pathways of infection have been demonstrated by Sulaiman and Murphy (1994, 1995) and Murphy et al. (1997b).
Results from “graveyard tests” with a number of bamboo species in Dehra Dun, India, showed generally a low durability (Class III) with only slight differences. For example, Bambusa nutans and B. polymorpha are less resistant than Bambusa tulda and Dendrocalamus hamiltonii (Liese 1959, Kumar et al. 1994). Laboratory experiments with a number of Basidiomycetes indicated Dendrocalamus merrillianus as “perishable”, Schizostachyum lima and S. diffusum as “resistant”, Bambusa vulgaris and Gigantochloa aspera5 as “moderately resistant” and Schizostachyum zollingerii as “very resistant” (de Guzman 1978). Possible influences of structure on the natural resistance have not been investigated so far.
BEETLES
Bamboo culms as well as bamboo products are very vulnerable to powder post beetles, mostly Dinoderus brevis, D. celluris, D. minutus and Lyctus spp. The attack, which may start as soon as the culm is felled, is related to the presence of starch in the parenchyma. Bamboos harvested during summer are more rapidly destroyed than those felled after the rainy period as the latter has less starch present. Sulthoni (1996) confirmed the close relation between the traditional harvesting time in East Java (April-May) and the low starch content during this period (see page 87). However, even at the same time of the year considerable differences exist regarding the susceptibility of different species, with Bambusa vulgaris always attacked intensively. In Colombia, this species is much faster deteriorated than the common Guadua angustifolia. The traditional preservation method of soaking fresh culms in muddy water reduces the starch content by bacterial degradation and thus reduces their susceptibility to beetle attack.
Flowered bamboo, with starch completely depleted, has a higher resistance to beetles. It may be noted that standing culms often become brittle, without any obvious sign of biological deterioration.
5 Syn. Dendrocalamus asper
158 the anatomy of bamboo culms
Termites are hardly influenced by the starch content. It is reported that Dendrocalamus longispathus is more resistant to termites than Dendrocalamus strictus (Kumar et al. 1984; Liese 1959, 1981, 1997).
Treatability
The low durability of bamboo in an exposed environment requires preservation with chemicals for certain uses in which long life is required. The efficiency of such a treatment depends much on the penetration of the chemicals into the culm tissue. The anatomical structure bamboo culm, however, makes it difficult to treat effectively as it is more resistant to chemical penetration than wood (Liese 1985, 1997). The penetrability of the preservative is limited by the following anatomical characteristics:
At the cut end of a culm, the metaxylem vessels are the main avenues of penetration and run in a strongly axial direction. They are isolated from each other by parenchyma in the internodes and connected only at the nodal diaphragm (see page 93). The vessels are very small at the periphery of a culm wall and become larger in the middle and inner part. The vessel lumina on a transversal section amount to only 5-8%, in comparison with 70% lumina in softwoods and about 30% for diffuse hardwoods (see page 51).
Access to the vessels gets reduced as soon as the culm is harvested and seasoned. As a wound reaction, slime and tyloses develop from the neighbouring parenchyma cells, and move into the vessels blocking the lumina (see page 142).
There are no radial pathways, like the medullary ray cells in wood, in the culm tissue. The horizontal movement of preservative from the vessels into the neighbouring tissue of parenchyma and fibres is only by diffusion and therefore a slow process.
Radial penetration through the outer culm wall is resisted by the skin with its epidermis and waxy apposition. Also, diffusion from the pith cavity is hindered by the solid sclerenchymatous tissue (see page 19). The diffusion through the culm leaf scars at the nodes is negligible for preservation purposes. A minimal intake is possible at the nodal ridge by the tissue opened by cutting-off the branches.
159 implications of structure
As a consequence of the above factors, preservative treatment by the immersion method has only limited efficiency, and takes a long time owing to the low penetration of the liquid preservative through the outer wall. To improve penetration, scraping off the resistant outer skin as well as the inner wall has been proposed. But this would require a bore to be drilled through the impermeable diaphragm.
A permeability index related to anatomical characters has been compiled from diffusion results with small samples (Wu et al. 1992). Investigations by Wu (1994) have shown a faster diffusion rate longitudinally, and a slightly higher radial rate than tangentially. This may be caused by the greater structural heterogeneity of vessels and fibres in the radial than the tangential direction.
The nodes with their distorted structure (see page 92) may hinder penetration, especially while soaking air-dry culms (Fig. 126 - see page 126) in which axial diffusion may be blocked at the nodal area. The nodal influence on penetration was described also by Younus-uzzaman (1990).
The best treatment can be achieved by using the water-filled vessels of a freshly cut culm as transportation channels. A simple procedure is the butt-end treatment, wherein the bottom part of a fresh bamboo culm with branches and leaves still attached is placed in a barrel containing the preservative. The ongoing transpiration by the leaves leads to an uptake of the preservative.
The most effective treatment is the modified Boucherie process — sap displacement of a fresh culm by applying the preservative at one end under a moderate pressure of about 1 bar. The process was originally invented by Purushotham et al. (1954) and further refined by Liese (1959, 1989, 1990). It is applied now on a practical scale by, the Environmental Bamboo Foundation, Indonesia (EBF 1994), the Costa Rica National Bamboo Project (González and Gutiérrez 1996, Liese, et al. 1998), and in Australia where a description was provided by Cusack (1997). The Figs. 127 and 128 (see pages 126 & 127) show the culm clamp at the lower culm end and the outflowing preservative at the top end. The pressure has to be maintained until nearly all vessels are filled with the preservative and this is influenced by the vessel area of the species (see page 53). The smaller vessels at the outer culm portion will require a longer pressure time. Even
160 the anatomy of bamboo culms
then, only 5-8% vessel volume of the total culm will be treated. For a good protection of the culms, a diffusion of the preservative from the vessels into the surrounding tissue is required.
The culms have to be treated as early as possible after harvest, on the day of their harvest itself. Even then, both ends should be cut up to the next node in order to remove the blockage of vessels that might have already occurred (see page 142). This removal is also required after an intermediate water storage for keeping the culms fresh.
As an early contribution to effects of structure on bamboo treatability, Suzuki (1952) found that water storage of culms prevent the blocking of parenchyma pits. Although this effect on the pits is not likely, his hypothesis could be explained as the formation of slime and tyloses, which block the vessel passage, is prevented in water-stored bamboo. Further experiments with water storage of bamboo culms up to four months showed a considerably higher absorption during the diffusion process and pressure processes (Singh and Tewari 1978), because not only vessel blocking was prevented, but also pits of parenchyma cells were disintegrated by bacterial action, as in the case of pit membranes of tracheids in conifers.
Physical and Mechanical Properties
The physical and mechanical properties of a bamboo culm are strongly correlated with its anatomical structures, as shown by several investigations by Janssen (1981), Espiloy (1992, 1994), Liese (1987b), Widjaja and Risyad (1987), Abd. Latif et al. (1990, 1992), Abd. Latif and Mohd. Zin (1992), Sattar et al. (1991), Abd. Latif and Liese (1995), and many others. Only a few examples are given here; for detailed explanations, the works cited above may be referred to.
To a large extent, the mechanical properties of a culm are determined by the specific gravity, which varies approximately from 0.5 to 0.9 g/cm . Specific gravity depends mainly on the fibre content, and also on fibre diameter and cell wall thickness, and therefore varies considerably within a culm and also between species (Zhou 1981). It increases during the maturation of the culm from 1 to 3 years owing to the thickening
161 implications of structure
of the fibre walls, but slightly also during later years (see page 130). The outer part of the culm with its denser distribution of fibres has a far higher specific gravity than the inner part. This concentration of fibres at the outside is to be seen as, in engineering terms, maximizing the radius by gyration (Cusack 1997).
The specific gravity increases along the culm from bottom to top owing to the thinner culm wall with a higher frequency of vascular bundles. The decreasing thickness of the culm wall has an association with mechanical strength, especially for the inner part which then contains less parenchyma and more fibres. At the base, the bending strength of the outer part is 2-3 times higher than that of the inner part.
Shrinkage is influenced by the stage of fibre maturation and the density of vascular bundles. Older culms are more dimensionally stable than younger ones. The radial and tangential shrinkages decrease with the height of the culm since the top portion has a higher number of vascular bundles and lower initial moisture content. Nodes have a great influence on the culm’s mechanical strength. They show higher specific gravity, a lower volume shrinkage and lower tensile strength than the internodes because of shorter fibres and distorted vascular bundles (see page 91) (Kabir et al. 1996).
Fibre length has a positive correlation with the modulus of elasticity (MOE) and compression strength. The fibre wall thickness correlates positively with compression strength parallel to grain, stress at proportional limit and modulus of elasticity (MOE), but negatively with the modulus of rupture (MOR) (Abd. Latif 1993, 1995). The similar pattern of fibre length and cell wall thickness relationship with mechanical properties is due to the strong correlation between the fibre dimensions (except for the lumen diameter).
The fractural behaviour of a culm is different from that of wood; no spontaneous fracture occurs through the whole culm, the cracks becoming deflected in the direction of fibres and thus reducing the disadvantageous effect at the sites of strength loss.
The fine structure of the fibre wall influences the fractured appearance of bamboo tissue (Fig. 129) (Parameswaran and Liese 1976). Ruptured fibre walls show both the splintering and the cross fracture types. The cross fracture could be classified as having flat, smooth face or ridged face. The flat face fracture occurs predominantly in the
162 the anatomy of bamboo culms
F Pa
F
129
Fig. 129: Tensile fracture of an internode with different breakage modes of fibres (F) and parenchyma (Pa) - Dendrocalamus latiflorus (25X) broad lamellar zones, although the narrow ones also exhibit this type (Fig. 130, 131). The ridged face fracture appears only occasionally, showing a tendency for spiral arrangement of the fibrils (Fig. 132). There is also a separation of the lamellae in tension, the weakest regions corresponding to the narrow lamellae. An intercellular separation
163 130
10µm
Fig. 130: Flat, smooth fracture of fibre walls shearing at the middle lamella region - Dendrocalamus latiflorus
131
10µm
Fig. 131: Flat, smooth fracture of a polylamellate fibre wall with separation between broad lamella in regions of narrow lamella - Dendrocalamus latiflorus
164 the anatomy of bamboo culms
132
10µm
Fig. 132: Ridged type fracture of a polylamellate fibre wall with a tendency for spiral arrangement of broken fibrils - Dendrocalamus latiflorus
occurs only rarely; the exposed wall areas display shear failure in the outermost wall layers, probably in the first lamellar layer adjacent to the middle lamella (Fig. 130).
On the basis of the fine structure of the polylamellate fibre walls combined with the lignified polylamellate parenchyma walls, the extremely high tensile strength of about 3 800 daN/mm² in the peripheral culm region can be explained. The alternating fibrillar orientation in the two lamellae types contributes partially to a nullification of the anisotropic properties, which would otherwise be detrimental. Because of the presence of such polylamellate fibres at the statically efficient sites, the peripheral culm zone presents a highly reinforced area (Parameswaran and Liese 1981).
165 implications of structure
Processing
In recent decades, a large number of investigations have dealt with the anatomical, chemical, physical and mechanical properties of bamboo. However, only a few reports were concerned with the ultimate goal — an improved and wider utilization of bamboo products (Liese 1992b, 1996). Some observations were made on aspects such as the influence of certain characteristics on the utilization (Espiloy 1992), and the machining characteristics and the recovery rate of Malaysian bamboo (Abd. Latif 1993, 1995).
The influence of age on processing was often considered a general parameter (Abd. Latif 1993). The recovery rate of four bamboo products — toothpicks, “satay” sticks, skewer and chopsticks — from 1-3 years old culms of two sympodial species showed that cross-cutting, splitting and shaving are easy at a younger age. Finished products, however, are observed to be of low quality, fibrous, easily bent and with considerable shrinkage. This is probably because of the presence of lesser lignified fibres and parenchyma walls, with their low basic density and shear strength values. In order to obtain medium to high quality products and recovery rates of more than 50%, usage (limited to the basal and middle portions only) of at least 2-year-old culms is suggested. Culms of higher age are preferred for bamboo furniture since they show less shrinkage and splitting, the worst danger for a furniture producer besides beetle attack. The high demand for bamboo often results in its premature felling, which reduces the further productivity of the mother plant. Also, the immature culms are prone to splitting, shrinkage, breakage and biological attack.
A few examples may illustrate the many influences of anatomical structure on processing and utilization. A surface decoration is often demanded for indoor use, as in Japan. This improvement of appearance by the application of lac or dyes is hindered considerably by the chemical composition of the culm epidermis, necessitating the use of special treatment processes (Kawamura and Katani 1990). For outdoor use, conservation of the characteristic green colour of the culms is important because the economic value of the products increases with a stable colour. Investigations with Dendrocalamus species have shown that treatment with Boliden K 33 or Thanalith C gives the best green colour-fastness and mildew resistance (Chang 1997). These
166 the anatomy of bamboo culms
preservatives, however, are highly toxic with considerable problems for their final disposal.
Sympodial and monopodial bamboos differ considerably in the structure of their vascular bundle types (see page 26). The basic differences affect a number of properties and, consequently, also the processing. Sympodial bamboos may be less suitable for obtaining a smooth surface, as required for parquet, because the isolated fibre bundles may stick out when sanding the surface. Chopsticks are made mainly from monopodial species with only fibre sheaths, like Phyllostachys. The different fibre arrangement and percentage affect a number of properties, such as density, strength, bending behaviour, shrinkage and splitting. A detailed comparison of all these interrelationships can be of considerable importance for processing qualities, as shown by the fibre morphology studies by Ma et al. (1993) on 26 species of 9 genera and by Zhang et al. (1995) on 34 species of Phyllostachys. An interesting example of practical use of fibre length is the Vietnamese ladies’ hat, which are made from Neohouzeaua dulloa, a species with especially long fibres (up to 4 mm) (Fig. 133 - see page 128).
To overcome certain inherent deficiencies — such as low natural resistance and cracking — bamboo polymer composites have been considered (Lawniczak 1991; Liao and Peng 1992). Monomers do penetrate through the lumina of the cells into the tissue, but penetration of the cell wall is limited to only a few substances with low molecular weight. The results given by timber cannot be expected of bamboo since on a cross section, only 6-8% metaxylem vessels are open for penetration. Therefore, a rather limited application of polymer impregnation appears likely, as far as the stabilization of furniture parts is concerned (Liese 1994). Often bamboo culms have a high level of starch and sugar content in their parenchyma cells and this influences the setting of cement in cement-bonded particle boards. For the intended wider application of cement- bamboo structures a pretreatment and chemical additives have to be considered, besides the seasonal changes of the carbohydrates within the culm (Rahim et al. 1996).
The silica content amounts up to 5% and more depending on species, and affects the cutting and pulping properties. Most silica appears situated in the cortex region, but more knowledge about its location would be useful for processing technologies. Since
167 implications of structure
the ash content is usually associated with the amount of silica, the selection of species with a lower amount is significant for the manufacture of products such as furniture, structural components and skewers (Abd. Razak and Abd. Latif 1994).
The nodal portion of a culm has shorter fibres and lower holocellulose content, but higher content of extractives, pentosans, lignin and ash than the internodal portion. For splitting and matting, the node causes a serious problem of reduced elasticity and uniformity of bamboo strips. Muller (1996) described the techniques of Indonesian craftworkers in splitting long weaving strips (iratan) because of the special structure of the nodal region. Bamboo culms for weaving are chosen from species that are easy to split and are not old enough to become brittle (preferably below two years).
168 the anatomy of bamboo culms
Glossary
air canal: Intercellular canal in the rhizome of certain taxa. anastomosis: Cross-connection of vessels and sieve tubes in the diaphragm. annular thickening: Ring-like thickening of parallely oriented microfibrils in the protoxylem lacuna. callus: Wound-induced tissue formed by the cambium of dicotyledonous trees to overgrow damaged/wounded tissue. companion cell: Sister cell of a sieve tube cell that retains the nucleus and dense cytoplasm. compartmentalization: Cellular reactions to limit the damaging effects of invading air following wounding. conducting tissue: The part of vascular bundle that transports water (metaxylem vessel) and soluble carbohydrates (phloem). cortex: Outer part of a culm (also called bamboo green or rind) or rhizome, between epidermis and ground tissue. culm: The aerial axis emerging from buds of the subterranean system, divided into nodes and internodes. culm sheath: The tubular leaf, inserted at a node and covering part of the culm. diaphragm: A transversal tissue partition of the culm at the node, containing intensive interconnections of vessels and sieve tubes. epidermis: The outermost layer of a culm or rhizome, often with thickened and cutinized outer wall. fibre: Long cells with lignified walls, generally dead, providing mechanical support for the culm as fibre sheath or fibre bundle. fibre, septate: A fibre with transverse walls across the lumen.
169 glossary
fibre bundle: A group of fibres that forms a part of the vascular bundle, and isolated by parenchyma from the metaxylem and phloem. fibre cup: A group of fibres that forms a part of the vascular bundle, attached to the metaxylem, phloem and protoxylem; syn. fibre sheath. fibre sheath: See fibre cup. glomerule: Spindle-shaped cluster of numerous filiform cells arranged in a storied pattern with subunits. ground tissue: The tissue that forms the bamboo culm; consists of parenchyma cells and vascular bundles. hypodermis: The layers beneath the epidermis, comprising thick-walled sclerenchymatous cells. immature: Referring to young culms in which the lignification that strengthens the tissue is not yet complete. inclusions: Depositions of organic or inorganic nature in parenchyma cells, found within the cell wall or as appositions. internode: The part of the culm or rhizome that lies between two nodes. lacuna: Inner space of a hollow culm; syn. pith cavity. lamella: A thin layer as part of the cell wall. leptomorph: Slender, elongated type of rhizome with buds at the nodes, resulting in single-stemmed culms; syn. monopodial. lignification: Formation of a polymer within the cell wall that provides strength to the culm. lining cavity: Inner tissue of a culm at the lacuna. lumen: A hollow space that exists in a single non-living cell or in a hollow stem. maturation: The process of lignification of the cell wall that strengthens the culm. metaphloem: Conducting tissue for the downward transportation of assimilates; consists of sieve tubes and companion cells; syn. phloem. metaxylem: Conducting tissue for the upward transportation of water with minerals; consists of vessels.
170 the anatomy of bamboo culms
microfibril: A threadlike component of the cell wall consisting of cellulose molecules and visible in the electronmicroscope. mitochondria: Ingredients of cytoplasm; act as the “power plant” of the cell. moisture content: The weight of water in the culm, expressed as a percentage of its oven-dry weight. monopodial: See leptomorph. neck: The constricted basal part of the segmented axes of a bamboo plant. node: A segmentation of the culm or rhizome, from where branches or roots originate. At the node, a diaphragm divides the culm. pachymorph: A short, thick rhizome proper, typical of clump-forming bamboos; syn. sympodial. parenchyma: Brick-shaped, generally alive cells with simple pits that store and distribute food materials. perforation plate: The connecting end-wall between two vessel members, originally without any perforation but later developing multiple openings. perforation plate, reticulate: A perforation plate with multiple perforations in a net- like pattern. perforation plate, scalariform: A perforation plate with multiple, elongate and parallel perforations; the remnants of the plate between perforations are called bars. perforation rim: The remnant of a perforation plate that form a border around a simple perforation. phloem: See metaphloem. pit: An opening in the cell wall for interconnection between cells; consists of pit cavity and pit membrane. pit border: The over-arching part of the secondary cell wall. pit membrane: The part of the intercellular layer and primary wall that limits a pit cavity externally. pith cavity: See lacuna. pith ring: Cellular layers surrounding the pith cavity.
171 glossary
plastids: Ingredients of cytoplasm; contain pigments necessary for photosynthesis. pore: A term for the cross-section of a vessel. primary wall: The first-formed cell wall layer with a typical criss-cross orientation of microfibrils. protective layer: A special cytoplasmic layer of a parenchyma cell between the plasma membrane and the pit membrane; involved in the formation of tylosis. protophloem: The first-formed primary phloem cells, formed as part of the vascular bundle. protoxylem: The first-formed primary xylem cells, formed as part of the vascular bundle; characterized by annular thickenings. rhizome: The segmented, complex, subterranean stem system (the “root stock”) of a bamboo plant; present in two basic types — monopodial (leptomorph) and sympodial (pachymorph). Runkel ratio: The indicator for the pulping quality of fibres, expressed as two times the fibre wall thickness divided by the fibre lumen diameter. sclereid: A cell with thick, lignified walls as a strengthening element. sclerenchyma: A tissue composed of sclerenchyma cells. These are variable in form and size, often with thick, lignified secondary walls. secondary wall: Lamellae laid down on top of the primary wall during the differentiation and further ageing of a cell, with the microfibrils in parallel orientation. septum: The transverse wall across the lumen of a fibre. sheath scar: The mark left on the culm after the abscission of a culm sheath. sieve plate: The connection between sieve tubes for the transport of assimilates. sieve tube: The food conducting cell of the phloem arranged as an axial series. silica body: Globular or amorphous conglomeration of siliceous materials of various size and shape; generally present as inclusions in parenchymatous cells. silica cell: An epidermal, short cell filled by a single silica body. stomata: Cell complexes in the epidermis that facilitate air exchange, such as in respiration.
172 the anatomy of bamboo culms
suberin: Fatty substance in a cell wall layer deposited on an earlier layer. sympodial: See pachymorph. tabashir: The siliceous deposit found in the lacuna of certain sympodial bamboo species. terminal layers: Cell layers on the outer and inner parts of the culm, such as the cortex on the outside and the pith ring on the inside. tracheid: An imperforate wood cell of the xylem with connecting pits. tylosis: The outgrowth from a parenchyma cell through the pit cavity into a vessel, leading to its blockage. vascular bundle: Consists of two metaxylem vessels, protoxylem and phloem, surrounded by fibre sheaths and, in sympodial taxa, accompanied by fibre bundles. vessel: Large cells arranged in axial series for water conduction. wall lamella: Lamellar part of the cell wall, laid down during maturation and further ageing. wart: An apposition of the inner cell wall. xylem: The water-conducting tissue in plants, serving also as a supporting tissue, characterized by the presence of tracheary elements.
Note: Some terms have been adapted from Chapman 1997; Dransfield and Widjaja 1995; IAWA 1964; McClure 1966.
173 the anatomy of bamboo culms
References
Abd. Latif, M. 1993. Effects of age and height on the machining properties of Malaysian bamboo. Journal of Tropical Forest Science, 5(4), 528-535. Abd. Latif, M. 1995. Some selected properties of two Malaysian Bamboo species in relation to age, height, site and seasonal variation. Ph.D. thesis, Faculty Forestry, UPM, 281 pp. Abd. Latif, M.; Khoo, K.C.; Jamaludin, K.; Abd. Jalil, H. A. 1994. Fibre morphology and chemical properties of Gigantochloa scortechinii. Journal of Tropical Forest Science, 6, 397-407. Abd. Latif, M.; Khoo, K.C.; Nor Azah, M.A. 1994. Carbohydrates in commercial Malaysian Bamboo. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 227-229. Abd. Latif, M.; Liese, W. 1995. Utilization of bamboo. In Abd. Razak O.; Abd. Latif, M.; W. Liese; Norini, H. ed., Planting and utilization of bamboo in Peninsula Malaysia. FRIM Research Pamphlet No. 118. Forest Research Institute Malaysia, Kuala Lumpur. pp. 50-102. Abd. Latif, M.; Mohd. Zin, J. 1992. Culm characteristics of Bambusa blumeana and Gigantochloa scortechinii and its effects on physical and mechanical properties. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 118-128. Abd. Latif, M.; Wan, T.; Wan, A.; Fauzidah, A. 1990. Anatomical features and mechanical properties of three Malaysian bamboos. Journal of Tropical Forest Science, 2, 227-234. Abd. Razak, M.A.; Abd. Latif, M. 1994. Bambusa heterostachyum II: on the mechanical characteristics and proximate chemical analysis. Bamboo Journal, 12, 74-82.
175 references
Agrawal, S.P.; Luxmi Chauhan. 1990. Culm and leaf epidermis of Indian bamboos, Part III, Melocanna baccifera (Roxb.) Kurz. Indian Forester, 116, 815-818. Alam, M. K.; Dransfield, S. 1981. Anatomy of Melocalamus compactiflorus. Bano Biggyan Patrika, 10, 1-11. Alvin, K.L.; Murphy, R.J. 1988. Variation in fibre and parenchyma wall thickness in culms of the bamboo Sinobambusa tootsik. IAWA Bulletin. n.s., 9, 353-361. André, J.P. 1996. Investigations on the vascular organization of the bamboos (Phyllostachys sp.) by application of the microcasting method. IAWA Journal, 17, 233. Azmy, H. M. 1996. Effects of fertilizing and harvesting intensity on natural stands of Gigantochloa scortechinii. In Ramanuja Rao, I.V.; Widjaja, E. ed., Bamboo, People and the Environment, Vol. 1. Propagation and Management. Proceedings of the Vth International Bamboo Workshop, Ubud, Bali, Indonesia, 19-22 June 1995. International Network for Bamboo and Rattan, New Delhi, India. pp. 86-95. Banik, R.L. 1993. Morphological characters for culm age determination of different bamboos of Bangladesh. Bangladesh Journal of Forest Science, 22, 18-22. Behnke, H.D. 1990. Sieve elements in internodal and nodal anastomoses of the monocotyledon liana Dioscorea. In Behnke, H.D.; Sjolund, R.D. ed., Sieve elements. Springer, Berlin, Germany. pp. 161-178. Beri, R.M.; Karnik, M.G.; Pharasi, S.C. 1967. Note on the Dogwa bamboo wax. Indian Forester, 93, 188-189. Bisen, S.S.; Mishra, G.P.; Sharma, S.C. 1988. Scanning electron microscopic studies on culm and leaf epidermis of Indian bamboos. Indian Forester, 114 (10), 656-669. Blanchette, R.A.; Briggs, A.R. ed. 1992. Defence mechanisms of woody plants against fungi. Springer Series on Wood Science, Springer, Berlin, Germany. 458 pp. Brandis, D. 1907. Remarks on the structure of bamboo leaves. Transactions of Linnaen Society, Series II, 7, 69-92. Braun, H.J. 1957. Die Leitbündelbecken in den Nodien der Dioscoreaceae mit besonderer Berücksichtigung eines neuen Typs assimilatleitender Zellen. Ber. Dtsch. Bot. Ges., 70, 305-322. Chang, S. T. 1997. Comparison of the green colour fastness of Ma bamboo (Dendrocalamus spp.) culms treated with inorganic salts. Journal of Japanese Wood Research Society, 43(6), 487-492.
176 the anatomy of bamboo culms
Chapman, G. ed. 1997. The bamboos. Linnaean Society, London, UK. Academy Press. 368 pp. Chen, Y.; Qin, W.; Li, X.; Gong, J.; Nimanna. 1987. The chemical composition of ten bamboo species. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6- 14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 110-113. Cusack, V. 1997. Bamboo rediscovered. Earth Garden Books, Trentham, Australia. 95 pp. de Bary, A.1877. Anatomie der Vegetationsorgane der Phanerogamen und Farne. Leipzig. de Guzman, E.D. 1978. Resistance of bamboos to decay fungi. Research Terminal Report, PCARR 283, UPLB College of Forestry, Laguna, the Philippines. 9 pp. Dhamodaran, T.K.; Mathew, G.; Gnanaharan, R ; Nair, K.S. 1986. Relationship between starch content and susceptibility to insect borer in the bamboo reed Ochlandra travancorica. Entomology, 11, 215-218. Ding, Y. 1998. Systematic studies on Phyllostachys. Ph.D. thesis, Forestry University, Nanjing, China, 120 pp. Ding, Y.L.; Zhao. 1994. Studies on the comparative anatomy of bamboo leaves and its significance for bamboo systematic taxonomy. Journal of Nanjing Forestry University, 18(3), 1-6. Ding, Y.L.; Grosser, D.; Liese, W.; Hsiung, W.Y. .1993. Anatomical studies on the rhizome of some monopodial bamboos. Chinese Journal of Botany, 5(2), 122-129. Ding, Y.L.; Liese, W. 1995. On the nodal structure of bamboo. Journal of Bamboo Research, 14(1), 24-32. Ding, Y.L.; Liese, W. 1997. Anatomical investigations on the nodes of bamboo. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 265-279. Ding, Y.L.; Tang, G.G.; Chao, C.S. 1996. Anatomical studies on the rhizome of some pachymorph bamboos. In Ramanuja Rao, I.V.; Widjaja, E. ed., Bamboo, people and the environment, Vol. 1, Propagation and management. Proceedings of the Vth International Bamboo Workshop, Ubud, Bali, Indonesia, 19-22 June 1995. International Network for Bamboo and Rattan, New Delhi, India. pp. 121-131. Ding, Y.L.; Tang, G.G.; Chao, C.S. 1997a. Anatomical studies on the culm neck of some pachymorph bamboos. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 285-292.
177 references
Ding, Y.L; Weiner, G.; Liese, W. 1997b. Wound reactions in the rhizome of Phyllostachys edulis. Acta Botanica, 39, 55-58. Dransfield, S.; Widjaja, E.A. ed. 1995. Bamboos. Plant Resources of South-East Asia No. 7. Backhuys Publishers, Leiden, the Netherlands. 189 pp. EBF (Environmental Bamboo Foundation). 1994. Training manual for bamboo preservation using the Boucherie system: trainer’s guide. EBF, Bali, Indonesia. 24 pp. Espiloy, Z. 1987. Physico-mechanical properties and anatomical relationships of some Philippine bamboos. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6-14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 257-264. Espiloy, Z. 1992. Properties affecting bamboo utilization. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 139-142. Espiloy, Z. 1994. Effect of age on the physio-mechanical properties of some Philippine Bamboo. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 180-182. Fujii, T. 1985. Cell-wall structure of the culm of Azumanezaza (Pleioblastus chino Max.). Mokuzai Gakhaishi, 31(11), 865-872. Ghosh, S.S.; Negi, B.S. 1959. Anatomical features of bamboo used for paper manufacture. Cell. Res. Symp. Council of Scientific and Industrial Research, New Delhi, India. pp. 139-148. Ghosh, S.S.; Negi, B.S. 1961. Occurrence, distribution and other characteristics of septate fibres in monocotyledons with special reference to Dendrocalamus strictus Nees. In Proceedings of the 48th Industrial Science Conference, Part III. pp. 277-278. Ghosh, S.S.; Negi, B.S. 1960. Anatomy of Indian bamboos, Part I. Epidermal features of Bambusa arundinacea Willd., B. polymorpha Munro, B. vulgaris Schrad., Dendrocalamus membranaceus Munro, D. strictus Nees and Melocanna bambusoides Trin. Indian Forester, 86(12), 719-727.
178 the anatomy of bamboo culms
Gnanaharan, R. 1994. Physical and strength properties of Dendrocalamus strictus grown in Kerala, India. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 188-192 Gonzales, G.; Gutierez, J. 1997. Bamboo preservation at the Costa Rican National Bamboo Project. In Ganapathy, P.M.; Janssen, J.A.; Sastry, C.B. ed., Bamboo, People and the Environment, Vol. 3, Engineering and Utilization. Proceedings of the Vth International Bamboo Workshop, Ubud, Bali, Indonesia, 19-22 June 1995. International Network for Bamboo and Rattan, New Delhi, India. pp. 121-129. Grosser, D. 1971. Beitrag zur Histologie und Klassifikation asiatischer Bambusarten. Mitt. BFH, Reinbek, 85, 1-321. Grosser, D.; Liese, W. 1971. On the anatomy of Asian bamboos, with special reference to their vascular bundles. Wood Science and Technology, 5, 290-312. Grosser, D.; Liese, W. 1973. Present status and problems of bamboo classification. J. Arnold Arboretum, 54, 293-308. Grosser, D.; Liese, W. 1974. Verteilung der Leitbündel und Zellarten in Sproßachsen verschiedener Bambusarten. Holz Roh-Werkstoff, 32, 473-482. Grosser, D.; Zamuco, G.I. 1973. Anatomy of some bamboo species in the Philippines. The Philippine Journal of Science, 100(1), 57-72. Höster, H.-R.; Liese, W. 1967. Über fransenförmige Gefäßdurchbrechungen in der Sproßachse von Dendrocalamus strictus. Ber. Dt. Bot. Ges., 80(2), 103-108. Holtum, R.E. 1956. The classification of bamboo. Phytomorphology, 6, 73-90. Hsieh, R.S.; Wu, S.C.; Wang, H.H. 1986. : Studies on the tissue structure of leptomorph rhizome and pachymorph rhizome bamboos. Forest Product Industries, 5, 49-61. Hsieh, J.S.; Wu, S.C. 1994. The ultrastructure of Taiwan giant bamboo and moso bamboo. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 199-204. Hsiung, W.Y.; Quiao, S.Y.; Li, Y.F. 1980. The anatomical structure of culms of Phyllostachys pubescens Mazel ex H. de Lehaie. Acta Botanica Sinica, 22(4), 343-348.
179 references
Hsiung, W.Y.; Ding, Z.F.; Li, Y.F. 1980. Intercalary meristem and internodal elongation of bamboo plants. Acta Silva Sinica, 2, 81-89. IAWA (International Association of Wood Anatomists). 1964. Multilingual glossary of terms used in wood anatomy. Mitteilungen der Schweizerischen Anstalt für das Forstliche Versuchswesen, 40 (1), 186 pp. Imai, T.; Majima, S.; Fujita, M.; Saiki, H. 1995. Cellular structures in culm internodes of three Phyllostachys species - Madake, Hachiku and Mosochiku. Bulletin of Kyoto University of Forestry, 67, 147-157. Itoh, T. 1990. Lignification of bamboo (Phyllostachys heterocycla Mitf.) during its growth. Holzforschung, 44, 191-200. Janssen, J.J.A. 1981. The relationship between the mechanical properties and the biological and chemical composition of bamboo. In Higuchi, T. ed., Bamboo production and utilization. Proceedings of the XVII IUFRO World Congress, Kyoto, Japan, 6-17 September 1981. Kyoto University, Kyoto, Japan. pp. 27-32. Jiang, X. 1992. The microstructure of main Chinese bamboos. Chinese Academy of Forestry, Beijing, China. 192 pp. Jiang, X.; Li, Q. 1983. Preliminary observations on vascular bundles of bamboos native to China. Journal of Sichuan Agricultural College, 1, 57-68. Jones, L.P.H.; Milne, A.A.; Sanders, J.V. 1966. Tabashir, an opal of plant origin. Science, 151, 464-466. Joseph, K.V. 1958. Preliminary studies on the seasonal variation in starch content of bamboos in Kerala State and its relation to beetle borer infestation. Journal of Bombay Natural History Society, 55, 221-227. Kabir, M.F.; Bhattacharjee, D.K.; Sattar, M.A. 1996. Physical properties of node and internode of culm and branch of Dendrocalamus hamiltonii. In Ganapathy, P.M.; Janssen, J.A.; Sastry, C.B. ed., Bamboo, people and the environment, Vol. 3, Engineering and utilization. Proceedings of the Vth International Bamboo Workshop, Ubud, Bali, Indonesia, 19-22 June 1995. International Network for Bamboo and Rattan, New Delhi, India. pp. 180-184. Kawamura, N; Katani, K. 1990. The modification of culm epidermis. In Proceedings of the 40th Japanese Timber Society Conference. 4.1.
180 the anatomy of bamboo culms
Kawase, K.; Sato, K.; Imagawa, H.; Ujie, M. 1986. Studies on utilization of Sasa bamboos as forest resources - pulping of young culms and histological change of cell structure of the culms in growing process. Research Bulletin of the College of Forestry, Hokkaido University, Japan. 43(1), 73-98. Keng, P.C.; Wen, T.H. 1991. A preliminary study on bamboo taxonomy based on vegetative characters. In Rao, A.N.; Zhang, X.P.; Zhu, S.L. ed., Selected papers on recent bamboo research in China. Bamboo Information Centre, Beijing, China. pp. 72-83. Kitamura, H. 1962. Studies on the physical properties of bamboos. IX - on the fibre content. Journal of the Japanese Wood Research Society, 8(6), 249-252. Kitamura, H. 1975. Physical and mechanical properties of bloomed bamboo culms - Phyllostachys heterocycla var. pubescens. Bulletin of the College of Agriculture, Utsunomly University, Japan, 9, 37-47. Kovacs, D.; Azarae, I. 1995. Depredations of a bamboo shoot weevil. Nature Malaysiana, 19, 115-122. Kumar, S.; Dobriyal, P.B. 1992. Treatability and flow path studies in bamboo. Part I - Dendrocalamus strictus Nees. Wood and Fibre Science, 24, 113-117. Kumar, S.; Shukla, K.S.; Dev, I.; Dobriyal, P.B. 1994. Bamboo preservation techniques - a review. INBAR Technical Report No. 3. International Network for Bamboo and Rattan, New Delhi, India; Indian Council of Forestry Research and Education, Dehra Dun, India. 59 pp. Lakshmana, A.C. Culm production of Bambusa arundinacea in natural forests of Karnataka, India. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 100-103. Lawniczak, M. 1991. Bamboo-polymer composite new construction material. Lecture delivered at the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. Lee, C.L.; Chin, T.C. 1960. Anatomical studies on some Chinese bamboos. Acta Botanica Sinica, 9(1), 76-95. Lessard, G.; Chouinard, A. ed. 1980. Bamboo research in Asia. Proceedings of a workshop held in Singapore, 28-30 May 1980. International Development Research Centre, Ottawa, Canada. 228 pp.
181 references
Liao, K.F.; Peng, S.F. 1992. Studies on the manufacture of bamboo and plastic combinations II. Bull. Exp. For. Nat. Chung Hsiung University, 4, 26-49. Liese, W. 1959. Bamboo preservation and soft-rot. FAO Report to Government of India, No. 1106, 37 pp. Liese, W. 1980. Anatomy of bamboo. In Lessard, G.; Chouinard, A. ed., Bamboo research in Asia. Proceedings of a workshop held in Singapore, 28-30 May 1980. International Development Research Centre, Ottawa, Canada. pp. 161-164. Liese, W. 1980. Preservation of bamboos. In Lessard, G.; Chouinard, A. ed., Bamboo research in Asia. Proceedings of a workshop held in Singapore, 28-30 May 1980. International Development Research Centre, Ottawa, Canada. pp. 165-172. Liese, W. 1985. Bamboos - biology, silvics, properties, utilization. Dt. Ges. Techn. Zusammenarbeit (GTZ), Nr. 180, TZ Verlagsges., Roßdorf 1, 132 pp. Liese, W. 1987a. Research on bamboo. Wood Science and Technology, 21, 189-209. Liese, W. 1987b. Anatomy and properties of bamboo. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6-14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 196-208. Liese, W. 1989. Progress in bamboo research. Journal of Bamboo Research, Hangzhou, China, 8(2), 2-16. Liese, W. 1989. Bamboo preservation in Costa Rica. UN HABITAT Progress Report No. 1. 37 pp. Liese, W. 1990a. Bamboo preservation in Costa Rica. UN HABITAT Progress Report No. 2. 16 pp. Liese, W. 1990b. Bamboo preservation in Costa Rica. UN HABITAT Progress Report No. 3. 11 pp. Liese, W. (1992a): The structure of bamboo in relation to its properties and utilization. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 95-100.
182 the anatomy of bamboo culms
Liese, W. 1992b. Production and utilization of bamboo and related species in the year 2000. In Proceedings of the IUFRO All Division V Conference on Forest Products, Nancy, 23-28 August 1992, Vol. 2. pp. 743-751. Liese, W. 1994. Biological aspects of bamboo and rattan for quality improvement by polymer impregnation. Folia Forestalia Polonica, Series B, 25, 43-56. Liese, W. 1995. Anatomy and utilization of bamboo. European Bamboo Society Journal, Meise, Belgium, 1, 5-12. Liese, W. 1996. Structural research on bamboo and rattan for their wider utilization. Bamboo Research, 15, 1-14. Liese, W. 1997. The protection of bamboo against deterioration. In Jena, P.K. ed., Proceedings of the 2nd International Conference on Non- Conventional Building Materials (NOCMAT-97), Bhubaneswar, India, 17-19 June 1997. Natural Resources Development Foundation, Bhubaneswar, India. pp. 111-117. Liese, W.; Ding, Y.L. 1991. Structure and functions of the nodes in bamboo. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 213-217. Liese, W.; Dujesiefken, D. 1996. Wound reactions of trees. In Maramoresch, K.; Raychaudhuri, S.P. ed., Forest trees and palms diseases and control. Oxford & IHB Publication Company, New Delhi, India. pp. 20-42. Liese, W.; Gutiérrez, J.; González, G. 1998 Preservation of bamboo for the construction of houses for low income people. Profc. V International Bamboo Congress, San Jose, Costa Rica, in press. Liese, W.; Grosser, D. 1971. Über das Vorkommen von anomalem Gewebe in der Sproßachse von Monokotyledonen. Holz Roh-Werkst, 29, 339-344. Liese, W.; Grosser, D. 1972. Untersuchungen zur Variabilität der Faserlänge bei Bambus. Holzforsch. Holzverwert, 26(6), 202-211. Liese, W.; Grover, P.N. 1961. Untersuchungen über den Wassergehalt von indischen Bambushalmen. Ber. Deut. Bot. Gesellschaft, 74, 105- 117. Liese, W.; Mende, C. 1969. Histometrische Untersuchungen über den Aufbau der Sproßachse zweier Bambusarten. Holzforsch. Holzverwert, 21(9), 113-117
183 references
Liese, W.; Weiner, G. 1996. Ageing of bamboo culms. Wood Science and Technology, 30, 77-89. Liese, W.; Weiner, G. 1997. Modifications of bamboo culm structures due to ageing and wounding. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 313-322. Luxmi Chauhan; Bisen, S.S.; Agrawal, S.P. 1988. Leaf epidermis of Indian bamboos. Part 1 - Dendrocalamus Nees. Indian Forester, 144(10), 684-692. Luxmi Chauhan; Rao, R.V.; Agrawal, P.S.; Dayal, R. 1992. Leaf and culm epidermal studies of Indian bamboo - an appraisal. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 87-94. Ma, I.; Han, H.; Ma, N. 1993. Fibre morphology of some scattered bamboos and main properties of physics and chemistry. Journal of Zhejiang Forestry College, 10, 361-367. Majima, S.; Fujita, M.; Saiki ,H. 1991. The cell wall maturation in moso bamboo (Phyllostachys pubescens Mazel) and the occurrence of peroxidase related to lignification. Bulletin of Kyoto University, Kyoto University Press, 63, 236-244. McClure, F.A. 1925. Some observations on the bamboo of Kwangtung (China). Lingnan Agricultural Review, 3, 40-47. McClure, F.A. 1966. The bamboos - a fresh perspective, Harvard University Press, Cambridge, USA. 347 pp. Metcalfe, C.R. 1956. Some thoughts on the structure of bamboo leaves. Botanical Magazine, Tokyo, Japan, 69, 391-400. Metcalfe, C.R. 1960. Anatomy of the monocotyledons. I. Gramineae - Bambuseae. Oxford University Press, USA. pp. 578-585. Mohiuddin, M.; Alam, M.K. 1992. Pith behaviour in the culm of Dendrocalamus giganteus Munro (Bambusoeides: Poaceae). Bangladesh Journal of Forestry Science, 2, 54-59. Muller, L. 1996. Vascular bundle arrangement, an aid to bamboo recognition. American Bamboo Society Newsletter, 17, 6-9. Muller, L. 1997. Three bamboo passengers on Gondwanan raft “Australis”. American Bamboo Society Newsletter, 18, 4-10.
184 the anatomy of bamboo culms
Munro, C. 1868. A monograph of the Bambusaceae. Transactions of Linnaean Society, London, UK. 26. Murphy, R.J.; Alvin, K.L. 1992. Variation in fibre wall structure of bamboo. IAWA Bulletin, n.s., 13, 403-410. Murphy, R.J; Alvin, K.L. 1997a. Fibre maturation in bamboos. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 293- 303. Murphy, R.J.; Alvin, K.L. 1997b. Fibre maturation in the bamboo Gigantochloa scortechinii. IAWA bulletin, 18, 147-156. Murphy, R.J.; Sulaiman, O.; Alvin K.L. 1997a. Ultrastructural aspects of cell wall organization in bamboos. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 305-312. Murphy, R.J.; Sulaiman, O.; Alvin, K.L. 1997b. Fungal deterioration of bamboo cell walls. In Chapman, G., ed., The bamboos. Linnaean Society, London, UK. pp. 323-332. Nomura, T. 1993. Growth of moso bamboo (Phyllostachys heterocycla (Carr.) Mitford) - structural and dimensional change of bamboo tissue along with ageing of seedling bamboo. Bamboo Journal, 11, 54-62. Ota, M.; Sugi, S. 1953. On the arrangement of the vascular bundle in the bamboo stem. Report of Kyushi University of Forests, 1, 79-86. Parameswaran, N. 1978. Sieve element plastids in bamboo. Experientia, 34, 1015. Parameswaran, N.; Liese, W. 1969. On the formation and fine structure of septate wood fibres of Ribes sanguineum. Wood Science and Technology, 3, 272-286. Parameswaran, N.; Liese, W. 1973. Electron microscopy of multiperforate perforation plates. Holzforschung 27, 181-186. Parameswaran, N.; Liese, W. 1975. On the polylamellate structure of parenchyma wall in Phyllostachys edulis. IAWA Bulletin, 4, 57-58. Parameswaran, N.; Liese, W. 1976. On the fine structure of bamboo fibres. Wood Science and Technology, 10, 231-246. Parameswaran, N.; Liese, W. 1977a. Structure of septate fibres in bamboo. Holzforschung, 31, 55-57. Parameswaran, N.; Liese, W. 1977b. Occurrence of warts in bamboo species. Wood Science and Technology, 11, 313-318.
185 references
Parameswaran, N.; Liese, W. 1978. A note on the fine structure of protoxylem elements in bamboo. IAWA Bulletin, 2/3, 29-32. Parameswaran, N.; Liese, W. 1980. Ultrastructural aspects of bamboo cells. Cellulose Chem. Techn., 14, 587-609. Parameswaran, N.; Liese, W. 1981. The fine structure of bamboo. In Higuchi, T. ed., Bamboo production and utilization. Proceedings of the XVII IUFRO World Congress, Kyoto, Japan, 6-17 September 1981. Kyoto University, Kyoto, Japan. pp. 178-183. Pattanath, P.G. 1972. Trend of variation in the fibre length in bamboos. Indian Forester, 98, 241-243. Pattanath, P.G.; Ramesh Rao, K. 1969. Epidermal and internodal structure of the culm as an aid to identification and classification of bamboos. In Recent advances in the anatomy of tropical seed plants - monograph. pp. 179-196. Plank, H.K.; Hageman, R.H. 1951. Starch and other carbohydrates in relation to the powder-post beetle infestation in freshly harvested bamboo. Journal of Economic Entomology, 44, 73-75. Preston, R.D.; Singh, K. 1950. The fine structure of bamboo fibres. I. Optical properties and x-ray data. Journal of Exp. Botany, 1, 214-226. Purushotham, A.; Sudan, S.K.; Vidya, S. 1954. Preservative treatment of green bamboos under low pneumatic pressures. Indian Forestry Bulletin No. 178, 21 pp. Rong, P.R. 1985. Comparative observation on the anatomical structure of culms of Phyllostachys pubescens and of its several varieties. Journal of Bamboo Research, 4(2), 89-97. Qiao, S. 1991. Observation on vascular bundles of 52 bamboo species. Bamboo Research, 10(1), 84-100. Raechal, L.; Curtis, J.D. 1990. Root anatomy of the bambusoideae (Poaceae). American Journal of Botany, 77(4), 475-482. Rahim, S.; Abd. Latif, M.; Jamaludin, K. 1996. Cement-bonded boards from Bambusa vulgaris. Bangladesh Journal of Forest Science, 25 (1&2), 8-14. Rao, A.N. 1987. Anatomical studies on certain bamboo growing in Singapore. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6-14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 209-226.
186 the anatomy of bamboo culms
Samapuddhi, K. 1959. A preliminary study on the structure and some properties of some Thai bamboos. Royal Forest Department Bulletin No. 30, Bangkok, Thailand. 19 pp. Sattar, M.A.; Kabir, M.F.; Battacharjee, D.K. 1994. Effect of age and height position of muli (Melocanna baccifera) and borak (Bambusa balcooa) bamboos on their physical and mechanical properties. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 183-187. Schmitt, U.; Liese, W. 1995. Zur Feinstruktur von Wundreaktionen im Xylem von Laubbäumen. Drevasky Vyskum, 40, 1-10. Schwendener, S. 1874. Das mechanische Prinzip im anatomischen Bau der Monocotylen, Leipzig. 179 pp., 14 plates. Sekar, T. 1992. Histo-morphological and cyto-spectral studies on culms of some Indian Bambusaceae. Ph.D. thesis, University of Madras, 131 pp. Sekar, T.; Balasubramanian, A. 1994a. Occurrence of a new vascular bundle type in bamboo. Bamboo Information Centre-India Bulletin, 4, 41-42. Sekar, T; Balasubramanian, A. 1994b. Culm anatomy of Guadua and its systematic position. Bamboo Information Centre-India Bulletin, 4: 6-9. Sekar, T.; Balasubramanian, A. 1994c. Culm anatomy of Melocanna baccifera (Roxb.) Kurz. Bamboo Information Centre-India Bulletin, 4: 10-13. Sekar, T.; Balasubramanian, A. 1994d. Structural diversity of culm in Bambusa vulgaris. Journal of Indian Academy of Wood Science, 25(1&2), 25-31. Shanmughavel, P.; Francis, K. 1996. Effect of fertilization on Bambusa bambos plantation. Journal of Non-timber Forest Products, 3 (1&2), 23-25. Shibata, K. 1900. Beiträge zur Wachstumsgeschichte der Bambusgewächse. Journal of College of Science, Imperial University, Tokyo, 8, 427-496. Shigematsu, Y. 1940. On the length and the width of the fibre of kanchiku (Chimonobambusa marmorea). Journal of Japan Forestry Society, 22, 693-696. Shigo, A. 1984. Compartmentalization: a conceptual framework for understanding how trees grow and defend themselves. Annual Review of Phytopathology, 22, 189-214.
187 references
Shor, G. 1992. Tabasheer - The rock in the tree. American Bamboo Society Newsletter, 13, 12-13. Singh, B.; Tewari M.C. 1978. Studies on the effect of ponding on the preservative treatment of bamboos by pressure and diffusion process. Journal of Indian Academy of Wood Science, 9(1), 46-49. Singh, P. 1990. Current status of pests of bamboos in India. In Ramanuja Rao, I.V.; Gnanaharan, R; Sastry, C.B., ed., Bamboos: Current Research. Proceedings of the International Bamboo Workshop, Cochin, India, 14-18 November 1988. Kerala Forest Research Institute, Kerala, India; International Development Research Centre, Ottawa, Canada. pp. 190-194. Strasburger, E. 1891. Über den Bau und die Verrichtungen der Leitungsbahnen. Verl. G.Fischer, Jena. 1000 pp., 5 plates. Sulaiman, O.; Murphy, R. 1994. Soft rot decay in bamboo. Material u. Organismen, 28(3), 167-195. Sulaiman, O.; Murphy, R. 1995. Ultrastructure of soft rot decay in bamboo cell walls. Material u. Organismen, 29(4), 241-253. Suzuki, Y. 1952. Studies on bamboo (VII): Stopping pit pore in bamboo parenchyma cell and its effect on gas and liquid permeability into bamboo. Bulletin of Tokyo University of Forestry, No. 43, 151-156. Soeprayitno, T.; Tobing, T.L.; Widjaja, E. 1990. Why the Sundanese of West Java prefer slope-inhabiting Gigantochloa pseudoarundinacea to those growing in the valley. In Ramanuja Rao, I.V.; Gnanaharan, R; Sastry, C.B., ed., Bamboos: Current Research. Proceedings of the International Bamboo Workshop, Cochin, India, 14-18 November 1988. Kerala Forest Research Institute, Kerala, India; International Development Research Centre, Ottawa, Canada. pp. 215-217. Sulthoni, A. 1987. Traditional preservation of bamboo in Java, Indonesia. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6-14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 349-357. Sulthoni, A. 1996. Shooting period of sympodial bamboo species: an important indicator to manage culm harvesting. In Ramanuja Rao, I.V.; Widjaja, E. ed., Bamboo, People and
188 the anatomy of bamboo culms
the Environment, Vol. 1, Propagation and Management. Proceedings of the Vth International Bamboo Workshop, Ubud, Bali, Indonesia, 19-22 June 1995. International Network for Bamboo and Rattan, New Delhi, India. pp. 96-100. Takenouchi, Y. 1926. On the rhizome of Japanese bamboos. Transactions of Natural History Society, Formosa. 16. Takenouchi, Y. 1931. Systematisch-vergleichende Morphologie und Anatomie der Vegetationsorgane der japanischen Bambus-Arten. Mem. Fac. Sci. Agr., Taihoku Imp. Univ., III (1), 1-60. Tamolang, F.N.; Valbuena, A.B.; Lomibao, A.B.; Artuz, A.E.; Kalaw, C.; Tongacan, A. 1958. Fibre dimensions of certain Philippine broadleaved and coniferous woods, palms and bamboos II .Tappi, 41, 614-621. Tamolang, F.; Lopez, F.; Semana, J.; Casin, R.; Espiloy, Z. 1980. In Lessard, G.; Chouinard, A. ed., Bamboo research in Asia. Proceedings of a workshop held in Singapore, 28-30 May 1980. International Development Research Centre, Ottawa, Canada. pp. 189-200. Tewari, D.N. 1992. A monograph on Bamboo. International Book Distributors, Dehra Dun, India. 498 pp. Tono, T.; Ono, K. 1962. The layered structure and its morphological transformation by acid treatment. Journal of Japanese Wood Research Society, 8, 245-249. Tran, Xvan, Thiep. 1978. Report on forest resources for paper and fibre production in Vietnam. Forest Survey Planning Institute, Hanoi, Vietnam. Ueda, K. 1960. Studies on the physiology of bamboo. Bulletin of Kyoto University of Forestry, No. 30, 167 pp. Valade, I.; Dahlan, Z. 1991. Approaching the underground development of a bamboo with leptomorph rhizomes: Phyllostachys viridis (Young) McClure. Journal of American Bamboo Society, 8(1&2), 23-42. Van der Molen, G.F.; Beckmann, C.H.; Rodehorst, E.:1977: Vascular gelation: a general response phenomenon following infection. Physical and Molecular Plant Pathology, 11, 95-100. Velasquez, G.; Santos, J. 1931. Anatomical study of the culm of five Philippine bamboos. Natural and Applied Science Bulletin, University of the Philippines, 1(4), 281-296.
189 references
Wang, Y.C.; Hsieh, J.S.; Wu, S.C. 1994. Structural variability of vascular bundles of some exotic bamboo species. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 222-226. Wei, X.Z.; He, X.Q.; Hu, Y.X.; Linn, J.X. 1998. The comparative anatomy of four bamboo culms in Guangxi. Journal of Bamboo Research, 17, 23-30. Weiner, G.; Liese, W. 1995. Wound response in the stem of the Royal palm. IAWA Bulletin, n.s., 16, 433-442. Weiner, G.; Liese, W. 1996. Wound reactions in bamboo culms and rhizomes. Journal of Tropical Forest Science, 9, 379-397. Weiner, G.; Liese, W.; Schmitt, U. 1997. Cell wall thickening in fibres of the palms Rapis excelsa (Thunb.) Henry and Calamus axillaris Becc. In Donaldson, L.A; Singh, A.P.; Butterfield, B.G.; Whitehouse, L. ed., Recent advances in wood anatomy. New Zealand Forest Research Institute, Rotorua, New Zealand. pp. 191-197. Wen, T.H.; Chou, W.W. 1984. A report on the anatomy of the vascular bundle of bamboos from China (1), Journal of Bamboo Research, 3, 1-21. Wen, T.H.; He, X.L. 1989. The morphology of fruits and starches in bamboo and their systematic value. Acta Phytotaxonomica Sinica, 27, 365-377. Widjaja, E.A.; Risyad, Z. 1987. Anatomical properties of some bamboo utilized in Indonesia. In Rao, A.N.; Dhanarajan, G.; Sastry, C.B. ed., Recent Research on Bamboo. Proceedings of the International Bamboo Workshop, Hangzhou, China, 6-14 October 1985. Chinese Academy of Forestry, Beijing, China; International Development Research Centre, Ottawa, Canada. pp. 244-246. Wu, S.C. 1994. Movement of water and chemical solution in ma bamboo and moso bamboo. Mem. Coll. Agric. N.T.U., Taipei, 34(2), 111- 123. Wu, S.C.; Hsieh, J.S. 1990. The structural variation of recently introduced bamboos in Taiwan. Forest Products Industry, 9(1), 55-78. Wu, S.C.; Hsieh, J.S. 1991. Anatomical characteristics of Taiwan giant bamboo and moso bamboo. In Bamboo in Asia and the Pacific. Proceedings of the 4th International Bamboo Workshop, Chiangmai, Thailand, 27-30 November 1991. International Development Research Centre, Ottawa, Canada; Forestry Research Support Programme for Asia and the Pacific, Bangkok, Thailand. pp. 193-198.
190 the anatomy of bamboo culms
Wu, S.C.; Hsieh, J.S.; Liou, J.L. 1996. The anatomical properties of some bamboo species grown in mainland China (III). Quart. J. Exp. Forest, NTU, 10(2), 37-59. Wu, S.C.; Leu, J.L. 1987. The ultrastructure of vascular bundles of some Taiwan bamboo species. Quart. J. Exp. For., NTU, 1, 21-44. Wu, S.C.; Leu, J.; Hsieh, J.S. 1992. Relationships between anatomical characteristics and permeability properties in Taiwan grown bamboo species. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 101-111. Wu, S.C.; Wang, H.H. 1976. Studies on the structure of bamboos grown in Taiwan. Bull. Nat. Taiwan Univ., 16, 79 pp. Yao, X.S.; Xu, H. 1992. Anatomical analyses of native bamboo culms in China. In Zhu, S.; Li, W.; Zhang, X.; Wang, Z. ed., Bamboo and its use. Proceedings of the International Symposium on Industrial Use of Bamboo, Beijing, China, 7-11 December 1992. International Tropical Timber Organization; Chinese Academy of Forestry, Beijing, China. pp. 133-138 Younus-uzzaman, M. 1991. Anatomical features of Arundinaria falconeri and Sinobambusa tootsik with reference to their effects on penetration of preservatives. Journal of Forest Science, 20(1&2), 39-44. Zee, S.Y. 1974. Distribution of vascular transfer cells in the culm nodes of bamboo. Canadian Journal of Botany, 52, 345-347. Zhang, J.; Wang, R.; Ma, N.; Zhang, W. 1995. Fibre morphology and main physical and chemical properties of some bamboo wood of Phyllostachys. Forestry Research, 8, 54-61. Zhou, F.C. 1981. Studies on physical and mechanical properties of bamboo woods. Journal of Nanjing Technology College of Forest Products, 2, 1-32.
191 192 the anatomy of bamboo culms
Index a accessory tissue 94 adventitious roots 104, 107 ageing 58, 88, 130 air canal 104, 156 anastomoses 95 annular thickening 42, 105 ash 88, 156,168 b beetles 158 bending strength 140, 162 Boucherie process 160 cbud 104, 107 callus 142 cell wall 43, 62, 68, 75-78, 130-135, 144-149, 162-165 companion cell 22, 54, 105 conducting tissue 24, 51 cork cell 16 cortex 16, 104-107, 142, 153, 156 culm base 104, 107 culm neck 107 culm sheath 12, 130 culm shoot 104 culm stalk 107
193 index
culm wall 12, 22-26, 130-140 cutinization 21 cytoplasm 54, 67, 75, 144 d diaphragm 92, 95 edurability 156-159 elongation 104, 130 epidermis 16-18,88, 104, 143, 153, 166 f fertilization 104-141 fibre 22, 58-73, 95-98, 104-107, 130-136, 161-168 fibre bundle 26-35, 58, 95, 108 fibre length 58-62, 95-98, 167-168 fibre ring 109 fibre sheath (cap) 26-35, 58, 105, 108, 167 fibre strand = (see)fibre bundle fibre wall 62-69, 130-135 fibril 47, 62-68, 162 filiform cell 95 flexibility coefficient 62 gflowering 140 glomerule 95 ground tissue 22-26, 74, 92, 130-139, 153
194 the anatomy of bamboo culms h harvest 87, 130-141 ihypodermis 16, 104 inclusion 83-89, 155 inflorescence 108 internode 11-22, 92, 105-112, 142-149, 162 l lacuna 12-14, 88, 107 lamella 47, 62-69, 130-137 lamellation 16, 62-69, 130-137 lateral bud 103, 107 leptomorph 102 lignification 62-68, 136, 144-147 lignin 46, 62-69, 168 mlumen 50-54, 88, 143, 159 maturation 130-135, 161-162 metabolic residue 88 metaphloem = (see) phloem metaxylem 22-27, 47-54, 92, 95-96, 104-108, 132, 139, 141, 142-149, 154 metaxylem cap (sheath) = (see) fibre sheath microfibril 17, 47, 55, 62-68, 162 mitochondria 55, 82 moisture 83, 157, 160-161 modulus of elasticity 141, 162 modulus of rupture 141, 162 monopodial 28-30, 92, 95-97, 102-107, 156, 167
195 index n nodal ridge 92, 159 pnode 12, 61, 75, 92, 93-99, 104, 146, 160, 162, 168 pachymorph 102 parenchyma 12, 21, 22-35, 47, 65, 74-84, 95-99, 106, 135-138, 144-149, 153, 161 perforation plate 47-50 phloem 8, 34, 54-58, 94-95, 105, 140, 143-149, 154 phloem cap (sheath) 30, 71, 105 pit 42, 69, 75, 95 pit membrane 47, 69, 71, 76, 143, 161 pith cavity 19-22, 104, 159 pith cell 21 pith ring 19-20, 104, 143 plastid 54, 79, 146 polylamellate 43, 47, 62-68, 75, 131-137, 165 preservation 52, 126, 127, 159-160 property anatomical 140, 166 chemical 167 cutting 88, 167 mechanical 161-165 pulping 61, 62, 167 physical 161-165 strength 141 protophloem 54 rprotoxylem 26, 29-30, 42-46, 58, 95-96, 104, 131 resistance biological 156-159 rhizome 102-112, 148 Runkel ratio 62 root 104, 107
196 the anatomy of bamboo culms s sclereid 21 sclerified cell 95 sclerenchyma 14, 17 sclerenchyma sheath = fibre sheath septate fibre 70-73, 135, 147, 155 septum 71-73, 135, 147, 155 sheath scar 12, 109, 130, 157 shoot 42, 62, 92, 103 shrinkage 164 sieve tube 24, 54-56, 95, 105, 147 silica 16, 88, 156, 167 silica cell 16, 104 site condition 140 slenderness ratio 62 spicule 16 starch 43-46, 85-87, 98-99, 135-138, 144-147, 155, 158 stomata 16-17, 104 suberin 144-149 substance inorganic 30, 88, 156 organic 84-87, 155 phenolic 146-149 tsympodial 32-42, 94, 107-112, 156, 167 tabashir 88 taxonomy 105-112, 152-156 tensile strength 62, 140, 162, 167 terminal layer 14 thickening annular 42, 105 wall 72, 131-137, 144-149 tracheal cell 42 tyloses 43, 95, 105, 139, 143-149
197 index v vascular bundle 26-42, 62, 71, 93-95, 104, 136-137, 142, 147, 154 vessel 22-26, 47-54, 77, 85, 94, 104-108, 139, 143-149, 159-161, 167 area 50-54, 159 wsize 50-54 water storage 83, 160 wart 50-52, 68-70 wax 16, 83, 155 xwounding 141-149 xylem 22-26, 42, 47, 94, 108 xylem cap (sheath) 26, 28-30, 71, 105
198 the anatomy of bamboo culms
prof. walter liese - a biographical sketch
199 A BIOGRAPHICAL SKETCH prof. walter liese
If ever Germany was to have an ambassador at large for forestry, and for bamboo in particular, Prof. Walter Liese would eminently qualify for the post. His international assignments have carried him far and wide - from the lowlands of Bangladesh to the high mountains of Chile; from the humid forests of Indonesia and Vietnam to the arid zones of Nigeria and Tanzania; from the near shores of Portugal to the far shores of the Philippines. In his career as a wood biologist and forestry expert, which spans nearly five decades, Prof. Liese has stretched his faculties to their limits to become an institution in himself. Walter Liese was born in Berlin on 31 January 1926, when the Weimar Republic was eight years old and appeared stable and prosperous. His childhood and adolescent years were spent in Eberswalde, a small town south of Berlin where his father was Professor of Forest Botany. By the time he was seven years old, the Weimar Republic had collapsed and Adolf Hitler was in control of Germany. Like all other able-bodied German youth, Walter Liese was also drafted into war service at the age of 18. At the end of the military service, Walter Liese pursued his studies. He chose forestry as his main subject, probably influenced by his childhood images of lush forests near Eberswalde. He studied forestry from 1946 to 1950, first in Freiburg in the Black Forest and then in Hann.Münden at the Forest Faculty, University of Göttingen. In 1951 he graduated and began his career with a one-year study on root physiology at the Forest Research Institute in Düsseldorf. The year 1951 added another dimension in the history of botanical studies in Germany. Although palms and bamboos were botanically known through their earlier descriptions by Linné, all palms were classified as bamboos. Their structural characteristics came to be examined only much later, through the efforts of scientists like Hugo von Mohl (1845), Schwendener (1874), de Bary (1877), Strasburger (1891), Haberlandt (1924), and Solereder and Meyer (1928). Then, for some inexplicable reason, the anatomies of bamboos and palms were much neglected. It was only in 1951 that interest in these areas was revived in earnest. As destiny would have it, the seeds for this revival were sown through a chance meeting under favourable circumstances. In April 1951, Walter Liese had started working as a research scientist at the Forest Research Institute in Lintorf, near Düsseldorf. Dr Franz Erich Eidmann, then Head of
200 the anatomy of bamboo culms
the Institute, kindled Liese’s interest in bamboo. The discussion centred on the suitability of culms as pit props in coal mines. Liese, motivated by Dr Eidmann’s enthusiasm, carried out a series of experiments on the properties of bamboo for its use in mines. Liese also had contacts with Prof. Bodo von Borries of the Institute of Higher Microscopy in Düsseldorf, who was part of the team that developed the electron microscope. The apparatus was still a novelty then and awaiting newer applications. Liese made good use of the transmission electron microscope to study the structure of bamboo, the “new” material, and produced in 1951 the first electron micrographs on the fine structural details of the cell walls of bamboo fibres. This was followed in 1953, while working at the Institute of Forest Botany, University of Freiburg, by another series on structures in the cell walls in bamboo. These achievements brought both the researcher and the research subject into the limelight. Liese’s six-year sojourn (1953-59) at the University of Freiburg, where he had once been a student of forestry, launched his outstanding career as a wood biologist and bamboo scientist. The study of anatomical structure using advanced microscopy and other techniques, which began there as a curiosity that developed out of a chance opportunity, became a life-time’s passion. The latter half of the 1950s marked the beginning of Walter Liese’s presence in the international arena. Before joining the University of Freiburg, he had spent one year working in the wood preservation industry in Mannheim. This experience in wood preservation came of use in 1957, when Liese was contracted by the United Nations Food and Agriculture Organization (FAO) to India to study and propose an impregnation method to preserve bamboo from deterioration, and in 1958 to work on wood preservation in Indonesia. In 1958, barely eight years after his graduation, Liese was already a visiting scientist to the Commonwealth Scientific and Industrial Research Organization (CSIRO) in Melbourne, Australia. In 1962, while working at the Institute of Forest Botany, University of Munich, he was serving as a visiting scientist at the prestigious Harvard University in the United States. Later, when his fame as a top- order forestry expert spread, many other universities - Berkeley University of the United States, Canterbury University of New Zealand, Nanjing Forestry University of China, Universidad Austral of Chile and National University of Taiwan-China — followed suit.
201 prof. walter liese - a biographical sketch
Although his primary vocation as a wood biologist and forest botanist prompted Liese to move to Hamburg in 1963, taking up the position of Professor of Wood Science at Hamburg University, bamboo remained a source of fascination for him. His enthusiasm on the subject attracted several young scientists, and some of them became his research partners. During the Freiburg years, Prof. Liese carried out seminal work on the histometry of the cell elements in various bamboos, with special emphasis on tissue composition. The Munich years also saw several studies being carried out on bamboo, not only on anatomy but also on the permeation properties of bamboo culms.
Prof. Liese’s research on bamboo anatomy peaked during the Hamburg years (1963- 91) though he still continues to work as Professor emeritus. The first stimulus came from his association with Dr Dietger Grosser, who had the aptitude and patience to search for even the most minute details in anatomical studies. Together they presented an impressive array of histological studies on bamboo — the characterization of the four basic vascular bundle structures, and their relation to taxonomical classification; variability of fibre lengths in bamboos; distribution of vascular bundles and the cell types in bamboo culms etc. Prof. Liese’s joint work with Prof. Narayan Parameswaran added a competitive depth to bamboo research. Their initial research covered the fine structure of cell walls, especially of fibres and parenchyma cells. This was followed by studies on the occurrence of warty structures in certain bamboo species, fine structure of protoxylem elements, and ultrastructural aspects of bamboo cells, culms etc. Much of this research remains to date the most important contribution to the subject. In between and after these fruitful joint research associations, Prof. Liese has made several forays on his own and published research papers of excellence.
Although enamoured by the lure of bamboo, Prof. Walter Liese never allowed that to affect his other academic interests — wood biology, wood pathology and wood protection. He has delivered lectures in over 50 countries on these subjects, and has carried out research on a number of related areas such as: wood and bark anatomies; fine structure of wood; wood quality; wound reactions in trees and monocotyledons; micromorphology of wood degradation; physiology and enzymology of wood fungi; and promotion of wood utilization in developing countries. A prolific writer, Prof. Liese has to his credit well over 400 scientific papers (70 of which are on bamboo and
202 the anatomy of bamboo culms
20 on palms, mainly co-authored by Gudrun Weiner). He has also guided 70 diploma students and 35 doctoral students.
Apart from teaching at the Hamburg University, Prof. Liese also served as the Director of the Institute for Wood Biology and Wood Protection, and from time to time as the Executive Director of the Federal Research Centre for Forestry and Forest Products. During the Hamburg years, and after his official retirement in 1991, he lent his expertise to several international and national entities, including: the FAO Advisory Committee on Forestry Education (1966-90); the International Union of Forest Research Organizations (IUFRO — as President during 1977-1981 and in various other capacities from 1968 to 1995); the FAO/IUFRO Committee on Bibliography and Terminology (1964-73); the International Academy of Wood Science (as Fellow in 1966 and as Vice President during 1969-72); EUROSILVA, the European Research Cooperation on Tree Physiology (as Chairman of the Joint Steering Committee during 1988-93 and as Vice Chairman in 1994); Deutsche Gesellschaft für Holzforschung (as Chairman for Wood Protection during 1972-76); the Research Advisory Board of the Forest Research Institute, Malaysia (1989-90); etc.
Prof. Liese was instrumental in getting the International Development Research Centre (IDRC) of Canada interested in bamboo, and played an important part in the creation of the International Network for Bamboo and Rattan (INBAR). He is often referred to as the “grandfather of INBAR”.
During his IUFRO presidency Prof. Liese strongly advocated and spearheaded the involvement of developing countries in the organization, and helped focus IUFRO’s activities more on issues of tropical forestry. He was instrumental in initiating the call for action on tropical forestry, which later developed into the IUFRO Special Programme for Developing Countries. It was also during his presidency that IUFRO turned truly international.
International recognition of Prof. Liese’s expertise in his chosen fields was never found wanting. He was accorded honorary memberships of the Philippine Forest Research Society, Finland Society of Forestry, International Association of Wood Anatomists, Indian Academy of Wood Science, Society of American Foresters,
203 prof. walter liese - a biographical sketch
l’Académie d’Agriculture of France, IUFRO, Chinese Bamboo Association, Academia Italiana di Science Forestate, German Society for Wood Research, Polish Academy of Science and the European Bamboo Society, amongst others. In appreciation of his academic brilliance, Prof. Liese was awarded five honorary doctorates, including ones from the University of Sopron, Hungary; University of Zvolen, Czech Republic; University of Istanbul, Turkey; University of Poznan and University of Ljubljana, Slovenia. He also received numerous medals of merit for his achievements in forestry.
Prof. Liese is very highly regarded in Asian countries, especially China and India, not only for his research contributions but also for helping Asian scientists.
Although he retired from official engagements in 1991, Prof. Liese continues to contribute to the world of forestry with his profound knowledge and extensive experience.
204 Given its unrivalled position in terms of diversity, distribution and uses, coupled with the vital role it plays in the rural economies of several countries around the world, bamboo has emerged in recent years as potentially the most important non-wood forest resource to replace wood in construction and other uses. Concomitantly, the interest being shown in this invaluable natural resource since the 1980s has resulted in the accumulation of a considerable body of information through research on various aspects of bamboos, including the anatomy of the bamboo culm. There is however, no comprehensive publication available on the anatomy of bamboo culm, with the available literature being fragmented, scattered and inadequate. A comprehensive overview of current knowledge on the subject is thus an urgent necessity.
This landmark monograph by renowned wood biologist, forestry expert and bamboo specialist, Prof. Walter Liese, whose innovative work on the study of anatomical structure using advanced microscopy and other techniques has won him wide international acclaim, was written to address this need. It is the first attempt to synthesize information from studies on the subject, many of which have been contributed by Prof. Liese, spread over the past four decades. By identifying gaps in the current anatomical knowledge base of bamboo culm, it is expected to stimulate further research and to act as a prime mover for knowledge generation in the key areas of bamboo anatomy, growth and taxonomy.
210