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Neolamarckia cadamba
III-4
1Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
Neolamarckia cadamba (Roxb.) boSSeR, 1984
syn.: Nauclea cadamba R, Anthocephalus cadamba (R.) M., Samama
cadamba (R.) K, Anthocephalus chinensis (L.) A. R. ex W., A.
indicus A. R., A. morindaefolius K, Sarcocephalus cadamba (R.) K.
Common
bur-ower
tree,
Burower
tree,
Laran Family: Rubiaceae
Brunei: Bangkal
Indonesia: Jabon
Philippines: Kaatom
Bangkal
India: Kadam
Malaysia: Laran
(Sabah,
Peninsular
Malaysia);
Kelempayan
(Peninsular
Malaysia),
Selimpoh
(Sarawak)
<produktionstitel>Enzyklopädie der Holzgewächse</produktionstitel>
<kurztitel>Neolamarckia cadamba</kurztitel>
<DOI>ehg2017002</DOI>
<lieferung>70</lieferung>
<teilnummer>III</teilnummer>
<kapitelnummer>4</kapitelnummer>
Fig.1: Neolamarckia cadamba. Mature tree at Kota Marudu in the northeast of Sabah state, Borneo, Malaysia.
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Neolamarckia cadamba
III-4
2Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
Neolamarckia cadamba (R.) B or Laran is a de-
ciduous tropical tree species that is naturally and widely
distributed from South Asia, Southeast Asia to Australia
[3]. It has a long history of domestication and cultiva-
tion in its native region due to its usefulness in traditional
medicine and for general-utility lumber. It was also suc-
cessfully introduced into tropical and subtropical areas
outside of its native range. It is fast growing, has good
silvicultural characteristics (straight cylindrical bole), is
able to grow in diverse soil conditions and is free from
serious pests and diseases. The species is increasingly im-
portant for various wood-based industries as smallhold-
ers and plantation concessionaires expand their plantings
in the region.
Distribution
The laran tree is distributed naturally from Sri Lanka,
India, Nepal, Bhutan, Bangladesh, southern China, In-
do-China, throughout the Malesian region, Papua New
Guinea and Australia. It was introduced successfully
to African and Central American countries as orna-
mental and industrial plantation trees. Laran is a pio-
neer species which commonly grows in broad-leaved
primary and secondary forests at altitudes of between
100 and 1000 m a.s. l. on a variety of soil conditions.
It can grow on river banks and in the transitional zone
between swampy, permanently ooded areas and drier
loams, and in areas that are periodically ooded [22, 44,
54]. However, it can also grow in areas with as little as
200 mm of annual rain.
The tree is very light demanding and intolerant to frost
[20]. In Malaysia, commercial and trial plantings have
been established at Kanowit Sarawak, Sandakan dan
Tawau, Sabah and Setul Forest Reserve (F.R.), Batu Leng-
gong F.R., and within the Forest Research Institute of
Malaysia (FRIM). In Sabah, agroforestry trials of inter-
cropping oil palm and Laran were highly recommended
[26]. In Indonesia, cultivation of Laran was established
in west and east Java, south and east Kalimantan, across
Sumatra and Sulawesi, Sumbawa and Irian Jaya [29].
Fig.2: Natural distribution of Neolamarckia cadamba.
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Neolamarckia cadamba
III-4
3Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
Morphology
The large deciduous tree has a broad umbrella-shaped
crown, a straight cylindrical bole [25] and horizontally
spreading branches that are characteristically arranged in
tiers, attened becoming subterete and glabrescent [54].
The tree has a small buttress up to 2 m high, with the
height reaching up to 45 m. The stem diameter at breast
height can reach between 100 and 160 cm.
Leaves and Young Shoots
The leaves are simple, opposite, usually with domatia,
simple sessile or petiolate. The buds are conical, but ra-
phides are absent. The leaf shape is ovate, elliptic or ob-
long-elliptic while the apex is acute to acuminate. The
size on juvenile growth is 50–60 × 15–30 cm, on adult
growth, 15–25 × 7–12 cm [58]. The leaf is dark green,
shiny and glabrous adaxially, and pale yellowish-green,
glabrous to densely puberulent abaxially. The base is shal-
lowly cordate on juvenile growth, rounded or truncate on
adult growth. Veins are distinct underneath, with 8–12
pairs of secondary veins, and apparently without doma-
tia. The stipules are oblong or lanceolate, 12–20 mm long,
covering the buds, caducous, interpetiolar and triangular.
The midrib is prominently ridged below, greenish white;
and the petiole is 20–35 mm long, robust, glabrous and
cylindrical. The leaf blade dries thinly and leathery [25,
54]. In young fertilised trees, the leaves are larger, subor-
dinate at the base and acuminated at the apex, the stipules
are interpetiolar, narrowly triangular and deciduous [44].
Flowers, Fruits and Seeds
The inorescence consists of terminal, solitary, capitate
globose heads and bracteates, with peduncles that are
2–4 cm and rather stout. The owers are subsessile on a
glabrous receptacle, bisexual, monomorphic and actino-
morphic. The owering heads measure 35–45 mm across
the calyces and 40–60 mm across the corollas. The ow-
ers are small and orange or yellow in colour.
The calyx tube is tunnel-shaped with narrow lobes and
pubescent outside; the calyx limb is 5-lobed, and is some-
times spatulate. The corolla is yellow to white in colour,
salverform to funnelform, glabrous outside, and has ve
lanceolate lobes that measure 2.5 mm. The corolla tube is
10 mm in size, gamopetalous, narrow, saucer-shaped and
imbricated in the bud. There are ve stamens inserted in
the upper part of the corolla tube and partially exserted,
with short laments, basixed anthers and exserted style.
TS-Note:
pls check
Fig.3: Leaet of N. cadamba with signs of defoliators attack.
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Neolamarckia cadamba
III-4
4Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
The ovary portion is ellipsoid to obovoid in shape and
measures at 1.5 mm; its limb is 3–4 mm and partially to
deeply lobed; while the lobes are oblong to
spatulate ob-
tuse to a rounded shape. The 2-celled ovary lies in the bas-
al position, and sometimes 2–4 celled
ones are located in
the upper portion. There are several ovules in each cell on
the
axile, with simple or 2-forked placentas attached to
the third top septum. The stigma is spindle-shaped, cylin-
drical or fusiform, at the apex bid and exserted [44, 54].
The pollen grains are monads, radial-symmetric, and tiny.
Laran pollen morphological characters are reported as
the polar (P) axis diameter measured as 9.7 (9.2–10.6)
mm, the equatorial (E) diameter measured as 10.7 (9.9–
11.9) mm, and the shape of the pollen as oblate spheroidal
(P/E 0.93). Three compound apertures are found in each
grain, each comprised of a long ectocolpus, a lolongate
to slightly circular mesoporus and an endoaperture. The
ectocolpus ends are acute, the membrane granular; the
mesoporus is circular, distinct with protruding oncus; the
endoaperture is unknown, and the lumina of microretic-
ulations are irregularly polygonal. The sexine ornamen-
tation is striate-reticulate. The pollen wall ultrastructure
shows that the tectum is indistinct about 0.18–0.38mm
thick; the columellae are indistinct about 0.01–0.05mm
thick, and electron-dense material (lipidic and wider than
the columellae) occasionally occurs between columellae.
Nexine segregates into a foot layer and a very thin endex-
ine while the foot layer is parted from the endexine by a
single white line. The nexine is 0.11–0.18mm thick in
the mesocolpial region and thickens into costae around
the aperture. The intine is approximately 0.02–0.06mm
thick and forms a protruding oncus of ca. 1.6 mm in
diameter. The oncus is composed of a bi-layered ectin-
tine with an electron-dense outer layer and a thick elec-
tron-lucent inner layer; and isolated from the cytoplasm
of the pollen cell by intine material [21].
The fruiting head consists of the central axis, which be-
comes distinctly enlarged and eshy to brous. The ow-
ers are compacted on the thickened axis of the inores-
cence heads but not fused to each other. The owers are
fragrant and attract bees and other insects [10]. The fruit-
ing head is yellowish green, 30–40 mm in diameter, with
peduncles visibly thickened. The fruit is drupaceous, gla-
brous, consisting of ovaries and numerous small owers
[25], divided into four dehiscent valves, shaped as ellipsoid
to cylindrical or obconic, and the size is about 2–2.5 ×
~1 mm. The exocarp is somewhat eshy to membranous
or papery, the endocarp cartilaginous to bony, and the ca-
lyx limb is persistent [54]. The ripened fruits have been
reported to be edible in the raw state. The fruit weigh-
ing 100 g of the edible portion was reported to contain
40.02 mg Fe, 343.7 mg Ca, 191.7 mg Mg, 2.43 mg Zn,
TS-Note:
please check
spatulate
instead of
spatulatel
TS-Note:
please check
insertion of
“ones” (for
ovaries)
TS-Note:
please check
insertion of
“the”
Fig.4: Flower.
Fig.5: Bark of mature N. cadamba.
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Neolamarckia cadamba
III-4
5Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
2.36 mg Cr and 1.34 mg Cu [17]. The seeds are released
when the fruit splits apart when it has reached maturity,
and are easily dispersed by the wind or rain [20, 44].
The fruits can either be dried so that the eshy part can be
removed, or are macerated by soaking to allow the eshy
parts to be separated from the seed. The ne seeds can be
cleaned and graded by winnowing [10]. There are several
seeds, miniscule about 0.5–0.7 mm in size, compressed,
trigonal shaped with membranous testa but not winged;
the endosperm is eshy; and the embryo is small and cy-
lindrical shaped [44, 54]. The seeds mature when the fruit
changed colour to dark brown. Fresh seeds are non-dor-
mant as they germinate rapidly after sowing [5]. The seed-
lings show epigeal germination; the cotyledons are sessile,
herbaceous, acute, green, and the leaves are opposite [44].
Bark and Wood
The bark is grey, smooth and very light in young trees but
becomes dark grey, scabrous and longitudinally some-
times ssured with small ridges, cracked and coarsely
aky when old. Its outer surface is greyish brown and
white while the inner surface is light brown [15, 23, 41,
44]. One of the characteristics is the bark having a spe-
cic own taste and odour. The bark consists of the outer
rhytidome and the inner secondary phloem. The transi-
tion between rhytidome and secondary phloem is grad-
ual such that the line of demarcation between these two
zones is not distinct. The bark is hard and brous. Exfoli-
ation occurs through thick chunks. The bark also consists
of tannins and an astringent principle, which is due to the
presence of an acid similar to cincho-tannic acid [9].
The heartwood is white with a yellow tinge, darkening to
creamy yellow on exposure, and not clearly differentiated
from the sapwood. During hardwood formation, a sig-
nicant increase of specic gravity from near pith to near
bark can be found, which is due to the accumulation of
air in the closed cell system [12, 13, 22]. The growth rings
are moderately distinctive but not conspicuous. They are
usually 2–13 mm wide, delimitated by faint pink bands in
the transverse section. The large vessels are clearly visible,
forming conspicuous straight lines. According to the For-
est Research Centre of the Sabah Forestry Department,
parenchyma and rays are not distinctive to the naked eye.
It was recorded that the vessel proportion increases with
height while the ray proportion, bre diameter and bre
lumen diameter decrease. The bre length vertically was
found to be with no signicant trend [19]. Under the mi-
croscopic study, cell wall substance and vessel and ray
proportions increase from pith to bark while the bre
proportion is decreasing. The length and the wall thick-
ness of the bre increase from pith to bark, while the bre
diameter and the bre lumen diameter are the rst to in-
crease, followed by a decrease. Within-tree variations are
more consistent radially than vertically [19].
At a moisture content of 15%, the wood density is in
the range of 0.29–0.56 g/cm3 [44]. The Forest Research
Centre of the Sabah Forestry Department noted that the
moisture content increases with tree portion. It is due to
the higher percentage of sapwood at the top compared
to the bottom portion. Sapwood has lots of active cells
compared to heartwood, which is composed of more
dormant cells. The top part of the tree is classied by
the fastest growth rate, lower density, and higher cellu-
lose content, thus experiencing higher moisture content.
The timber, which is classied under Light Hardwood in
Malaysia, has a soft ne to medium texture and is light
with a creamy yellow colour of the wood. The wood is
odourless and tasteless.
However, the wood has been rated as non-durable. The
wood seasons with little degradation in the form of loose
knots, slight twist and collapse. Graveyard tests in In-
donesia show that the average life of wood in contact
TS-Note:
please check
change made;
or what are
“domain
cells” in the
original text?
TS-Note:
please check
change made
Fig.6: Fruit of Laran (left) and seeds.
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Neolamarckia cadamba
III-4
6Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
with the ground is less than 1.5years [23]. The wood
is susceptible to termite, Anobium and marine borer at-
tack. Anyhow, it can be treated using either the open tank
system or the vacuum-pressure system. The impregnation
with synthetic resins can increase the density and the
compressive strength. The timber air-dries rapidly, with
little or no degradation. The blue stain can be avoided by
treatment within 48 h or by submerging in water if it is
not to be worked up soon after cutting [22, 44].
Rooting Habit
The tree possesses buttress roots. The size of the root de-
pends on the tree height itself and tends to be greater
in shallow areas. Some well-grown trees of N.cadamba,
which may reach a height of 45 m with a stem diameter
of 100–160 cm, have been reported to have buttressed up
to 2 m high. Furthermore, K etal. [22] state
that the majority of the buttressed trees belong to the rst
storey layer of the mixed tropical forests. However, the
buttressed root for this species, which is a deciduous tree
occurring in the lower hills, is small [8].
Taxonomy
The genus of Neolamarckia is a small group in the Rubia-
ceae family, subfamily: Cinchonoideae. Synonyms are An-
thocephalus chinensis (L.) A. R. ex. W., Antho-
cephalus macrophyllus (R.) H., Nauclea cadamba
(R.), Neolamarckia cadamba (R.) B, Sarco-
cephalus cadamba (R.) K., Anthocephalus indicus
A. R. and Anthocephalus morindaefolius K [22,
55, 57]. There are many local names given which follow
by countries of origin. Common names in Indonesia are
galupai, galupai bengkal, harapean, johan, kalampain,
kelampai, kelempi, kiuna, lampaian, pelapaian, selapaian,
serebunaik (Sumatra); jabon, jabun, hanja, kelampeyan,
kelampaian (Java); ilan, kelampayan, taloh, tawa telan,
tuak, tuneh, tuwak (Kalimantan); bance, pute, loeraa,
pontua, suge manai, sugi manai, pekaung, toa (Sulawe-
si); gumpayan, kelapan, mugawe, sencari (Nusa Tengga-
ra); aparabire, masarambi (Papua) [23, 29]. The common
names of Neolamarckia in other countries are: bangkal,
kaatoan bangkal (Brunei); thkoow (Cambodia); kadam,
cadamba (England); koo-somz, sako, mai sa kho (Laos);
kelempayan, laran, selimpoh (Malaysia); mau-lettan-she,
maukadon, yemau (Myanmar); labula (Papua New Guin-
ea); kaatoan bangkal (Philippines); krathum, krathum-
bok, taku (Thailand); c[aa]y g[as]o, c[af] tom, g[as]o
tr[aws]ng (Vietnam) [22, 23, 44].
Genetics and Breeding
The genetic resources of Laran are secured as they are
widespread and abundant [44] in the natural forest.
Their economic importance saw the establishment of
seed stands in natural settings, trial and commercial plan-
tations. Several seed orchards and seedling seed orchards
have been established in Sabah since 2001, to serve as
seed sources to meet local seed demands [43]. Breeding
programmes are still at the early stages but are expect-
ed to progress as demand for the species increases where
plantation programmes are established. Thailand and In-
dia identied plus trees for ex situ conservation of Laran.
In recent years, molecular markers have been developed
and utilised to study genetic diversity, population structure,
breeding values and heritability of tree traits of Laran stands.
Genetic diversity measured using microsatellite markers in
Laran populations in Sarawak (six natural forests and two
plantations) was lower than in populations from Indone-
sia, with Shannon’s diversity index (I) values from 0.1399
to 0.2354. There was less genetic diversity within the popu-
lation [6, 53]. Genetic diversity of four Laran natural pop-
ulations in Indonesia based on amplied fragment length
polymorphism (AFLP) showed that the genetic variability
was structured according to their origin. Within-popula-
tion genetic diversity of Laran in the study was higher, with
I values of 0.1871 to 0.4894, and Nei’s genetic diversity
(He) values from 0.1489 to 0.3339, respectively. Analysis
of the population structure revealed that the genetic diver-
sity within populations (Hs = 0.2260) represented 73% of
the total genetic diversity (Ht = 0.3098). The proportion to
the total genetic diversity that was attributed to population
differentiation was high (Gst = 0.2707). Laran has small
owers pollinated by small insects, but restricted seed dis-
persal due to heavy fruits, so that the gene ow is relatively
low (Nm = 1.3473) [45]. At the population level, Laran
showed variation, high heritability values and genetic gain
in fruit weight, seedling height, root collar diameter, stur-
diness index, leaf number, leaf length and leaf width [46].
The genotypic variation and coefcients were higher than
the environmental variation and coefcients of Laran fruit
characteristics. These characteristics indicate strong genetic
control, and considerable scope exists for the exploitation
of heritable additive genetic components for future breed-
ing and improvement of Laran.
Genomic approaches have been utilised to explore the
molecular basis of wood formation in Laran. Genera-
tion and analysis of the genomic resources, namely a to-
tal of 6622 high-quality expressed sequence tags (ESTs)
for wood formation in Laran, have been performed via
high-throughput DNA sequencing of cDNA clones de-
rived from developing xylem tissues [16]. According to
TS-Note:
please check
lower-case
letter b
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Neolamarckia cadamba
III-4
7Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
P etal. [38], a total of 2405 ESTs were successfully
annotated with 7753 gene ontology (GO) terms that dis-
tributed among three main GO categories, which were
biological processes (2333), molecular functions (3056)
and cellular components (2364). L etal. [24] derived
EST-simple sequence repeat (SSR) markers from the Laran
EST database (NcdbEST), which were highly polymorphic
and transferable amongst ve cross-genera species. EST-
SSR markers showed that the Laran was predominantly
outcrossing. The NcdbEST is being used as baseline infor-
mation for the characterisation of full-length cDNA and
the genomic sequence of candidate genes, EST-SSR devel-
opment, single-nucleotide polymorphism (SNP) discovery
in candidate genes and association mapping of Laran. The
data can be used to develop genetic markers for the trait of
interest and marker-assisted breeding in Laran [38].
SNPs have been detected from Laran’s partial C4H
(3538 bp) and CAD (2354 bp) DNA sequences and have
then been associated with basic wood density parameters.
Association genetics revealed that 4 and 6 SNPs from C4H
and CAD genes, respectively, were insignicantly associat-
ed with the wood basic density of Laran (p<0.05). Vali-
dated genetic variation identied by the SNP markers will
facilitate the selection of Laran parental lines or seedlings
with optimal quality through the gene-assisted selection
(GAS) approach [47]. Full-length cellulose synthase (CesA)
cDNA was isolated from Laran and characterised in sili-
co, and showed its involvement in catalysing the cellulose
biosynthesis of the secondary cell wall rather than the pri-
mary cell wall. The full-length NcCesA1 DNA can serve as
a good candidate gene for association genetics studies that
lead to GAS in the Laran tree breeding programme [52].
Growth, Development and Yield
The N. cadamba tree is reported as a fast growing spe-
cies [36, 44]. In a 30-year rotation in Indonesia, the stand
attained a mean height of 38 m with an average diame-
ter of 65 cm and produced 350 m3/ha in the nal harvest
[44]. The wood volume of 7 cm in diameter can produce
23 m3/ha per year including thinning production. During
the rst 6–8years, the tree would normally have a faster
growth rate. When reaching the age of 10–15years, its
growth rate decreases. The trees grow best in exploited
and denuded areas, especially in logged-over areas. They
also often grow in secondary forests along riverbanks,
and in the transitional zone between swampy, perma-
nently and periodically ooded areas. The tree is also
considered as a light demanding species [49, 50]. The
seedlings are sensitive to drought, excessive moisture,
grazing and direct sun [40].
The trees can reach a height of 45 m with trunk diam-
eters of 100–160 cm. A study on a small farm in South
Kalimantan shows that trees younger than 5years of age
have a mean diameter ranging from approximately 6.0
to 16.4 cm with a maximum of 25.3 cm [22, 23], while
for trees older than 10years the mean diameter is 18.6
to 42.3 cm. The average height is 22 m and the mean di-
ameter is 40.5 cm. The diameter at breast height (DBH)
of trees younger than 5years old increases rapidly up to
8–18 cm. However, the growth rate is slow after 10years.
The mean height was recorded as 19.6 m for trees young-
er than 10years old, while ranging from 17.3 to 30 m
after 10years [22]. This difference is probably due to the
difference in site quality and owner management.
TS-Note:
please check
change made!!
Or signicant-
ly??
Fig.7: Branch formation. N. cadamba with horizontally arranged spreading branches.
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Neolamarckia cadamba
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8Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
Reproduction, Regeneration,
Propagation and Cultivation
By the age of 5years under plantation conditions, seed
production usually begins. The common owering peri-
od lasts for 2–5months based on the country of loca-
tion. In Indonesia, the owering period starts in April to
August, and the fruit matures in June to August. In Ma-
laysia, the owering period begins in June to September,
and the fruits mature in September to February. When
the fruit has changed its colour to dark brown, the seeds
are considered as mature. The fruits can be harvested by
climbing or by gathering from covers on the ground after
shaking the branches [23].
Some special techniques need to be considered, such as
air-drying, crushing and sieving during the seed prepa-
ration to extract the seeds from the fruits. The fruits are
soaked in open areas until rotten, followed by manual
grounding into a thick slurry, and consequently air-dried
and passed through a series of sieves [22]. This technique
can help improve the germination rate up to 98%. Other
than that, the seed also needs to be stored in dry, airtight
containers to ensure that it can last longer for up to 2years
and up to 6months at ambient temperature. When seeds
are stored properly in cool, airtight containers for about
2.5months, a higher germination rate up to 95% can be
obtained. Since their size is small, the seeds are rst mixed
with ne salt before sowing in seedbeds or paper pots.
Heavy rain and too much water should be prevented to
avoid the problem of damping-off [22]. It is suggested to
place the seedlings in well-ventilated conditions or to use
a mild fungicidal spray. The germination will take place
2–3weeks after sowing. The seedlings can be transplanted
to nursery beds or polythene/plastic bags when they reach
8–12weeks of age. A medium enriched with organic mat-
ter is recommended. After 6–7months, when the seedlings
are about 30–40 cm tall, they are ready to be transplanted
into the eld. Under proper care, seedlings can sometimes
be planted out when they are 10–15 cm tall [23]. The
planting of topped seedlings of about 1 cm in diameter can
give satisfactory results [44]. The planting distance (spac-
ing) is suggested at around 3–4 × 3–4 m. The seedlings are
highly susceptible to weeds. Therefore, the young seedlings
need to be weeded several times during the rst few years
from competing vegetation like climbers and plants caus-
ing shade, until the trees approach the canopy.
TS-Note:
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change made
Fig.8: Seedling.
TS-Note:
How old is the
seedling? How
tall is it in cm?
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Neolamarckia cadamba
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9Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
The replanting is often done twice during the rotation,
preferably during the rainy season after 1month of plant-
ing followed by replacing the dead seedlings at the end of
the second year [22]. Thinning practices and fertilisation
are also required. It was recorded that applying urea fer-
tiliser of about 15 g per plant in a ring around the seed-
lings results in much faster growth [22, 23, 43]. Thinning
practices encourage crown development; they are easier
to perform as long as the trees have straight stems with-
out defects and a small regular crown. Thinning practices
should be done early and frequently, depending on the
site quality. Pruning is unnecessary in N.cadamba plan-
tations as the species shows natural pruning, with dead
branches falling off [44].
Ecology
According to K et al. [22], N. cadamba is a
pioneer species that grows on deep, moist, alluvial sites;
besides, it is often found in secondary forests along river-
banks and the transitional zone of swampy, permanently
ooded and periodically ooded areas. Also, it can some-
times be observed in primary rainforests. The species
can grow in a variety of soil types. The productivity of
a 4-year-old planted stand tends to decrease in mean an-
nual volume as the elevation and distance from the water
source increase. Laran stands in Sabah and Sarawak had
better growth due to higher moisture conditions [2]. How-
ever, N.cadamba is hardly grown on leached and poorly
aerated soils. On the other hand, it can grow well on fer-
tile soils. K etal. [23] stated that for N.cadam-
ba in natural habitats, the maximum temperature varies
from 32 to 42°C while the minimum temperature varies
from 3 to 15.5°C. Besides, the mean annual rainfall for its
growing ranges from 1500 to 5000 mm. However, it can
also grow in dry areas with 200 mm rain per year.
Pathology
N.cadamba is mostly attacked by defoliators [48]; e. g.,
Arthroschista hilaralis (W ) (Lepidoptera) and
Daphnis hypothous (C 1780) (Lepidoptera) [18]
are the most common insect pests on N. cadamba [7].
The plant’s growth and health may be affected due to
defoliation. Defoliation causes a tree to become weak-
ened, and hence it can be easily attacked by other insects,
fungi or diseases, which eventually leads to death [8]. In
Indonesia, not only the defoliators but also the fungus
Gloeosporium anthocephali L & T (Leotiomy-
cetes) attacks N.cadamba and causes dieback and defoli-
ation, partially or entirely. Besides, white grubs also have
been recorded to damage the roots of 1- or 2-year-old
trees planted under the taungya system. Furthermore, in
Sarawak, Endoclita aroura (T 1958) (Lepidop-
tera), a stem borer, has attacked some of the 2-year-old
trees. Pesticides or insecticides can be used to control the
pests or insects; besides, fungicides can be used to protect
the tree from attack by fungi [39]. In addition, biological
control and suitable silvicultural techniques can be used
as pest or disease control measures.
Uses
N.cadamba is a lightweight hardwood with poor dura-
bility and thus cannot be utilised in an outdoor condi-
tion. The grain of the wood is commonly straight, with
ne and even texture. These characteristics result in ef-
cient and practical raw material of timber in the wood
industries. Having no interlocked grain, the timber is easy
to work with hand and machine tools, gives a beautiful
surface, cuts cleanly and is easy to nail. With the chemi-
cal and mechanical properties, it is, therefore, similar to
other light hardwood species with suitability for gener-
al-utility timber [19]. It can be used for light construction
work, beams, rafters, boxes, tea-chests, packing cases,
shuttering, ceiling boards, toys, wooden shoes, chopsticks
and pencils. Also, if the wood is properly seasoned, it is
suitable to be used as furniture materials. Other than this,
the wood log is peeled and applied for both face and core
veneer in plywood. It is also utilised in the manufacturing
of particle board, cement-bonded board and hardboard.
The wood produces a good sulphate pulp that is favour-
able for paper making [4]. This wood is also sometimes
mixed with other long-bred materials to produce low-
to medium-quality paper.
Laran is a medicinal plant traditionally used in the treat-
ment of various illnesses; especially its leaves and bark are
used by some native people [51]. Recently, scientic studies
have been initiated to screen the phytochemical constitu-
ents, explore their pharmacological values and test for the
biological activity of extracts from various parts of Laran
[11]. Previous phytochemical studies of this plant have
led to the isolation and identication of novel indole al-
kaloids. Two novel natural products, aminocadambinesA
and B, that have been reported could be further modied
by amino acids [28]; cadamine, cadambine, isocadambine
[9]; neolamarckinesA and B [42]; spirocadambine (con-
taining glucoalkaloids), dehydraisodihydrocadambine, ni-
trocadambinesA and B [28]; and 3b-isodihydrocadambine
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Neolamarckia cadamba
III-4
10 Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
[37]. A phytochemical screening conducted by Z etal.
[59] identied a total of 26 compounds with the major
chemical constituents of n-hexadecanoic acid (44.8 %),
hexadecanoic acid ethyl ester (17.9%) and the octadeca-
noic acid ethyl ester (11.71%) [60].
Some of the known compounds have been reported to
possess various biological activities and signicant phy-
topharmacological values [59]. The leaves and fruit ex-
tracts of Laran have signicant contents of phenolics
(48.0 mgGAE/gDW), avonoids (103.3 mgE/gDW) and
proanthocyanidins (106.0 mgCE/gDW), and exhibited
strong DPPH radical-scavenging and Fe2+-chelating ac-
tivities similar to standard antioxidants. Furthermore,
the antioxidant properties of the extracts were correlated
with their phenolic contents [14]. The methanolic extract
of Laran leaf had benecial effects in reducing elevated
blood glucose levels of hyperglycaemic mice, showing
maximum anti-hyperglycaemic activity at 400 mg Laran
leaf extract per kilogramme body weight, which was sim-
ilar to that of glibenclamide (10 mg/kg) [11]. The leaf ex-
tract also showed antifungal activity against Aspergillus
fumigatus F. and Candida albicans (C. P. R)
B, which is comparable to that of the standard
drug ketokanazole [37].
The bark of the plant is reported to possess tonic, bitter,
pungent, sweet, acrid, astringent, febrifugal, anti-inam-
matory, digestive, carminative, diuretic, expectorant, con-
stipating and antiemetic properties and is given to treat
fever and inammation of the eyes. Ethanolic extract of
Laran bark in low to high dosage showed antiepileptic
sedative activities in several experimental animal models
of mice and rats [33] and showed evidence of analge-
sic, anti-inammatory and antipyretic activity [31]. The
dried bark can be used to relieve fever and as a tonic,
whereas a leaf extract can serve as a mouth wash [16].
Laran root extract had signicant antimicrobial activity
on gram-positive and gram-negative bacteria, and an-
thelmintic activity in a dose-dependent manner on adult
Indian earthworms Pheretima posthuma (G 1959)
when compared to the reference drug piperazine citrate
as suggested by traditional practitioners [1, 31, 32].
Laran leaves have high forage quality for ruminants and
non-ruminants, comparable to the traditional Leucaena
leococephala and Medicago sativa (alfalfa hay, which is
used as the forage quality index). The nutritive value of
leaves harvested from 6-month-old seedlings of Laran
had 20.9% of crude protein (CP), 20.2% of crude bre
(CF), 2.6% of crude fat (EE), 9.7% of total ash (TA),
90.3% of organic matter, 36.7% of nitrogen-free extract
(NFE) and 0.03 % of mimosine. Laran leaves produce
294.2 kcal/100 g of gross energy and crude protein to
gross energy (P/E ratio). Crude protein, crude fat, raw en-
ergy, P/E ratio, organic matter and total ash in Laran were
higher compared to those of M.sativa. Laran is suitable for
forage due to its high protein and low mimosine contents
and therefore has a high potential for developing cost-ef-
fective livestock feed with high protein content [30, 59].
The species is well known as an ornamental plant [35]
that functions as a shade tree for dipterocarp line plant-
ing. N. cadamba is suitable for reforestation in water-
sheds and eroded areas and also in agroforestry systems
as a supply of timber for future uses [27].
Lastly, the tree is highly regarded religiously and cultural-
ly in India, being sacred to the Lord Krishna. Radha and
Krishna conducted their love play in the hospitable and
sweet-scented shade of the Kadamba tree [50, 56].
Further Information
The species also has promising potential for certain in-
dustries. The alkaloid 3b-isodihydrocadambine from the
extracts of Laran is an excellent eco-friendly corrosion in-
hibitor that increases the resistance of the system, acting
through a mixed type of corrosion inhibition mechanism,
and forms a protective layer over mild steel surfaces [37].
In some places, Laran is a common avenue tree, especial-
ly in India, together with species such as Ficus bengalen-
sis and Alstonia scholaris; it shows tolerance to vehicular
air pollution. Its leaves show less visible leaf injury such
as pigmentation, chlorosis, necrosis and burning effects,
which increased according to the increment in vehicular
load of selected areas [34].
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Neolamarckia cadamba
III-4
11Enzyklopädie der Holzgewächse – 70. Erg. Lfg. 02/17
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The Authors:
Dr. A R M
Dr. W L
M M
J K
Forestry Complex
Faculty of Science and Natural Resource
Universiti Malaysia Sabah
Jalan UMS, 88400 Kota Kinabalu, Sabah
Malaysia
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