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Phytomorphology ,

41(1), pp. 67-75, 1997


I ian Institute of Spices Research , P.O. Box 1701,
Calicut 673 012, Kerala, India

The vascul pattern of the underground rhizome of ginger was studied at
different stages f development . The internal structure of rhizome showed an inner
zone and an o er zone, separated by intermediate layers. Collateral vascular
bundles were s tattered and more in inner zone. The xylem vessels are with
scalariform thi ck enings and scalariform perforation plates.
Libriform fibres or
rarely fibre trac ids are met with . The phloem
consisted of sieve tubes companion
cells, parenchy a and fibres . Oil cells and canals are many .
Numerous starch
grains , varying n their shape, size and number ,
were present in both sides of
intermediate zone. The rhizome enlargement is by the activity
of : (i) primary
thickening meri tem and procambial cells in the ground parenchyma , (ii) the
actively dividin g ground parenchyma and (iii ) the secondary thickening meristem
in which the fus orm and ray initials are discrete . The presence of cambium like
layer was an im p ortant feature found in ginger.
Key Words : Ginger Zingiber officinale, fusiform and ray
initials, vascular pattern , rhizome

The developmer
studies on ginger ga
Tomlinson 1956), wl
Zingiberaceae and Shf
pattern of ginger and
ginger. Practically I
morphogenesis of gil
knowledge on the rhi2

41 anatomy of ginger rhizome is not clearly understood . Earlier

P only the general anatomical features (Solereder & Meyer 1930,
le Pillai et al. (1961 ) reported on the root apical organization of
u& Raju (1975) studied the general morphology ,
growth and branching
rmeric. Bell (1980 ) described , the vascular pattern of rhizomatous
p studies were carried out on the developmental morphology and
er rhizome . The present aim is to fill the existing gap in our
Pme structure , development and differentiation.
Material and Methods

Rhizomes of gi er were planted in pots. To study the initial stages of rhizome

development, samples ere collected at 20 day intervals,
cut into small pieces and fixed in
1. Received for publ
2. Department of B
The authors are grat
scheme on ` Developmental
providing facilities and enc

ation : August 19, 1996.

ny, University of Calicut, Kerala, India.
I to the Indian Council of Agricultural Research for the grant of an Ad-hoc research
mphology of rhizome of Ginger and Turmeric, and to Dr. K.V. Peter, Director, for



FAA. The m terials were processed for microtomy according to standard procedure
(Johansen 1940
For microscopical investigations serial sections were cut at 10 m and
stained in safran n-fastgreen for histological studies and PAS for histochentical studies as per
the standard pre Eedure (Krishnamoorthy 1988).
Ginger stet
form the aerial
The rhizon
bearing on the v
scale leaves that
meristems are of
The rhizon
thickness (early
These two regioi
resulting in an it
has more. Vase
inverted and irre
scalariform thick
are highly lignif
consists of sievt
parenchyma and
The young
width of 38.8m
and 3.2m wall t
displays an avers
width of sieve to]
and 80 .25 itm at
gradually increas
The oil cell,
are formed lysigf
changes of oil ca
cells near the iota
cortical layers (F
Rhizome en
early in the devel
young scale leaf 1

t is sympodially branched. The distal end of each sympodium turns erect to

ioot; continued growth of the rhizome is by means of lateral buds.
axis differs from an aerial axis in morphological features and has a direct
scular anatomy. The aerial pseudostem carries leaves. The rhizome bears
rotect the bud apex. From the rhizome, adventitious roots and lateral bud
ginated under appropriate conditions.
is the storage organ in ginger. Transection of a rhizome of 10-15mm
evelopmental stage) shows a cortex and a wide central cylinder (Fig. IA).
s are separated by a thin walled endodermoidal layer or intermediate layer,
ter zone and an outer zone. Both have vascular bundles but the inner zone
lar bundles are collateral, and scattered (Fig. 1B). They are occasionally
ularly distributed nearer the endodermoidal layer (Fig. 1B). Vessels show
ping and scalariform perforation plates (Fig. 1C,D). The libriform fibres
d. Occasionally fibre tracheids are also found (Fig. 2A). The phloem
tubes, five to seven companion cells per sieve-tube member, phloem
tloem fibres (Fig. 2B).
iizome (about 10mm in diameter) shows in cross section an average vessel
and 1.6 m wall thicknesses. The fibres have an average width of 12.8m
sicknesses. At maturity, a rhizome with 20 mm diameter (mature rhizome)
e of 42.32m vessel width and 2.02m wall thicknesses. The length and
e member is 76.82 m and 7.96 m, respectively, in 10 mm thick rhizome
1 8.86 Itm in 20 mm thick rhizome (Table 1). All these dimensions are
d proportionately with the growth of the plant.
and ducts are many in the ground parenchyma (Fig. 4 A,B). The oil canals
tously by the disintegration of an entire cell (Fig. 2C). The dimensional
als are given in Table 1. Large number of starch grains are found in the
mediate layer and in ground parenchyma of the central zone and the inner
irgement in ginger is by the activity of three meristematic zones. Very
pment of rhizome, a zone of meristematic cells is formed at the base of a
rimordium of developing rhizome. These meristematic cells develop into

Fig. 1A-D 1, endodermoidal layer; iz, inner zone; oz, outer zone;
pp, perforation plate; s, starch
grains; vb , vascular bi dle; x, xylem). A. T.s. of ginger rhizome showing a cortex
and a wide central cylinder.
x 40. B ,C. Ts. of rhi me showing endodermoidal layer, vascular bundles
and inverted xylem elements. B. x 200,
C. X 400. D. Section owing scalariform perforation plate and starch grains. x 200.



the primary thickening meristem (PTM) and procambial strands. The meristematic activity
is responsible for the initial width increase of the cortex (Fig. 2D). The second type is the
actively dividing ground parenchyma and the third type is the secondary thickening meristem
(STM) in which fusiform and ray initials are clearly visible (Fig. 3B). STM is observed just
below the endo ermoidal layer.
At a lowe r l evel in the rhizome from the shoot bud apex
, the PTM can be still identified.
The scattered va
scular bundles are developing from the PTM or procambial cells. These
groups of cells n be identifie d b y the
plane of cell division . (Fig . 2D) .
The avers tangential dimensions of procambial cell s o f b
oth young (10 mm) and
(20 mm rhizomes showed only slight variation (Table 1), and they can be identified
by the plane of division and deep stainability (Fig. 2D). The differentiation of procambial
cells into vascular tissue takes place at different stages of rhizome growth (Fig. 3A). This
type of growth as observed in many moncots. However, in ginger we observed a special
meristematic layer along with the intermediate or endodermoidal layer. This is a cambiumlike layer of cells, They are thin walled and consists of two layers (Fig. 3B). In certain loci
where vascular bundles develop, these cells are elongated with tapered ends and appear quite
similar to the fus drm initials
with an average length of 62.34m and 8.12m width in mature
stages (Table 1, Fig. 3C). Between these fusiform initials some cells show transverse
divisions to for ray initials with an average length of 21.36 Ain and 17.32m width in
mature rhizomes (Table 1, Fig.
3C). The presence of a cambium-like layer is an important
feature in rhizome development. From this layer inverted and irregularly distributed groups
of xylem and ph em are formed along the intermediate layer but it was not observed in the
cortex (Fig. 3B). The cells outer and inner to the cambial layers also get filled with starch
grains (Fig. 3D). The present study shows that the cambium-like layer is present in rhizome
of ginger.
Zimmermann & Tomlinson (1972)
concluded that the unique and consistent feature of
monocotyledonou vascular anatomy was the existence of a well developed inner system,
recognized prima i y not only on a developmental basis but also on topographic terms as the
system was wholl y within the central cylinder
. In addition, there was often an outer system,
largely restricted the cortex but not always clearly
demarcated from the inner system. Bell
(1980) pointed ou t
that the tissue at the interface between the cortex and the central cylinder
was of particula interest and in many different monocotyledons it was potentially
meristematic . Thi was apparent in the rhizome
, the intermediate zone becoming a functional
vascular cylinder .
Tomlinson (
1969) reported that this intermediate layer eventually
differentiated as th walled fibres and proposed the term fibrous layer
. The versatility of this
tissue and the var , g ways
in which it may get differentiated has resulted in a multiplicity

Fig. 2A
-D - (c companion cells; fi, fiber; od, oil ducts;
Pc, procambium; P1, phloexm; st ,
sieve tube;
x, xylem). A. Librifor fibre. x 400. B. L.s. of rhizome showing xylem and phloem elements.
showing lysigenous dev pment
200 C. Section
of oil ducts. x 400. D. T.s. of rhizome showing procambial groups. x 200.



Fig. 3A-D - (c, c bium; f, fusiform initials; Pc, procambium ; Ph, phloem;
r, ray initials ; s, starch grains;
x, xylem). A. L.s. of ic ome showing formation of cambium .
B. Long fusiform cells and ' short ray initials cells.
x 800 . C. L.s. of rhizo
showing endodermoidal region with fusiform and my initials . x 800 . D. L.s. showing
starch grains and cambia layers. x 800.




of names given
it by different authors. In the present study we prefer to retain the term
intermediate zon n line with Petersen (1892). Scahacht (1852) and de Bary (1877) used the
term cambial rin and cambial like instead of the
term intermediate zone. Sargent & Arber
(1915) reported
the roots are originating from the intermediate layer so they termed the
layer as root plat
Coetxee (1976) coined the term cambial band besides the intermediate
layer. Kumar (1 3) described that in certain Zingiberaceae
a meristematic , cambium- like
zone was present at separated the outer and inner system. In ginger, the present authors
could observe fu brm and ray initials, but it is not persistent for long, so termed it as
cambium-like lay

c, oil cell; od, oil duct). A . Oil cells. x 400.

B. L.s showing oil canal. x 400.


The earlier
of them pointed o
present authors so
justify the term in
develop by means
and the secondary
differentiates as a c
is caused by the c
the same species G
of primary and sec
girth increase in rh
meristemosity of c
and radial division
orderly divisions
prolonged duration
cambium, xylem an
Transections o
loci of cambium-lik
The enzymes respo
However, more stu

rkers had given many terms to the potentially meristematic layer but none
the essential featur es o f t h

e fusiform and ray initials of cambium. The

ceded in showin g p h otographic evidence of fusif
orm and ray initials to
mediate cambium-like layers.
Tomlinson (1970) reported that in
Dracaen a th e cortical system can
meristematic activity of an intermediate zone. Between the inner system
ssue of Dracaena
lies the `cambium' from which the secondary tissue
ttinuation of the primary . The growt h i
n thickness of rhizome inNutsedge
division and enlargement in the inner and o u t
er zone (Esau 1977). In
ford & Bayer (1995) reported that the rhizome development is by means
Nary thickenin g meri s t em. I
n ginger the present auth ors found that the
ome is by means of the initial divisive activity of the procambial cells,
Ibium-like cells, and of ground parench y ma . Th e f
irst i s by tangential
for a short span, the second for still more prolonged time by way of
fusiform and ray
initials, and third by multiplane cell-divisions for
From the Table I it is clear
that the dimensions of procambium,
phloem have increased proportionately with th e growth.
ginger rhizome showed, that starch accumulates in cells situated near the
tone , x y lem an d p hl oem. These cells ma
y be th e pri mary sink for starch.
ble for the conversion of sugar to starch could be more i n
th ese regions.
es are to be conducted on the biochemical reactions in these cell layers.

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monocotyledone; Bo
Pillai S K, Pillai A & Schi
Acad. Sci. 53 240-2
Sargent A E & Arber A 1


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