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National Herbarium & Botanic Garden
ANOMALOUS SECONDARY GROWTH IN THE STEMS OF THE LIANES MIKANIA CORDATA

(BURM.f.) ROBINS. (COMPOSITAE) AND PAULLINIA PINNATA LINN. (SAPINDACEAE)
Author(s): H. G. Schweikerdt, J. J. A. van der Walt, H. P. van der Schijff and H. G.
Schweickerdt
Source: Kirkia, Vol. 9, No. 1 (1973), pp. 109-138
Published by: National Herbarium & Botanic Garden
Stable URL: http://www.jstor.org/stable/23502205
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ANOMALOUS SECONDARY GROWTH IN THE STEMS OF
THE LIANES MIKANIA CORDATA (BURM.f.) ROBINS.
(COMPOSITAE) AND PA ULLINI A PINNATA LINN.
(SAPINDACEAE)
by
J. J. A. van der Walt, H. P. van der Schijff and

H. G. schweickerdt*
ABSTRACT
The stems of two liane species Mikania cordata (Burm.f.) Robins,

and Paullinia pinnata Linn, which exhibit anomalous thickening were
investigated. The investigation revealed the following:
1. Mikania cordata (Burm.f.) Robins.
Leaf trace bundles and stembound bundles are distinguishable
one from another in the internodes at the stem. When secondary
thickening commences, six leaf trace bundles are discernible, the
differentiation of which eventually results in the splitting of the
vascular cambium into twelve segments. From this stage, secondary
thickening is effected by means of a discontinuous vascular cambium.
2. Paullinia pinnata Linn.
Anomalous secondary thickening in the stem of this species
results in the formation of a compound vascular cylinder. At the
internodes there is a large vascular cylinder with three smaller
peripheral vascular cylinders. In turn, at successive nodes, one of
the three peripheral cylinders becomes incorporated into the central
cylinder, to re-emerge gradually above the node where it is again
recognizable as a distinct cylinder.
INTRODUCTION
The phenomenon of anomalous secondary growth is found in the

stems and roots of members of a large number of plant families.
As shown by de Bary (1884), Schenck (1893) and Haberlandt (1914)
the form of the anomalous secondary growth may be different in
different families. They also point out that anomalous secondary
growth commonly occurs in liane stems.
* This research was supported by the University of Pretoria and the South
African Council for Scientific and Industrial Research. The work partially
fulfilled the requirements for the M.Sc. degree of one of the authors,
J. J. A. van der Walt, at the University of Pretoria.
109
Kirkia [Vol.
In previous publications (van der Walt, Schweickerdt and van

der Schiff, 1969 & 1970) in which anomalous secondary growth in
stems of a number of indigenous liane species is discussed, it was
shown that a particular type of anomalous secondary growth occurs
in both woody and non-woody liane species, and further, that a
particular type of anomalous secondary growth occurs in some
tendril climbers and twiners.
The external morphology of the stems of both Mikania cordata

and Paullinia pinnata shows an apparent cable structure. This study
was undertaken to determine the nature and development of
anomalous secondary growth in the stems of these two liane species.
MATERIAL AND METHODS
For a complete anatomical study of the stem it was necessary to

collect stem segments of different thicknesses, including growing
points. As the stems usually increase in thickness very slowly, it
was only necessary to collect every fifth internode. The material
was fixed in 70 per cent F.A.A. and then dehydrated in a series of
solutions of ethyl alcohol-xylol of increasing strength following the
method described by Johansen (1940). Sections were made with a
rotary microtome and by hand. The microtome sections were
stained with haematoxylin and safranine and the hand sections with
Reactif Genevois. In order to be able to study cell types in three
dimensions, the tissues were macerated in Jeffrey's solution.
The diagrams were drawn freehand to scale.
MIKANIA CORDATA (BURM.f.) ROBINS.

EXTERNAL MORPHOLOGY OF THE STEM
The thickest stems of this twiner were about 25 mm in diameter.

The stems are flexible and on old stems shallow longitudinal grooves
occur; the depth and prominence of these grooves, which are more
marked in the twining stems (Figure 3) than in the non-twining
stems (Figure 2), vary from stem to stem. The flexibility of the
stems is, to a large extent, retained for the whole life span of the
plant. Old stems are covered with a cork layer in which marked
lenticels occur. At every thickened node (Figure 1-2) there are two
opposite leaves.
INTERNAL MORPHOLOGY OF THE STEM
Structure of the stem in the embryonic state

Transverse sections of the stem at this stage of development show
that just behind the apical meristem (Figure 4) tissues are already
110
9] Mikania cordata and Paullinia pinnata
differentiated. The following tissues can be distinguished at this

stage:
a) An epidermis (E) of a single layer of cells with trichomes (Th).
b) Cortex zones consisting of collenchymatous (KK) and
parenchymatous (PK) tissue; also secretory canals (Sek) between
the parenchyma elements, opposite the primary medullary rays (Ms),
which, according to Metcalfe and Chalk (1950), appear to be resin
canals.
c) A stele consisting of pericyclic caps (Pe), with component

elements much smaller than the surrounding cells ; also a single ring
of procambium bundles (Pkb) is present—the procambium bundles
are separated from one another laterally by broad, thin-walled
medullary rays (Ms).
d) The pith (M) consisting of thin-walled elements in between
which secretory canals also occur; the latter are arranged in a ring
and alternate with the protoxylem groups.
Tissue types in a slightly older but relatively young stem (Figures
5-10).
These consist of :
a) An epidermis of ordinary epidermal cells with thickened

cellulose walls (Figures 5-7, E) covered with a cuticle (K)—a
surface view of the epidermis (Figure 8) shows that the ordinary
epidermal cells are elongated; multicellular glands (Figure 5) and
multicellular hairs (Figure 6) are also present.
Stomata occur in the epidermis.
b) A collenchymatous cortex (Figure 7, KK) is adjacent to the
epidermis, but is interrupted beneath the guard cells by chloren
chyma. There is a gradual transition from hypodermal collenchyma
to deeper lying parenchyma (PK). Between the parenchyma cells
are conspicuous air spaces (Int.). The stone cells (St), which develop
on the outside of the pericyclic caps, are of a transitional type as
only the inner tangential walls are thickened and lignified. The resin
canals (Figure 9), which are found in the parenchymatous cortex,
consist of a central canal surrounded by epithelial cells (Ep). No
endodermis was observed.
(c) The stele (Figure 10) shows:

i) Pericyclic caps (Pe), which lie on the outside of the col
lateral vascular bundles, consisting of fibres (Pv) with thickened,
lignified walls in which a number of simple pits (ES) occur, and
ii) Primary phloem consisting of relatively large sieve tubes
(Sv), companion cells (BS) and phloem parenchyma (Fp). The walls
of all these elements are sparsely thickened with cellulose.
Ill
Kirkia [Vol.
-S«k
" "W..-' \ \\
o
o
A IV
Figure 4. Transverse section through the embryological stem
of Mikania cordata
-«
Figure 5.
Gland -I
Figure 6. Hair -I
Figure 7.Epidermis and cortex
2
1=?
Figure 8.
Surface view of epidermis
Figure 9. Resin canal
112
The vascular cambium (Vk) has already given rise to a certain

amount of secondary phloem (SF) and secondary xylem (SX). As
is characteristic of many liane stems, a so-called axial xylem cylinder
(AX) (Schenck, 1893) is formed; this consists partially of primary
and partially of secondary sclerenchymatous tissues. The secondary
axial xylem (SAX) consists largely of xylem fibres (Xv). In Figure 10
the first wood vessels (H) of the peri-axial xylem are already dis
tinguishable. These vessels are surrounded by thickened xylem
parenchyma cells (VXp).
The metaxylem vessels (Mx) are relatively large, with numerous

bordered pits (Hs) in their walls. The protoxylem vessels (Px), some
of which are obliterated, are surrounded by thin-walled proto
parenchyma (Pp). The latter are separated from the thin-walled
pith cells by a sheath of thick-walled elements (MV).
Origin of the cambia and secondary growth
a) Origin of the vascular cambium and the commencement of

secondary growth (Figures 11-12)
For a time, primary growth in thickness takes place, after which a

vascular cambium (Figure 11, Vk) is formed.
The first formed fascicular segments join with the interfascicular

segments to form a complete cambium cylinder.
At first this vascular cambium gives rise, centripetally over its

whole circumference, to thick-walled elements and the axial xylem
cylinder (Figure 12, AX) becomes differentiated. From the beginning
the fascicular segments give rise, on their outer sides, to secondary
phloem (SF), and the interfascicular segments to thin-walled vascular
ray cells (Vs). With the formation of the peri-axial xylem (PEX),
àfi irregularity appears in the activity of the vascular cambium. In
some vascular bundles (Figure 12, a, b, c, d, e and f) much less
secondary xylem is formed and the vessels become smaller but on
the other hand much more secondary phloem is formed.
b) Further secondary growth (Figures 13-15) \

With further secondary growth the vascular bundles mentioned
above a, b, c, d, e and f (Figures 13-14) become better defined, as
a result of the continued irregular activity of the cambium (Vk).
Eventually the cambium cylinder is broken up into segments at
these vascular bundles.
113
Kirkia [Vol.
) / V / \ 'i
Figure 10. Vascular
FigurelO. Vascular bundle
bundle
Figure 11. Transverse section of the stem of Mikania cordata showing

the commencement of secondary growth.
114
Transverse section of the stem

; of Mikania
of Mikania cordata
cordata showin
showing the
formation of periaxial xylem.
Figure 12
Figure 13
Figure 13
Transverse section of stem showing the discontinuous
Transverse
vascular cambiumsection of stem showing the discontinuous
cylinder
vascular cambium cvlinder
115
Kirkia [Vol
.ozm t ° i foJ/V/o W^j o

'O IrA/O Jo °jy °—
J
\Sn°< °H 0°° O °n°7
O
V\^nO(3\P °\ cpoy
A «
ocA~t
o ° ^y^o°0 ^Po
apdofb .cW^^r rair^*..3.
"_"c-"r
\ V e
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Oj
o
. *'$ ^WLHo0^''0"!
/o o//5o/toj H= oQ\\o \y <7\f-s>
SsfJ H ffia? ■
Figure 14.
Transverse section of stem showing the irregular activity of the

vascular cambium.
It will be noticed that the vascular bundles marked a-f are les
developed, and, further, are characterized by the presence of
relatively large amount of secondary phloem (SF); also the secondar
xylem of these vascular bundles (SX) is distinguishable from tha
of the remaining vascular bundles as there is much less of it and th
cells are smaller than in the normally developed bundles. As a resul
of this anomalous secondary thickening, shallow longitudini
grooves arise on the stem (Figure 15). The external morphology t
a certain extent reflects the internal morphology of the stem as th
grooves coincide with the less developed vascular bundles (a-f).
116
rax°°o) //
yo \ ry\J / /?%
/ <H u
4 payor// So o
0„Vk9KQci^
<*o t6W
o q%©
€M//»
PoS^dPO/
Figure 15. Transverse section of stem ofMikania cordata showing

tha Innnitn^tnol ornnv^c
Figure 15. Transverse section of stem ofMikania cordata showing

the longitudinal grooves
c) Leaf trace bundles (Figures 16-21)

After a number of nodes had been studied anatomically, it was
established that the vascular bundles a-f (as shown in Figures 13-15)
represent leaf trace bundles. These leaf trace bundles are a con
tinuation of bundles 4, 8 and 12; and 16, 20 and 24 in the internode
as reproduced in Figure 16. Leaf trace bundles, 4, 8 and 12 form
one complete leaf trace, while leaf trace bundles 16, 20 and 24 form
the complete leaf trace of the opposite leaf. Where the two groups
of leaf trace budles approach the node (Figures 17-19) they diverge
and each runs into one of the opposite leaf bases ; at the nodes the
opposite leaf bases form protrusions, so that the stem becomes
flattened laterally at these places.
The number of vascular bundles continuing in the stem is main
tained by the division of some of the remaining stem bundles. In
Figures 17-19 vascular bundles 13, 15 and 21 have given rise to
bundles 13a, 15a, and 21a respectively. In the following internode
the vascular bundles are again more or less radially symmetrically
117
Kirkia [Vol.
arranged (Figure 21). Six of these bundles, e.g. 22, 1 and 6; and 10,
14 and 18, in turn form the leaf traces for the next pair of opposite
leaves (Figure 21).
W,
I £2*\
c=>*\
1 I?, s1
•€pd.
Figure 16. Internode Figure 17. Beneath the node
ft#
Figure
Figure 18.
18. Just
Just beneath
beneath the
the node
node Figure
Figure 19.
19. Node
Node
0X>}
g%
.^3, W^7 gg
Figure 20. Node Figure 21. Internode

Figures 16-21. Schematic representation of the path of leaf traces through
the stem of Mikania cordata
118
In this manner at each node three leaf trace bundles diverge to

each of the opposite leaves to form the leaf trace.
d) The origin of the phellogen (Figure 22)
With the commencement of secondary thickening, cork formation
also begins. The phellogen (Fg) arises from the first layer of hypo
dermal cells which divide to form the phellem, the cell walls of which
are suberized.
Figure 22. Origin of the phellogen in the stem of Mikania cordata
s» I I
Figure 23.
Figure 23.Transverse section
Transverse section Figure
Figure 24.
24.
Longitudinal section
Figures 23 and 24. Secondary phloem in the stem of Mikania cordata
119
Kirkia [Vol.
Tissue types of an old stem (Figures 22-33)
(a) Phellem. This consists of a well developed cork layer in which

lenticels are conspicuous (Figure 14, Le).
(b) Cortex. During the thickening of the stem the cells of the cortex
tissue undergo considerable enlargement and apparently also multiply
considerably so that the collenchyma loses its identity. Resin canals
occur mainly opposite the vascular rays, but in some cases they also
form a network in the cortex and vascular rays. A stoney cell sheath
(Figure 15, St) which lends mechanical strength to the underlying
tissues, forms on the outside of the pericyclic caps (Pe).
(c) Stele, (i) Pericycle. The fibre caps (Pe) that occur on the
outside of the vascular bundles become separated laterally from
one another by thin-walled elements.
(iï) Phloem. The greater part of the primary phloem

(PrF) is obliterated. The secondary phloem (SF) (Figure 23) con
sists of sieve tubes (Sv) with companion cells (BS) and phloem
parenchyma (Fp). The walls of all these elements are thickened to a
degree, with cellulose. The sieve plates (Figure 24, Sp) occur in the
transverse walls of the sieve tubes.
(iii) Xylem. The secondary xylem (SX) of the stem

bundles (Figure 25) consists of relatively large vessels (H), xylem
fibres (Xv) and thickened xylem parenchyma (VXp). In the leaf trace
bundles only small xylem vessels (Figure 26) occur. Besides the
xylem fibres and thickened xylem parenchyma, large amounts of
unthickened xylem parenchyma (Xp) occur. All the xylem vessels
are typical pitted vessels (Figure 30). The xylem fibres (Figures
27-29) possess thickened lignified cell walls with numerous funnel
pits. These fibres are separated from one another by thin cellulose
septa (DW). The tips of some fibres are branched (Figure 27). In
the lignified walls of the thickened xylem parenchyma cells (Figures
31-32) are numerous simple pits (ES).
In some cases tyloses (Ti) are present in the vessels of the stem
vascular bundles which completely block the vessels (Figure 33).
(iv) Vascular rays. The broad, thin-walled, parenchy
matous rays between neighbouring vascular bundles are only inter
rupted by the axial xylem cylinder.
(v) Pith. Apart from the thickened sheath which forms
part of the axial xylem cylinder, the pith consists of parenchymatous
cells with a ring of resin canals.
120
DW
^ l/CM r- 07 OQ
Figures 27 - 29.
Ftguressections MaceraSxytem fib«s
Figures 25 and 26. Transverse sections
Macerated xylem fibres
Figure 33.
Figure 30.
Figure30- Figures
Segment
Segment of macerated
of macerated
31-32.
vessel. Figures
* 31 - 32.
xylem
xylem vessel.
Figures 25 - 33. Secondary xylem in the stem of Mikania cordata
RÉSUMÉ
After secondary thickening has taken place for a short time, the stem
bundles can be clearly distinguished from the six leaf trace bundles.
This differentiation of vascular bundles is the result of irregular
activity of the vascular cambium.
121
Kirkia [Vol.
Figure 34:34:
Figure Figure 35.
Primary state
Primary ofof
state stem
stem Commencement of secondary growth
Figure
Figure 36.
36.
Discontinuous
Discontinuous vascular
vascular cambium
cambium
Figure
Figure 37.
37. Longitudinal
Longitudinal grooves
grooves
Figures 34 - 37 Schematic representation of the anomalous secondary growth

in the stem of Mikania cordata
The secondary growth of the stem of this species can be regarded

as anomalous because:
(i) the secondary tissues are irregularly formed; and
ii) the vascular cambium is eventually subdivided into different
segments.
In figures 34-37 it is endeavoured to give a generalized sequence
of the growth processes in the successive internodia of the stem.
Figure 34. The stem with primary tissues
A single ring of collateral vascular bundles is present.
Figure 35. The stem after irregularity in the activity of the vascular
cambium has commenced.
122
For a short period the cambium divides normally to form

secondary tissues. At first an axial xylem cylinder is formed. With
the differentiation of the peri-axial xylem an irregularity arises in the
activity of the vascular cambium. In six bundles (leaf trace bundles)
less secondary xylem but more secondary phloem is formed than in
the remaining bundles.
Figure 36. The stem after the leaf trace bundles have become clearly
discernible and the vascular cambium has divided into
segments.
With the continued irregular activity of the vascular

cambium, the difference between the stem bundles and leaf trace
bundles becomes more apparent. The irregular formation of
secondary tissue eventually results in the breaking up of the vascular
cambium into 12 segments.
Figure 37. The stem with shallow longitudinal grooves.
The establishment of the leaf trace bundles results in the
formation of shallow longitudinal grooves on the surface of the stem
which coincide with the position of the leaf trace bundles within the
stem.
PAULLINIA PINNATA LINN.
EXTERNAL MORPHOLOGY OF THE STEM
The internodes of this tendril climber are relatively long (up to

+ 100mm). The axillary branched tendrils opposite the compound
leaves, represent highly modified lateral branches.
Although the old stems are hard and woody they remain
flexible. This suppleness must be ascribed to the compound nature
of the stele. In the internodes the central vascular cylinder is sur
rounded by three smaller peripheral vascular cylinders (Figure 38).
In turn at each succeeding node one of the peripheral cylinders fuses
with the central cylinder. This incorporation occurs gradually and
in some cases commences 30mm below the node. Above the node the
incorporated cylinder gradually re-emerges to form a distinct peri
pheral cylinder. The incorporation of the peripheral cylinders
occurs repeatedly but in strict rotation (Figure 39).
INTERNAL MORPHOLOGY OF THE STEM
The embryological state of the stem (Figures 40-42)

Shortly behind the growing point, the stem deviates from the
normal form, as even in the young stages lobes (a, b, c, d, e and f)
begin to develop.
123
Kirkia [Yol.
3 IAS 2
Figure 39. Schematic representation of the path of the peripheral

vascular cylinders in the stem of Paullinia pinnata (see text for
explanation).
124
The epidermis (Figure 40, E) with numerous trichomes can be

distinguished (the trichomes are not shown in the diagram). Secretory
cells (Ss) are present in the superficial, thin-walled tissue, while
protoxylem vessels (Px) occur in some of the lobes (a and b, Figure
40). It is at these vessels that vascular bundles develop later (Figure
41). Thus it is here in the lobes that the first differentiation of
vascular tissue occurs.
In the older stem (Figure 42) the primary collateral vascular

bundles of the different lobes can be distinguished in different stages
of development. Opposite lobe a, more towards the centre of the
stem, is a protoxylem vessel (Px) which represents the first vascular
element of the central vascular cylinder in this zone. Later, further
xylem vessels develop outwards at this single protoxylem vessel, to
Figure 40.
Figure 40.
Figure 41.
Figure 42.
Figures 40-42. Successive sections through stem of Paullinia pinnata
125
Kirkia [Vol .
T"jT,
Figure 43.
Surface view of epidermis Figure 44.
Gland y
Figure
Figure45.
45.Hair
Hair
K
-~E
oegp3
KK
V €>
j
PK o
^£l.Pv £S
>P
DP
fv
-ft
-Sv
• -BS
50 y
Figure
Figure 46.
46. Transverse
Transverse section
section Figure 47. Transverse section of the
of
of the
the epidermis
epidermis and
and phloem
phloem xylem and pith.
Figures 43 - 47. Tissues of the young stem of Paullinia pinnata
form eventually a primary collateral bundle (Figure 48, Vb). In
this way an irregular ring of vascular bundles, which later form the
central cylinder, develops gradually (Figure 49, AS). Although
during their passage through the internodes, the lobes a, c and e with
their vascular tissue (Figures 39 and 49), give the impression of
126
being independent units note onnected to the central vascular

cylinder, they are morphologically part of the vascular cylinder
as a whole as :
(i) the pericycle, which only develops later, forms a complete
sheath round the stele (Figure 50); and
(ii) the vascular tissue of each of the lobes a, c and e, joins the
central cylinder strictly in turn, to re-emerge later (Figure 39).
Figure 48. Vascular cambium in lobe
Figure 49. Vascular cambium in central cylinder
Figures 48 and 49 Transverse sections through the stem of Paullinia pinnata

illustrating schematically the commencement of secondary growth.
127
Kirkia [Vol.
Primary tissue types of the stem (Figures 43-47)

(a) Epidermis. The outermost tangential and radial walls of the
ordinary epidermal cells are thickened with cellulose (Figure 46, E)
while the epidermis is covered by a cuticle (K)—in surface view
(Figure 43) these cells appear more or less elongated. Guard cells
and two kinds of trichomes (one-celled hairs (Figure 45) and multi
cellular glands (Figure 44)) can be distinguished.
(b) Cortex (Figure 46). The cortex layers underneath the epidermis
are collenchymatous (KK). These layers are followed towards the
inside, by relatively large cells which represent a transition between
collenchyma and parenchyma. The parenchymatous zone (PK)
borders on the pericycle (P), and no endodermis can be distinguished.
Bordering on the stomata typical chlorenchyma cells are found which
reach to the guard cells. Many of the cortex cells contain secretions.
(c) Pericycle and vascular tissue. The pericycle forms a scleren
chyma sheath, consisting of fibres (Figure 46, Pv) and occasional
thin-walled cells (DP). In the lignified walls of the fibres are numer
ous simple pits (ES). Parenchyma and occasional fibres (Fv) of the
protophloem (Pf) border directly on the pericycle. In the meta
phloem, sieve tubes (Sv) with companion cells (BS) can be dis
tinguished. The metaxylem vessels (Figure 47, Mx) are surrounded
by thin-walled parenchyma cells, while the protoxylem vessels (Px)
are embedded in thin-walled protoparenchyma.
The primary vascular tissue is separated from the thin-walled
parenchymatous pith by a sheath of thick-walled cells (MV). The
latter consists of cells with lignified walls with numerous simple pits.
Origin of the cambia (Figures 48-52)

(a) The vascular cambium and secondary growth. Cambial activity
begins in the collateral vascular bundles of the lobes (Figure 48, Vk).
These newly formed cambium segments give rise to secondary
vascular tissue. Peripheral vascular bundles (PVb) are present in
lobes a, c and e (Figure 49). The bundles in the remaining lobes,
together with the bundles which became differentiated opposite the
peripheral bundles, form an irregular vascular bundle ring of the
central cylinder (AS). Interfascicular cambial segments are gradually
differentiated in this central cylinder, so that the vascular cambium
eventually forms a complete cylinder.
The cambium of both the fascicular and interfascicular segments
gives rise to secondary vascular elements. Thus, in the older stem
(Figure 50) a xylem cylinder (SX) surrounded by a phloem cylinder
(SF) is formed. The vascular rays are only one to three cells wide.
The cells of the phloem rays are thin-walled but those of the xylem
rays are thick-walled so that the xylem cylinder appears relatively
solid even in this young stage of the stem.
128
The transverse section of the stem (Figure 50) is near a node

and thus only two peripheral vascular bundles (in lobes a and e)
are present, the third peripheral bundle (in lobe c) having been
incorporated in the central vascular cylinder. The vascular cambia
Figure 50.
Figure 50. Transverse
Transverse section
section through
through stem
stem near
near aa node
node
(schematic representation)
(schematic representation)
129
Kirkia [Vol.
-£
♦to
OG
—PrF
SF
Vk
-SX
■—PrX
-Vs
- -M
XV
Vk
SF
■frtL
-SX ^
Vs
-PrFl
PV*
Figures 51 and 52.Transverse sections through the internodes

of the stem of Paullinia pinnata (schematic)
of the peripheral bundles gradually spread out to form an arch (in

lobes a and e, Figure 50) which will eventually form a complete
cylinder (Figure 51).
130
In an older internode (Figure 51) the three peripheral vascular

cylinders lying to the outside of the central vascular cylinder (AS)
are easily distinguishable as the cambium of each of these is present
in the form of a complete cylinder so that secondary vascular tissues
are formed in the normal way. The marked excentric structure of
each of these peripheral vascular cylinders is the result of the
excentric development and origin of the vascular cambium cylinder
concerned.
The peripheral cylinders are anomalous in structure as the proto

parenchyma on the pith side is bounded partially by primary and
partially by secondary xylem with the result that secondary thicken
ing in the internodes of the stem takes place through four different
vascular cambia which, for a time, are independent of each other.
Each of the four cambia forms secondary vascular tissue. The
secondary phloem of the peripheral vascular cylinders is separated
from the primary phloem of the central cylinder by thin-walled
ground tissue (Figure 52, DG).
(b) Origin of the phellogen (Figure 52). A phellogen develops from
the first hypodermal cell layer and gives rise to phellem (F). The
first phellogen and cork cells are found in the lobes with peripheral
cylinders. Gradually the phellogen develops into a complete cylinder
over the whole circumference of the stem.
Fusion of the peripheral vascular cylinder with the central vascular

cylinder (Figures 53-58).
Figure 53 shows one of the peripheral vascular cylinders in the
internode below a node.
Nearer the node (Figure 54) the incorporation of this cylinder
with the central cylinder begins as a local irregularity in the activity
of the vascular cambium of both cylinders. That part of the cambium
of the peripheral cylinder which lies nearest the central cylinder gives
rise to less secondary tissue than the remaining part. Eventually
there is no further differentiation of secondary phloem between the
peripheral and the central cylinders.
The part of the cambium of the central vascular cylinder which
lies opposite the peripheral cylinder is the first to show a decrease in
activity. During this process the cambium is eventually interrupted
and as a result a longitudinal groove appears in the xylem of the
central cylinder.
As a result of further irregular activity of the cambium of the
peripheral cylinder and its eventual interruption (Figure 55), the
cylinder gradually takes on the form of a collateral vascular bundle.
The longitudinal groove of the central cylinder in which primary
ground tissue is present, becomes wider and deeper (Figure 56),
131
Kirkia [Vol.
Figure 53. Internode Figure 54. Beneath internode
Figure 55. Nearer to node Figure 56. Beginning of node
Figure 57. Beginning of node Figure 58. Node
Figure 59." Above node

59. Above node Figure
Figure 60.
60. Internode
Internode
Figures 53-60 Schematic representation of the incorporation and
re-emergence of the periphejral cylinders in the stem of Paullinia pinnata.
132
Figure
Figure61.
61.
Transverse section
Transverse section
through
throughphloem
phloem
It
YTFigures
1 Figures
/f| 62 62
andand
63. 63. ||; |
Longitudinal
Longitudinal section
section through
through phloem
phloem
Figures 65-66.
Figures 65-66. Figures
Figures 67-68
67-68
Xylem vessels Tracheids
Xylem vessels Tracheids
c\
i*
DW
ow
ti r ^
Figures71
Figures 71toto72
72
Figure
Figure64.64.
Transverse sectionsection Fion™. J?
Transverse Ill m
Xylemparenchyma
parenchyma
Xylem
through
through secondary xylem. xylem.Figures
secondary v , 69-70
°"-70
Xylem
Xylem fibre
fibre
Figures 61 - 72. Tissues of an old stem of Paulinnia pinnata
while the peripheral bundle moves towards the central cylinder.

Eventually the interrupted ends of the two cambium cylinders fuse
to form a common vascular cambium.
The strip of thin-walled, primary ground tissue becomes thinner

until the peripheral bundle fuses with the central cylinder and
eventually becomes incorporated in it (Figure 57).
133
Kirkia [Vol.
In the node (Figure 58) the incorporation is completed and the

former peripheral cylinder forms part of the central cylinder.
The re-emerging of the peripheral vascular bundle from the central

vascular cylinder (Figures 59-60)
Above the node the incorporated vascular bundle once again
gradually re-emerges laterally from the central cylinder (Figure 59),
the steps in its re-emergence occurring in precisely the reverse order
from those describing its incorporation with the central cylinder.
Eventually the peripheral cylinder can again be distinguished as a
separate entity in the next internode (Figure 60).
Tissue types in the old stem (Figures 61-72)

(a) Phellem. A well developed cork layer with obvious lenticels, is
present.
(b) Cortex. With the increase in diameter of the surface of the stem
as a result of secondary thickening the primary tissues undergo
marked cell stretching. For this reason the cortex cells are larger
and their walls thinner than in the primary state. Most cells contain
secretions.
(c) Stele (i) Pericycle. According to Metcalf and Chalk (1950),

a common sclerenchymatous pericycle sheath surrounding a com
pound stele, is characteristic of the Sapindaceae. This sheath con
sists largely of fibres with a few stone cells. The latter developed
slowly from the thin-walled cells present in the primary stem. The
sclerenchymatous sheath keeps pace with the increase in diameter
and surface area of the stem either by the stretching of these thin
walled elements or, in some cases, by local divisions.
(ii) Phloem. The secondary phloem (Figure 61) consists
of sieve tubes (Sv) with companion cells (BS), phloem parenchyma
(Fp), secretory cells (L) and phloem fibres (Fv).
The sieve plates (Figure 62, Sp) are restricted to the transverse
walls of the tubes. Although the companion-cells (Figure 62, BS)
are much narrower than the sieve tubes, they are of the same length.
In some phloem parenchyma cells, calcium oxalate crystals are found
(Figure 62, Ke), and in others, compound and simple starch grains
(Figure 63, Set) are present. The secretory cells (Figure 62, L) are
thin-walled, elongated elements. Numerous funnel pits (Ts) are
present in the thickened, lignified cell walls of the phloem fibres
(Figure 63, Fv), and these elements are septated by thin cellulose
transverse walls (DW).
(iii) Xylem. The secondary xylem (Figure 64) consists
of the following thick-walled elements : vessels (H), tracheids (Tr),
xylem fibres (Xv) and xylem parenchyma (VXp).
134
The size of the pitted vessels varies (Figures 65-66). As in the

vessels, bordered pits (Figure 68, Hs) occur in the walls of the
tracheids. The xylem fibres (Figure 69) with numerous funnel pits
(Figure 70, Ts) in their walls, are septated. Some elongated xylem
parenchyma cells (Figure 71) may also be septated, and numerous
simple pits (ES) occur in their walls; these cells mostly contain
starch and/or secretions.
(iv) Vascular rays. Although the vascular rays are
relatively narrow they are still obvious and most of the cells contain
starch and/or secretions. The cells of the xylem rays are thick-walled
with large intercellular spaces (Figure 72, Int). The phloem rays are
a continuation of the xylem rays but consist of thin-walled elements.
(v) Pith. In the medullary tissue two zones can be dis
tinguished namely the sheath of thick-walled pith cells bordering on
the xylem, and a central parenchymatous zone.
(vi) Types of secretions. On grounds of a colour differ
ence, two secretions can be distinguished which, according to
Metcalf and Chalk (1950), are saponine and tannic acid compounds.
The vascular elements of the central and peripheral cylinders
only differ quantitatively and not qualitatively.
RÉSUMÉ
The compound vascular cylinder, as found in Paullinia pinnata,
is considered by Schenck (1893), to be a highly specialized liane
structure as even during the differentiation of the primary tissues,
deviations from the more normal form occur.
The condition summarized below, only applies to the internode

since at each node one of the peripheral cylinders, in turn, becomes
incorporated in the central cylinder. Above the node this peripheral
cylinder gradually re-emerges from the central cylinder so that
eventually it can be recognized again as a separate cylinder. Al
though transverse sections of the internodes give the impression of a
cable structure, this is not completely correct.
Figure 73. The stem in the primary state.
The stem is lobed from the apical meristem and it is in these
lobes that the first vascular elements become differentiated. In each
of the six lobes a primary collateral vascular bundle is differentiated.
Figure 74. The differentiation of the central vascular cylinder.
In three of the lobes only single vascular bundles are formed
which can be called future peripheral vascular bundles. The vascular
bundles in the remaining lobes, together with bundles which are
differentiated later towards the inside of the peripheral bundles, form
the central vascular cylinder.
135
Kirkia [Vol.
Figure 75. A complete vascular cambium cylinder of the central

cylinder is formed.
The fascicular and interfascicular cambium segments of the
central cylinder gradually join together sideways to form a complete
0
E 0
PrF 9
"1.--PrX
\,CC5 "'AS.. CD
-M
[PrX
PVbl vk
PrF
Figure 73
Figure 73 Figure
Figure74
74
Primary state of stem
Primary state of stem Central vascular
Central vascular cylinder
cylinder
Figure
Figure75.
75.Cambuim
Cambuimofofcentral
centralcylinder
cylinder
Figure
Figure 76.
76. Four
Four vascular
vascular cylinders
cylinders
Figures 73 - 76. Schematic representation of anomalous secondary growth
in the stem of Paullinia pinnata
136
cambium cylinder. As secondary growth occurs in the peripheral

bundles the vascular cambium gradually takes on an arch shape and
eventually forms a cylinder.
Figure 76. Secondary thickening occurs in the four vascular
cylinders by means of the four cambia.
In the internodes of the stem secondary thickening occurs by
means of four temporarily independent cambia. Each of these
cambia gives rise to secondary vascular tissue.
DISCUSSION
Although the stem of Mikania cordata to a certain extent re

sembles that of Paullinia pinnata in its external morphology, the
apparent cable structure is brought about by a different type of
anomalous secondary thickening. Neither of them is a true cable
structure as even in Paullinia pinnata the peripheral cylinders from
time to time become incorporated in the central cylinder.
In Mikania cordata the anomaly only appears after secondary
growth has begun whereas in the stem of Paullinia pinnata the
anomalous structure has its origin in the apical meristem. The two
liane species differ further from one another in that the latter has a
hard, woody stem while the former has a relatively soft stem as a
result of the presence of much thin-walled parenchymatous tissue.
As in the case in the stems of most of the lianes that exhibit
anomalous secondary growth, a solid xylem cylinder is not formed
in either Mikania cordata or Paullinia pinnata—a characteristic
which apparently increases the flexibility of the stem.
LIST OF ABBREVIATIONS
AX = Axial xylem cylinder Pe = Pericycle cap

AS = Central vascular cylinder PEX = Peri-axial xylem
BL = Leaf trace bundle Pf = Protophloem
Bg = Leaf gap PK = Parenchymatous cortex
BS = Companion cell Pkb = Procambium bundle
DG = Thin-walled ground tissue Pp = Protoparenchyma
DP = Thin-walled pericycle Pr = Perforation
DW = Thin cellulose wall PrF = Primary phloem
E = Epidermis PrX = Primary xylem
Ep = Epithelium cell Pv = Pericycle fibre
ES = Simple pit PVb = Peripheral vascular bundle
F = Phellem PVs = Peripheral vascular cylinder
Fg = Phellogen Px = Protoxylem
Fp = Phloem parenchyma SAX = Secondary axial xylem
Fs = Phloem ray Sek = Secretory canal
Fv = Phloem fibre Set = Starch grain
H = Vessel SF = Secondary phloem
Hs = Bordered pit Sp = Sieve plate
137
Kirkia [Vol.
Ino =
Intemode Ss =
Secretory cell
Int =
Intercellular space St =
Stone cell
K =
Cuticle Sv =
Sieve tube
KK =
Collenchymatous cortex SX =
Secondary xylem
Ke =
Calcium oxalate crystal Th =
Trichome
Kr =
Gland Ti =
Tylosis
Ko =
Cortex Tr =
Tracheid
L =
Tannic acid containing cell Ts =
Funnel pit
Le =
Lenticel Vb =
Vascular bundle
M =
Pith Vk =
Vascular cambium
Ms =
Medullary ray Vs =
Vascular ray
MV --
Pith cells with thickened walls VXp =

Thickened xylem parenchyma
Mx =
Metaxylem Xp =
Xylem parenchyma
No =
Node Xs Xylem ray
P =
Pericycle Xv Xylem fibre
REFERENCES
De Bary, A. (1884). Comparative anatomy of the vegetative organs

of phanerogams and ferns. Oxford, Clarendon Press.
Haberlandt, G. (1914). Physiological plant anatomy. London,
Macmillan and Co.
Johansen, D. A. (1940). Plant microtechnique. New York, McGraw

Hill Book Company.
Metcalfe, C. R. & Chalk, L. (1950). Anatomy of the dicotyledons.
Oxford, Clarendon Press.
Schenck, H. (1893). Beitrage zur Biologie und Anatomie der Lianen.
Jena, Yerlag von Gustav Fischer.
Van der Walt, J. J. A., Schweickerdt, H. G. & van der Schijff, H.P.
(1969). Afwykende Sekondêre Diktegroei in die Stingels van die
Liane Cyphostemma anatomicum (C.A.Sm.) Wild & Drummond
(Vitaceae) en Adenia gummifera (Harv.) Harms (Passifloraceae).
Tydskr. Natuurwet., 9, 2-3: 89-123.
Van der Walt, J. J. A., Schweickerdt, H. G. & van der Schijíf, H. P.
(1970). Afwykende Sekondêre Diktegroei in die Stingels van die
Liane Cocculus hirsutus (L.) Diels (Menispermaceae) en Pyrena
cantha grandiflora Baill. (Icacinaceae). Tydskr. Natuurwet. 10»
2: 173-199.
H. G. Schweickerdt & H. P. van der Schijfï J. J. A. van der Walt

Department of General Botany Department of Botany
University of Pretoria University of Stellenbosch
SOUTH AFRICA SOUTH AFRICA
138
Mïkania cordata (Burm.f.) Robins)
VI
m
1
Figure 1 Figure 2. Figure 3.
Figure 1. Portion of stem of Mikania cordata (± 2 cm in diameter) with

thickened nodes and longitudinal grooves
Figure 2. Portion of non-twining stem of Mikania cordata
Figure 3. Portion of twining stem of Mikania cordata
(± 1.3 cm in diameter)
Mm
.ft " '
' i Sr*©8
BlIE . <|3®J|Pl
■ ; *, , < msmm
wk i.
ilNiw
mm i |£ <- ^WpWWWpWMMWB
Jf^V- •'. ;/"

S"~ -
I
i
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Figure 38. Paullinia pinnata Linn.

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National Herbarium & Botanic Garden

ANOMALOUS SECONDARY GROWTH IN THE STEMS OF THE LIANES MIKANIA CORDATA

J. J. A. van der Walt, H. P. van der Schijff and

The stems of two liane species Mikania cordata (Burm.f.) Robins,

The phenomenon of anomalous secondary growth is found in the

In previous publications (van der Walt, Schweickerdt and van

The external morphology of the stems of both Mikania cordata

MATERIAL AND METHODS

For a complete anatomical study of the stem it was necessary to

MIKANIA CORDATA (BURM.f.) ROBINS.

The thickest stems of this twiner were about 25 mm in diameter.

INTERNAL MORPHOLOGY OF THE STEM

Structure of the stem in the embryonic state

differentiated. The following tissues can be distinguished at this

c) A stele consisting of pericyclic caps (Pe), with component

a) An epidermis of ordinary epidermal cells with thickened

(c) The stele (Figure 10) shows:

Figure 7.Epidermis and cortex

The vascular cambium (Vk) has already given rise to a certain

The metaxylem vessels (Mx) are relatively large, with numerous

Origin of the cambia and secondary growth

a) Origin of the vascular cambium and the commencement of

For a time, primary growth in thickness takes place, after which a

The first formed fascicular segments join with the interfascicular

At first this vascular cambium gives rise, centripetally over its

b) Further secondary growth (Figures 13-15) \

Figure 11. Transverse section of the stem of Mikania cordata showing

Transverse section of the stem

.ozm t ° i foJ/V/o W^j o

Transverse section of stem showing the irregular activity of the

Figure 15. Transverse section of stem ofMikania cordata showing

Figure 15. Transverse section of stem ofMikania cordata showing

c) Leaf trace bundles (Figures 16-21)

Figure 20. Node Figure 21. Internode

In this manner at each node three leaf trace bundles diverge to

Figure 22. Origin of the phellogen in the stem of Mikania cordata

Tissue types of an old stem (Figures 22-33)

(a) Phellem. This consists of a well developed cork layer in which

(iï) Phloem. The greater part of the primary phloem

(iii) Xylem. The secondary xylem (SX) of the stem

Figures 25 - 33. Secondary xylem in the stem of Mikania cordata

Figures 34 - 37 Schematic representation of the anomalous secondary growth

The secondary growth of the stem of this species can be regarded

For a short period the cambium divides normally to form

With the continued irregular activity of the vascular

PAULLINIA PINNATA LINN.

EXTERNAL MORPHOLOGY OF THE STEM

The internodes of this tendril climber are relatively long (up to

INTERNAL MORPHOLOGY OF THE STEM

The embryological state of the stem (Figures 40-42)

Figure 39. Schematic representation of the path of the peripheral

The epidermis (Figure 40, E) with numerous trichomes can be

In the older stem (Figure 42) the primary collateral vascular

being independent units note onnected to the central vascular

Figure 48. Vascular cambium in lobe

Figure 49. Vascular cambium in central cylinder

Figures 48 and 49 Transverse sections through the stem of Paullinia pinnata

Primary tissue types of the stem (Figures 43-47)

Origin of the cambia (Figures 48-52)

The transverse section of the stem (Figure 50) is near a node

Figures 51 and 52.Transverse sections through the internodes

of the peripheral bundles gradually spread out to form an arch (in

In an older internode (Figure 51) the three peripheral vascular