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 SIPHONoSTELE WITH
SIPHONOSTELE WITH
ONE GAP AT A THREE LEAF AT A
SIPHONOSTELE WITHOUT NODE (UNILACU STRANDS NODE (TRILACUNAR
GAPS
NAR NODE) JOINED INTO NODE)
ONE IN THE
PETIQLE
ONE
OFTWO
BRANCH
TRACES
~LEAF
TRACES
THREE LEAF C
A B TRACES EACH ASSOCIATED
WITH A SEPARATE GAP
Fig. 38.2. Primary vascular system. A, Selaginella-three-dimensional view,
siphonostelic type without
leaf-gaps; B, Nicotiana-unilacunar node, siphonostelic with leaf gap; C, Salix-trilacunar node,
siphonostelic type with leaf gaps. (After Esau).
There are three common types of nodes in the
dicotyledons. The node with a single gap and
a
single trace to a leaf is known as unilacunar; the node with three gaps and three traces to a leaf
(one median and two lateral) is known as trilacunar; and the node with
several to many gaps and
traces to a leaf is known as multilacunar. The most
accepted concept
condition is primitive in the dicotyledons and that the unilacunar and
is that the trilacunar
the multilacunar have been
derived from it. Several monocotyledonous plants leaves with sheathing bases and nodes
possess
with a large number of leaf traces separately inserted around the
stem. In ferns the number of
traces to leaf varies from one to many, but they are
a
always associated with a single gap. In
gymnosperms a unilacunar node is common.
The leaf trace relationships at the nodes are thought to be of
phylogenetic importance, and
therefore. nodal anatomy is concerned with the study of
systematics and phylogeny of
angiosperms.
Nodal Anatomy in Wheat (Monocot) Stem
In the wheat stem the course of the vascular bundles
sheath is almost parallel. Near the node the leaf sheath is
through the internode and the leaf
maximum thickness just above its union with the stem. On considerably thick and attains its
the other hand, the stem has the
smallest diameter above the junction with the leaf sheath.
The stem is hollow in the internode
solid at the node. The sheath remains and
open on one side at higher levels, just near the node.
Massive collenchymatous bundle
caps are present in the bundles of leaf sheath.
Just beneath the junction of the leaf sheath and
stem the
smaller of the leaf traces are
prolonged in the
peripheral part of the axis, and the
cylinder of strands. The internodal bundles larger leaf traces become part of the inner
located above the leaf insertion assume,
just above the
 NODES
ABOVE NODES

A
SPIRAEA
UNILACUNAR
NODE
LEAF GAP
MEDIAN TRACE
PETIOLE

}LEAF
GAP

B
SALIX
TRILACUNAR
NODE
LATERAL TRACES
BRASSICA LEAF BASE
TRILACUNAR
NODE 2

RUMEX
MULTILACUNAR

NODE

leat has one leaf trace and one leaf gap

dicotyledons. A. Spiraea-each node):
Fig. 38.3. Nodal anatomy of each leaf has three leat traces and
three leaf gaps (trilacunar
Salix- node); D, Aumex- many
(unilacunar node); B, three leat gaps per lear (trilacunar
traces and node).
C, Brassica- three leaf
(multilacunar
leaf gaps per leaf
many
leaf traces and
are reoriented toward a more peripheral
course (Fig. 3S. 7 C, D), and variously
node horizontal and oblique
a These horizontal and oblique bundles
(Fig. 38.7 D, E). the
below it and the bundles from
position in the node and traces

their number
reduces.
Ine large lear bundles of the
of the next lower
oranch and coalesce, and the
inner cyiinder
leaf make
of the the bundles are leaf traces from the
approximately halfof
insertion t
nternode above the cylinder
nternode (Fig 38.7 E). in this half of the bundles are 1from the internode above the insertion of
other eaf traces. The most conspicuous
lea,
nearest leaf above and the bundles are mostly
peripheral es in the nodal regions.
bundles
the leaf (Fig. 38.7
. E). The transverse
is the presence of
character of grass
gra stems
 LEAF TRACE
LEAF GAPS
OR LACUNAE

XYLEM

PHLOEM

LEAF
TRACE

A B

in transection; B, nodal anatomy of

Fig. 38.4. Nodal anatomy. A, nodal anatomy of Picea (a conifer) traces to leaves, and
Adiantum (a fern) in transection. Both possess alternate leaf arrangement, single
on both sides of xylem.
Single leat gaps at the nodes. A, has some secondary growth; B, has phloem

APICAL MERISTEM
xYLEM PHLOEM

LEAF BUNDLE
VASCULAR CYLINDER
OF MAIN AXIS

-GAP

-LEAF
TRACE
A C
BRANCH TRACES

Fig. 38.5. Nodal anatomy. The diagrams depict the primary vascular system of plant with an opposite
(decussate) leaf arrangement in L.S. (A) and . s . (5 and C). The branches present in the axils of the
leaves are inflorescences. The leat trace and orancn traces in its axil are associated with one common
gap. The two branch traces ot level C are united into a tubular vascular cylinder in B.
 679

SUBTENDING
LEAF
AXILLARY
BUD
BRANCH GAP

BRANCH C
TRACES EA VASCULAR
CYLINDER OF
MAIN AXIS

MEDIAN
LEAF GAP

D E
Fia, 38.6. Branch
traces and gaps.
Diagrams showing vascular connection betwe en an axillary branch
(bud) and the main axis in Salix. Vascular
system of bud is shown in
leaves of bud are oPposite. Ihe branch gap and the median gap of theblack in the diagram. First twO
subtending leaf are contluent.

SHEATH

STEM

COLLENCHYMATOUS 3
TISSUE VASCULAR BUNDLESS
A VASCULAR
BUNDLES

CAVITY IN PITH E
C D NODAL REGION BELOW NODE
ABOVE NODE
in stem are depicted in
of sheath
and
their traces
Fig .i. Nodal anatomy of Triticum stem. Bundles
its continuation
througn node Is hatched.
A-E, transectionsof
d e c r e a s e s in thickness.
Vascular tissue of internode and increases and stem
node, sheath
in at various levels. Toward
680
Branch Gaps system o f
Branch Traces and branches the vascular
is also derived from the
to lateral
vascular supply less often, o n e bundle. These strands are known

The primary bundles,

and
form of two
1947).
usually in the
Dicotvledons
and MacDaniels,
the main axis, (Eames to
traces or
traces
ramular the v a s c u l a r system of the branch
branch traces, connecting
as
two branch main stem
commonly have of the
connection
axillary shoot with the
gymnosperms the appendages are
monocotyledons the main axis and
that of the main stem. In extended within
branch t r a c e s are t w o traces, these bundles
strands. The branch p o s e s s e s
of many When the
when o n e trace o c c u r s , this
consists
vascular system.
primary cylinder;
tied together by a
complete a
vascular
s t r u c t u r e with the openinp
short distance, forming h o r s e - s h o e shaped
unite within a form of a as the
cross-sectional closure of the opening
possesses the is formed by the
strand usually of the branch
vascular cylinder
downward, and the
associated with the
trace is
branch traces passes out. of a branch
the outward passage of the trace,
In most of
vascular plants
around and above
the point of departure
Branch gaps
formation of a break
in the vascular cylinder accompanies a
branch trace.
which always the gaps do not
known as branch gap, However, in protosteles
This opening is which possess a pith.
and extend for
are present in
all vascular plants lower than leat gaps
a r e commonly
Branch gaps
occur because there is no pith.
distances in the axis.
greater

CLOSING OF LEAF GAPS in the secondary body

not perpetuate
the nodal s t r u c t u r e do
characterize t i s u e s in continuity
The features which forms vascular
of the leaf gap and
A cambium develops
in the parenchyma of the gap. The parenchyma
is known as closing
the gap. This phenomenon the inner portion
with those bordering cambium and those in
the first to change into
of the gap are
cells near the margin
XYLEM IYEAR

LEAF TRACE

PHLOEM

CORTEXX

LEAF TRACE A
LEAF XYLEM
GAP

LEAF
SCAR

LEAF
TRACE

B D E
of
Fig. 38.8. Closing of leaf gaps by secondary growth. A and B TS. and L.S.. through nodal regio
stems in first year of growth: C-E, T.S. and L.S. through stems several years old: D and E.
depict stages in closing of gaps and rupture of leaf trace (E)
 nge
e later.
later. This process takes place gradually, and the such
is
thin the secondary body until and cambium is differentiatedparenchyma maintanc *
gap
throughout the entire tangential
the gap.
width of
In the leat trace itselt complicated changes take place during secondary growth. Ihe prima
or
wlem is buried by secondary tissues while the phloem is pushed outward. The upper partthat
the
and crosses the plane of the cambium. The part of
the cambium
e trace
the trace diverges
div outwardly trace and the
th in the tissue between the
differentiates
above trace gap region, produces vascular
the trace and
vascular cylinder This tissue which increases in pressure upon
amount, exerts a
vascular cylinder. into cambiumn
causes itS rupture. The break is
filled with parenchyma which is changed this
ultimately with that formed in the gap. After
of the trace
the cambium ot the lower part break becomes
and connects
the end of the trace below the
formed some secondary tissues, outward, and in
cambium has 38.8 E). The upper
severed end is carried
Since the
in secondary xylem (Fig. the activity of the periderm.
embedded with the cortex, by the trace
thrown off, together outward, the
buried part of
be
time it may the trace phloem
the trace itself pushes
within
cambium
consists of xylem only.