The Shoot System: Primary Stem Structure - 1

Shoot System
The shoot system comprises the leaves and s tems of plants. Leaves are located
at nodes on the stem; the distance along the stem between nodes is known as an
internode. Shoots develop from s hoot meristem, which contains the apical
shoot tip meristem from the epicotyl of the embryo, leaf primordia, and bud
primordia, which are embryonic lateral shoot systems. Flowers, the reproductive
organs of angiosperms, are modified shoots. Shoot meristems are located at the
growing tips of stems, and in buds.
Shoot Meristems
The apical meristem is divided into two regions: t unica and c orpus. The tunica
meristem cells divided in a plane the produces additional surface meristem. The
corpus meristem cells under the tunica divide in a plane that adds bulk to the shoot
meristem. Generally there are two tunica layers and one corpus layer.

As a shoot grows, buds are laid down by shoot meristem in the axils of leaf
primordia. The repeating units of leaf and bud primordia are called p hytomeres.
These buds are dormant meristems that are activated at some later time in
growth.
Stem tissues are produced from the same three derivative meristems as root
tissues are:
Protoderm is responsible for Epidermis
Ground Meristem differentiates into ground tissues
Procambium produces the vascular tissues

Coleus stem tip, l.s.

Shoot tip, xs

SEM of shoot tip

The Shoot System: Primary Stem Structure - 2

The arrangement of derivative meristems is different in the stem from the root
meristem zone. In the shoot meristem, procambium forms a cylinder of cells with
ground meristem to the interior and to the exterior of the procambium cylinder.
As expected, the protoderm is the outer layer of derivative meristem. Single
central strands of procambium extend out into the leaf primordia from the shoot
meristem. These are called leaf traces, and leave a vascular gap in the stem
tissue at those points.
The positioning of new leaf primordia, and hence branches, is regulated by the
inhibition effect of existing primordia so that leaf and branching patterns generally
spiral along the stem. Growth regulators in the shoot tip and leaf primordia are
responsible for the inhibition (see later).
Stem Functions
The stem is the axis of the shoot system that provides mechanical s upport
for and serves as the attachment site for leaves and reproductive
shoots.
Stems elevate leaves for photosynthesis and position reproductive shoots
for optimal access to pollinators and dispersal agents.
Stems conduct water and minerals from roots to the leaves and conduct
solutes from leaves to storage and use sites.
Stems are responsible for the overall growth (height and girth) of the plant
from the primary and secondary shoot meristems
There is much variation in stem types and stem anatomy, although there are some
common features, which we will discuss. Most s econdary growth of plant shoot
systems (increase in girth) occurs in stems. (Leaves are generally primary growth
structures.) Our discussion of stems will include both internal anatomy and
external features of primary and secondary growth in stems.
Primary Growth of Stems
There are three basic primary growth patterns in stems:
Early growth of most herbaceous eudicot stems and some woody eudicots
Early growth of some woody eudicots
Monocot stems

Most woody eudicots

Herbaceous eudicots

Monocots

The Shoot System: Primary Stem Structure - 3

While the arrangement of tissues in stems differs from roots, the tissues are the
same, and by now, should be becoming familiar.
Typical Eudicot Stem Primary Growth Patterns
Epidermis
Cuticle for protection
Some stomata for gas exchange
Trichomes are common glands, prickles and hairs
Cortex region
Few layers of collenchyma for flexible strength
Parenchyma layers
Some sclerenchyma may be found in cortex, too
Pith
The pith is comprised of parenchyma tissue with many intercellular spaces.
Some plants may have a hollow pith. An endodermis layer forms interior
to the vascular tissue in these hollow-stemmed plants.
Pith rays, extensions of pith between vascular bundles radiate from the pith
to the cortex. Pith rays are difficult to observe in plants that have a
cylinder of vascular tissue in primary growth.

Helianthus stem, xs

The Shoot System: Primary Stem Structure - 4

Vascular Tissue
Pattern 1: Ring of discrete vascular bundles with pith rays
Vascular bundles may have a fiber bundle cap (which is called a phloem
fiber cap because it is adjacent to the phloem) or a simple sheath of
sclerenchyma surrounding the entire vascular bundle. Individual
bundles are separated from each other by pith rays, parenchyma
tissue that extends from the interior pith region of the stem to the
cortex.
Primary P hloem is found toward the exterior of the vascular bundle.
(Some vascular bundles also have phloem on the inside of the xylem, too.
Cucurbita stem vascular tissue is an example of this.)

Primary X ylem is found toward the interior of the vascular bundle.

When there is secondary growth, there will be a layer of p rocambium
retained between the primary xylem and primary phloem. Such vascular
bundles are said to be open vascular bundles, because they can
proceed to secondary growth.
Eudicots that lack secondary growth have c losed vascular bundles,
and no procambium remains between the primary xylem and primary
phloem.

Eudicot open vascular bundle

Closed vascular bundle

The Shoot System: Primary Stem Structure - 5

Pattern 2:
Cylinder of Vascular Tissue
Some eudicots that have secondary growth, particularly woody plants, may
have a complete cylinder of vascular tissue in primary growth separating the
cortex from the pith. Within the cylinder, whose interior and exterior
borders are typically formed by a layer of sclerenchyma cells, are
radiating rows of ground parenchyma that separate alternating xylem and
phloem vascular bundles. It can be difficult to distinguish the bundles
because there are no bundle sheaths or fiber caps.

Primary stem in Tilia

There is a reasonable amount of variation in secondary growth in eudicots.
Woody plants have extensive secondary growth. Most herbaceous eudicots
have open bundles and exhibit some secondary growth, although most of this
secondary growth is for support rather than for conduction. The transition
in stem growth pattern from primary to secondary is discussed with
secondary growth.

The Shoot System: Primary Stem Structure - 6

Monocot Stem Variations
Most monocots are reasonably small, herbaceous plants. There are some notable
exceptions, however, such as the palms, which attain great height and dimension.
Most monocots have no secondary growth, even perennial monocots. The common
monocot families are the lily, grass and orchid families.
Some distinctive monocot stem features:
Monocot vascular bundles are closed. No procambium remains so generally
monocots have no cambium (and no increase in girth)
There may be 2 or 3 layers of sclerenchyma beneath the epidermis layer for
strength and support of the stem structure. Some parenchyma cells may
also develop thickened walls in monocot stems as they mature.
Vascular bundles are "scattered" in appearance throughout the ground
parenchyma, so there is no distinction between cortex and pith. Vascular
bundles have an anastomosing, or an interwoven pattern in longitudinal
section. The parenchyma cells between vascular bundles are just referred to
as ground tissue. This is the third vascular bundle pattern common in
primary growth of stems.
Most monocot vascular bundles contain two large xylem vessels toward the
interior of the bundle, and some smaller vessels between them. The very
first vessels formed usually collapse from stretching during growth, leaving
an air space. As a result the vascular bundle often takes on the appearance
of a "clown face". Phloem is always located toward the epidermis layer, and
contains no fibers. A sclerenchyma bundle sheath surrounds each vascular
bundle.

Monocot stem, xs

Monocot vascular bundle

The Shoot System: Primary Stem Structure - 7

Special Variations in some Monocots
Hollow Stems
In some monocot stems a central pith cavity develops from cells of the ground
tissue that are destroyed during early growth. This gives an appearance of a ring
of vascular bundles, similar to eudicot patterns, surrounding the hollow central
cavity.
Intercalary Meristems
Many grasses have meristem layers at the bases of nodes, which provide for nonapical growth and enlargement of cells throughout the plant stems. (which is why
we have to mow lawns).
Thickened Meristems
Some monocots achieve great dimensions without secondary growth. Although we
will discuss this more when we discuss secondary growth and woody plants, one
way in which a plant can have a larger girth is to produce a meristem that
proliferates laterally just below the apical meristem. This thickened meristem is so
large that the true apical meristem with its leaf primordia appears sunken into this
area. The thickened meristem is procambium, which produces huge numbers of
vascular bundles within the stem. This region of proliferating procambium is known
as the m eristematic cap.