BIOL 1507E Woody Stem and Root Structure

Summary
A. The Twig - A Young Woody Stem
B. Introduction to Woody Stem Anatomy
1. Gymnosperm
a) Xylem cell types in a gymnosperm
b) Cross section of the gymnosperm
2. Angiosperm
a) Xylem cell types in an angiosperm
b) Cross section of the angiosperm
C. Cross section of the Woody Angiosperm Root
D. Cross section of the Woody Gymnosperm Root

The woody plant form has existed on earth for millions of years. It predominated
at a time when world climate was universally mild. As the climate fluctuated, and
drought and cold spells became common, the tree’s way of life was not as well suited for
survival as that of the herb. Many were destroyed.

In today’s climate, woody plants living in zones with drastic seasonal differences
survive the cold or dry season by becoming dormant. However, their woody framework
still stands vulnerable to wind, ice, snow, or drought. The herb escapes these forces by
not making any permanent woody framework. Instead, it is short-lived and quickly
produces the seeds of the next generation.

Herbs are fast growing plants. Trees have a slower growth rate and net
photosynthetic rate than those of herbs. Trees are slow to reproduce, typically requiring
years of growth before they first flower and produce the seeds of another generation.
Most of their energy is spent on building the woody framework and making a new set of
leaves each year.

A. The Twig - A Young Woody Stem

Twigs are the youngest part of a tree or shrub. Examine a leafless twig taken
from a dormant tree. Draw a representation of the twig in the indicated box and label
your drawing with the following highlighted terms.

A terminal bud occurs at the tip of a twig and is responsible for its growth in
length. If your twig has branches, each branch will also have a terminal bud. Note the
bud scales. These are modified leaves. They enclose and protect the embryonic leaves
and apical meristem within the bud. Depending on the bud type or the plant species, the
bud may also contain embryonic flowers.

At the onset of the growing season, bud scales flare out and fall off as the young
stem and leaves grow. The internodes between successive leaves elongate, thereby
increasing the twig length. Some plants have a solitary terminal bud. Others, like oaks,
ashes and maples have two or more lateral buds clustered closely about the terminal bud,
giving the appearance of several terminal buds. Still others, like sycamore have no
terminal buds; nearby lateral buds bring about twig elongation.
BIOL 1507E Woody Stem and Root Structure
A terminal bud scale scar is a band of closely grouped scars (rings) encircling
the twig. Each such band of scars marks the position of the terminal bud of a previous
year. The scars are those left after the bud scales fell off. Indicate with a bracket one
season of growth on your twig diagram.

A leaf scar is a scar left on the twig where the petiole or base of the leaf was once
attached. The scar is a corky layer, sealing off the living tissues beneath. Leaf scars are
of many different shapes and sizes, depending on the species, and are a key feature used
to identify woody plants. No new leaves will arise from these leaf scars.

Continue to examine one of the leaf scars on your specimen. Note the pinhead-
size, corky bumps within the leaf scar. These are the scars of the broken ends of the
vascular strands (bundles) that passed between leaf and stem and are therefore called
vascular bundle scars. The numbers and arrangement of these vascular bundle scars
vary with different species. They, like leaf scars, are useful characters for identifying
woody plants during their dormant season when leaves are absent.

The leaf scar occurs at a position on the stem called the node. The node is
defined as that region on a twig where one or more leaves are borne. The portion of the
twig between any two successive nodes is the internode. Indicate an internode on your
diagram. Compare the lengths of the internodes of one growing season with those of
another growing season. Are they equal in length each year? YES NO.

Note the bud on the upper margin of each leaf scar. Since this bud occurs on the
side of the twig, it is called a lateral bud. When the leaf is present, this bud is in the axil
(upper angle) between the petiole and the stem, so it is also known as an axillary bud.
Lateral (axillary) buds are made of the same parts as a terminal bud.

Lateral buds are formed at the same time as the leaves of a twig. However they
do not grow when the leaves grow since they are inhibited by a hormone from the
terminal bud. As the twig lengthens, the distance between lateral buds and the terminal
bud increases. This results in less inhibition by the terminal bud, and lateral buds then
may grow and produce branches. Sometimes lateral buds fail to grow for an indefinite
period and become buried in the bark as the tree ages. These are called latent buds.
Under certain conditions they may start growth and form small leafy shoots on old large
limbs.

Examine the surface of your twig. Note that all over the surface there are
rounded, pinhead-like or linear, slit-like markings. These are the lenticels. Lenticels are
raised areas of loose cork. In trees with deeply furrowed barks they are not visible. They
are avenues for entry of air into the woody parts just as stomata admit air into leaves. A
heavy growth of moss, lichens, or vines on the trunk of a tree can interfere with its
lenticels and in extreme cases the tree may die.
BIOL 1507E Woody Stem and Root Structure

Woody twig
BIOL 1507E Woody Stem and Root Structure
B. Introduction to Woody Stem Anatomy
A woody plant is a permanent structure that yearly increases its leafy canopy
through primary growth. Supporting this canopy requires a reinforcement of the woody
framework. Secondary growth (growth that increases width) provides this reinforcement.
As the plant grows more massive through secondary growth, most of its cells become
buried by the added layers and cannot be supplied with oxygen. It would seem then that
there might be a physiological limit to the size that a woody plant can grow. It turns out
that the plant has no such limit. This dilemma has been “solved” by the way plants
evolved. Only the outer few layers of cells are alive, while those nearer the centre are in
an inactive state. Most of these inactive cells remain structurally intact and furnish the
additional support needed each year.

The woody stem has three basic parts, the bark, the vascular cambium and the
wood (Figure 1). These parts will be discussed in detail in the following sections.

1. Gymnosperm stem
Gymnosperms such as pines, hemlocks and spruce are ancient plants that bear
‘naked’ seeds not enclosed in any specialized chamber. Internally their stems are similar
to those of the woody angiosperms (flowering plants), but they are generally much
simpler and more homogeneous. They have no vessels or sieve tubes (they have less
advanced sieve cells), relatively little parenchyma, and many lack fibres. For this reason
you will find it easier to begin your study of woody stems with a gymnosperm.

a) Xylem cell types in a gymnosperm

Examine a prepared slide of macerated wood of a gymnosperm, pine (Pinus sp.).

This will enable you to see the general shape of the water-conducting tracheids and their
sidewalls; you will also see the ray parenchyma cells.

Locate a tracheid. A tracheid is an elongate, slender, needle-like cell with long,

tapering end walls. The cell is dead, lacks a protoplast (living component of a plant cell)
and has reinforced walls. Note the circular pits on the (radial) sidewalls. Pits are
permeable cavities in the wall. In the plant, tracheid cells overlap each other, with the
tapered end walls of one cell touching the radial wall of another. Water passes from
tracheid to tracheid through pores in the pits. This zigzag routing of water means that the
water moves slowly compared to the rates typical of angiosperms in which water moves
through large tube-like vessels.

Find a ray parenchyma cell. This is a parenchyma cell with a living protoplast
that stores foods. In the plant, the ray parenchyma cells border the tracheids at right
angles and connect with them through pits. They are much shorter cells than tracheids,
their length being only one to three times the width of a tracheid. Their end walls are not
tapered.
BIOL 1507E Woody Stem and Root Structure
b) Cross section of the woody stem of a gymnosperm

Examine a prepared slide showing a cross section of a three-year-old (or older)

stem of pine (Pinus sp.). Use your textbook or atlas as a guide to help you find the
tissues and cells. The main purpose is for you to see that the arrangement is an efficient
design. It integrates five different functions: support, conduction, storage, protection and
growth.

In ordinary use, the term “wood” means any hard part of a tree or shrub. But
botanically, wood is secondary xylem only. Water is conducted in the tracheid system.
Focus on the central portion of the stem where you can easily recognize concentric rings
of very orderly arranged cells. These are the tracheids that are more rectangular than
round in cross section. The rings are known as annual rings or growth rings, and each
represents one year’s formation of xylem. Each year, the vascular cambium adds a new
ring around that of the previous year. Note the gradation in size of the cells in any one
ring. Tracheids at the inner margin of each ring are much larger than those at its outer
edge. The larger tracheids were formed early in the growing season and are known as
earlywood or springwood. The smaller tracheids, formed later in the growing season,
are known as latewood or summerwood. The sharp contract between the last formed
latewood cells of one growing season and the first formed earlywood cells of the
following season delineates the boundary of a growth ring.

Most of the water moves up the plant in the outermost ring, that is, the current
season’s tracheids. The more recent of the previous years’ annual rings contain water,
but the plant does not rely on them for its current needs.

Switch to high power and examine the walls of the tracheids. Note that the walls
are relatively thick. These are the secondary walls that give the cells a rigid
reinforcement. They are also encrusted with a material known as lignin that makes them
hard and dense.

1. Which has the thicker walls compared to the overall cell size, latewood or earlywood?
2. Which do you suppose has the greater strength, latewood or earlywood?
3. Tracheids thus perform two functions: (a)____________________________

(b)____________________________

Note the single-file rows of thin-walled cells that radiate intermittently across the
growth rings somewhat like the spokes of a wheel. These radial rows are called rays.
Each ray is a sheet of parenchyma cells that extends vertically in the stem. Ray cells
store sugars, fats and water. In the spring these stored materials are transferred to the
actively dividing cambium or to the apical meristems.

Note also that many of these rays extend into the phloem that lies outside the
xylem. Rays in the phloem have the same structure and function as those in the xylem.
BIOL 1507E Woody Stem and Root Structure
As the tree enlarges, new rays are constantly formed by the cambium so that the density
of rays is maintained approximately the same.

Focus on the centre of the stem within the innermost growth ring. This central
core is the pith. Originally it contained parenchyma cells, and these may or may not still
be present in the stem section you are examining. The life span of the pith cells varies
with the species. With age, pith cells commonly accumulate tannins and crystals (waste
products). In many plants, the pith becomes hollow or chambered.

Switch to low power and observe the small cavities scattered throughout the stem.
These are the resin canals or resin ducts (not found in all gymnosperms). They extend
lengthwise in the stem. Switch to high power and note that the cavities are lined with
parenchyma cells. The parenchyma cells secrete a resinous substance that in pines is
known as oleoresin. Its usefulness to the plant is unknown. It may possibly repel or
attract insects. Turpentine and resin, which are manufactured from oleoresin, are still
used in some high-quality paints, though they have been largely replaced by synthetics.

The vascular cambium increases the girth of the stem by forming layer after
layer of xylem and phloem around the original primary cylinder. In the temperate zones
of the world cambial growth is cyclic with periods of activity alternating with periods of
relative rest. Cambial activity is under the control of hormones that reactivate it each
spring. Some of these hormones are known to come from the developing buds and
leaves.

Focus on the latewood of the outermost growth ring in the xylem. Immediately
outside this latewood you will find a band of flattened looking cells. This is the vascular
cambial zone a band of unexpanded meristematic cells. It is about four to eight cells
thick. It consists of one vascular cambium layer of self-perpetuating cells, called
cambial initials, flanked by layers of dividing daughter cells. Daughter cells on the
outside of the cambial layer are called phloem initials and develop into phloem cells;
those on the inside of the cambial layer are called xylem initials and develop into xylem
cells.

Bark is defined as all those tissues that lie outside of the vascular cambium. It
consists of three tissues: phloem, cortex (which may or may not be present), and the
periderm (present only in older stems). The phloem is considered to be the inner bark
and the periderm the outer bark.

Focus on the region just outside the vascular cambial zone. Here are angular,
thin-walled cells in somewhat orderly rows. Scattered among them are larger, round cells
filled with dark-colored material. Cells farther out are flattened and in a zigzag pattern.
If you have located tissue fitting these descriptions you have found the phloem. Locate
the following phloem cell types.

Sieve cells are vertically elongate cells, which in this cross section are the
angular, thin-walled cells in orderly rows. Sieve cells conduct the products of
BIOL 1507E Woody Stem and Root Structure
photosynthesis to actively growing buds and young leaves or to roots or storage rays.
Sieve cells are named for their numerous sieve areas, the porous zones in their walls.
Strands of protoplasm extending through these pores connect functioning adjacent sieve
cells and ray parenchyma. Transport in sieve cells is thought to occur in the form of a
flowing solution of material within the cytoplasm. The cytoplasm itself doesn’t flow.
Exactly how this is done is still unresolved.

Observe the outer regions of the phloem. These outer, older layers may still be
alive but are not all actively functioning. Many of the sieve cells are crushed and the rays
are distorted into a wavy shape. Normally only the innermost (youngest) layer of phloem
is functional. The outer, older layers are incorporated into the outer bark, which may
eventually be sloughed off.

Locate the round, thin-walled cells filled with dark-colored material. These are
the phloem parenchyma cells. When alive, these cells store organic products transferred
to them from adjacent or nearby sieve cells or other parenchyma. When these products
are mobilized for growth, they are recirculated back through sieve cells or through ray
parenchyma to the cambium and developing buds and leaves.

Focus on the inner (youngest) region of the phloem and find the large, radially
elongated cells with prominent nuclei. The length of one of these cells spans about five
of the adjacent sieve cells seen in cross section. These are the ray parenchyma cells.
They are lined up in a transverse row, called a ray. Can you identify any ray in the outer
crushed, nonfunctioning phloem? YES NO. Crushed rays give the old phloem a zigzag
pattern. Phloem rays are similar in form and function to xylem rays.

The cortex is primary tissue, produced by the apical meristem. In a young, non-
woody stem, one of the functions of the cortex is support, but as the plant ages, this
function is accomplished by the wood produced by secondary growth. Certain of its cells
may become meristematic again. These meristematic cells become organized into the
cork cambium (phellogen). As the tree gets even older, however, it is commonly old
phloem cells that become cork cambium.

The cork cambium (phellogen) produces cork cells (phellem) towards the
outside of the stem and thick-walled parenchyma cells, phelloderm towards the inside.
These three tissues, phellogen, phellem, and phelloderm, comprise the outer bark - the
periderm. You will not be able to identify the individual components of the outer bark
but rather you will identify the outermost layers that comprise the periderm. Mature cork
cells are dead and approximately square in outline with thick walls. They are usually
arranged compactly with no intercellular spaces. What advantage is this?

Cork cell walls are impregnated with suberin, a fatty material that makes them
impervious to water. Cork also has the thermal insulating qualities. If your pine stem
section came from a stem only three years old, the amount of periderm will be negligible.
BIOL 1507E Woody Stem and Root Structure
Prepare a LOW POWER PLAN of the cross section of Pinus sp. stem including the
following labels: periderm, phloem, vascular cambium, xylem, parenchyma ray,
pith, resin duct. A pie section drawing is appropriate for this slide. Draw it on the
upper half of the page.

Look at the tree trunk cross section under the dissection microscope.
Distinguish the dark brown bark from the lighter wood which is secondary xylem. The
bark is made up of periderm (outer bark) and phloem (inner bark) which are not
distinguishable one from the other. If you see purple lines in the bark it would be cork
cambium (phellogen). The vascular cambium, which is found between the phloem and
xylem, is not distinguishable on the trunk cross-section. Look at the lighter wood area.
You should be able to see xylem cells which are mostly tracheid. Observe the annular
growth rings. These are created by the difference in size between the xylem cells of
early or springwood and the late or summerwood. Count the annular rings to
determine the approximate age of the tree. Use the pointer and not a pen or pencil so as
not to mark the trunk cross section. Note that all of the rings are not the same size. Why
do you think this might be?

2. Angiosperm stem
The general organization of a woody angiosperm stem is essentially the same as
that of a gymnosperm. The main difference between them is that angiosperms have
many more specialized cells, particularly those cells concerned with water and organic
nutrient conduction and with support. In gymnosperm wood, tracheids provide much of
the support, in addition to conducting water. In angiosperm wood, more fibres are
present for support and, besides tracheids, there are vessel members (cells) that are
specialized for rapid water flow. The sieve tube cells of angiosperms can move organic
materials faster than the sieve cells of gymnosperms.

a) Xylem cell types in an angiosperm

Examine a prepared slide of macerated wood of an angiosperm, oak (Quercus

sp.). Note the tracheids as seen in the gymnosperm but look around to find the larger
vessel elements or vessel members which are mostly found in angiosperms. The vessel
members are larger than the tracheids and have thick lignified walls at maturity which
also have pits. The vessel elements have large openings at each end and are arranged end
to end to form large pipelines for the movement of water. Tracheids are tapered at each
end and water moves laterally through the pores.

b) Cross section of the woody angiosperm (eudicotyledon) stem

Examine a slide showing the cross section of some woody angiosperm stem such
as basswood (Tilia sp.). Tilia sp. is not representative of all woody angiosperms. Find
the phloem, which is the darker stained region near the outer part of the stem. It is made
BIOL 1507E Woody Stem and Root Structure
up of triangular-shaped sections. Those triangular sections that “point” inward are the
phloem rays. Other rays, which are only a few cells wide, form narrow rows of
parenchyma cells as they cross the phloem region in general. Note that all the rays in the
phloem are continuous with rays in the xylem (which is the homogeneous-looking central
core with the growth rings).

Edged between the inward-pointing phloem rays are other triangular sections that
“point” outward. In these sections are bands of heavily stained cells with very thick walls
and pin-point-like lumens (the central part of the cell bounded by the walls). These
heavily stained cells are the phloem fibers. Alternating with these bands of phloem
fibres are narrower bands of larger cells with thinner walls. These large, angular cells
with very little cytoplasm and with thin walls are the sieve tube members. They are
wide, columnar cells connected end to end in a longitudinal series to form sieve tubes.
The end walls of sieve tube members are not completely open as in vessel members but
have large porous end areas, called sieve plates. Sieve plates are not commonly seen on
these slides. Any one of the several sieve tube members that make up a sieve tube can
deliver materials at a rate of some five times its own volume per second. Hence the flow
through the entire tube is very fast.

Find the companion cells. These are smaller cells usually found at the corners of
sieve tube members, hence the name “companion cell”. They have thin walls,
considerable cytoplasm, and a nucleus that occupies most of the cell. The nuclei of
companion cells are thought to exert some control over the normal functioning of the
sieve tube members that have lost their nuclei in the process of cell maturation. The
absence of a nucleus in such a physiologically active cell as the sieve tube member is
inexplicable.

The centre of the cross section is occupied by a pith and three or more concentric
layers of xylem. Earlywood and latewood are easily recognized. A vascular cambial
zone separates the xylem and phloem. Basswood is a softwooded angiosperm because
its xylem has many more vessels than fibres. The structural difference between the
various xylem cells are not readily apparent in Tilia sp.. It is therefore, not as good an
example as oak for an introductory study of angiosperm wood (xylem tissue). Hence we
do not ask that you study the basswood xylem in detail. Simply note that, as in all woody
dicotyledons angiosperms and gymnosperms of the temperate zone, the xylem is in
concentric growth rings and ray parenchyma radiate across them.

Examine the outermost portion of a three-year-old Tilia sp. stem. The cortex is
the zone of loosely arranged, relatively large parenchyma cells found just external to the
outermost band of phloem fibres. Depending on how old the stem is, the cortex may
have been replaced by the periderm. Is the cortex a primary or a secondary tissue?

The periderm is the dense band of cells just under the stem surface. All the
periderm cells are rectangular, with their long axes paralleling the circumference of the
stem. From the phloem outward, the periderm consists of (a) four to six layers of
phelloderm cells that are thick-walled and filled with cytoplasm; the phelloderm is a
secondary cortex formed by the cork cambium; (b) one to two layers of thin-walled,
BIOL 1507E Woody Stem and Root Structure
nucleated cells filled with cytoplasm and making up the cork cambial zone (phellogen);
and (c) three to four scale-like layers of dead cork cells (phellum).

Prepare a LOW POWER PLAN of the cross section of Tilia sp. stem including the
following labels: periderm, phloem ray, phloem fibers, vascular cambium, xylem,
parenchyma ray, pith. A pie section drawing is appropriate for this slide. Use the
bottom half of the page with the gymnosperm stem drawing.

C. Cross section of the Woody Angiosperm

(Dicotyledon) Root
Study a prepared slide of a cross section of a woody root of an angiosperm such
as pear (Pyrus sp.). This sample section is a young woody root only a few years old.
When this root was less than one year old, it had the same structure as the buttercup root.
Here, though, secondary xylem has been laid down by the cambium in concentric annual
rings similar to those in stems. The regularity of the rings in these very young woody
roots is, however, not very clear cut and may be difficult for you to recognize. Secondary
phloem lies outside the xylem and is not as distinct as the clusters of primary phloem you
saw in younger roots. Cortex is eventually replaced by cork and/or wood, but how soon
this happens varies with different species.

Prepare a LOW POWER PLAN of the cross section of Pyrus sp. root including the
following labels: periderm, phloem, vascular cambium, xylem, parenchyma ray. A
pie section drawing is appropriate for this slide. Do the drawing on the upper half of the
back side of the stem drawings.

D. Cross section of the Woody Gymnosperm Root

Study a prepared slide of a pine (Pinus sp.) root. Note the two large resin canals
near the centre. Between these is the primary xylem. Surrounding the primary xylem is
the zone of secondary xylem with, at least one annual growth ring, and scattered resin
canals. Find the ring of small cambium cells (stained light blue) outside the xylem.
Locate the secondary phloem (with tiers of cells stained blue) and the rays of parenchyma
that transect the phloem and secondary xylem. The outside of the root is surrounded by a
young periderm (red) that has arisen in the cortex. The loose fragments are the remains
of the epidermis.

Prepare a LOW POWER PLAN of the cross section of Pinus sp. root including the
following labels: periderm, phloem, vascular cambium, xylem, parenchyma ray,
and resin ducts. A pie section drawing is appropriate for this slide. Do the drawing on
the bottom half of the page below the angiosperm root drawings.

1. What similarities and what differences do you find between all four sections?
BIOL 1507E Woody Stem and Root Structure

Figure 1 Parts and functions of the components of a woody stem

Use the following table to identify the nine different plant cells types and fill in their
characteristics. In the Location column write down in which plant part you have found
these tissues and cell types. Use this table to help you study.
BIOL 1507E Woody Stem and Root Structure

Tissue Cell type Characteristics Function

type
Dermal 1.

. 2.

Ground 3.

4.

5.

Vascular 6.

7.

8.

9.