C H A P T E R

Stems

Overview
Learning Outcomes
External Form of a Woody Twig
Origin and Development of Stems
ECOLOGY: Standing in Fields of Stone
Tissue Patterns in Stems
Steles
Herbaceous Dicotyledonous Stems
Woody Dicotyledonous Stems
ECOLOGY: Dendroclimatology
Monocotyledonous Stems
Specialized Stems
Rhizomes
Runners and Stolons
Tubers
Bulbs
Corms
Cladophylls
Other Specialized Stems
Wood and Its Uses
Properties of Wood
SUMMARY
REVIEW QUESTIONS
DISCUSSION QUESTIONS
ADDITIONAL READING
LEARNING ONLINE

Tree trunks of giant sequoia (Sequoiadendron

giganteum) found only in Sequoia National Park
within the Sierra Nevada mountain range of
California; they are the bulkiest trees in the world.
A member of a related species, coastal redwood
(Sequoia sempervirens), which grows only along
the Pacific Coast, is the tallest tree in the world.
Together, these two tree species share the title of
State Trees of California.
© Doug Sherman/Geofile
OVERVIEW

After a brief introduction, this chapter discusses the origin and development of stems. Structures such as the apical meristem and the tissues
derived from it, including leaf gaps, cambia, secondary tissues, and lenticels are included. This general discussion is followed by notes on the
distinctions between herbaceous and woody dicot stems and monocot stems. This section covers annual rings, rays, heartwood and sapwood,
resin canals, bark, laticifers, and vascular bundles.
Next, there is a survey of specialized stems (rhizomes, stolons, tubers, bulbs, corms, cladophylls, and others). The chapter concludes
with a discussion of the economic importance of wood and stems.

Learning Outcomes
1. Name the tissues that develop from shoot apices and the 4. Describe the composition of wood and its annual rings,
meristems from which various tissue types are derived. sapwood, heartwood, and bark. Explain how a log is sawed
State the differences between primary tissues and secondary for commercial use.
tissues. 5. Provide descriptions of rhizomes, stolons, tubers, bulbs,
2. List and describe the function of each of the following: corms, cladophylls, and tendrils.
vascular cambium, cork cambium, stomata, lenticels. 6. List at least 10 human uses of wood and stems in general.
3. Describe the differences in the development of dicot and
monocot stems.

A
brief check of furnishings and tools around the house External Form
and garage—or even the house itself—soon reveals
that much of what we use daily and take for granted,
of a Woody Twig
from pencils and pianos to newspapers and brooms, has A woody twig consists of an axis with attached leaves (Fig. 6.1).
some wood content. Most of that wood comes from plant If the leaves are attached to the twig alternately or in a spiral
stems. In fact, stems have been an integral part of human around the stem, they are said to be alternate, or alternately
life ever since cave dwellers first used wooden clubs to kill arranged. If the leaves are attached in pairs, they are said to
for food. be opposite, or oppositely arranged, or if they are in whorls
Grafting, which usually involves artificially uniting (groups of three or more), their arrangement is whorled. The
stems or parts of stems of different but related varieties area, or region (not structure), of a stem where a leaf or leaves
of plants, has been practiced by humans for hundreds of are attached is called a node, and a stem region between nodes
years. The careful matching of certain tissues is critical is called an internode. A leaf usually has a flattened blade and
to its success, as is seen in the discussion of grafting in in most cases is attached to the twig by a stalk called the petiole.
Chapter 14 and in Appendix 4. To understand how and Each angle between a petiole and the stem contains a
why grafts may or may not be successful and to identify bud. The angle is called an axil, and the bud located in the
which parts of stems are useful, we first need to examine axil is an axillary bud. In flowering plants (angiosperms),
the structure of stems and learn the basic functions of the axillary buds may become branches, or they may contain tis-
various tissues. sues that will develop into the next season’s flowers. Most
Unlike animals, some plants have indeterminate growth buds are protected by one to several bud scales, which fall
(i.e., they can grow indefinitely), with the meristems at their off when the bud tissue starts to grow.
tips increasing their length and other meristems increas- There often (but not always) is a terminal bud present at
ing their girth for hundreds or even thousands of years. In the tip of each twig. A terminal bud usually resembles an axil-
stems, the cells produced by the meristems usually become lary bud, although it is often a little larger. Unlike axillary buds,
the familiar, erect, aerial shoot system with branches and terminal buds do not become separate branches, but, instead,
leaves. In certain plants, such as ferns or perennial grasses, the meristems within them normally produce tissues that make
this shoot system may develop horizontally beneath or at the twig grow longer during the growing season. The bud
the surface of the ground; in other plants, the stem may be scales of a terminal bud leave tiny scars around the twig when
so short and inconspicuous as to appear nonexistent. In a they fall off in the spring. Counting the number of groups of
number of plants, stems are modified in ways that allow bud scale scars on a twig can tell one how old the twig is.
specialized functions, such as climbing or the storage of Sometimes other scars of different origin also occur
food or water. on a twig. These scars come from a leaf that has stipules at

86
 Stems 87

terminal
bud

bundle axillary
scar bud
leaf scar

node

internode

lenticel

blade
petiole
bud scale scars

stipules
(a) (b)

Figure 6.1 A woody twig. (a) The twig in its winter condition. (b) The twig as it appeared the summer before.

the base of the petiole. Stipules are paired, often somewhat meristem is dormant before the growing season begins. It is
leaflike, appendages that may remain throughout the life of protected by bud scales of the bud in which it is located and to
the leaf. In some plants, they fall off as the buds expand in a certain extent by leaf primordia (singular: primordium),
the spring, leaving tiny stipule scars. The stipule scars may the tiny embryonic leaves that will develop into mature leaves
resemble a fine line encircling the twig, or they may be very after the bud scales drop off and growth begins. The apical
inconspicuous small scars on either side of the petiole base. meristem in the embryonic stem of a seed is also dormant
Deciduous trees and shrubs (those that lose all their leaves until the seed begins to germinate.
annually) characteristically have dormant axillary buds with When a bud begins to expand or a seed germinates, the
leaf scars left below them after the leaves fall. Tiny bundle cells of the apical meristem undergo mitosis, and soon three
scars, which mark the location of the water-conducting and primary meristems develop from it (see Figs. 4.1 and 6.2).
food-conducting tissues, are usually visible within the leaf The outermost of these primary meristems, the protoderm,
scars. There may be one to many bundle scars present, but more gives rise to the epidermis. Although there are exceptions,
often than not, there are three. The shape and size of the leaf the epidermis is typically one cell thick and usually becomes
scars and the arrangement and numbers of the bundle scars are coated with a thin, waxy, protective layer, the cuticle. A cyl-
characteristic for each species. One can often identify a woody inder of strands constituting the procambium appears to the
plant in its winter condition by means of scars and buds. interior of the protoderm. (The procambium produces water-
conducting primary xylem cells and primary phloem cells that

Origin and Development have several functions, including the conduction of food.)
The remainder of the meristematic tissue, called ground
of Stems meristem, produces two tissues composed of parenchyma
cells. The parenchyma tissue in the center of the stem is the
There is an apical meristem (tissue in which cells actively pith. Pith cells tend to be very large and may break down
divide) at the tip of each stem, and it is this meristem that con- shortly after they are formed, leaving a cylindrical, hollow area.
tributes to an increase in the length of the stem. The apical Even if they do not break down early, they may eventually be
KEY THEME : ecology Standing in Fields of Stone

T he combination of high altitude; dry, cool air; low rainfall;

high winds; and poor soil has provided sustenance for the
oldest known living species on the earth! These ancient
warriors, whose great age was unknown until 1953, are the bris-
tlecone pines that flourish atop the arid mountains of the Great
Basin from Colorado to California. The oldest—almost 5,000
years old—is located in the Ancient Bristlecone Pine Forest high
in the White Mountains of California. Some of the trees standing
today were seedlings when the great pyramids were built, were
middle-aged trees during the time of Christ, and today are hoary
patriarchs standing in fields of stone.
There are two species of bristlecones (Box Figure 6.1), one
living in the westernmost regions (Pinus longaeva) and the other
inhabiting the eastern regions (Pinus aristata). The trees do not Box Figure 6.1 An ancient bristlecone pine (Pinus aristata)
from the White Mountains of California. © Stephen Saks/Getty Images
grow very tall, with none over 60 feet and many much shorter.
Typical of their girth is the bristlecone named the “Patriarch,” which bristlecone over 4,000 years old is nearly 4 feet in diameter, but
is just over 36.5 feet around, but it is a relative youngster at 1,500 its functional conducting tissue is a ring only 10 inches wide.
years. With the short summer growing season high in these Other characteristics provide bristlecones with survival
mountains, bristlecones typically grow 1/100th of an inch or less advantages. Because their dense, resin-filled wood renders
in diameter in any given year. The trees stand in isolation, each their sites inhospitable for colonization by pathogenic fungi or
appearing as a sentinel, overlooking an otherwise barren rock- bacteria, they are relatively free from these attacks. These trees
strewn landscape. Their needles are remarkable. While occurring are often struck by lightning, but with the absence of ground
five per bundle and about 1 to 1.5 inches in length, they can live cover and decaying leaf litter, fire rarely spreads from tree to tree.
for 20 to 30 years before being cast off. This extraordinary leaf As with other trees, the age of these living trees is
longevity gives the trees a stable photosynthetic output and determined by an instrument called an increment borer, a
sustains the tree during years of unusual stress when producing type of drill that is inserted into the tree trunk at its widest girth.
new leaf tissue would be difficult. The trees must generate new Using a hollow drill bit, a linear core of wood is obtained that
leaves and cones as well as produce enough reserves for the long can be “read” for the number of annual rings. In this way, the
winter months, all on scant annual precipitation of about 10 inches. age of a tree can be obtained without cutting the tree down to
Ecologists have long noted the peculiar distribution and examine the annual rings revealed in the cross section of the
growth habits of certain plants. Why do plants grow where they stump. Each year, a tree will add a layer of wood to its trunk, and
do? What adaptations permit them to survive in life-threatening these layers become the annual rings that can be observed in a
environments? The bristlecones’ age and habitat offer several cross section of the trunk. During spring growth of wood, large-
insights. These trees grow in places on earth where no other diameter water-conducting cells are formed; later in the summer,
plant can grow. One answer appears to be the type of soil in the water-conducting cells produced have a smaller diameter.
which they are anchored. Stands of bristlecone pine grow on This difference in appearance between early (spring) wood and
outcrops of dolomite, an alkaline limestone substrate low in late (summer) wood is sufficient to make each growth increment
nutrients but of higher moisture content than the surrounding distinctive, and thus counting of the rings possible.
sandstone. The granite and sandstone formations surrounding In 1957, the bristlecone pine “Methuselah” was discovered
the dolomite outcroppings support sagebrush and Limber pine, and determined to be 4,723 years old. Methuselah remains
but not bristlecones. At these altitudes, the radiant sunlight is today as the world’s oldest living organism. But what about
extreme. Dolomite reflects more sunlight than other rocks and tomorrow? After surviving nearly five millennia, Methuselah is
thus keeps the root zones cooler and more moisture-laden being protected against a more insidious enemy. Standing in
during the important growing season. its field of stone without a marker because of fear of vandalism,
Another survival tactic is revealed by taking small core Methuselah serves as a reminder that the human species can be
samples of the wood. These trees have large amounts of die-back just as destructive as any microbe.
(deadwood that is no longer functional), reducing the amount of
tissue that the leaves need to supply with food. For example, one D . C. Scheirer

88
 Stems 89

leaf
primordium
procambium
protoderm

apical
meristem
bud
primordium
ground
meristem

base of leaf
encircling stem

Figure 6.2 A longitudinal section through the tip of a Coleus stem. ×800. Photomicrograph by G.S. Ellmore

crushed as new tissues produced by other meristems add to the

girth of the stem, particularly in woody plants. The other tis-
sue produced by the ground meristem is the cortex. The cor-
tex may become more extensive than the pith, but in woody
plants, it, too, eventually will be crushed and replaced by new
tissues produced from within. The parenchyma of both the pith
and the cortex function in storing food or sometimes, if chloro-
plasts are present, in manufacturing it.
All five of the tissues produced by this apical meristem
complex (epidermis, primary xylem, primary phloem, pith, bud gap
and cortex) arise while the stem is increasing in length and axillary bud
are called primary tissues. As these primary tissues are pro-
duced, the leaf primordia and the bud primordia (embryonic
buds in the axils of the leaf primordia) develop into mature
leaves and buds (Fig. 6.2). As each leaf and bud develop,
leaf
a strand of xylem and phloem, called a trace, branches off leaf gap
from the cylinder of xylem and phloem extending up and leaf scar leaf gap
down the stem and enters the leaf or the bud. As the traces vascular
tissue
branch from the main cylinder of xylem and phloem, each
trace leaves a little, thumbnail-shaped gap in the cylinder of bark
vascular tissue. These gaps, called leaf gaps and bud gaps,
are filled with parenchyma tissue (Fig. 6.3).
A narrow band of cells between the primary xylem and Figure 6.3 A portion of a stem showing leaf gaps and bud
the primary phloem may retain its meristematic nature and gaps in the cylinder of vascular tissue.
90 Chapter 6

time
same
o f s t e l o ca ti o n
m
a t a l o r r oo
a te r t
te m t ime
gs t
un
yo r ro o surface of stem or root
o immature phloem cell

cell of vascular cambium

new cell produced

by cambium
center of stem or root
immature xylem cell

Figure 6.4 An illustration of how a cell of the vascular cambium produces new secondary phloem cells to the outside and new secondary
xylem cells to the inside. Note, in cross section, that the cambium gradually becomes shifted away from the center as new cells are produced.
Phloem is produced before xylem in secondary growth.

become the vascular cambium, one of the two lateral meri- of gases (e.g., oxygen, carbon dioxide) can continue through
stems. The vascular cambium is often referred to simply as spaces between the cells. Lenticels occur in the fissures of
the cambium. The cells of the cambium continue to divide the bark of older trees and often appear as small bumps on
indefinitely, with the divisions taking place mostly in a plane younger bark. In birch and cherry trees, the lenticels form
parallel to the surface of the plant. The secondary tissues conspicuous horizontal lines.
produced by the vascular cambium add to the girth of the Differences between the activities of the apical meristem
stem instead of to its length (Fig. 6.4). and those of the cambium and cork cambium become appar-
Cells produced by the vascular cambium become tra- ent if one drives a nail into the side of a tree and observes
cheids, vessel elements, fibers, or other components of second- it over a period of years. The nail may eventually become
ary xylem (inside of the meristem, toward the center), or they embedded as the stem increases in girth, but it will always
become sieve tube members, companion cells, or other com- remain at the same height above the ground, as the cells that
ponents of secondary phloem (outside of the meristem, toward increase the length of a stem are produced only at the tips.
the surface). The functions of these secondary tissues are
the same as those of their primary counterparts—secondary
xylem conducts water and soluble nutrients, while secondary Tissue Patterns in Stems
phloem conducts, in soluble form, food manufactured by photo
synthesis throughout the plant.
In many plants, especially woody species, a second cam-
Steles
bium arises within the cortex or, in some instances, devel- Primary xylem, primary phloem, and the pith, if present,
ops from the epidermis or phloem. This is called the cork make up a central cylinder called the stele in most younger
cambium, or phellogen. The cork cambium produces box- and a few older stems and roots. The simplest form of stele,
like cork cells, which become impregnated with suberin, a called a protostele, consists of a solid core of conducting
waxy substance that makes the cells impervious to moisture. tissues in which the phloem usually surrounds the xylem.
The cork cells, which are produced annually in cylindrical Protosteles were common in primitive seed plants that are
layers, die shortly after they are formed. The cork cambium now extinct and are also found in whisk ferns, club mosses
may also produce parenchyma-like phelloderm cells to the (see Chapter 21), and other relatives of ferns. Siphonosteles,
inside. Cork tissue makes up the outer bark of woody plants; which are tubular with pith in the center, are common in ferns.
it functions in reducing water loss and in protecting the stem Most present-day flowering plants and conifers have
against mechanical injury (see the discussion of periderm in eusteles in which the primary xylem and primary phloem
Chapter 4). are in discrete vascular bundles, as discussed in the section
Cork tissue cuts off water and food supplies to the epi- “Herbaceous Dicotyledonous Stems.”
dermis, which soon dies and is sloughed off. In fact, if the Flowering plants develop from seeds that have either
cork were to be formed as a solid cylinder covering the entire one or two “seed leaves,” called cotyledons, attached to their
stem, vital gas exchange with the interior of the stem would embryonic stems (see Chapters 8 and 23). The seeds of pines
not be possible. In young stems, such gas exchange takes and other cone-bearing trees have several (usually eight) coty-
place through the stomata, located in the epidermis (see ledons. The cotyledons usually store food needed by the young
Figs. 7.6 and 9.13). As woody stems age, lenticels (see Fig. seedling until its first true leaves can produce food themselves.
4.14) develop beneath the stomata. As cork is produced, the Flowering plants that develop from seeds having two coty-
parenchyma cells of the lenticels remain, so that exchange ledons are called dicotyledons (usually abbreviated to dicots),
 Stems 91

epidermis epidermis

cortex
cortex phloem
fibers
phloem-
conducting
vascular cells
bundle vascular
pith cambium
xylem

pith

(a) (b)

Figure 6.5 (a) A cross section of an alfalfa (Medicago) stem. ×40. The tissue arrangement is typical of herbaceous dicot stems. (b) An
enlargement of a small portion of the outer part of the stem. ×400. (a-b) © James E. Bidlack

while those developing from seeds with a single cotyledon are The most conspicuous differences involve the secondary
called monocotyledons (abbreviated to monocots). Dicots and xylem, or wood, as it is best known (Fig. 6.6). Some tropical
monocots differ from one another in several other respects; the trees (e.g., ebony), in which both the vascular cambium and
differences in stem structure are noted in the following sections, the cork cambium are active all year, produce an ungrained,
and a summary of these and other differences in these two uniform wood. The wood of most trees, however, is pro-
groups of flowering plants is given in Table 8.1. duced seasonally. In trees of temperate climates, virtually all
growth takes place during the spring and summer and then
ceases until the following spring.
Herbaceous Dicotyledonous Stems When the vascular cambium of a typical broadleaf tree
In general, plants that die after going from seed to maturity first becomes active in the spring, it usually produces rela-
within one growing season (annuals) have green, herba- tively large vessel elements of secondary xylem; such xylem
ceous (nonwoody) stems. Most monocots are annuals, but is referred to as spring wood. As the season progresses,
many dicots (discussed next) are also annuals. the vascular cambium may produce vessel elements whose
The tissues of annual dicots are largely primary, diameters become progressively smaller in each succeed-
although cambia (plural of cambium) may develop some sec- ing series of cells produced, or there may be fewer vessel
ondary tissues. Herbaceous dicot stems (Fig. 6.5) have dis- elements in proportion to tracheids produced until tracheids
crete vascular bundles composed of patches of xylem and (and sometimes fibers) predominate.
phloem. The vascular bundles are arranged in a cylinder that The xylem that is produced after the spring wood, and
separates the cortex from the pith, although in a few plants which has smaller or fewer vessel elements and larger num-
(e.g., foxgloves), the xylem and the phloem are produced as bers of tracheids, is referred to as summer wood. Over a
continuous rings (cylinders) instead of in separate bundles. period of years, the result of this type of switch between the
The procambium produces only primary xylem and early spring and the summer growth is a series of alternating
phloem, but later, a vascular cambium arises between these concentric rings of light and dark cells. One year’s growth
two primary tissues and adds secondary xylem and phloem of xylem is called an annual ring. In conifers, the wood
to the vascular bundles. In some plants, the cambium consists mostly of tracheids, with vessels and fibers being
extends between the vascular bundles, appearing as a nar- absent. Annual rings are still visible, however, because the
row ring, producing not only the conducting tissues within first tracheids produced in the spring are considerably larger
the bundles but also the parenchyma cells between them. In and lighter in color than those produced later in the growing
other plants, the cambium is not in an uninterrupted cylinder season. Note that an annual ring normally may contain many
but is instead confined to the bundles, each of which has its layers of xylem cells and it is all the layers produced in one
own small band of cambium between the xylem and phloem. growing season that constitute an annual ring—not just the
dark layers.
The vascular cambium produces more secondary xylem
Woody Dicotyledonous Stems than it does phloem. Xylem cells also have stronger, more
In the early stages of development, the primary tissues of rigid walls than those of phloem cells and are less subject to
stems of young herbaceous dicots, woody dicots, and cone- collapse under tension. As a result, the bulk of a tree trunk
bearing trees are all arranged in a similar fashion. In woody consists of annual rings of wood. The annual rings not only
plants, however, obvious differences begin to appear as soon indicate the age of the tree (because, normally, only one is
as the vascular cambium and the cork cambium develop. produced each year), but they can also tell something of the
92 Chapter 6

climate and other conditions that occurred during the

tree’s lifetime (Fig. 6.7). For example, if the rainfall dur-
ing a particular year is higher than normal, the annual ring
for that year will be wider than usual because of increased
growth. Sometimes, caterpillars or locusts will strip the
leaves of a tree shortly after they have appeared. This usu-
ally results in a narrow annual ring, because very little
growth can take place under such conditions.
If there is a fire that doesn’t result in the death of
the tree, it may be possible to determine when the fire
occurred, because the burn scar may appear next to a
given ring. The most recent season’s growth is next to
the vascular cambium, and one need only count the
rings back from the cambium to determine the actual
year of the fire.
It is not necessary to cut down a tree to determine its
age. Instead, botanists and foresters can employ an incre-
ment borer to find out how old a woody plant is. This
device, which resembles a piece of pipe with a handle on
one end, removes a plug of wood from the tree perpen-
dicular to the axis, and the annual rings in the plug can
then be counted. The small hole left in the tree can be
treated with a disinfectant to prevent disease and covered
up without harm to the tree.
A count of annual rings has produced some red faces
on at least one occasion. The Hooker Oak, which was
named in honor of Sir Joseph Hooker, a famous British
botanist who once examined it, was located in the com-
munity of Chico, California. Until its demise in 1977,
thousands of visitors from all over the world visited the
huge oak, which provided enough shade for 9,000 people
on a summer day. A plaque indicating the tree to be over
1,000 years old was located beneath the tree. A count of
rings after its death, however, revealed that the Hooker
Oak was less than 300 years old.
When a tree trunk is examined in transverse, or cross
section, fairly obvious lighter streaks or lines can be seen
radiating out from the center across the annual rings (see
Figure 6.6 A cross section of a portion of a young linden (Tilia) Figs. 6.6 and 6.8). These lines, called vascular rays,
stem. ×300. © Kingsley Stern

KEY THEME : ecology

Dendroclimatology
Dendroclimatology is the science of determining climate history by June of that year. During a good growing season, a tree lays down
examining annual growth rings of trees. These annual rings provide a wide ring of growth, with dense wood in late summer. During
a temporal record of environmental conditions and can be used the summer of 1783, annual rings were normal at the beginning
to date important events in history. For example, Alaskan Inuit oral of the summer, but the density of the wood was unusually low in
history tells of a disaster described as “the time that summertime late summer. This indicates that summer ended quickly, before the
did not come.” Many people were said to have died as a result of trees could produce dense late summer wood. The Laki volcanic
this cold period and the famine that followed. Dendroclimatology eruption in Iceland resulted in low temperatures throughout the
data have shown that, in fact, a dramatic climate event occurred in Northern Hemisphere in 1783, and apparently had an especially
1783, probably as a result of a large volcanic eruption in Iceland in severe effect on the people of northwest Alaska.
 Stems 93

cork
cork cambium
phelloderm
cortex
primary phloem

broad phloem ray

primary phloem
narrow phloem ray

vascular cambium secondary phloem

annual ring of xylem

broad xylem ray

secondary xylem
narrow xylem rays (wood)

primary xylem

pith

Figure 6.6 Continued.

consist of parenchyma cells that may remain alive for 10 or

more years. Their primary function is the lateral conduction
of nutrients and water from the stele, through the xylem and
phloem, to the cortex, with some cells also storing food. Any
part of a ray within the xylem is called a xylem ray, while its
extension through the phloem is called a phloem ray. In trees
such as basswood (Tilia), some of the phloem rays, when
observed in cross section, flare out from a width of two or
three cells near the cambium to many cells wide in the part
next to the cortex (see Fig. 6.6).
In radial section, rays may be from two or three cells to
50 or more cells deep, but the majority of rays in both xylem
Figure 6.7 Colored scanning electron micrograph of a cross-section and phloem are one or two cells wide. Ray cells can be seen
through a small stem. Note the series of concentric rings radiating out in cross section if a woody stem is cut or split lengthwise
from the center (top right) of the stem. The apparent “holes” are tracheary along a ray (Fig. 6.8). Another view of rays (in tangential
elements, and vary in size depending on environmental conditions each section) is obtained when the stem is cut at a tangent (i.e.,
year, and hence form the annual rings. © Clouds Hill Imaging Ltd./Science Source cut lengthwise and off center) (see Fig. 6.17).
94 Chapter 6

transverse surface
vessel

ray

tangential
bark surface

tracheid

fiber vessel
sieve-tube member
phloem vascular cambium
ray

radial surface xylem

Figure 6.8 A three-dimensional, magnified view of a block of a woody dicot.

As a tree ages, the protoplasts of some of the parenchyma

cells that surround the vessels and tracheids grow through the
pits in the walls of these conducting cells and balloon out into
the cavities. As the protoplasm continues to expand, much of
the cavity of the vessel or tracheid becomes filled. Such pro-
trusions, called tyloses (singular: tylosis), prevent further con-
duction of water and dissolved substances. When this occurs, heartwood
resins, gums, and tannins begin to accumulate, along with
pigments that darken the color of the wood.
This older, darker wood at the center is called heart-
wood, while the lighter, still-functioning xylem closest to sapwood
the cambium is called sapwood (Fig. 6.9). Except for giving
strength and support, the heartwood is not of much use to
the tree, because it can no longer conduct materials. A tree (a)
may live and function perfectly well after the heartwood has
rotted away and left the interior hollow. It is even possible to
remove part of the sapwood and other tissues and apparently
not affect the tree very much, as has been done with giant
trees, such as the coastal redwoods of California, where
holes big enough to drive a car through have been cut out heartwood
without killing the trees (Fig. 6.10). sapwood
Sapwood forms at roughly the same rate as heartwood
vascular
develops, so there is always enough “plumbing” for the cambium
vital conducting functions. The relative widths of the two phloem
types of wood, however, vary considerably from species to
species. For example, in the golden chain tree (a native of cork
Europe and a member of the Legume Family), the sapwood
is usually only one or two rings wide, while in several North
(b)
American trees (e.g., maple, ash, and beech), the sapwood
may be many rings wide. Figure 6.9 (a) A cross section of a 39-year-old larch tree
Pines and other cone-bearing trees have xylem that con- (Larix decidua) showing the darker heartwood to the inside and
sists primarily of tracheids; no fibers or vessel elements are lighter sapwood to the outside. (b) Longitudinal section showing
produced. Because it has no fibers, the wood tends to be softer heartwood and sapwood in relation to other tissues of a tree trunk.
 Stems 95

resin canals

Figure 6.11 Resin canals in a portion of a pine (Pinus) stem.

×100. © James E. Bidlack

The parenchyma cells of the cortex to the outside of the

phloem also function only briefly because they, too, become
crushed or sloughed off. Before they disappear, however, the
cork cambium begins its production of cork, and because new
xylem and phloem tissues produced by the vascular cambium
arise to the inside of the older phloem, the mature bark may
consist of alternating layers of crushed phloem and cork.
The younger layers of phloem nearest to the cambium
Figure 6.10 This coastal redwood is thriving despite the removal transport, via their sieve tubes, sugars and other substances
of its lower heartwood and a little of its sapwood. Cork cells of bark in solution from the leaves where they are made to various
are dead and their removal would not a ect growth. © Kingsley Stern parts of the plant, where they are either stored or used in the
process of respiration (discussed in Chapter 10). The pres-
than wood with fibers and is commonly referred to as softwood, ence of sugar in the phloem was recognized in the past by
while the wood of woody dicot trees is called hardwood. Native Americans. Some stripped the young phloem and
In many cone-bearing trees, resin canals are scattered cambium from Douglas fir trees and used the dried strips as
not only through the xylem but throughout other tissues food for winter and in emergencies.
as well. These canals are tubelike and may or may not be Specialized cells or ducts called laticifers are found
branched; they are lined with specialized cells that secrete in about 20 families of herbaceous and woody flowering
resin (discussed in Chapter 22) into their cavities (Fig. 6.11). plants. These cells are most common in the phloem but
Although resin canals are commonly associated with cone- occur throughout all parts of the plants. The laticifers, which
bearing trees, they are not confined to them. Tropical flow- resemble vessels, form extensive branched networks of
ering plants, such as olibanum and myrrh trees, have resin latex-secreting cells originating from rows of meristematic
ducts in the bark that produce the soft resins frankincense cells. Unlike vessels, however, the cells remain living and
and myrrh of biblical note. may have many nuclei.
While the vascular cambium is producing secondary Latex is a thick fluid that is white, yellow, orange, or
xylem to the inside, it is also producing secondary phloem to red in color and consists of gums, proteins, sugars, oils,
the outside. The term bark is usually applied to all the tissues salts, alkaloidal drugs, enzymes, and other substances. Its
outside the cambium, including the phloem. Some scientists function in the plant is not clear, although some believe it
distinguish between the inner bark, consisting of primary aids in closing wounds. Some forms of latex have consid-
and secondary phloem, and the outer bark (periderm), con- erable commercial value (see the discussion under “The
sisting of cork tissue and cork cambium. Despite the pres- Spurge Family” in Chapter 24). Of these, rubber is the most
ence of fibers, the thin-walled conducting cells of the phloem important. Amazon Indians utilized rubber for making balls
usually are not able to withstand for many seasons the pres- and containers hundreds of years before Pará rubber trees
sure of thousands of new cells added to their interior, and the were cultivated for their latex. The chicle tree produces a
older layers become crushed and functionless. latex used in the making of chewing gum. Several poppies,
96 Chapter 6

notably the opium poppy, produce latex-containing, impor- sclerenchyma cells beneath the epidermis and the thicker-
tant drugs, such as morphine and its derivative, known as walled parenchyma cells, all contribute to giving the stem
heroin. Other well-known latex producers include milk- the capacity to withstand stresses resulting from summer
weeds, dogbanes, and dandelions. storms and the weight of the leaves and the ears of corn as
they mature.
Monocotyledonous Stems In wheat, rice, barley, oats, rye, and other grasses, there
is an intercalary meristem (discussed in Chapter 4) at the
Most monocots (e.g., grasses, lilies) are herbaceous plants base of each internode; like the apical meristem, it contrib-
that do not grow tall. The stems have neither a vascular cam- utes to increasing stem length. Although the stems of such
bium nor a cork cambium and thus produce no secondary plants elongate rapidly during the growing season, growth is
vascular tissues or cork. As in herbaceous dicots, the surfaces columnar (i.e., there is little difference in diameter between
of the stems are covered by an epidermis, but the xylem and the top and the bottom) because there is no vascular cambium
phloem tissues produced by the procambium appear in cross producing tissues that would add to the girth of the stems.
section as discrete vascular bundles scattered throughout the Palm trees, which differ from most monocots in that
stem instead of being arranged in a ring (Fig. 6.12). they often grow quite large, do so primarily as their paren-
Each bundle, regardless of its specific location, is ori- chyma cells continue to divide and enlarge without a true
ented so that its xylem is closer to the center of the stem cambium developing. Several popular house plants (e.g.,
and its phloem is closer to the surface. In a typical monocot, ti plants, Dracaena, Sansevieria) are monocots in which
such as corn, a bundle’s xylem usually contains two large a secondary meristem develops as a cylinder that extends
vessels with several small vessels between them (Fig. 6.13). throughout the stem. Unlike the vascular cambium of dicots
The first-formed xylem cells usually stretch and collapse and conifers, this secondary meristem produces only paren-
under the stresses of early growth and leave an irregularly chyma cells to the outside and secondary vascular bundles to
shaped air space toward the base of the bundle; the remnants the inside.
of a vessel are often present in this air space. The phloem Several commercially important cordage fibers (e.g.,
consists entirely of sieve tubes and companion cells, and the broomcorn, Mauritius and Manila hemps, sisal) come from
entire bundle is surrounded by a sheath of thicker-walled the stems and leaves of monocots, but the individual cells are
sclerenchyma cells. The parenchyma tissue between the vas- not separated from one another by retting (a process that uti-
cular bundles is not separated into cortex and pith in mono- lizes the rotting power of microorganisms thriving under moist
cots, although its function and appearance are the same as conditions to break down the thin-walled parenchyma cells) as
those of the parenchyma cells in cortex and pith. they are when fibers from dicots are obtained. Instead, during
In a corn stem, there are more bundles just beneath the commercial preparation, entire vascular bundles are scraped
surface than there are toward the center. Also, a band of free of the surrounding parenchyma cells by hand; the indi-
sclerenchyma cells, usually two or three cells thick, devel- vidual bundles then serve as unit “fibers.” If such fibers are
ops immediately beneath the epidermis, and parenchyma treated with chemicals or bleached, the cementing middle
cells in the area develop thicker walls as the stem matures. lamella between the cells breaks down. Monocot fibers are not
The concentration of bundles, combined with the band of as strong or as durable as most dicot fibers.

bundle
sheath cell

phloem sieve tube

member
companion
cell

vessel
element
xylem
air space

ground
tissue
(parenchyma)

Figure 6.12 A cross section of a monocot (corn—Zea mays) Figure 6.13 A single vascular bundle of corn (Zea mays)
stem. ×20. © James E. Bidlack enlarged. ×400. © Kingsley Stern
 Stems 97

Specialized Stems so far, but all stems have nodes, internodes, and axillary buds;
these features distinguish them from roots and leaves, which
Although most higher plants have an erect shoot system, many do not have them. The leaves at the nodes of these special-
species have specialized stems that are modified for various ized stems are often small and scalelike. They are seldom
functions (Fig. 6.14). The overall appearance of specialized green, but full-sized, functioning leaves may also be produced.
stems may differ markedly from that of the stems discussed Descriptions of some of the specialized stems follow.

stolon

typical leaf
each eye
is a node
scalelike leaf
at each node

adventitious roots
tuber
rhizome

runner
adventitious
roots

stem tendril

axillary bud

flattened stem

fleshy leaves

cladophyll

papery leaves

stem
stem

old corm
bulb

corm

Figure 6.14 Types of specialized stems.

98 Chapter 6

Rhizomes Corms
Rhizomes (Fig. 6.14) are horizontal stems that grow below Corms resemble bulbs but differ from them in being com-
ground, often near the surface of the soil. They superfi- posed almost entirely of stem tissue, except for the few papery,
cially resemble roots, but close examination will reveal scalelike leaves sparsely covering the outside (Fig. 6.14).
scalelike leaves and axillary buds at each node, at least dur- Adventitious roots are produced at the base, and corms, like
ing some stage of development, with short to long inter- bulbs, store food. The crocus and the gladiolus are examples of
nodes in between. Adventitious roots are produced all along plants that produce corms.
the rhizome, mainly on the lower surface. As indicated in
Chapter 5, the word adventitious refers to structures aris-
ing at unusual places, such as roots growing from stems, or
Cladophylls
leaves or buds appearing at places other than leaf axils and The stems of butcher’s broom plants are flattened and appear
tips of stems. A rhizome may be a relatively thick, fleshy, leaflike. Such flattened stems are called cladophylls (or
food-storage organ, as in irises, or it may be quite slender, clad-odes or phylloclades) (Fig. 6.14). There is a node bearing
as in many perennial grasses or some ferns. very small, scalelike leaves with axillary buds in the center of
each butcher’s broom cladophyll. The feathery appearance of
Runners and Stolons asparagus is due to numerous small cladophylls. Cladophylls
also occur in greenbriers, certain orchids, prickly pear cacti
Runners are horizontal stems that differ from rhizomes in (Fig. 6.15), and several other lesser-known plants.
that they grow aboveground, generally along the surface; they
also have long internodes (Fig. 6.14). In strawberries, runners
are usually produced after the first flowers of the season have Other Specialized Stems
appeared. Several runners may radiate out from the parent The stems of many cacti and some spurges are stout and
plant and within a few weeks may grow 1 meter (3 feet) or fleshy. Such stems are adapted for storage of water and food.
more long. Adventitious buds appear at alternate nodes along Other stems may be modified in the form of thorns, as in
the runners and develop into new strawberry plants, which the honey locust, whose branched thorns may be more than
can be separated and grown independently. In saxifrages and 3 decimeters (1 foot) long, but all thornlike objects are not
some other house plants, runners may produce new plants at
intervals as they grow out and hang over the edge of the pot.
Stolons are similar to runners but are produced beneath
the surface of the ground and tend to grow in different direc-
tions but usually not horizontally. In Irish potato plants,
tubers are produced at the tips of stolons.
Some botanists consider stolons and runners to be
variations of each other and prefer not to make a distinction
between them.

Tubers
In Irish or white potato plants, several internodes at the tips of
stolons become tubers as they swell from the accumulation
of food (Fig. 6.14). The mature tuber becomes isolated after
the stolon to which it was attached dies. The “eyes” of the
potato are actually nodes formed in a spiral around the thick-
ened stem. Each eye consists of an axillary bud in the axil of a
scalelike leaf, although this leaf is visible only in very young
tubers; the small ridges seen on mature tubers are leaf scars.

Bulbs
Bulbs (Fig. 6.14) are actually large buds surrounded by
numerous fleshy leaves, with a small stem at the lower end.
Adventitious roots grow from the bottom of the stem, but
the fleshy leaves comprise the bulk of the bulb tissue, which
stores food. In onions, the fleshy leaves usually are sur-
rounded by the scalelike leaf bases of long, green, above- Figure 6.15 The flattened stems of prickly pear cacti (Opuntia)
ground leaves. Other plants producing bulbs include lilies, are cladophylls on which the leaves have been reduced to spines.
hyacinths, and tulips. © Kingsley Stern
 Stems 99

necessarily modified stems. For example, at the base of the about 0.12. Woods with specific gravities of less than 0.50
petiole of most leaves of the black locust is a pair of spines are considered light; those with specific gravities of above
(modifed stipules; stipules were mentioned in the discussion 0.70 are considered heavy. Among the heaviest woods are
of twigs and are discussed further in Chapter 7). The prick- the South American ironwood and lignum vitae, with spe-
les of raspberries and roses, both of which originate from cific gravities of over 1.25. Lignum vitae, obtained from
the epidermis, are neither thorns nor spines. Tiger lilies pro- West Indian trees, is extremely hard wood and has been
duce small, aerial bulblets in the axils of their leaves. used instead of metal in the manufacture of main bearings
Climbing plants have stems modified in various ways for drive shafts of submarines because it is self-lubricating
that adapt them for their manner of growth. Some stems, and less noisy.
called ramblers, simply rest on the tops of other plants, but
many produce tendrils (see Fig. 6.14). These are specialized Durability
stems in the grape and Boston ivy, but are modified leaves or
leaf parts in plants like peas and cucumbers. In Boston ivy, A wood’s ability to withstand decay caused by organisms
the tendrils have adhesive disks. In English ivy, the stems and insects is referred to as its durability. Moisture is needed
climb with the aid of adventitious roots that arise along the for the enzymatic breakdown of cellulose and other wood
sides of the stem and become embedded in the bark or other substances by decay organisms, but the seasoning process
support material over which the plant is growing. usually reduces the moisture to a level below that neces-
sary for the fungi and other decay organisms to survive.
Other natural constituents of wood that repel decay organ-
isms include tannins and oils. Wood with a tannin content
Wood and Its Uses of 15% or more may survive on a forest floor for many years
after diseases of the phloem and other causes have toppled
The use of wood by humans for fuel, shelter, weapons, and it. Among the most durable of American woods are cedar,
other purposes dates back into antiquity, and present uses catalpa, black locust, red mulberry, and Osage orange. The
are so numerous that it would be impossible to list in a work least durable woods include cottonwood, willow, fir, and
of this type more than the most important ones. Before dis- basswood.
cussing the economic importance of wood, let’s take a brief
look at its properties.
Types of Sawing
In a living tree, up to 50% of the weight of the wood
comes from the water content. Before the wood can be used, Logs are usually cut longitudinally in one of two ways: along
seasoning reduces the moisture content to 10% or less, either the radius or perpendicular to the rays (Fig. 6.17). Radially
by air-drying it in ventilated piles or stacks or by drying it in cut, or quartersawed, boards show the annual rings in side
special ovens known as kilns. The seasoning has to be done view; they appear as longitudinal streaks and are the most
gradually and under carefully controlled conditions, or the conspicuous feature of the wood. Only a few perfect quar-
timber may warp and split along the rays, making it unfit for tersawed boards can be obtained from a log, making them
most uses. The dry part of wood is composed of 60% to 75% quite expensive. Boards cut perpendicular to the rays (tan-
cellulose and about 15% to 25% lignin, an organic substance gentially cut boards) are more common. In these, the annual
that makes the walls of xylem cells tough and hard. Other rings appear as irregular bands of light and dark alternat-
substances present in smaller amounts include resins, gums, ing streaks or patches, with the ends of the rays visible as
oils, dyes, tannins, and starch. The proportions and amounts narrower and less conspicuous vertical streaks. Lumber cut
of these and other substances determine how various woods tangentially is referred to as being plain-sawed, or slab cut.
will be used (Fig. 6.16). Slabs are the boards with rounded sides at the outside of the
log; they are usually made into chips for pulping.
Properties of Wood
Knots
Density Knots are the bases of lost branches that have become
The density of wood is among its most important physical covered, over a period of time, by new annual rings of
properties. Technically, the density is the weight per unit wood produced by the cambium of the trunk. They are
volume. The weight is compared with that of an equal vol- found in greater concentration in the older parts of the
ume of water. If the weight of a volume of wood equals the log toward the center, because in the forest, the lowermost
weight of the same volume of water, it is said to have a spe- branches of a tree (produced while the trunk was small
cific gravity of 1.0. The specific gravity is less than 1.0 if in girth) often die from insufficient light. When a branch
the wood weighs less than the water and greater than 1.0 if dies and falls off, the cambium at its base also dies, but
the wood weighs more than water. Because of the consider- the cambium of the trunk remains alive and increases the
able air space within the cells, most woods have a specific girth of the tree, slowly enveloping the dead tissue of the
gravity of less than 1.0. The range of specific gravities of branch base until it may be completely buried and not vis-
known woods varies from 0.04 to 1.40, the lightest com- ible from the surface. Knots usually weaken the boards in
mercially used wood being balsa with a specific gravity of which they occur.
100 Chapter 6

Hemlock. This relatively soft, light, straight- White Pine. A softwood widely used in home
grained, resin-free wood, with its uniformly long construction and for virtually everything from
fibers, is becoming one of the most important masts and matches to boxes and crates. Its soft,
species for paper pulp. It is also used for structural uniform texture and straight grain cuts easily in
lumber and plywood, and for boxes, barrels, and every direction, polishes well, and warps
concrete forms. or swells little.

Red Spruce. A favorite for violin sounding boards Bald Cypress. Because it is weather-resistant
because of its high resonant qualities. A softwood, without treatment, this wood was widely used for
it is easy to work, and is light in relation to its cross ties in the early days of railroading. Today
strength and sti ness. These qualities also make it is used for water tanks and other applications
it eminently suitable for ladder rails, canoe paddles, requiring prolonged contact with water.
and oars.

White Oak. Makes good barrels because the White Ash. Perfect for baseball bats, tennis
wood is resilient, durable, and impermeable to racquets, oars and long tool handles. This
liquids. This hardwood, which is about twice as hardwood’s major virtues are straight grain,
dense as white pine, has many other uses ranging sti ness, strength, moderate weight, good
from flooring to fine cabinet work. bending qualities, and capacity for wearing smooth.

Figure 6.16 Uses of common North American woods.

 Stems 101

Wood Products
In the United States and Canada, about half of the wood
produced is used as lumber, primarily for construction; the
sawdust and other waste formed in processing the boards are
converted to particle board and pulp. A considerable amount
of lumber goes into the making of furniture, which may
be constructed of solid wood or covered with a veneer. A
veneer is a very thin sheet of desirable wood that is glued to
cheaper lumber; it is carefully cut so as to produce the best
possible view of the grain.
The next most extensive use of wood is for pulp, which
among other things is converted by various processes to
paper, synthetic fibers, plastics, and linoleum. In recent
years, it has been added as a filler to commercial ice cream
and bread. Some hardwoods are treated chemically or heated
under controlled conditions to yield a number of chemicals,
such as wood alcohol and acetic acid, but other sources of
these products are now usually considered more economical.
Charcoal, excelsior, cooperage (kegs, casks, and barrels),
railroad ties, boxes and crates, musical instruments, bowling
pins, tool handles, pilings, cellophane, and Christmas trees
are but a few of the additional wood products worth billions
of dollars annually on the world market (see Fig. 6.16).
In developing countries, approximately half of the tim-
ber cut is used for fuel, but in the United States and Canada,
a little less than 10% is currently used for that purpose. In
colonial times, wood was the almost exclusive source of heat-
ing energy. In the early 2000s, Brazil’s major cities were still
using scrub timber from the surrounding forests to energize

Figure 6.17 How the surfaces of plain-sawed, quartersawed,

Figure 6.16 Continued. and transversely sawed wood appear. (See also Fig. 6.8.)
102 Chapter 6

their utilities, but rapidly depleting supplies and problems aids in protection. Lenticels in the bark permit gas
related to the greenhouse effect (discussed in Chapter 25) exchange.
have pointed to the need for alternate sources of energy. Many 9. Primary vascular tissues and the pith, if present,
types of coal are wood that has been compressed for millions constitute the stele. Protosteles have a solid core of
of years until nearly pure carbon remains. The formation of xylem, usually surrounded by phloem; siphonosteles
coal and other fossils is discussed in Chapter 21. Although the are tubular, with pith in the center; eusteles have the
world’s supply of coal is still plentiful, the rate at which this vascular tissues in discrete bundles.
fossil fuel is being consumed makes it obvious that resources
10. Dicotyledons (dicots) are plants whose seeds have
will eventually be exhausted unless we find renewable or less
two seed leaves (cotyledons), while monocotyledons
destructive alternatives to meet our energy demands.
(monocots) have seeds with one seed leaf. Herbaceous
Some of the vast array of secondary products from
dicots have vascular bundles arranged in a ring in
stems, including dyes, medicines, spices, and foods, are dis-
the stem.
cussed in later chapters and in the appendices.
11. Woody dicots have most of their secondary tissues
arranged in concentric layers. The most conspicuous
tissue is wood (secondary xylem). In broadleaf
SUMMARY
trees, spring wood usually has relatively large vessel
members, while summer wood has smaller vessels
1. Stem structure and function need to be examined to and/or a predominance of tracheids.
understand practical uses of stems.
12. An annual ring is one year’s growth of xylem. A tree’s
2. The shoot system of plants is usually erect, but some age and other aspects of its history can be determined
stems may be horizontal or modified for climbing or from annual rings. Rays, which function in lateral
food/water storage. conduction, radiate out from the center of the trunk.
3. Leaves of woody twigs may be arranged alternately, Older wood toward the center (heartwood) ceases to
oppositely, or in a whorl. Nodes are stem regions where function when its cells become plugged with tyloses.
leaves are attached; internodes occur between nodes. Younger, functioning wood (sapwood) is closer to the
Most leaves have petioles and blades. Axillary buds surface. A tree’s functions are not particularly affected
occur in leaf axils. Most buds are protected by bud by the rotting of its heartwood.
scales. Terminal buds occur at twig tips. Terminal bud 13. The wood of cone-bearing trees consists primarily
scales, when they fall, leave bud scale scars that help of tracheids, and resin canals are often present. The
one determine the age of the twig. wood of conifers has no fibers or vessels and is called
4. Stipules are paired appendages present at the base of some softwood, while the wood of woody dicots is called
leaves; when they fall off, they leave small scars on the hardwood. In woody plants, older tissues composed of
twig. When whole leaves fall, they leave leaf scars on the thin-walled cells become crushed and functionless, and
twig, with tiny bundle scars within the leaf-scar surfaces. some are sloughed off.
5. Each stem has an apical meristem at its tip that 14. Laticifers are latex-secreting cells or ducts found in
produces tissues resulting in increase in length. various flowering plants. The latex of some plants has
Leaf primordia develop into mature leaves when considerable commercial value.
growth begins. Three primary meristems develop 15. Monocot stems have scattered vascular bundles and
from an apical meristem: the protoderm gives rise no cambia. The parenchyma tissue is not divided into
to the epidermis; the procambium produces primary pith and cortex. Each vascular bundle is surrounded
xylem and primary phloem; and the ground meristem by a sheath of sclerenchyma cells. Numerous bundles
produces pith and cortex. and a band of sclerenchyma cells and thicker-walled
6. As leaves and buds develop from primordia, traces of parenchyma cells just beneath the surface of monocot
xylem and phloem branch off from the main cylinder, stems aid in withstanding stresses.
leaving leaf gaps or bud gaps. 16. Palm trees are monocots that become large because their
7. A vascular cambium, producing secondary tissues, may parenchyma cells continue to divide. Other monocots
arise between primary xylem and phloem. Secondary develop a secondary meristem that produces parenchyma
xylem cells include tracheids, vessel elements, and cells and secondary vascular bundles. Grasses have
fibers. Secondary phloem cells include sieve-tube at the base of each internode intercalary meristems
members and companion cells. that contribute to rapid increases in length. Several
8. In many plants, a cork cambium producing cork and commercially important cordage fibers are obtained
phelloderm cells develops near the surface of the from monocots.
stem. Cork cells, which are part of the outer bark 17. Specialized stems include rhizomes, stolons, tubers,
(periderm), have suberin in their walls. Suberin is bulbs, corms, cladophylls, and tendrils. Such stems may
impervious to moisture, and the outer bark, therefore, have adventitious roots.
 Stems 103

18. The dry part of wood consists primarily of cellulose 3. Do climbing plants have any advantages over erect
and lignin. Resins, gums, oils, dyes, tannins, and starch plants? Any disadvantages?
are also present. Properties of wood that play a role in 4. If two leaves are removed from a plant and one is
its use include density, specific gravity, and durability. coated with petroleum jelly, while the other is not,
19. Logs are usually cut longitudinally along the radius the uncoated leaf will shrivel considerably sooner
(quartersawed) or perpendicular to the rays (tangentially than the coated one. Would it be helpful to coat
cut, plain-sawed, or slab cut). Knots are bases of lost the stems of young trees with petroleum jelly?
branches that have become covered over by new wood; Explain.
they usually weaken the boards in which they occur. 5. Suggest some reasons for heartwood being preferred to
20. About half the timber produced in the United States sapwood for making furniture.
and Canada is used as lumber. Sawdust and waste
are converted to particle board and pulp for paper,
synthetics, and linoleum. Other timber is used for ADDITIONAL READING
cooperage, charcoal, railroad ties, boxes, tool handles,
Bowers, B. G. 2000. A color atlas of plant structure. Ames, IA:
and so forth. Developing countries use a greater
Iowa State Press.
proportion of their timber for fuel.
Carlquist, S. J. 2001. Comparative wood anatomy: Systematic,
ecological and evolutionary aspects of dicotyledon wood,
2d rev. ed. New York: Springer-Verlag.
REVIEW QUESTIONS Desch, H. E. 1996. Timber: Structure, properties, conversion
and use, 7th ed. Binghampton, NY: Haworth Press Journal
1. What is the function of bud scales? Co-Editions.
2. How can you tell the age of a twig? Dickison, W. C. 2000. Integrative plant anatomy. San Diego, CA:
Academic Press.
3. Distinguish among procambium, vascular cambium,
Evert, R. F. 2006. Esau’s plant anatomy: Meristems, cells, and
and cork cambium. tissues of the plant body: Their structure, function, and
4. How can you tell, when you look at a cross section of a development, 3d ed. Somerset, NJ: John Wiley & Sons.
young stem, whether it is a dicot or a monocot? Fahn, A. 1990. Plant anatomy, 4th ed. Elmsford, NY: Pergamon
5. What are laticifers? Press.
Lewin, M., and I. S. Goldstein (Eds.). 1991. Wood structure and
6. An Irish or white potato is a stem, but a sweet potato is composition. New York: Dekker, Marcel.
a root. How can you tell? Mauseth, J. D. 1988. Plant anatomy. Menlo Park, CA: Benjamin/
7. Distinguish among corms, bulbs, and tubers. Cummings.
8. If you were looking at the top of a wooden desk, Metcalfe, C. R., and L. Chalk (Eds.). 1988–1989. Anatomy of the
how could you tell if the wood had been radially or dicotyledons, 2 vols. Fair Lawn, NY: Oxford University Press.
tangentially cut (quartersawed or plain-sawed)? Perlin, J. A. 1991. A forest journey: The role of wood in the
development of civilization. Cambridge, MA: Harvard
9. What differences are there between heartwood and University Press.
sapwood? Porter, T. 2007. Wood: Identification and use. New York: Sterling.
10. What is meant by the specific gravity of wood? Schweingruber, F. H. 1993. Trees and wood in dendrochronology.
New York: Springer-Verlag.
Schweingruber, F. H. 2007. Wood structure and environment.
DISCUSSION QUESTIONS New York: Springer-Verlag.

1. If the cambium of a tropical tree were active all year LEARNING ONLINE
long, how would its wood differ from that of a typical
temperate climate tree?
2. A nail driven into the side of a tree will remain at
exactly the same distance from the ground for the life of Visit our website at http://connect.mheducation.com for additional
the tree. Why? information and learning tools.