University of Al-Mustansiriyah/ College of Science/ Department of

Biology
Course : Botany
Lecture: 6

TISSUE PATTERNS IN STEMS

Steles
Primary xylem, primary phloem, and the pith, if present, make up a central cylinder
called the stele in most younger and a few older stems and roots. The simplest form
of stele, called a protostele, consists of a solid core of conducting tissues in which
the phloem usually surrounds the xylem. Protosteles were common in primitive
seed plants that are now extinct
and are also found in whisk ferns, club mosses , and other relatives of ferns.
Siphonosteles, which are tubular with pith in the center, are common in ferns. Most
present-day flowering plants and conifers have eusteles in which the primary xylem
and primary phloem are in discrete vascular bundles. Flowering plants develop
from seeds that have either one or two “seed leaves,” called cotyledons attached to
their embryonic stems . The seeds of pines and other cone-bearing trees have
several (usually eight) cotyledons. The cotyledons usually store food needed by the
young seedling until its first true leaves can produce food themselves.
Flowering plants that develop from seeds having two cotyledons are called
dicotyledons (usually abbreviated to dicots), while those developing from seeds
with a single cotyledon are called monocotyledons (abbreviated to monocots).
Dicots and monocots differ from one another in several other respects, a summary
of these and other differences in these two classes of flowering plants is given in
Table below.

Some Differences Between Dicots and Monocots

DICOTS MONOCOTS
Seed with two cotyledons (seed leaves Seed with one cotyledon (seed leaf)
. Flower parts mostly in fours or fives or . Flower parts in threes or multiples of
multiples of four or five three
Leaf with a distinct network of primary . Leaf with more or less parallel primary
veins veins
Vascular cambium, and frequently cork Vascular cambium and cork cambium
cambium, present absent
Vascular bundles of stem in a ring Vascular bundles of stem scattered
Pollen grains mostly with three Pollen grains mostly with one aperture
apertures (thin areas in the aperture
wall—see Figures 23.6 and 23.7)

Herbaceous Dicotyledonous Stems

In general, plants that die after going from seed to maturity within one growing
season (annuals) have green, herbaceous (nonwoody) stems. Most monocots are
annuals, but many dicots (discussed next) are also annuals. The tissues of annual
dicots are largely primary, although cambia (plural of cambium) may develop some
secondary tissues. Herbaceous dicot stems (Fig. 6.5) have discrete vascular

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bundles composed of patches of xylem and phloem. The vascular bundles are
arranged in a cylinder that separates the cortex from the pith, although in a few
plants (e.g., foxgloves), the xylem and the phloem are produced as continuous rings
(cylinders) instead of in separate bundles.
The procambium produces only primary xylem and phloem, but later, a vascular
cambium arises between these two primary tissues and adds secondary xylem and
phloem to the vascular bundles. In some plants, the cambium extends between the
vascular bundles, appearing as a narrow ring, producing not only the conducting
tissues within the bundles but also the parenchyma cells between them. In other
plants, the cambium is not in an uninterrupted cylinder but is instead confined to
the bundles, each of which has its own small band of cambium between the xylem
and phloem.

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.

Woody Dicotyledonous Stems

In woody plants obvious differences begin to appear as soon as the vascular
cambium and the cork cambium develop. The differences involve the secondary
xylem, or wood (Fig. 6.6). Some tropical trees (e.g., ebony), in which both the
vascular cambium and the cork cambium are active all year, produce an uniform
wood. The wood of most trees, however, is produced seasonally. In trees of
temperate climates, virtually all growth takes place during the spring and summer
and then ceases until the following spring. When the vascular cambium of a tree
first becomes active in the spring, it usually produces relatively large vessel
elements of secondary xylem; such xylem is referred to as spring wood. As the
season progresses, the vascular cambium may produce vessel elements whose
diameters become progressively smaller in each succeeding series of cells
produced, or there may be fewer vessel elements in proportion to tracheids
produced until tracheids (and sometimes fibers) predominate.
The xylem that is produced after the spring wood, and which has smaller or fewer
vessel elements and larger numbers of tracheids, is referred to as summer wood.
Over a period of years, the result of this type of switch between the early spring and
the summer growth is a series of alternating concentric rings of light and dark cells.
One year’s growth of xylem is called an annual ring. In conifers, the wood consists
mostly of tracheids, with vessels and fibers being absent. Annual rings are still

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visible, however, since the first tracheids produced in the spring are considerably
larger and lighter in color than those produced later in the growing season.
The annual rings not only indicate the age of the tree (since normally only one is
produced each year), but they can also tell something of the climate and other
conditions that occurred during the tree’s lifetime (Fig. 6.7). For example, if the
rainfall during a particular year is higher than normal, the annual ring for that year
will be wider than usual because of increased growth.
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 Figs. 6.6 and 6.8). These lines, called vascular rays, 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, but its extension through the phloem is called a
phloem ray.
The term bark is usually applied to all the tissues outside the cambium, including
the phloem.
Despite the presence of fibers, the thin-walled conducting cells of the phloem are
not usually able to withstand for many seasons the pressure of thousands of new
cells added to their interior, and the older layers become crushed and functionless.
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 since 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.

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Figure 6.6 A cross section of a portion of a young linden (Tilia) stem. ca. 300.

Monocotyledonous Stems
Most monocots (e.g., grasses, lilies) are herbaceous plants that do not attain great
size. The stems have neither a vascular cambium nor a cork cambium and thus
produce no secondary vascular tissues or cork. As in herbaceous dicots, the
surfaces of the stems are covered by an epidermis, but the xylem and phloem
tissues produced by the procambium appear in cross section as discrete vascular
bundles scattered throughout the stem instead of being arranged in a ring (Fig.
6.12). Each bundle, regardless of its specific location, is oriented so that its xylem
is closer to the center of the stem and its phloem is closer to the surface. In a
typical monocot such as corn, a bundle’s xylem usually contains two large vessels
with several small vessels between them (Fig. 6.13). The first-formed xylem cells
usually stretch and collapse under the stresses of early growth and leave an
irregularly shaped air space toward the base of the bundle; the remnants of a vessel
are often present in this air space. The phloem consists entirely of sieve tubes and
companion cells, and the entire bundle is surrounded by a sheath of thicker-walled
sclerenchyma cells. The parenchyma tissue between the vascular bundles is not
separated into cortex and pith in monocots, although its function and appearance
are the same as those of the parenchyma cells in cortex and pith. In a corn stem,
there are more bundles just beneath the surface than there are toward the center.
Also, a band of sclerenchyma cells, usually two or three cells thick, develops
immediately beneath the epidermis, and parenchyma cells in the area develop
thicker walls as the stem matures. The concentration of bundles, combined with the
band of sclerenchyma cells beneath the epidermis and the thicker walled
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parenchyma cells, all contribute to giving the stem the capacity to withstand
stresses resulting from summer storms and the weight of the leaves and the ears of
corn as they mature.

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

1. Rhizomes
Rhizomes (Fig. 6.14) are horizontal stems that grow below ground, often near the
surface of the soil. They superficially resemble roots, but close examination will
reveal scalelike leaves and axillary buds at each node, at least during some stage of
development, with short to long internodes in between. Adventitious roots are
produced all along the rhizome, mainly on the lower surface. The word adventitious
refers to structures arising at unusual places, such as roots growing from stems, or
leaves or buds appearing at places other than leaf axils and tips of stems. A rhizome
may be a relatively thick, fleshy, food-storage organ, as in irises, or it may be quite
slender, as in many perennial grasses or some ferns.

2. Runners and Stolons

Runners are horizontal stems that differ from rhizomes in that they grow above
ground, 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
appeared. Several runners may radiate out from the parent plant, and within a few
weeks may grow up to 1 meter (3 feet) or more long. Adventitious buds appear at
alternate nodes along the runners and develop into new strawberry plants, which
can be separated and grown independently. In saxifrages and some other house

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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 directions 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.

3. 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 thickened 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.

4. 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 surrounded by the scalelike leaf bases of
long, green, aboveground leaves. Other plants producing bulbs include lilies,
hyacinths, and tulips.

5. Corms
Corms resemble bulbs but differ from them in being composed almost entirely of
stem tissue, except for the few papery, scalelike leaves sparsely covering the
outside (Fig. 6.14). Adventitious roots are produced at the base, and corms, like
bulbs, store food. The crocus and the gladiolus are examples of plants that produce
corms.

6. Cladophylls
The stems of butcher’s broom plants are flattened and appear leaflike. Such
flattened stems are called cladophylls (or cladodes or phylloclades) (Fig. 6.14).
There is a node bearing very small, scalelike leaves with axillary buds in the center
of each butcher’s broom cladophyll. The feathery appearance of asparagus is due to
numerous small cladophylls. Cladophylls also occur in greenbriers, certain orchids,
prickly pear cacti (Fig. 6.15), and several other lesser-known plants.

7. Other Specialized Stems

The stems of many cacti and some spurges are stout and fleshy. Such stems are adapted for storage
of water and food. Other stems may be modified in the form of thorns, as in the honey locust, whose
branched thorns may be more than 3 decimeters (1 foot) long, but all thornlike objects are not
necessarily modified stems. For example, at the base of the petiole of most leaves of the black locust
is a pair of spines (modifed stipules; stipules were mentioned in the discussion of twigs). The
prickles of raspberries and roses, both of which originate from the epidermis, are neither thorns nor
spines. Tiger lilies produce small, aerial bulblets in the axils of their leaves. Climbing plants have
stems modified in various ways that adapt them for their manner of growth. Some stems, called
ramblers, simply rest on the tops of other plants, but many produce tendrils (see Fig. 6.14). These
are specialized stems in the grape and Boston ivy but are modified leaves or leaf parts in plants like
peas and cucumbers. In Boston ivy, the tendrils have adhesive disks. In English ivy, the stems climb

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with the aid of adventitious roots that arise along the sides of the stem and become embedded in the
bark or other support material over which the plant is growing.

Figure 6.15 The flattened stems of prickly pear cacti (Opuntia) are cladophylls on which
the leaves have been reduced to spines.

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 (Leaves)
Leaves of flowering plants are associated with leaf gaps (as illustrated in Fig. 6.3),
and all have an axillary bud at the base. Leaves may be simple or compound. A
simple leaf has a single blade, while the blade of a compound leaf is divided in
various ways into leaflets (see Fig. 7.4). A compound leaf has a single axillary bud
at its base, with the leaflets having no such buds. Pinnately compound leaves have
the leaflets in pairs along an extension of the petiole called a rachis, while
palmately compound leaves have all the leaflets attached at the same point at the
end of the petiole.

LEAF ARRANGEMENTS AND TYPES

Many of different species of plants that produce leaves can be distinguished from
one another by their leaves alone. Leaves are attached to stems at regions called
nodes, with stem regions between nodes being known as internodes.
The arrangement of leaves on a stem in a given species of plant generally occurs in
one of three ways. In most species, leaves are attached alternately or in a spiral
along a stem, with one leaf per node, in what is called an alternate arrangement. In
some plants, two leaves may
be attached at each node, providing an opposite arrangement. When three or more
leaves occur at a node, they are said to be whorled. The arrangement of veins in a
leaf or leaflet blade (venation) may also be either pinnate or palmate. In pinnately
veined leaves, there is one primary vein called the midvein, which is included
within an enlarged midrib; secondary veins branch from the midvein. In palmately
veined leaves, several primary veins fan out from the base of the blade. The
primary veins are more or less parallel to one another
in monocots (Fig. 7.5) and diverge from one another in various ways in dicots (see
Fig. 7.9). The branching arrangement of veins in dicots is called netted, or
reticulate venation. In a
few leaves (e.g., those of Ginkgo), no midvein or other large veins are present.
Instead, the veins fork evenly and progressively from the base of the blade to the
opposite margin. This
is called dichotomous venation (Fig. 7.4K).

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Figure 7.4 Types of leaves and leaf arrangements. A. Palmately compound leaf of a
buckeye. B. Opposite, simple leaves of a dogwood. C. Pinnately compound leaf of a black
walnut. D. Alternate, simple but lobed leaves of a tulip tree. E. Parallel-veined leaf of a
grass. F. Palmately veined leaf of a maple. G. Whorled leaves of a bedstraw. H. Pinnately
veined, lobed leaf of an oak. I. Linear leaves of a yew. J. Globe-shaped succulent leaves of
string-of-pearls. K. Fan-shaped leaf of a Ginkgo tree, showing dichotomous venation.

INTERNAL STRUCTURE OF LEAVES

If a typical leaf is cut transversely and examined with the aid of a microscope,
three regions stand out: epidermis, mesophyll, and veins (referred to as vascular
bundles in our discussion of roots and stems) (Fig. 7.6). The epidermis is a single
layer of cells covering the entire surface of the leaf. The epidermis on the lower
surface of the blade can sometimes be distinguished from the upper epidermis by
the presence of tiny pores called stomata. Except for guard cells, the upper
epidermal cells for the most part do not contain chloroplasts, their
function being primarily protection of the delicate tissues to the interior. A coating
of waxy cutin (the cuticle—see Fig. 4.6) is normally present. In addition to the
cuticle, many plants produce other waxy substances on their surfaces (Fig. 7.7).
Presumably, the wax affords added protection to the leaves. Different types of

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glands may also be present in the epidermis. Glands occur in the form of
depressions, protuberances, or appendages either directly on the leaf surface or on
the ends of hairs (see Fig. 4.13A). Glands often secrete sticky substances.

STOMATA
The lower epidermis of most plants generally resembles the upper epidermis, but
the lower is perforated by numerous tiny pores called stomata (Fig. 7.8). Each pore
is bordered by two sausage- or dumbbell-shaped cells that usually are smaller than
most of the neighboring epidermal cells. These guard cells contain chloroplasts.
Guard cell walls are distinctly thickened but quite flexible on the side adjacent to
the pore. As the guard cells inflate or deflate with changes in the amount of water
within the cells, their unique construction causes the stomata to open or close.

MESOPHYLL AND VEINS

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Most photosynthesis takes place in the mesophyll between the two epidermal
layers, with two regions often being distinguishable. The uppermost mesophyll
consists of compactly
stacked, barrel-shaped, or post-shaped parenchyma cells that are commonly in two
rows. This region is called the palisade mesophyll and may contain more than 80%
of the leaf’s chloroplasts. The lower region, consisting of loosely arranged
parenchyma cells with abundant air spaces between them, is called the spongy
mesophyll. Its cells also have numerous chloroplasts. Parenchyma tissue with
chloroplasts is called chlorenchyma. Chlorenchyma tissue is also found in the outer
parts of the cortex in the stems of herbaceous plants as well as in leaves.
Veins (vascular bundles) of various sizes are scattered throughout the mesophyll
(Fig. 7.9). They consist of xylem and phloem tissues surrounded by a jacket of
thickerwalled parenchyma cells called the bundle sheath. The veins give the leaf
its “skeleton.” The phloem transports sugars and other carbohydrates throughout
the plant. Water is brought up to the leaf by the xylem.

SPECIALIZED LEAVES
1. Shade Leaves
because leaves in the shade receive less total light needed for photosynthesis, they
tend to be larger than their counterparts in the sun (Fig. 7.11). In addition, since
they receive less intense sunlight and heat, they are thinner, and have fewer well-
defined mesophyll layers and fewer chloroplasts. They also do not have as many
hairs.

2. Leaves of Arid Regions

Because of the limited availability of water, wide temperature ranges, and high light
intensities, plants growing in arid regions have developed adaptations that allow
them to thrive under such conditions. Many have thick, leathery leaves and fewer
stomata, or stomata
that are sunken below the surface in special depressions, all of which reduce loss of
water through transpiration. They also may have succulent, water-retaining leaves
or no leaves at all , or they may have dense, hairy coverings. Pine trees, whose
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water supply may be severely restricted in the winter when the soil is frozen, have
some leaf modifications similar to those of desert plants. The modifications include
sunken stomata, a thick cuticle, and a layer of thick-walled cells (the hypodermis)
beneath the epidermis (Fig. 7.12).

3. Leaves of Aquatic Areas

The submerged leaves of plants that grow in water usually have considerably less
xylem than phloem, and the mesophyll, which is not differentiated into palisade and
spongy layers, has large air spaces.
4. Tendrils
There are many plants whose leaves are partly or completely modified as tendrils.
These modified leaves, when curled tightly around more rigid objects, help the
plant in climbing or in supporting weak stems. The leaves of garden peas are
compound (Fig. 7.14), and the terminal leaflets are reduced to whip like strands
that, like all tendrils, are very sensitive to contact. Members of the Pumpkin Family
(Cucurbitaceae), which includes squashes, melons, and cucumbers, produce tendrils
that may be up to 3 decimeters (1 foot) long. As the tendrils develop, they become
coiled like a spring.

5. Spines, Thorns, and Prickles

The leaves of many cacti and other desert plants are modified as spines. This
reduction in leaf surface correspondingly reduces water loss from the plants, and
the spines also tend to protect the plants from browsing animals. In such desert
plants, photosynthesis, which would otherwise take place in leaves, occurs in the
green stems. Most spines are modifications of the whole leaf, in which much of the
normal leaf tissue is replaced with sclerenchyma, but in a number of woody plants
(e.g., mesquite, black locust), it is the stipules at the bases of the leaves that are
modified as short, paired spines.

6. Storage Leaves
Desert plants may have succulent leaves (i.e., leaves that are modified for water
retention). The adaptations for water storage involve large, thin-walled parenchyma
cells without chloroplasts to the interior of chlorenchyma tissue just beneath the
epidermis. These nonphotosynthetic cells contain large vacuoles that can store
relatively substantial amounts of water. The fleshy leaves of onion, lily, and other
bulbs store large amounts of carbohydrates, which are used by the plant during
rapid growth early in the subsequent growing season.

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 7. Flower-Pot Leaves
Some leaves of Dischidia (Fig. 7.16), an epiphyte (a plant that grows, usually non-
parasitically, on other plants) from tropical Australasia, develop into urnlike
pouches that become the home of ant colonies. The ants carry in soil and add
nitrogenous wastes, while moisture collects in the leaves through condensation of
the water vapor coming from the mesophyll through stomata. This creates a good
growing medium for roots, which develop adventitiously from the same node as the
leaf and grow down into the soil contained in the urnlike pouch. In other words, this
plant not only reproduces itself by conventional means but also, with the aid of
ants, provides its own fertilized growing medium and flower pots and then
produces special roots, which “exploit” the situation.

Figure 7.16 A flower-pot leaf of Dischidia. The hollow leaf has been sliced lengthwise to
reveal the roots inside that have started to grow down from the top.

8. Window Leaves
There are plants belonging to the Carpetweed Family (Aizoaceae) that have unique
adaptations to living in dry, sandy areas. Their leaves, which are shaped like ice-
cream cones, are about 3.75 centimeters (1.5 inches) long (Fig. 7.17) and are buried
in the sand; only the dimesized wide end of a leaf is exposed at the surface. This
exposed end is covered with a relatively transparent, thick epidermis with few
stomata and a waxy cuticle. There is a mass of tightly packed, transparent water-
storage cells below the exposed end; these allow light coming through the
“windows” to penetrate to the chloroplasts in the mesophyll, located all around the
inside of the shell of the leaf. This arrangement, which keeps most of the plant
buried and away from drying winds, allows the plant to thrive under circumstances
that most other plants could not tolerate.

9. Reproductive Leaves
Some of the leaves of the walking fern are most unusual in that they produce new
plants at their tips. Occasionally, three generations of plants may be found linked

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together. The succulent leaves of air plants (Fig. 7.18) have little notches along the
leaf margins in which tiny plantlets are produced, complete with roots and leaves,
even after a leaf has been removed from the parent plant. Each of the plantlets can
develop into a mature plant if given the opportunity to do so.

10. Floral Leaves (Bracts)

Specialized leaves known as bracts are found at the bases of flowers or flower
stalks. In the Christmas flower (poinsettia), the flowers themselves have no petals,
but the brightly colored floral bracts that surround the small flowers function like
petals in attracting pollinators (Fig. 7.19).
11. Insect-Trapping Leaves
Highly specialized insect-trapping leaves have intrigued humans for hundreds of
years. Insectivorous plants grow mostly in swampy areas and bogs of tropical and
temperate regions. In such environments, certain needed elements, particularly
nitrogen, may be deficient in the soil. Some of these elements are furnished when
the soft parts of insects and other small organisms trapped by the specialized leaves
are broken down and digested. All the plants have chlorophyll and are able to make
their own food. It has been demonstrated that they can develop normally without
insects if they are given the nutrients they need. Ex. 1: Pitcher Plants that have
larger and cone-shaped or vaselike (Figure 7.18). Ex. 2: Sundews, the roundish to
oval leaves are covered with up to 200 upright glandular hairs that look like
miniature clubs. There is a clear, glistening drop of sticky fluid containing digestive
enzymes at the tip of each hair (Figure 7.19). Ex. 3: Venus’s Flytraps, It catches its
prey—chiefly insects and arachnids with a trapping structure formed by the
terminal portion of each of the plant's leaves and is triggered by tiny hairs on their
inner surfaces. When an insect or spider crawling along the leaves contacts a hair,
the trap closes if a different hair is contacted within twenty seconds of the first
strike. (Figure 7.20, Figure 7.21).

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Figure 7.18 Pitcher Plants

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 Figure 7.19 Sundews.

Figure 7.20 Venus flytrap trigger hairs

Figure 7.21 Venus flytrap trigger hairs

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