Chapter II: Morphology of Higher Plants.

II. Morphology of higher plants:

When one thinks of a plant, the image that comes to mind is typically that of an organism with
stems, leaves, and roots. Biologists refer to these as Cormophytes, constituting over 70% of
terrestrial plants. They feature a cluster of leafy stems, with or without roots, collectively termed
the cormus.

II.1. Roots

II.1.1. Definition

The root, a vital plant organ usually developing in the soil, plays a pivotal role in the plant's
nutrition by absorbing water and minerals from its surroundings. With numerous branches, it
facilitates the plant's anchoring in the soil. Roots exhibit positive geotropism and negative
phototropism.

a• Geotropism: The orientation of the growth of certain organs (stem, root) concerning gravity,
with positive geotropism indicating growth from top to bottom and negative geotropism from
bottom to top.

b• Phototropism: The phenomenon of attraction or repulsion of growing organs concerning light

(natural or artificial), where positive phototropism implies attraction and negative phototropism
implies repulsion.

I1.1.2. Branching

The primary root gives rise to secondary lateral branches, which, in turn, generate tertiary lateral
branches, and so forth. The finest branches, known as rootlets, are where absorption is most active.
Collectively, they form the root system or root structure. Root morphology becomes more intricate
as the plant develops, branching into pivot, tuberous, and fasciculated roots (Fig. 2).

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 Chapter II: Morphology of Higher Plants.

Figure 2 – Lateral branching roots.

II.1.3. Different Parts of a Root

In a root system, from bottom to top, the following distinctions are made (fig. 3):

• Root Cap: The conical-shaped root cap is located at the root's tip, shielding the root meristem
and aiding in soil penetration. The root cap continuously sheds and regenerates.

• Zone of Elongation: This is a short, smooth, and clear zone where the differentiation of root
tissues occurs, facilitating the longitudinal growth of the organ.

• Branching Zone: Also known as the suberous zone, it is the longest and most massive part of
the root, where all the ramifications of the root system develop.

• Root Hair Zone (Piliferous Zone): The root hair zone encompasses the segment of the root
housing absorbent hairs. This region is crucial for the absorption of water and mineral salts. Despite
the brief lifespan of these hairs, the dimensions and location of the root hair zone remain
remarkably consistent.

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 Chapter II: Morphology of Higher Plants.

Figure 3 – Various parts of the root.

II.1.4. Root Systems

Root systems are categorized based on the prominence of the main root relative to its lateral
branches:

• Taproot System: In this system, the primary root, or taproot, has greater length and diameter
compared to its lateral roots. Example: Clover root, Wild carrot.

• Fibrous Root System (fasciculate): Distinguishing the main root from lateral roots becomes
challenging in this system, as all roots tend to have more or less equal diameters. This type is
commonly observed in Monocotyledonous Angiosperms. Example: Iris root, Wheat root.

• Adventitious Root System: This type of root system does not originate from the radicle
(embryonic root) but develops from various plant parts, such as stems or leaves. The adventitious
roots may have different sizes and can serve various functions like support or absorption. Example:
Onion, Mint.

II.1.5. Other roots (Modified Roots):

a- Tuberous Roots: Selected Examples: Salsify - Asphodel - turnip - carrot, radish, and cultivated
beets, dahlia, ...

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 Chapter II: Morphology of Higher Plants.

It is noteworthy that the taproot of Salsify becomes enormous due to the accumulation of reserves.
It forms a large mass or tuber (fig. 4); hence the expression: "tuberous taproot." In the case of
asphodel, dahlia, and some other plants (fig. 4), the root system consists of "fasciculated tuberous
roots."

Figure 4 – From the left to the right: Tuberous taproot of the Salsify and Fasciculate tuberous
root of the Asphodel.

b- Haustorial Roots: Selected Examples: Haustorium of cistanche on tamarisk. Other examples:

Dodder,...

Haustorium of dodder on stems of other plants (in stems), They firmly attach to their host using
organs that penetrate either into a stem (dodder) or into a root (cistanche): haustorial roots. A cross-
section in the attachment region shows no cap or absorbing hairs.

c- Climbing Roots: Exemplified by the climbing ivy. On one side of the ivy stem, we have already
observed numerous adventitious roots. They lack a cap, piliferous zone, or branching. They may
end with a flattened part that adheres strongly to the support. These are referred to as climbing
roots.

Note: Comparing the roots of various plants allows the distinction between taproots and
fasciculated roots, roots normally located at the base of the stem and adventitious ones, and various
types of modified roots.

II.1.6. Functions of the roots

Not all roots exactly resemble the type we have just described; there are those adapted to different
environments or specific functions. Here are some of these related adaptations:
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 Chapter II: Morphology of Higher Plants.

a- Fixation Role: The downward-oriented growth of roots firmly anchors the plant in the soil.
Some plants, in addition to their underground root system, have aerial adventitious roots
(Adventitious means additional) at the stem level, serving as a means of fixation.

b- Absorption Function: Maximum water and mineral absorption occurs in the root hair zone,
thanks to the absorbent hairs. In parasitic plants, the main root transforms into a haustorium, an
organ that penetrates the vascular tissues of the host plant to extract sap (Example: mistletoe).
Epiphytic plants have hanging aerial roots that absorb atmospheric water through perforations in
their walls. These plants lack absorbent hairs and do not parasitize the host plant (Example: Orchids
and Philodendron).

c- Storage Function: Some roots undergo tuberization due to the accumulation of reserves
(starch, or others). These nutritional substances accumulate in various tissues (Examples: phloem
for Carrot, cortical parenchyma for Asphodel).

Note: Associations between a plant's roots and other living organisms (Bacteria, Fungi) can exist,
such as nodules and mycorrhizae (fig. 5):

- Mycorrhizae are symbiotic relationships that form between fungi and plants. The fungi colonize
the root system of a host plant, providing increased water and nutrient absorption capabilities while
the plant provides the fungus with carbohydrates formed from photosynthesis.

- Nodules in the plant roots contain symbiotic bacteria within the nodules, producing nitrogen
compounds that help the plant to grow and compete with other plants. When the plant dies, the
fixed nitrogen is released, making it available to other plants, and this helps to fertilize the soil.

Figure 5 – Plant-Organism Associations; (A) Nodules (association bacteria and plant) on plant
roots, (B) Mycorrhizae (a plant-fungus symbiosis).
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II.2. Stems

II.2.1. Definition

The stem, easily observable in plants, is typically aerial and serves as the organ that carries leaves,
the reproductive apparatus (flower), as well as fruits and seeds. It plays crucial roles in both
conduction and support. The connection between the stem and the root is established at the collar,
a transitional zone between these two vital plant organs. Positive phototropism and negative
geotropism are characteristic behaviors exhibited by the stem.

II.2.2. Stem Structure: The caulinar apparatus, comprising the collective stems, is typically aerial.
Each stem is composed of a succession of internodes and nodes. The internodes decrease in height
as they approach the apical bud. Leaves are inserted at the nodes. At their axils, one or more lateral
buds are found, and their development will give rise to either lateral stems or flowers. To observe
buds, let's select plants where they are sufficiently large (such as oak, loquat, fig tree, ...). A bud
can be described as a cluster of tiny leaves closely pressed against each other. The outermost
"leaves" can be leathery. Those internally become progressively smaller and tender. The last ones,
simple protuberances, surround the apex.

II.2.3. Different Components of the Stem

The stem is comprised of several distinct parts (Fig.6):

-Main Axis: Often referred to as the main stem, the main axis of the stem bears leaves, buds, and
leafy branches.

-Nodes and Internodes: Nodes denote the points where leaves or leafy branches attach (or in other
explanation: the part of the stem where the leaves and lateral buds are attached to the stem is called
a node). Internodes, the segments between successive nodes, are regions where the stem
experiences longitudinal growth.

-Buds: Buds consist of very young leaves closely packed together. Moving toward the center of
the bud, the leaves gradually diminish in size and encircle a vegetative point (meristematic zone).
The outermost leaves are hairy and coated with a substance that imparts impermeability, serving
a protective function (fig. 7). Buds play a crucial role in the apical elongation of the stem and the
development of leaves and branches.

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Figure 6 – Different Parts of the Stem.

Figure 7 – Different components of the bud.

II.2.4. Morphological Types

II.2.4.1. Herbaceous Stems

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Herbaceous stems are characterized by their slender diameter, green hue, and delicate flexibility.
In this herbaceous type, both the main stem and any accompanying branches do not experience
substantial thickening. Herbaceous stems can manifest in various forms (fig. 8):

• Upright: The main axis stands vertically.

• Creeping: These elongated stems have a significantly reduced cross-section and minimal
supporting tissues. They sprawl horizontally across the ground.

• Climbing: Elongated stems that ascend in height by winding around a support or utilizing tendrils
and hooks for attachment.

Figure 8 – Herbaceous Stems: A: Upright stem (sunflower), B: creeping stem (strawberry Plant)
and C: climbing stem (ivy).

II.2.4.2. Woody Stems

The stems of woody plants are characterized by a large diameter (thick), a dark color (brown), and
hardness. They are perennial, living for years, and even centuries in the case of trees. The branching
pattern of woody stems allows us to distinguish the following main morphological types:

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• Arboreal Growth Form: This growth form applies to trees, shrubs, and small trees. The main
stem, known as the trunk, is thick and exhibits branches spreading out from a certain height.

• Bushy Growth Form: It involves a collection of woody stems forming a tuft, emerging from the
base, among which a distinct trunk cannot be distinguished.

II.2.4.3. Other types of stems

There are also other specific types of stems (fig. 9):

• Stipe: It is an unbranched cylindrical axis marked by scars from the bases of leaves (fallen
fronds), and its thickness is not due to the presence of woody tissues (completely absent).

• Culm: It is a hollow stem, except at the nodes where there is a thin partition called a
diaphragm. The culm is a slender and resistant stem that can reach several meters in height
(Example: reed and bamboo).

• Underground stems: These are stems that live in the soil, deeply modified by the environment
in which they are found. Their color is similar to that of roots, they bear buds, non-chlorophyllous
scale leaves, and adventitious roots.

Figure 9 – Other stem types; (A) Stipe (palm tree), (B) Culm (bamboo), and (C) Underground
stems (Potato).

II.2.4.4. Functions of the stem

Not all stems exactly resemble the type we just described; some are adapted to different
environments or specific functions. Here are some of these related adaptations:
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• For sap conduction: This is characterized by the prominence of conducting tissues and the
reduction of cortical parenchyma. The significance of conducting tissues is clearly visible in
climbing stems (very long) and woody stems (very thick).

• For chlorophyll assimilation function: Cladodes are flattened stems that morphologically
resemble leaves, for example, the cladode of the prickly pear.

• For reserve function: Some stems, often underground, can hypertrophy and accumulate
reserve substances (starch and carbohydrate), for example, potato tubers.

• For drought resistance: Succulent plants are a diverse group of plants with thick leaves and
stems that store water. The majority of succulents are native to hot, arid climates, but with the
right care, they can be grown almost anywhere. This adaptation reduces evapotranspiration and
protects the species against herbivores.

Note: There are plants without stems; they are called stemless plants. Some examples include
mosses and certain types of algae.

II.3. Leaves

Leaves are lateral extensions of the stem, attached at the nodes (Fig. 10). The vessels within the
leaf veins transport water and minerals essential for photosynthesis. Stomata facilitate the exchange
of gases. Leaves are categorized into three types:

- Deciduous: Shed during autumn.

- Evergreen: Persist for several years.

- Marcescent: Change color in autumn but typically fall only upon regrowth.

N.B: The difference between deciduous and marcescent leaves: In the oaks, appears to be the key
difference between normal deciduous leaves and marcescent leaves. For at around this time, normal
leaves (deciduous) on the tree fall, but the marcescent leaves persist on the branches through winter
unless they are broken off by wind, snow, or other mechanical forces.

- Leaves can be described based on various characteristics:

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a- The shape of the blade (lamina), which can be singular (as in a simple leaf) or divided into
multiple parts (in the case of a compound leaf); the outline of the blade; the degree of incision;
and the shape of its base or apex.

b- Additional features include leaf attachment to the stem (phyllotaxy), vein arrangement in
the blade, and the presence or absence of hairs (pubescence).

c-Leaves vary in characteristics related to the petiole, blade (lamina), veins, and their arrangement
on the stem.

d- A leaf undergoes defined growth over time, comprising three parts: the blade, the petiole, and
the sheath.

Figure 10 – The structure of the leaf.

II.3.1. Leaf Composition

II.3.1.1. The Blade (Limbus):

Derived from "limbus" meaning edge, it presents itself as a flattened blade, green in color (due to
chlorophyll), with a dark green upper (ventral) and a paler lower (dorsal) side. The outer edge of
the leaf is called the margin.

*The blade can be entire, crenate, toothed, lobed (pinnately or palmately lobed), dissected
(pinnately or palmately dissected), or laciniate (Laciniate: The leaf possesses incised margins. The
teeth of margin slightly bent towards base of the leaf. Such type of leaf is termed as laciniate).

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*As for the general shape of the blade, it can be peltate, round, oval, oblong, triangular, or
lanceolate. Leaves can be simple or compound. The morphological variation of the blade and the
arrangement of veins allow distinguishing different leaf types:

a- A simple leaf: It has a single blade continuous at the end of an unbranched petiole.

b- Compound leaves: The leaf is composed of several leaflets; pinnate, bipinnate, tripinnate,
palmate, trifoliate, and pedate leaves.

II.3.1.2. Veins (Nervures):

Nervation considers the arrangement of the important veins (vessels carrying sap) that traverse the
blade (Fig. 11).

- Uninervate leaves: The leaf has a single vein, characteristic of narrow-bladed leaves, as seen in
needle-like leaves of Rosemary.

- Parallel-nerve leaves: Veins start from the base of the blade without converging, following sub-
parallel paths; characteristic of monocotyledonous leaves, like in the corn leaf.

- Pennate-nerve leaves: The main vein divides the blade into two equal parts, and secondary veins
depart laterally on both sides along its entire length.

- Pedate nervation: The blades have 3 radiating veins, but others are branches of lateral veins,
always oriented toward the base of the leaf.

- Reticulate nervation: Veins form a quadrilateral network within the blade.

II.3.1.3. Petiole: From "petiolus" meaning little foot, it is the intermediary between the blade and
the stem, traversed by sap-conducting vessels. In some plants, the petiole is absent, resulting in
sessile leaves. Based on the leaf's insertion on the stem, we distinguish decurrent, clasping,
sheathing, perfoliate, petiolate, and sessile leaves (Fig. 11).

II.3.1.4. Phyllotaxis: It is the arrangement of leaves along the stem. To minimize leaf overlap on
the stem and competition for light, the meristem successively places leaf primordia in different
directions from the previous one.

a- A leaf is alternate when only one leaf appears at each node.

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b- Two leaves situated on the same node and arranged at 180° are called opposite.

c- When three or more leaves attach to a node, this arrangement is termed whorled.

II.3.1.5. The Sheath

The sheath is the widening of the base of the petiole (fig. 11). It is a green blade located at the base
of the petiole, with variable shape and size:

- Stipules: When present, they are usually in pairs of two.

- Ochrea: Corresponds to a fusion of stipules, it represents a membranous sheath at the base of the
petiole, completely surrounding the stem.

- Ligule: Mostly found in grasses, it corresponds to a doubling of the blade at the point where it
attaches to the sheath.

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Figure 11 – Characterization of leaves based on the shape of the apex and the base of the blade,
as well as venation.

II.3.2. Morphological Variations of Leaves and Adaptations

- Spine leaves: Leaves partially spiny like Holly leaves (example: Ilex aquifolium), with spiny
edges of the blade, or leaves completely transformed into spines; as seen in Cactus leaves.

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- Tendril leaves: The leaf can be partially or entirely transformed into tendrils to provide support
function. In some plants, the entire blade is transformed into a tendril, and stipules perform the
function of the leaf.

- Phyllodes: A phyllode is a flattened petiole resembling a blade in shape or at least a leaf;

Example: Acacia heterophylla.

II.4. Flowers

- A flower is a complex structure composed of specialized parts, emerging from an axis located at
the axil of a leaf, known as the floral axis.

- The lower portion of this axis is known as the peduncle, which supports the flower. At the end of
the peduncle, there's often a swollen structure called the floral receptacle.

- The floral receptacle serves as a platform for the insertion of various floral parts. It is here that
sepals, petals, stamens, and carpels are attached, forming the complete flower.

- Additionally, a bract, a modified leaf-like structure, may be found at the base of the floral
receptacle, providing further support or protection to the developing flower.

- Flowers have been historically significant in botany, serving as key elements for recognition and
classification by botanists. Their diverse structures and arrangements have played a crucial role in
understanding plant taxonomy and evolutionary relationships.

- A flower can have both male (stamens) and female (carpels) reproductive organs, making it
hermaphroditic, or it can possess only male or only female reproductive organs, leading to the
classification of unisexual flowers.

- There are unisexual species, possessing only a gynoecium (pistillate flowers), or possessing only
an androecium (staminate flowers). Sterile flowers, without stamens or carpels, may also occur
(fig. 12).

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Figure 12 – Longitudinal section and floral parts of a hermaphrodite flower.

• N.B: Carpel acts as a female part of a flower and helps in the production of seeds and
their dispersal. Pistil act as a female part of a flower but does not produce seed. Carpel
generally consists of ovary, stigma, and style. The pistils are the union of one or more
carpels.

- In relation to the possession of reproductive organs, flowers can be categorized into:

1- Unisexual flowers: These are categorized as either male flowers, containing only stamens (the
male reproductive organs), or female flowers, containing only carpels (the female reproductive
organs), examples: flower of Watermelon, and Cucumber.

2- Dioecious plants: These bear individuals that exclusively produce either male or female flowers.
This results in separate male and female plants within a species, example: flowers of date palm.

3- Monoecious plants: These have individuals that bear both male and female flowers on the same
plant. This allows for self-pollination or cross-pollination between flowers of the same plant,
example: flower of Corn.

- In a hermaphrodite flower, four types of floral parts are usually present, arranged in whorls
on the floral receptacle:

II.4.1. Sterile Parts - Perianth:

• Perianth

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The perianth consists of floral envelopes: calyx + corolla, forming the sterile part of the flower.
The characteristics of the perianth are often closely related to the type of pollination.

a- In wind-pollinated plants, perianth parts, if present, are often reduced, not hindering the
transport of pollen to the flower's reproductive organs.

b- In insect-pollinated plants, flowers have larger perianth parts, brightly colored with often an
attractive fragrance.

➢ When perianth parts are not differentiated into petals and sepals, they are called tepals
(when sepals and petals are of identical shape and color). The perianth can be persistent or
deciduous, accrescent (referring to a part of the flower that continues its growth after
fertilization) or not. The number of perianth parts (trimerous, tetramerous, pentamerous, or
polymorphic), their shapes, relationships (free or fused), symmetry (relative to an axis or
plane), are all characteristics used in systematics.

A- Calyx

The outer envelope of the flower, usually green in color (chlorophyllous parts), covering most often
the base of the corolla and consisting of all the sepals. These sepals can be free to the base
(dialysepalous calyx) or more or less fused together (gamosepalous calyx) (fig. 13). In some
flowers, the calyx is colored like the petals, and they are then called petaloid sepals or tepals.

B- Corolla

The ensemble formed by the petals of a flower. Generally colorful (non-chlorophyllous parts). The
corolla functions to attract pollinating insects. It can be persistent or deciduous. The petals can be
free (dialypetalous corolla) or more or less fused together (gamopetalous corolla) (fig. 13).

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Figure 13 – Welding of perianth parts (from left to right; dialypetalous, gamopetalous

dialysepalous, gamosepalous).

- Classifying Perianth Variations (Distinctions and Categories):

1- Actinomorphic flower: when it is regular, meaning it exhibits radial symmetry (with respect to
an axis) (fig. 14).

2- Zygomorphic flower: when it is irregular and shows a plane of symmetry (with respect to a
plane) (fig. 14).

3- Asymmetric flower: when it lacks any plane of symmetry.

Dialypetalous corolla Gamosepalous corolla

(from left to right: actinomorphic and zygomorphic flowers).

Figure 14 – Symmetry of flowers.

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II.4.2. Fertile Parts (Reproductive Parts):

a. Androecium

The androecium comprises the stamens, constituting the male part of an angiosperm flower. The
stamen, borne by the floral receptacle and positioned between the corolla and the pistil. A stamen
typically consists of a slender elongated portion called the filament, ending in an enlarged part
called the anther. The anther is the site of pollen production (male gametophyte), within cavities
known as pollen sacs (microsporangia), usually four in number, often fused into two locules
separated by a connective. The stamens, collectively forming the androecium, may be free to the
base or more or less fused to each other or to other parts of the flower (petals, pistil). Flowers with
only stamens and no pistil are called staminate flowers. At maturity, the anther opens, releasing
pollen, which enables fertilization (fig. 15).

Figure 15 – Structure of a stamen and cross-section of a mature anther.

b. Gynoecium

Also known as the pistil, the female reproductive organ of the plant, formed by one or more
carpels of the same flower, either free or fused together (partially or entirely). Each carpel consists,
from base to apex (fig. 16), of:

- An enlarged portion (the ovary) containing one or more ovules;

- A style extending from the ovary;

- A stigma capping the style, retaining pollen, serving as the passage for the pollen tube into the
ovary cavity. The style and stigma take on various forms.

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- A significant portion of the gynoecium persists after fertilization and develops into fruit. The
gynoecium may sometimes be reduced to a single carpel.

Figure 16 – The gynoecium.

- Placentation Types (Variations in Ovule Position Inside the Ovary): (fig. 17)

1- Axile placentation: ovules are attached to the center of an ovary formed of several locules.

2- Parietal placentation: ovules are inserted on the peripheral wall of a non-compartmentalized

ovary.

3- Central placentation: in a unilocular ovary resulting from the fusion of several closed carpels
whose septa have resorbed, only a central column remains to which ovules are attached.

Figure 17 – Various types of placentation: Axile, Parietal, and Central.

- The arrangement of ovaries and floral parts shows three cases (fig. 18):

✓ Hypogynous: describes a flower whose floral parts (sepals, petals, and stamens) are
inserted beneath the ovary, which is said to be superior.
✓ Epigynous: describes the organs (stamens, floral envelopes) of a flower with an inferior
ovary, situated above the gynoecium.

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✓ Perigynous: describes a flower whose floral envelopes and androecium are inserted around
the semi-inferior ovary, free at the bottom of the hollow receptacle.

Figure 18 – Arrangement of ovaries and floral parts.

II.5. Flower Inflorescence

Flower inflorescence refers to how flowers are organized or clustered on the stem of a flowering
plant. This arrangement can vary greatly among different plant species and can be an important
characteristic for plant identification.

➢ Flower Inflorescence types:

a- Definite Inflorescences (Cymose): In definite inflorescences, the main axis of the inflorescence
terminates in a single flower. This means that the primary stem ends in a flower bud, which
eventually develops into a flower (fig. 19). Examples of definite inflorescences type: include
Raceme, Catkin, Spike.

b- Indefinite Inflorescences (Racemose): In indefinite inflorescences, the main axis of the

inflorescence terminates in a bud, which continues to elongate and produce more flowers along the
axis. This results in a continuous or branching pattern of flowering (fig. 19). Examples of indefinite
inflorescences type: include uniparous cyme, biparous cyme and multiparous cyme.

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Figure 19 – Examples of defined and undefined inflorescences.

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