Lebanese University

Lebanese University Faculty of Sciences

Faculty of Sciences

B1100
Plant Histology
Department of Life and Earth Sciences

Fall Semester
2020/2021

©opyright Reserved
The following book was prepared and revised by the following instructors

Prepared by: Revised by:

Dr. Romeo AL BERSAOUI Dr. Hanadi ISMAIL
Dr. Bouchra DOUAIHY Dr. Jihad NOUN
Dr. Myrna MEDLEJ-HASHIM
Dr. Randa SAOUD

Copyright Reserved for the Lebanese University ©2018

This material is only for academic use by the students at the Faculty of Sciences in the
Lebanese University, it should not be distributed for purchase or photocopy anywhere
under the threat of legal prosecution.

8102© ‫حقوق الطبع والنشر محفوظة للجامعة اللبنانية‬

‫إ ن هذه النسخة موضوعة بتصرف طالب كلية العلوم في الجامعة اللبنانية و لهدف اكاديمي فقط ال غير وعليه يمنع‬
‫توزيع اي نسخة (ورقية او الكترونية) الي جهة اخرى او التصوير والبيع في كافة المكتبات تحت طائلة المالحقة‬
.‫القانونية‬
Table of Contents
I. Introduction
1 Plants versus animals .......................................................................................................................... 3
2 The cell wall......................................................................................................................................... 4
2.1 The primary cell wall .................................................................................................................... 5
2.2 The secondary cell wall ................................................................................................................ 7
2.3 Pits and plasmodesmata .............................................................................................................. 8
3 Classification of plant tissues .............................................................................................................. 9
II. The plant tissues
1 Meristematic tissues ......................................................................................................................... 10
1.1 Primary or apical meristems ...................................................................................................... 10
1.2 Secondary or lateral meristems ................................................................................................. 11
2 Differentiated tissues ........................................................................................................................ 12
2.1 Protective (dermal) tissues ........................................................................................................ 12
2.1.1 Epidermis ............................................................................................................................ 12
2.1.2 Periderm ............................................................................................................................. 16
2.2 Ground tissues............................................................................................................................ 18
2.2.1 Parenchyma ........................................................................................................................ 18
2.2.2 Collenchyma........................................................................................................................ 21
2.2.3 Sclerenchyma ...................................................................................................................... 22
2.3 Conducting or vascular tissues ................................................................................................... 23
2.3.1 Xylem................................................................................................................................... 23
2.3.2 Phloem ................................................................................................................................ 25
2.4 Secretory tissues ........................................................................................................................ 27
2.4.1 Secretion cells isolated within the parenchyma tissue ...................................................... 27
2.4.2 Secretory epidermis and glandular hairs ............................................................................ 28
2.4.3 Glands and canals ............................................................................................................... 28
2.4.4 Laticiferous tissue ............................................................................................................... 29

2
 I. Introduction
1 Plants versus animals
The differences between animals and plants are not in the basic molecular characteristics, but they are
rather related to cell and tissue functions of higher order.

Plants are differentiated from animals by:

 Their ability to manufacture food through photosynthesis.

 Their unlimited type of growth in very localized regions of growth.

On the cellular level the plant cells differ from the animal cells by (Fig. 1):

 The presence of cell walls.

 The absence of centriole in plant cells.
 The presence of different types of plastids (e.g. Chloroplast, chromoplast, amyloplast, leucoplast).
 The presence of a large central vacuole in differentiated plant cells that generally occupies around
90% of the cell volume. It is a large vesicle filled with many different substances in solution.

Figure 1 Typical animal and plant cells.

3
Some animals and some plants are unicellular, but
most organisms are multicellular, and organized on
the basis of tissues, organs and systems.

The plant body includes two major parts or organ

systems (Fig. 2):

 The root
 The shoot :
o Stems
o Leaves
o Reproductive organs (flowers and
cones)

Figure 2 The different parts of a vascular plant body.

2 The cell wall

The plant cells have a strong, porous, rather rigid cell wall. It is the outer boundary of plant cells, consisting
of non-living structure, secreted by the living part of the cell. Although it is a product of the protoplasm,
it is not considered as part of it.

The cell wall constitutes a kind of exoskeleton providing mechanical support and protection required for
the plant cell. It also provides transport functions between cells through plasmodesmata. Unlike the cell
membrane, substances can freely pass through the cell wall without selectivity.

During cell division, before cell wall development, granules appear between daughter cells. These
granules containing hemicellulose and pectin fuse together to form the cell plate which will then develop
into the middle lamella (intercellular layer common to both cells). The viscous, gelatinous nature of pectin
gives the middle lamella its role as a cement connecting the cell walls of adjacent cells (Fig. 3).

4
 Figure 3 Diagram showing the development of a new cell wall
(A). Transverse section showing the cell walls and middle lamella
of three adjacent plant cells (B). .

2.1 The primary cell wall

The primary cell wall is the first part of the cell wall laid down against the middle lamella by the protoplast
of a young growing cell. It is rather plastic and capable of extension as the cell grows. It consists of
carbohydrates (90%) and hydrated proteins:

 Mainly cellulose: a linear and unbranched polysaccharide macromolecule consisting of 300 to more
than 3,000 glucose subunits linked to each other (Fig. 4).

Figure 4 Cellulosic chain

5
  Hemicellulose: a mixture of neutral polysaccharides including pentoses (xylose and arabinose) and
hexoses (the most frequent being the galactose, mannose and glucose) macromolecules.
 Some pectin: a gel mixture consisting of acidic polysaccharides.
 Small amount of glycoproteins.

The skeleton is made up of cellulose molecules linked together by hydrogen bonds to form microfibrils,
which in turn are twisted together to form macrofibrils (Fig. 5). The macrofibrils are grouped into cellulose
fibers that form an irregular meshwork constituting the bulk of the cell wall. Hemicellulose, pectin and
the glycoproteins form an amorphous matrix in which the cellulose fibers are embedded (Fig. 6).

The primary wall becomes more rigid as additional cellulose molecules are added after the cell has reached
its mature size. The primary cell wall is the only cell wall of undifferentiated cells as well as some
differentiated cells, and of cells which growth has not been achieved yet. It is capable of growing in length
(by stretching of the network of already formed microfibrils) and in width (by cell synthesis of new
constituants: microfibrils and matrix). The primary wall participates, therefore, by growing, to the
elongation of young cells, which is necessary for the growth in length of plant organs.

Figure 5 The cellulose fiber structure

6
Figure 6 Primary walls (a) Surface view of the primary wall of a carrot (Daucus carota) cell, showing cellulose
microfibrils cross-linked by an intricate web of matrix molecules. (b) Schematic diagram showing how the
cellulose microfibrils are cross-linked into a complex network by hemicellulose molecules.

2.2 The secondary cell wall

The cells of the soft tissues of the plant have only primary walls. In harder parts of the plant and cells with
relatively thick walls, a secondary wall, usually of three layers, is produced and deposited between the
primary wall and the plasma membrane, after cell growth is over (Fig. 7). The secondary wall can increase
in width but not in length.

The secondary wall is much thicker than the

primary wall and is composed of a succession
of compact layers or lamellae. These layers are
basically made of cellulosic material. Cellulose
fibrils lie parallel to each other and are generally
oriented at angles of 60° to the fibrils of the next
lamella. This arrangement gives strength to the
cell wall unlike the primary cell wall in which
fibrils are arranged in an irregular network. In
addition to cellulose, other materials may also Figure 7 Diagram showing the organization of the cellulose
be present in the secondary wall layers mainly microfibrils and the three layers (S1, S2, S3) of the secondary
wall. The different orientations of the three layers strengthen
lignin. Lignin is a complex insoluble polymer the secondary wall.
responsible for hardness and decay-resisting
quality of many woods. It adds mechanical strength to the cell wall. Most cells with secondary walls are

7
devoid of living material at maturity. They are dead empty cells which main function is structural strength,
mechanical support and protection.

2.3 Pits and plasmodesmata

Cell walls are not generally completely uninterrupted boundaries around the cells. There are often tiny
holes in the walls through which delicate protoplasmic connections between adjacent cells may run. These
connections are called plasmodesmata. Intercellular exchange of materials takes place through the
protoplasmic connections. They are usually located in areas called pits where the cell wall is very thin and
no secondary wall is deposited (Fig. 8).

Figure 8 Primary pit-fields, pits, and plasmodesmata (a) Cells with primary walls and primary pit-fields,
which are thin areas in the walls. As shown here, plasmodesmata commonly traverse the wall at the
primary pit-fields. (b) Cells with secondary walls and numerous simple pits. (c) A simple pit-pair. (d) A
bordered pit-pair.

8
3 Classification of plant tissues
Plant histology is the study of plant tissues. A plant tissue is a set of cells serving the tissue function.
Differentiated adult tissues are derived from undifferentiated tissues, the meristems. The process of the
transformation of meristem into differentiated tissues is called differentiation. We distinguish in adult
plants different types of differentiated tissues classified according to their function.

Plant tissues of higher land plants (vascular plants) can be classified into two major categories (Fig. 9):

1. Meristematic tissue
1.1. Primary meristems
1.2. Secondary meristems
2. Differentiated tissues:
2.1. Protective (dermal) tissues:
2.1.1. Epidermis
2.1.2. Periderm
2.2. Ground tissues:
2.2.1. Parenchyma
2.2.2. Collenchyma
2.2.3. Sclerenchyma
2.3. Vascular tissues:
2.3.1. Xylem
2.3.2. Phloem
2.4. Secretory tissues Figure 9 The distribution of the ground tissue system, vascular tissue system, and
dermal tissue system in the (a) leaves, (b) stems, and (c) roots of a herbaceous
dicot such as Arabidopsis.

Each of these tissue types is found in all vascular plants except for the secretory tissues that are plant
specific.

9
 II. The plant tissues
1 Meristematic tissues
The term meristems originates from the Greek term “meristos” = divisible. The meristematic cells are in
a state of rapid division, they form undifferentiated tissues (embryonic), which will turn into differentiated
tissues. Some of these cells however remain meristematic and divide repeatedly. These are the initials and
serve to perpetuate the meristems.

There are two types of meristems differing from one another by their location, cytology and role in the
formation of different tissues: Primary and secondary meristem.

1.1 Primary or apical meristems

Apical meristems are found at or near the growing tips of plant organs, such as roots or shoots (Fig. 10).
They insure the growth in length. This type of growth is called primary growth. The cells are small,
isodiametric and contiguous (without intercellular air spaces). Stained sections of roots or stem tips show
primary meristematic cells to have a thin primary cellulosic wall, a large nucleus, a cytoplasm containing
small spherical or arranged in a fine network vacuoles and many mitochondria and no differentiated
plastids (proplastids) (Fig. 11). The tissues produced by the primary meristems are called primary tissues.

Figure 11 Meristematic cells in the apical

Figure 10 The position of apical meristems in a plant. meristem of the Epicea: Picea excelsa.

10
1.2 Secondary or lateral meristems

The secondary meristems (also called cambia) are

localized in the older parts of some roots and stems
ensuring their growth in width (Fig. 12). This type of
growth is called secondary growth. The secondary
meristematic cells have thin primary cell wall, large
vacuoles pushing the nucleus towards the periphery of the
cytoplasm, and no differentiated plastids. The cells have
a rectangular shape and are regularly disposed (Figs. 13,
14). The tissues produced by the cambia are produced after
the primary tissues have matured and are called secondary
tissues.

Figure 12 Localization of the plant meristems

Figure 13 Cytology of a cambium cell. Figure 14 Cambium in a vascular bundle.

There are two types of cambia:

 The vascular cambium, often referred to simply as the cambium, produces secondary tissues that
function primarily in support and conduction.
 The cork cambium which lies outside of the vascular cambium, between the cork and the
phelloderm, which it produces.

11
2 Differentiated tissues
2.1 Protective (dermal) tissues

They form the protective outer covering of the plant body. These tissues protect plant organs from external
factors such as climatic variations, mechanical injury, and pathogens attack... These tissues are resistant
and are impermeable to water. There are two types of protective tissues: epidermis and periderm.

2.1.1 Epidermis
The epidermis is a primary tissue originating from the apical meristem and is found in all plants. In some
it will be replaced by cork and phelloderm in the older plant parts.

2.1.1.1 The epidermal cells

The epidermis is usually one cell thick but may be thicker in
plants living in very dry habitats. The epidermis cells are variable
and irregular in shape (Fig. 15). They interlock tightly like pieces
of a puzzle, with no intercellular space (to be a good barrier).
They are living cells and generally have a very large vacuole and
only a thin layer of cytoplasm. They lack chloroplasts except in
aquatic plants. Often their outer and side walls are thicker than
the inner wall (Fig. 16).

Figure 15 Epidermal cells of different plants.

Figure 16 Transverse section of the epidermis

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Epidermal cells on the aerial parts of the plant secrete a waxy, water-resistant layer, the cuticle, on their
outer surface. It is composed of the fatty material (cutin) and waxes. The cuticle thickness varies,
depending on the age and species of plants. It is very thin and even absent in plants living in humid
environments but very thick in plants living in dry conditions. The cuticle, with the thick outer wall, helps
in protection against water loss, mechanical injury, and invasion by fungi and bacteria.

In the piliferous layer, at a short distance from the

root tip, the epidermal cells of the roots have no
cuticle and function in water absorption. These cells
produce tubular hairlike extensions called root hairs.
The root hairs greatly increase the absorptive surface
area (Fig. 17).

Figure 17 Root tip

2.1.1.2 The stomata

Among the epidermal cells of leaves and young green stems may be found numerous pores, called
stomata1 (singular: stoma), surrounded by pairs of crescent-shaped cells called guard cells. Below each
stoma is a space called sub-stomatal chamber, which is an air chamber (Fig. 18).

Stomata function in allowing gas exchange between the interior of the leaf and the atmosphere and in
regulating the evaporation (transpiration) of most of the water entering the plant at the roots.

Guard cells have thicker cell walls towards the pores and they contain chloroplasts and are capable of
undergoing changes in size and shape dependent upon changes in their turgor pressure. Increased turgor
causes them to open, and vice versa (Fig. 19).

1
The term “stoma” is applied in some references to the pore and the two guard cells.

13
Figure 18 Leaf anatomy (a) organization of the three tissue systems (b) surface view of a Tradescantia leaf (c)
Palisade and spongy regions of mesophyll within a leaf of a lilac.

Figure 19 How stomata open and close. As turgor pressure in the guard
cells increase, the thinner outer walls stretch more than the thicker inner
walls, causing the stoma to open.

14
2.1.1.3 Hydathodes (water pores)

In some plants, especially those living in humid habitats, excess water is expelled through special pores
called hydathodes. These are located at leaf border, are always open, are not associated with chlorophyllian
guard cells and sub-stomatal chamber. This loss of water in liquid form is called guttation (Fig. 20).

Figure 20 Left: Longitudinal view of a hydathode of the leaf of saxifrage (Saxifraga lingulata). Right: Guttation
droplets, seen here at the margins of a leaf of lady’s mantle (Alchemilla vulgaris).

2.1.1.4 The epidermal hairs

Epidermal hairs (trichomes) are elongated outgrowths of one or more epidermal cells. They may be
unicellular or multi-cellular (simple or branched), glandular or non-glandular (Fig. 21). These may protect
the plant against biotic and abiotic factors e.g. insects, high sunlight intensity etc…

15
Figure 21 Trichomes (a) Flat. (b) and (c) Branched trichomes, (d), (e) and (f) Glandular trichomes. (g) and (h)
Simple unbranched, nonangular trichomes.

2.1.2 Periderm
As the stems and roots of woody plants with active lateral meristems increase in diameter, the epidermis
is slowly replaced by the periderm. The periderm is mostly composed of cork cells produced by the cork
cambium and some parenchyma cells forming the phelloderm (Fig. 22).

The cork is a protective tissue composed of many layers of dead flat cells with suberified waterproof
primary wall. Gas exchange takes place through lenticels (pockets of loosely arranged parenchyma cells
that are not impregnated with suberin and appears as dots and stripes on the bark surface) (Fig. 23).

16
Figure 22 A. Diagram showing origin of the first cork cambium and the first layer of cork. B. Light micrograph
showing cork cambium and newly formed cork in an elderberry stem.

Figure 23 Left: Cross section of a portion of young stem of elderberry showing the lenticel. Right: Wine cork

17
2.2 Ground tissues

2.2.1 Parenchyma
Parenchyma is a simple tissue found in all plants. It can be primary or secondary, consisting of parenchyma
cells with various functions and shapes containing well-developed vacuoles. The various types of
parenchyma have in common usually a thin primary cellulosic cell wall with intercellular air spaces.
Parenchyma cells are living cells at maturity and are capable of mitotic divisions (the capacity of
parenchyma to multiply is especially important for tissue repairs when a plant is damaged or wounded).
According to their functions we can distinguish: chlorophyll parenchyma, reserve parenchyma, and
aerenchyma.

Parenchyma cells can be also found as a part of conductive tissues (xylem and phloem).

2.2.1.1 Chlorophyll parenchyma (chlorenchyma)

Chlorophyll parenchyma is abundant in the aerial parts of the plant (leaves, young stems, fruit) for which
it gives the green color. The presence of chloroplast implies a role in photosynthesis. The cells are
detached from each other at their corners forming intercellular spaces and sometimes air cavities insuring
gas flow (Fig. 24). In leaves, it is called the mesophyll, surrounded by the epidermis and crossed by the
veins.

Figure 24: Chlorophyll parenchyma A) with intercellular spaces B) with air cavities

In monocotyledon plant leaves (wheat, corn ...), the mesophyll is homogeneous (Fig. 25) whereas in
dicotyledon plant leaves (roses, daisies …), it is heterogeneous (Fig. 26):

18
  Upper or ventral side: the palisade parenchyma consists of one or more layers of elongated
chloroplastic cells, clamped against each other.
 Lower or dorsal side: the spongy parenchyma formed by rounded cells separated by air cavities
communicating with the stomata.

Figure 25 Transverse section of a portion of a maize (monocotyledon plant) leaf.

Figure 26 Transverse section in a leaf of a dicotyledon plants.

19
2.2.1.2 Reserve parenchyma
Reserve parenchyma is located in the plant’s roots, stems and seeds.
Devoid of chloroplasts, the cells contain different reserve substances such
as starch grains (Fig. 27) (e.g. potato tubers), sucrose (beet tuberous root),
oils (seeds), proteins (wheat grain, legumes), cellulose (date seeds)…
Reserve parenchyma can also store water (aquifer parenchyma)
in leaves and stems of succulent plants.

Figure 27 Starch granules in a

cell of a Ricinus communis seed.

2.2.1.3 Aerenchyma
In aquatic plants, the intercellular spaces are quite extensive and full of air and form a network throughout
the entire plant. This type of parenchyma tissue is called aerenchyma. The aerenchyma is responsible for
air circulation between the roots and shoots (Fig. 28).

Figure 28 Aerenchyma in a cross section of a semiaquatic Makaloa Stem

(Cyperus laevigatus)

20
2.2.2 Collenchyma
Collenchyma is a simple primary supportive tissue located on the periphery, underneath the epidermis, of
aerial organs only. It is composed of several layers of living cells with no air spaces (Fig. 29). Although
they are more elongated, they are structurally similar to parenchyma cells (have a large vacuole surrounded
by a peripheral thin layer of cytoplasm, absence of secondary cell wall), except that their cellulosic primary
cell wall is thicker than that of parenchyma cell.

Figure 29 Cross section in a plant stem showing the collenchyma tissue

laying between the epidermis and the parenchyma tissues.

Three forms of collenchyma are recognized based on the types of cellulose thickenings (Fig. 30):

 Annular collenchyma : uniformly

thickened cell walls
 Lamellar collenchyma: thickenings are
on the tangential wall.
 Angular collenchyma: thickenings are
at the intercellular contact points.

Figure 30 The three forms of collenchyma (A) and

(D) Angular collenchyma, (B) Lamellar
collenchyma, (C) Annular collenchyma.

21
Collenchyma is an important supporting and strengthening tissue in young plants, in the stems of non-
woody older plants and in leaves. It is plastic and stretches irreversibly with the growth of the organ in
which it occurs. It provides flexible support for both growing organs and mature organs such as leaves
and floral parts.

2.2.3 Sclerenchyma
Sclerenchyma is a dead supportive simple tissue that can have primary or secondary origin. It is usually
found in old organs throughout the plant body. The uniformly very thick, heavily lignified secondary wall
gives strength to the sclerenchyma cell body. Sclerenchyma is devoid of air spaces and is stiffer and harder
than collenchyma (Fig. 31).

Figure 31 Sclerenchyma at vascular bundles of a sunflower

stem. Ed Reschke/Photolibrary/Getty Images

We distinguish:

 Fibers: elongated fusiform flexible cells. Fibers occur in roots, stems, seeds, leaves and fruits as
supporting elements. These fibers are used in the textile industry.
 Stone cells (sclereids): Short cells of different shapes with very thick lignified walls. They are
common in the shell of nuts and in the hard part of seeds and in the flesh of hard fruits. They form
the cores of apples and produce the gritty texture of pears.

22
2.3 Conducting or vascular tissues

They are primary or secondary complex tissues ensuring the sap circulation in the different parts of higher
vascular plants. There are two types of conducting tissues: the xylem and the phloem.

2.3.1 Xylem
The xylem ensures the circulation of the crude sap, a very dilute aqueous solution of mineral salts, from
the roots to the aerial parts of the plant. This tissue can be primary or secondary tissue (wood). The wood
insures a mechanical support of the whole plant. The xylem is a complex tissue consisting of conducting
elements (vessels and tracheids), parenchyma cells and fibers.

2.3.1.1 The conducting or tracheary elements

 The tracheids: primitive conducting elements found in all vascular plants. They are elongated
cells with tapered ends. Their heavily lignified secondary cell wall gives them a supporting role in
addition to their sap conduction role. Pits may occur
anywhere on the cell wall, but they are often
particularly numerous on the tapered end of the cell.
The sap circulation occurs through the pits. Tracheids
of the first matured primary xylem are stretched
during the development of the plant and become non
functional. Their secondary walls in primary xylem
are usually in the form of rings (annular tracheids) or
spirals (spiral tracheid). It will be replaced by a
secondary xylem. Tracheids of secondary xylem arise
after all lengthwise growth has ceased, and they are
not stretched. Their secondary walls are thicker and
more continuous, being interrupted only by numerous
pits (pitted tracheids) in Gymnosperms (Figs. 32, 33
and 34). In Pteridophytes the secondary walls are
ladder-like.

Figure 32 Pitted tracheids.

23
Figure 33 Tracheids and vessels.
© Pearson Education, Inc.,
publishing as Benjamin
Cummings.

Figure 34 Different lignin depositions in tracheids

shown in cross (a) and longitudinal (b) sections.
Biology of Plants 7th edition © W.H. Freeman and
Company.

 The vessels: highly specialized conducting elements of the most evolved vascular plants
(angiosperms) made up of individual dead cells called vessel elements or vessel cells. These dead
cells form large diameter cavities bounded by a lignified cell wall. During the differentiation
process vessels are formed by the breaking down of the transverse wall forming the perforation
plate. If the transverse walls are extensively perforated they form compound perforation plates
whereas if they are completely dissolved they form a simple perforation plate. As in tracheids,
vessels can be annular, spiral, reticulate or pitted.

24
2.3.1.2 The fibers
The fibers have a supporting role along with the tracheids (Fig. 35).

Figure 35 Transverse section of xylem fibers and vessel (A), longitudinal section of a fiber (B).

2.3.1.3 The parenchymal cells

They are living cells playing a role in lateral conduction of sap (ray parenchyma) or in storage.

2.3.2 Phloem
Phloem is the conducting tissue of the elaborated sap (organic nutrients) produced by photosynthesis
throughout the plant. It can be primary or secondary complex tissue. It comprises the conducting sieve
elements, companion or albuminous cells, fibers and parenchyma cells (Fig. 36).

2.3.2.1 Conducting elements

This is the most important structural and functional element of the phloem. It consists of sieve tubes
associated with companion cells in angiosperms (flowering plants), and of sieve cells associated with
albuminous cells in gymnosperms and pteridophytes.

25
2.3.2.2 Sieve cells and sieve tubes
The sieve cells are superposed elongated living
cells with cellulosic cell walls. The sieve tubes are
formed of superposed elongated living sieve tube
elements with cellulosic cell walls.

The sieve cells of pteridophytes and

gymnosperms have their longitudinal and
transverse walls perforated with pores
interrupting the cell walls and the middle lamella
allowing the flow between adjacent cells. The
sieve tube elements in angiosperms also have their
longitudinal walls perforated with pits but their
transverse walls are thickened with larger pores,
forming the sieve plate. Sieve plate allows the
flow of small and large molecules, such as
proteins, from one sieve tube element to another. Figure 36 Longitudinal section in a phloem showing the
sieve tube elements, companion cells and parenchyma
Unlike tracheids and vessels of xylem, their nuclei cells.
disintegrate at maturity but their cytoplasm
remains and is very active in conducting food materials in solution throughout the plant.

2.3.2.3 Companion cells

Companion cells are specialized, elongated parenchymatous cells. They retain both their nuclei and their
cytoplasm at maturity. They are closely associated with the sieve tubes communicating through numerous
plasmodesmata. Their nucleus controls both their own cytoplasm and the cytoplasm of the adjoining sieve
element after the nucleus of the latter has disintegrated. The function of a sieve cell lasts a short time.
When the sieve cell dies, the companion cell divides longitudinally to give two cells, one of which will
replace the dead sieve cell and the other cell gives the companion cell.

26
2.3.2.4 The fibers
In phloem, fibers are less abundant than in the xylem.

2.3.2.5 The parenchymal cells

They are living cells playing a role in lateral conduction of sap or in storage.

2.4 Secretory tissues

The secretory tissue is plant specific and localized in specific parts of the plant body. This tissue consists
of epidermal or parenchyma cells which synthesize substances representing the secondary metabolic
products of plants (essential oils, tannins, resin, latex ....). They can accumulate the products synthesized
in their cells or secrete them in cavities.

2.4.1 Secretion cells isolated within the parenchyma tissue

Secretion cells or secretion sacs accumulate in their vacuoles the product they secrete like the tannin in
the secretory cells of rose stems (Fig. 37) and the essential oils in the secretory cells of camphor leaves.

Figure 37 Tannin cells in the pith of a rose stem.

27
2.4.2 Secretory epidermis and glandular hairs
Epidermal cells can develop and accumulate in their cytoplasm essential oils (epidermis of rose petals)
(Fig. 38.a). Multicellular trichomes can also
be secretory (glandular hairs). They
accumulate in their terminal cells essential oils
like in thyme, oregano, lavender ...
(Fig. 38.b).

Figure 38 Secretory epidermis (a) and hairs (glandular

trichomes) (b).

2.4.3 Glands and canals

Glands are globular cavities situated in the parenchyma of leaves, stems and fruits (oranges, tangerines,
lemons). They are surrounded by cells which excrete the products they elaborate in the glands (Fig. 39).
When these cavities are not restricted in location, but extend all the length of an organ; they are tube-like
spaces, elongated in the longitudinal plane of the organ (leaf, petal, and stem), they are called secretion
canals or ducts (Fig. 40). In cross section, they appear as circular cavities surrounded by one or two layers
of small secretory cells (Example: pine leaves secreting resin).

Figure 39 Gland (globular cavity) in the peel of an orange. Figure 40 Pinus resin duct (cross section).

28
2.4.4 Laticiferous tissue
Laticiferous tissue is a living tissue containing secretory tubes (laticifers) elaborating the latex; a milky
white (e.g. in lettuce, fig) or colored (e.g. brown yellow in cannabis, yellow in celandine and red in
bloodroot) viscous liquid (very complex mixture containing water, sugar, mineral salts, organic acids,
enzymes, etc.). Some forms of latex are commercially very important: rubber, chewing gums, medicinal
drugs… The laticifers are limited by a cellulosic primary wall. Their cytoplasm forms a thin layer
surrounding a large central vacuole containing latex.

There are two types of laticifers:

• Non-articulated laticifers: each laticifer is formed by a single large cell, very elongated and
containing many nuclei (coenocyte) which may reach several meters or even ten meters long. These
laticifers remain independent and will not anastomose.

• Articulated laticifers: Unlike non-articulated laticifers that are derived from a single cell, each
articulated laticifer is a structure made of individual laticiferous cells arranged in rows. Transverse walls
that separate the various individual cells may persist or be perforated or be dissolved (Fig. 41). These
laticifers can remain independent or anastomose at certain points.

Figure 41 Articulated laticifers – A: transverse walls persisted (Convolvulaceae) – B: transverse walls are
perforated (Chelidonium) – C: transverse walls are completely dissolved (Asteraceae)
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References
 Introductory Plant Biology by Kingsley R. Stern, Wm. C. Brown (WCB) Publishers, 9th edition, 1994.
 Botany , An Introduction to Plant Biology by James D. Mauseth, Jones & Bartlett Publishers, 3rd
edition, 2003.
 Biology of Plants by Peter H. Rave, Ray F. Evert & Susan E. Eichhorn, W. H. Freeman and Company
Publishers 8th edition, 2013.
 Biology by Eldra P. Solomon, Linda R. Berg & Diana W. Martin, Thomson Brooks/Cole Publishers,
8th edition, 2008.

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