EARTH AND LIFE SCIENCE

How Plants Survive

This module will introduce you to the function and form of plants. It
will also focus on how plants develop and grow. An understanding of
plant form and function is integral to understanding the natural world
since plants are essential for the survival of all life on earth. As
members of the biosphere, plants govern the oxygen exchange on the
planet, and are thus important subjects for study.

Plant Structure
Plants have structural adaptations to their environment. However, in
addition to this, plants have developed a specific morphology, or
external form, that they accumulated through natural selection. For
instance, cacti have become so specialized for the desert environment
that their leaves have become spines, and their stems are little more
than photosynthetic organs. The adaption of leaf morphology has
added to the success of cacti in the desert environment because the
surface areas of their leaves are reduced, which means that they lose
less water. Both genetic and environmental factors influence form in
both plants and animals. However, the effect of the environment in
greater in plants. Consequently, the morphology of plants vary widely
among species compares to animals.

Plant Organs, Tissues, and Cells

Like animals, plants have organs composed of different tissues, which

are, in turn, composed of different types of cells. A tissue is a group of
cells with a common structure, function, or both. An organ consists of
several types of tissues that, together, carry out particular functions for
the organism.

The three basic plant organs are roots, stems, and leaves. The basic
morphology of most vascular plants reflect their evolutionary history
as terrestrial organisms that inhabit and draw resources from below
ground and above ground. Plants need to absorb water and minerals
from below the ground surface and CO2 and light from above the
ground. The ability to acquire these resources resulted in three distinct
organs which are morphological features- leaves, stems, and roots.

These organs form a shoot system and a root system, with the former
consisting of stems and leaves. With very few exceptions, angiosperns
and other vascular plants rely on both these systems for survival.

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Roots are usually non-photosynthetic and they starve unless

photosynthates, which are sugars and carbohydrates imported from the
shoot system, arrive at the roots. Conversely, the shoot system depends
on the water and minerals that roots absorb from the soil.

Roots

Roots are multicellular organs that anchor vascular plants in the soil.
They also absorb water and minerals, and they often store
carbohydrates. Most eudicots and gymnosperms have taproot systems.
A taproot system consists of a main vertical root, the taproot, that
develops from the embryonic root. The taproot gives rise to lateral
roots, which are also called branch roots. In many angiosperms, the
taproot stores carbohydrates and sugars that the plant will consume
during flowering and fruit production. For this reason, many crops,
such as carrots and beets, are harvested before they flower. Taproot
systems generally penetrate deep into the soil and are well-adapted to
accessing sources of water that are far from the ground surface.

In seedless vascular plants, as well as in most monocots, such as

grasses, the embryonic root dies and does not give rise to the main
root. Instead, many small roots grow from the stem. These roots are
said to be adventitious, which is a term that describes a plant organ
that grows in an unusual location, such as roots arising from stems and
leaves. Each small root forms its own lateral roots. The result is a
fibrous root system. This is generally a mat of thin roots spreading out
below the soil surface, and no root functions as the main root. Fibrous
root systems usually do not penetrate the ground deeply and are
therefore best for shallow soils or regions where rainfall is light and
does not moisten the soil much below the surface layer. Most grasses
have this type of root system, which does not penetrate far underneath
the soil surface. These shallow roots hold the topsoil in place, which
means that they make excellent ground cover for preventing soil
erosion.

Although the entire root system helps anchor the plant into the soil,
most plants absorb minerals and water primarily near the tips of roots.
This is where vast numbers of root hairs are located, and these increase
the surface area of roots enormously. Root hairs are short-lived and
constantly being replaced. A root hair is a thin, tubular extension of a
root epidermal cell. It should not be confused with a lateral root, which
is a multicellular organ. Despite their large surface area, root hairs
contribute little to the anchoring of plants to the soil. Many plants have
modified roots. Some of these arise from roots, while others are

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adventitious, developing from stems. In rare cases, roots may also

develop from leaves.

Stems

A stem is an organ that consists of alternating systems of nodes. These

are the points at which leaves are attached, and internodes, which are
the segments of stem between the nodes. In the upper angle formed by
the leaf and stem (axil) is an axillary bud. The axillary bud is a
structure that can form a lateral shoot, commonly called a branch.
Most axillary buds of a young shoot are dormant. Thus, the elongation
of a young shoot is usually concentrated near the shoot tip. This
consists of an apical bud, or terminal bud, with developing leaves and
compact series of internodes and nodes.

The proximity of the axillary buds to the apical buds is partly

responsible for their dormancy. The inhibition of axillary buds by an
apical bud is called apical dominance. Through the concentration of
resources on elongation, the evolutionary adaptation of the apical
dominance increases the plant’s exposure to light. If an animal, for
instance, eats the end of the shoots, the axillary buds break their
dormancy, and they start growing. A growing axillary bud gives rise to
a lateral shoot, complete with its own apical buds, axillary buds, and
leaves. This is the reason why pruning trees and shrubs will increase
the number of leaves. Some plants have stems that have additional
functions, such as for food storage and asexual reproduction. These
modified stems, which include rhizomes, bulbs, stolons, and tubers,
are often mistaken for roots.

Leaves

In most vascular plants, the main photosynthetic organ is the leaf.

However, green stems may also perform photosynthetic functions.
Leaves vary extensively in their morphology, but generally consist of a
flattened blade and a stake, which is the petiole, and which joins the
leaf to the stem at a node. Grasses and many other monocots lack
petioles. Instead, the base of the leaf forms a sheath that envelops the
stem.

Monocots and eudicots differ in the arrangement of veins, which is the

vascular issue of leaves. Most monocots have parallel major veins that
run the length of the blade. Eudicots generally have a branched
network of major veins.

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In identifying angiosperms according to structure, taxonomists

generally rely on the floral morphology, but they also use variations in
leaf morphology. Many leaves are compound or doubly compound.
This structural adaptation may enable large leaves to withstand strong
winds with less tearing. It may also confine some pathogens that
invade the leaf to a single leaflet, rather than allowing them to spread
to the entire leaf. Almost all leaves are specialized for photosynthesis.
However, some species have leaves that allow them to perform special
functions, such as protection, support, or reproduction.

Dermal, Vascular, and Ground Tissue

Each of the three plant organs have dermal, vascular, and ground
tissues. These three categories of tissues form a tissue system, which is
a functional unit connecting all of the organs of the plant. Although
each tissue system is continuous throughout the plant, specific
characteristics of the tissues and their spatial relationships to one
another vary in different organs.

The dermal tissue system is the outer protective layer of the plant, or
the covering. It forms the first line of defense against pathogens and
physical damage. In plants that are not woody, it is usually a single
tissue called an epidermis, which is a layer of tightly packed cells. In
leaves and most stems, the cuticle, which is a waxy coating on the
epidermal surface, helps prevent the loss of water. In woody plants, the
protective tissue is called a periderm. This replaces the epidermis in
the older regions of the stems and roots. In addition to protecting the
plant from pathogens and water loss, the epidermis has specialized
characteristics in each organ. For instance, a root hair is an extension
of an epidermal cell near the tip of the root. Trichomes which are
hairlike outgrowths of the shoot epidermis, reduce water loss and
reflect excess light. They can also provide defense against insects by
secreting stickly fluids and toxic compounds. For example, trichomes
on aromatic leaves such as mint secret oils that protect plants from
herbivores.

The vascular tissue system carries the long-distance transport of

materials between the shoot and root systems. The two types of
vascular tissue are xylem and phloem. The xylem transports water and
dissolved minerals. The phloem transports sugars (usually from the
leaves) to where they are needed (usually the roots and other sites of
growth). The vascular tissue of a root or shoot system is collectively
called a stele. The arrangement of the stele varies, depending on the
organ and the species. In angiosperms, for examples, the root stele is a
solid vascular cylinder of phloem and xylem, whereas the stele of

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leaves and stems consists of vascular bundles, which are separate

strands composed of either phloem or xylem. Both the xylem and the
phloem consist of differentiated cell types for transport or support.

Tissues that are neither dermal nor vascular are part of the ground
tissue system. Ground tissue that is internal to the vascular tissue is
called pith. Ground tissue that is external to the vascular tissue is
called cortex. The ground tissue system is not just a filler. It includes
cells that are specialized for functions such as storage, photosynthesis,
and transport.

Common Types of Plant Cells

Cells in plants are specialized according to structure and function.

Cellular differentiation may consist of changes in the both the
cytoplasm and its organelles in the cell wall.

Parenchyma cells have primary walls that are relatively thin and
flexible, and most of them lack secondary walls. Parenchyma cells
generally lack a central vacuole when they are maure. These cells are
the least specialized structurally. Parenchyma cells perform most of
the metabolic functions of the plant. They synthesize and store various
organic products. For instance, photosynthesis occurs in the
chloroplasts of parenchyma cells in the leaf. Some parenchyma cells in
the roots and stems have plastids which store starch. The fleshy tissue
of most fruits is composed of parenchyma cells. Most parenchyma
cells retain the ability to divide and differentiate into other types of
plant cells under particular conditions. For instance, during wound
repair, parenchyma cells can differentiate. Thus, it is possible for
scientists to grow an entire plant from a single parenchyma cell.

Collenchyma cells are grouped either into cylinders or strands that

help support the young parts of the plant shoot. These cells have thick
primary walls, thicker than parenchyma cells, although the walls are
unevenly thickened. Young stems and petioles often have collenchyma
cells just below their epidermis. They also lack secondary walls.
Lignin, which is a hardening agent, is often absent from their primary
walls. Therefore, these cells are able to provide support without
restricting the growth of the plant. During maturity, these cells are
living and flexible, allowing them to elongate the leaves and stems that
they support.

Sclerenchyma cells, on the other hand, also function as supporting

elements in the plant. However, they have thick secondary walls that
are strengthened by lignin. They are much more rigid than

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collenchyma cells. Mature sclerenchyma cells cannot elongate, and

they are found in regions of the plant that have stopped growing.
Sclerenchyma cells are also specialized for support and are dead at
maturity. However, they produce secondary walls before the
protoplasts (the living the part of the cell) dies. The remaining walls
serve as skeletons, which can support the plant for hundreds of years.
There are two types of sclerenchyme cells: sclereids and fibers. Both
of these are specialized for support and transport. Sclereids are shorter
than fibers and irregular in shape. They have thick, lignified secondary
walls. Sclereids allow the shells of nuts to be hard. Fibers are usually
arranged into threads. They are long, slender, and tapered.

Water-Conducting Cells of the Xylem

There are two types of water conducting cells: tracheids and vessel
elements. Both of these cell types are tubular and elongated. They are
also dead at maturity. Tracheids are found in the xylem of nearly all
vascular plants. In addition to tracheids, most angiosperms, as well as
a few gymnosperms, have vessel elements. When the living contents
of the plant’s tracheids and vessel elements disintegrate, the thickened
walls of the cells remain behind. These form a living conduit through
which water can flow. The secondary walls of tracheids and vessel
elements are often interrupted by pits, which are thinner regions where
only primary walls are present. Thus, water can migrate laterally
through pits.

Tracheids are long, thin cells with tapered ends. Water moves from
cell to cell mainly through pits, where it does not have to cross thick
secondary walls. The secondary walls of tracheids are hardened
through lignin, and this prevents the collapse of the cell during water
transport.

Vessel elements are generally wider, shorter, and thin-walled. They are
also less tapered that tracheids. They are aligned from end-to-end,
forming long micropipes called vessels. The end walls of the vessel
elements have perforation plates that enable water to flow freely
through the vessels.

Sugar-Conducting Cells of the Phloem

Unliked the water-conducting cells of the xylem, the sugar-conducing

cells of the phloem are alive at functional maturity. In seedless
vascular plants and gymnosperms, sugars and other nutrients are
transported through long, narrow cells called sieve cells. In the phloem
of angiosperms, these nutrients are transported through sieve tubes,

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which consist of chains of cells called sieve-tube elements, or sieve-

tube members.

Although alive, sieve-tube elements lack ribosomes, a nucleus, a

distinct vacuole, and cytoskeletal elements. The reduction in the
contents of the cells allows nutrients to pass through the cell more
easily. The end walls between sieve-tube elements are called sieve
plates, which have pores that facilitate the flow of fluid form cell to
cell along the sieve tube. Alongside each sieve-tube element is
nonconducting cells called a companion cell. This is connected to the
sieve-tube elements by numerous channels, the plasmodesmata. The
nucleus and ribosomes of the companion cell not serve not only the
cell itself, but also the nearby sieve-tube element. In some plants, the
companion cells in leaves also help load sugars into the sieve-tube
elements, and then the sugars are transported to other regions of the
plant.

Plant Growth and Development

In contrast to animals, which only grow during the embryonic and
juvenile periods, plants grow throughout their life, which is termed as
indeterminate growth. At any given time, a plant has parts that are
mature, embryonic, or developing. Except for plants that are in their
dormant periods, most plants grow continuously. Some plants organs
undergo determinate growth, such as most leaves, thorns, and flowers.
That is, they stop growing once they reach a certain size.

Although plants continue to grow throughout their lives, they also die.
As was discussed in previous chapters, plants may be annuals, biennals
or perennials.

Plants are capable of indeterminate growth because they have

perpetually embryonic tissues called meristems. These two main types
of meristems: apical meristems and lateral meristems. Apical
meristems are located at the tips of roots and shoots and in the axillary
buds of shoots. They provide additional cells that allow the plant to
grow in length, in a process known as primary growth. Primary growth
allows roots to extend through soil and shoots to increase their
exposure to light. The herbaceous, or non-woody, plants, primary
growth produces most, if not all, of the plant body. Woody plants,
however, also grow in girth and in parts of stems and roots that no
longer grow in length. This growth in thickness, known as secondary
growth, is caused by the activity of lateral meristems, which are called
the vascular cambium and the cork cambium. These cylindes of
dividing cells extend along the length of stems and roots. The vascular

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cambium adds layers of vascular tissue called secondary xylem (wood)

and secondary phloem. The cork cambium, on the other hand, replaces
the epidermis with a thicker, tougher, epiderm.

The cells within the meristems divide relatively often, generating

additional cells. Some of these new cells remain in the meristem and
produce more cells, while other differentiate and are incorporated into
tissues and organs of growing plants. Cells that remain as sources of
new cells are called initials. The new cells that are displaced from the
meristem are called derivatives, since they continue to divide until the
cells they produce become specialized within developing tissues.

Primary Growth in Roots and Shoots

Primary growth is a growth in length, produced by apical meristems.

The primary plant body is the result of this growth. In herbaceous
plants, it is usually the entire plant. In woody plants, it consists of only
the youngest parts, which are not yet woody. Although apical
meristems lengthen both shoots and roots, there are differences in the
primary growth of these two systems.

Primary Growth of Roots

The tip of the root is covered by a root cap, which protects the delicate
apical meristem as the root pushes through the abrasive soil during
primary growth. The root cap also secretes a polysaccharide slime that
lubricates the soil around the tip of the root. Growth occurs behind the
tip in three zones of cells. These cells are at successive stages of
primary growth. Away from the tip, they are zones of cells division,
elongation, and differentiation.

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There are no sharp boundaries between the three zones, and they grade
together. The zone of cell division includes its derivatives and root
apical meristem. In this region, new root cells are produced. Behind
the tip of the root is the zone of elongation. This is where root cells
elongate. Sometimes, these cells elongate to more than 10 times their
length. In this zone, cell elongation allows the tip to penetrate farther
into the soil. Even before the root cells start lengthening, they may
begin specializing in structure and function. In the zone of
differentiation, which is the zone of maturation, cells complete the
differentiation process and become specific cell types.

The primary growth of a root is when its epidermis is produced, as

well as the ground tissue and vascular tissue. In most roots, the stele is
a vascular cylinder. It is a solid core of xylem and phloem.

In the roots of most eudicots, the xylem has a star-like appearance. The
phloem occurs the indentations between the arms of the star. In
monocots, the central core is composed of parenchyma cells. This core
surrounded by a ring of xylem and then a ring of phloem. This central
region is called a pith, but it is different from a stem pith.

The ground tissue of roots, which consists mostly of parenchyma cells,

fills the cortex. The cortex is the region between the vascular cylinder
and the epidermis. The ground tissue has cells that store carbohydrates.

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Their plasma membranes allow for the absorption of water and

minerals. The cortex has an innermost layer called the endodermis.
This is a cylinder that is one cell thick and forms the boundary with the
vascular bundle.

Lateral roots come from the pericycle, which is the outermost layer in
the vascular cylinder. It is adjacent to, and just inside, the endodermis.
A lateral root pushes through the epidermis and the cortex until it
emerges from the main root. The lateral root cannot come from near
the root’s surface because the vascular system must be continuous with
vascular cylinder at the center of the established root.

Primary Growth of Shoots

A dome-shaped mass of dividing cells at the shoot tip is the shoot

apical meristem. Finger-like projections along the sides of the apical
meristem are where leaves develop, the leaf primordia. Axillary buds
develop from islands of merismatic cells left by the apical meristem at
the bases of the leaf primordia. These buds can form lateral roots at
some later time.

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The internodes are shaped close together within leaf primordia because
the internodes are very short. Most of the elongation of the shoot is
due to the lengthening of the internode cells underneath the shoot tip.
In grasses, and other plants, some of the leaf cells are produced by
areas of merismatic tissue that is separated from the apical meristem.
These areas are called the intercalary meristems, and remain at the
base of leaf blades and stem internodes. This type of morphological
feature helps grasses tolerate grazing because the elevated part of the
leaf blade can be removed without interfering with growth.

The Tissue Organization of Stems

As part of the continuous dermal tissue system, the epidermis covers

the stem. As vascular bundles, vascular tissue runs the length of the
stem. Unlike lateral roots, which arise from the vascular tissue deep
within a root and thus disrupt the vascular cambium, cortex, and
epidermis as they emerge, lateral roots develop from axillary bud
meristems on the surface of the stem. These lateral roots do not disrupt
other tissues. The stem’s vascular bundles converge with the root’s
vascular cylinder in a zone of transition near the surface of the soil.

In most eudicot species, the vascular tissue is composed of vascular

bundles that are arranged in a ring. The xylem contained within each

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vascular bundle is situated beside the pith, and the phloem in each
bundle is situated beside the cortex. In most monoct stems, the
vascular bundles are scattered throughout the ground tissue. They do
no form rings. In the stems of both eudicots and monocots, the ground
tissue consists mainly of parenchymal cells. However, collenchyma
cells underneath the epidermis strengthen many stems. Sclerenchyma
cells, especially fiber cells, also provide support in parts of the stem
that have stopped elongating.

Tissue Elongation of Leaves

The epidermal barrier of leaves is interrupted by stomata, which allow

gas exchange between the surrounding air and the photosynthetic cells
inside the leaf. Stomata also function to regulate the CO2 uptake for
photosynthesis and they are major avenues for the evaporative loss of
water. The stomatal complex is composed of the pore flanked by two
guard cells, and the latter regulate the opening and closing of the pore.

Sandwiched between the upper and lower epidermal layers is the

ground tissue of the leaf, the mesophyll. Mesophyll contain mainly
parenchymal cells that are specialized for photosynthesis. The leaves
of many eudicots have two areas: the spongy mesophyll and the
palisade mesophyll. The palisade mesophyll consists of one or more
layers of elongated parenchyma cells located on the upper part of the
leaf. On the other hand, the spongy mesophyll is located below the
palisade mesophyll. The parenchyma cells of the spongy mesophyll
are loosely arranged and contain a labyrinth of spaces through which
CO2 and oxygen circulate. The air spaces are large near the somata,
where gas exchange with the environment occurs.

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The vascular tissue of each leaf is continuous with the vascular tissue
of the stem. Connections from the vascular bundles in the stem, leaf
traces, pass through petioles and into the leaves. In the vascular bundle
are veins, which branch out. This network brings the xylem and
phloem near the photosynthetic tissue. The photosynthetic tissue
receives water and minerals, and brings the products of photosynthesis
to the phloem. The vascular structure also reinforces the shape of the
leaf. Each vein is enclosed by a bundle sheath, consisting of one or
more layers of cells, which are usually parenchyma cells. Unlike stems
and roots, leaves rarely undergo secondary growth.

Secondary Growth

The secondary plant body consists of the tissue produced by the

vascular cambium and the cork cambium. The former adds secondary
xylem and secondary phloem, increasing the vascular flow and support
for the shoot system. On the other hand, the cork cambium produces a
tough, thick covering that consists mainly of cells with wax that
protect the stem from water loss and from invasion by insects, fungi,
and bacteria. Monocots rarely have secondary growth, unlike eudicots
and gymnosperms. Primary growth and secondary growth occur
simultenously.

The vascular cambium is a cylinder of merismatic cells, and it is often

only one cell thick. It increases in circumference and also adds layers
of secondary xylem to its interior and secondary phloem to its exterior.
Each layer has a larger diameter than the previous layer. In this way,
the vascular cambium thickens stems and roots.

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In woody plants, the vascular cambium consists of a continuous

cylinder of undifferentiated parenchyma cells, which are located
outside the pith, and primary xylem. The inside of the cortex has the
primary phloem. In a woody root, the vascular cambium forms the
exterior of the primary xylem and the interior of the primary phloem
and the pericycle.

As merismatic cells divide, they increase the circumference of the

vascular cambium. They also add secondary xylem to the inside of the
cambium and secondary phloem to its outside. Some initials are
elongated and are oriented with their long axis parallel to the axis of
the stem and root. They produce cells, such as the tracheids, vessel
elements, fibers of the xylem, as well as companion cells. The other
initials are shorter and are oriented perpendicular to the axis of the root
or stem. The produce vascular rays, which are radial files of cells that
connect the secondary xylem with the secondary phloem. These rays
move water and nutrients between the secondary xylem and the
phloem. They also store carbohydrates and aid in wound repair.

As secondary growth continues over many years, layers of secondary

xylem, which is wood, accumulates. This consists mainly of tracheids,
vessel elements, and fibers. Gymnosperms only have tracheids,
whereas angiosperms have both tracheids and vessel elements. The
walls of secondary xylem have lignin, which makes them hard. As the
tree grows, the older layers of secondary xylem no longer transports
minerals and water (which is a solution called xylem sap). These
layers are called heartwood because they are closer to the core of the
stem or root. The newest, and outermost layers of the secondary xylem
still transport xylem sap, and are therefore known as sapwood. Since
each new layer of secondary xylem is larger in circumference than the
previous one, growth enables the xylem to transport more xylem sap
each year, which supports an increasing number of leaves. Only the
youngest secondary phloem, which is closest to the vascular cambium,
functions in sugar transport. The older secondary phloem is sloughed
off, which is why secondary phloem does not accumulate as
extensively as secondary xylem.

The Cork Cambium and the Production of Periderm

During the first few stages of secondary growth, the epidermis is

pushed outward, causing it to dry, split, and fall of the root or stem. It
is replaced by two tissues produced by the first cork cambium, which
is a layer of dividing cells that arises in the outer cortex of the stems
and in the outer layer of the pericycle in roots. One of these tissues,
which is called the phelloderm, is a thin layer of parenchymal cells

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that forms to the interior of the cork cambium. The other group of cells
form to the exterior of the cork cambium. As these cells mature, they
deposit suberin, which is a waxy material. The cork tissue functions as
a barrier that prevents the stem or root from water loss, pathogens, and
physical damage. Each cork cambium and the tissue it produces
comprises a layer of the periderm.

Most of the periderm is impermeable to water due to the presence of

suberin, unlike the epidermis. In most plants, therefore, water and
minerals are absorbed mostly in the younger parts of the plant. The
older parts of the roots anchor the plant and transport water and solutes
between the soil and the shoots. Located on the periderm are small,
raised areas called lenticels. It enables plants with woody stems to
exchange gases with the environment. Lenticels often appear as slits.

The thickening of a stem or root often splits the first cork cambium.
The first cork cambium often differentiates into cork cells and loses its
merismatic activity. A new cork cambium forms on the inside,
resulting in another layer of periderm. The bark contains all the tissues
that are external to the vascular cambium. Its components are, from the
inside, the secondary phloem, the most recent periderm, and all the
older layers of periderm.

Glossary
Dicots: dicotyledons, where embryonic seeds have two cotyledons.

Monocots: monocotyledons, or flowering plants (angiosperms) which contain

only one cotyledon.

Videos and Resources

Plant Parts and Functions

Plant Nutrition and Transport

Plant Tissues

Plant Anatomy

Practical Plant Anatomy

Water Transport in Plants: Anatomy and Physiology

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References
Reece, J. B., Urry, L. A., Cain, M. L., Wasserman, S. A., Minorsky, P.
V., & Jackson, R. B. (2011). Campbell biology (p. 379). Boston:
Pearson.

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