Topic X Plant

Physiology:
Transport and
Photosynthesis

LEARNING OUTCOMES
By the end of this topic, you should be able to:

1.

Describe the form of various components (parts and tissue) of a

plant;

2.

Explain how the individual components function and how their

function is related to one another;

3.

Explain the fundamental processes of plant physiology, which

include transport and photosynthesis;

4.

Discuss how plant physiology has developed to enhance plant

survival in specific environments; and

5.

Connect all the above information into a complete picture of plant

form and function.

INTRODUCTION

Plants are multicellular autotrophs that have come to cover most of the earths
surface and play a vital role not only in maintaining life but in shaping most of
the physical structure our environments. There are over 200,000 species of
flowering plants, ranging in size from minute floating forms to giant trees.
Though plants may not look particularly similar at first sight, they share a basic
unit of structure that can be observed when the plants grow, reproduce and
manufacture their food. This topic will briefly examine the physiology of
vascular plants by discussing their form and relate this to the function of the
various tissues present in a typical vascular plant today. It will also briefly

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explain the mechanisms of transport in plants and the fundamentals of the

process of photosynthesis.

9.1

PLANT FORM AND STRUCTURE

This section attempts to explain how a plant is built by describing the different
types of cells, tissues and organs that make up an adult plant body. The plant
body can basically be considered as a series of repeating parts built up above and
below the ground. This basic division of the plant body into an above-ground
and below-ground component gives us the first division of a plant body into the
shoot system and the root system. The shoot system generally consists of a stem
that bears leaves, flowers and fruit that contain seeds. The shoot system serves to
hold the leaves (organs of photosynthesis) in a position that exposes them to
light. The root system forms a network beneath the ground surface. This anchors
the plant in the ground and absorbs water and nutrients from the ground.
The above ground shoot of plants can vary in structure, with plants being either
herbaceous or woody. Herbaceous plants have soft non-woody shoots while
woody plants produce a hard secondary tissue reinforced with lignin that we call
wood. Plants can further be divided according to life-span, with three groupings,
namely the annuals, the biennials and the perennials. Annuals are plants that
complete their life cycle (grow, reproduce and die) within a period of less than a
year. They are often highly seasonal, appearing in one season and then
disappearing until that season is repeated. Biennial plants take about two years
to complete their life cycle, while perennial plants have the potential to live for
more than two years. Woody plants are generally all perennials, with some living
for hundreds or thousands of years.

9.1.1

Plant Tissue

Plants have three basic types of tissue each of which extends throughout the
entire plant body. These three tissue types are:
1.

Dermal Tissue
This tissue forms the external cover or skin of the plant and protects the
plant from damage and from excessive water loss.

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2.

Ground Tissue
Most of the plant body is made up of ground tissue. This tissue has a
variety of functions including storage, photosynthesis and support of the
plant.

3.

Vascular Tissue
This is the conducting tissue that is responsible for transport within the
plant body and also helps in strengthening the plant body. Vascular tissue
is made up of phloem and xylem. Xylem conducts water and dissolved
minerals from the roots to sites of photosynthesis. Phloem conducts
carbohydrates and other plant metabolites (like hormones and amino acids)
from sites of manufacture to sites for utilisation or storage. Xylem is made
up primarily of dead cells, while phloem is made up primarily of living
cells (see Figure 9.1).

Plant organs like leaves are made up of all three tissue types together. Generally,
plant tissue is all interconnected. Tissue from the roots extends into the stem.
Each of the tissue types is made up of different tissue each of which is in turn
made up of different cell types specialised to a particular function (see Table 9.1).
Table 9.1: The Three Types of Tissue in Plants, with Examples of Cell Types
and Some Functions
Tissue Type
Ground
Tissue

Vascular
tissue

Tissue
Parenchyma

Cell Types
Parenchyma cells

Collenchyma
Sclerenchyma

Collenchyma cells
Sclerenchyma cells (or
fibres)
Tracheids

Xylem

Vessel elements

Phloem

Dermal
tissue

Epidermis

Periderm

Fibres
Xylem parenchyma
Sieve tube elements
Companion cells
Fibres
Phloem parenchyma
Epidermal cells
Guard cells
Trichomes
Cork cells
Cork cambium cells
Cork parenchyma

Main Functions
Storage, secretion,
photosynthesis
Support
Support, strength
Conduction of water, nutrients,
support
Conduction of water, nutrients,
support
Support, strength
Storage
Conduction of sugars, support
Control of sieve tube elements
Support, strength
Storage, sugar movement
Protective cover
Regulate stomata opening
Variety of functions
Protective covering
Meristematic
Storage

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Structure of the Root

The root system of a plant is normally located beneath the ground (some plants
like epiphytic orchids and bromeliads have their roots exposed to the air but the
structure and function of their roots remain the same. It functions to absorb water
and nutrients from the environment and to anchor the plant to its substrate (see
Figure 9.2). Like the rest of the plant body, the root is made up of layers of tissue
and a few specialised parts. These can be summarised as follows:

Epidermis: This is the external covering of the plant root. It absorbs water
and nutrients from the soil and protects the root from damage and from
penetration by pathogens and predators.

Root hair: This is thin microscopic extension of a root epidermal cell. It serves
to increase root surface area so as to increase the amount of water and
nutrients absorbed.

Root cap: This is a hardened mass of tissue that protects the growing apical
part of the root and serves to push the root through the soil without
damaging the growing root cells.

Cortex: This is the space filling tissue between the root epidermis and the
vascular tissue. It gives the root bulk and acts to conduct water and minerals
from the epidermis to the vascular tissue.

Epidermis: this is a thin layer of cells that surrounds the vascular tissue and
separates the vascular tissue from the cortex. It serves to control flow of
materials through the cortex.

Vascular tissue: consisting of phloem and xylem. This is the conductive tissue
and is arranged in a specific pattern (see Figure 9.1 (a)), with the xylem
forming an X-shape and the phloem filling the spaces between the arms of
the X.

Figure 9.1 (a): Cross section through a typical plant root showing the arranged
of the vascular tissue. Note the X-shape of the xylem tissue

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Buttress tissue: is found in many of the trees of tropical rain forests to support
the trees under windy conditions. The tissues are formed from the lower part
of the tree trunk, extend out horizontally on or just beneath the floor of the
soil. There are shallow root systems of the buttress trees which are more
adaptable in the rainforest soils. See Figure 9.1 (b).

Figure 9.1 (b): Shallow root system of buttress tissue

Source: C & D Frith Australian Tropical Rainforest Life

Root form depends on the type of plant. Roots can be fibrous, where a network of
roots originates from a central point and radiates randomly in every direction
with no central or main root (see Figure 9.2). This is like the roots found on
grasses. The root system can also consist of a taproot, where there is a single
main root that grows vertically into the ground and from this main root,
branches radiate away into the soil around the root. Roots can be modified to
serve a secondary function. In addition, roots can grow outside the root zone.
These are called adventitious roots (an example is the roots you see hanging of
the branches and stems of figs (banyan) trees. Other types of roots include:

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Figure 9.2: Examples of root structure in plants. Note the fibrous roots of
monocots, the tap roots of dicots, and an example of a storage root

Prop roots Produced on the lower part of the stem such as those seen on

Pandanus sp., help to support the plant.

Aerial roots Often found on epiphytic plants like orchids.

Pneumatophores Found in plants in swampy flooded areas (such as

mangrove), these roots extend above the water surface to facilitate oxygen
uptake for the submerged root.

Food storage roots Common in our familiar root crops, these roots xylem
tissue is modified to store large amounts of carbohydrates.

Water storage roots Common in plants that grow in arid areas.

Buttress roots Common in tropical rain forest trees, these roots increase the
base area of the tree to improve stability.

9.1.3

Structure of the Stem

The stem is the emergent part of the plant. It functions to hold the leaves in a
position that optimises their exposure to the sun. Leaves are attached either
directly to the stem, or onto branches which are technically just extensions of the
stem. The area where the leaf is attached to the stem is called the node, while the
area between two nodes is called the internode. Internally, the stem is made up of

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the same tissue that makes up the root arranged in overlapping layers. On the
outside is the epidermis. Beneath the epidermis is the cortex layer. Embedded
within the cortex layer are the vascular bundles consisting of phloem and xylem.
In the middle of the stem is the pith. Stems show primary and secondary growth.
Primary growth occurs throughout the plant and the growth and differentiation
of tissue that is responsible for establishing the body form of the plant. Secondary
growth occurs in some plants and serves to increase the girth (width) of the stem
by adding tissue continuously. This is what is responsible for the massive stems
of plants like trees. Just like roots, stems can be modified into different forms.
Examples include bulbs and tubers (swollen underground stems like in onions,
potatoes), runners (stems that extend along the ground and produce new plants),
and tendrils (modified stems that twist and aid in climbing).

9.1.4

Structure of the Leaves

Leaves are the primary sites of photosynthesis in land plants. They also play a
vital role in plant gaseous exchange and respiration. Leaves are simply an
extension of shoot tissue but are determinate structures, meaning they stop
growing once they reach maturity. External leaf structure is composed of a
flattened leaf blade and a slender stalk called the petiole. Throughout the leaf,
vascular tissue is distributed in veins that can be seen on the leaf surface. Leaf
blades come in variety of forms, from simple leaves that have undivided blades
(even though they may have indentations or lobes), to compound leaves in which
the leaf blade is divided into leaflets. Leaves can be arranged alternately on a
stem, or they may be opposite in arrangement. The external shape and structure
of the leaf is a compromise between exposing a maximum area to sunlight and
minimising water loss and overlapping (see Figure 9.3).
Internally, the leaf consists of different tissue similarly layered to the rest of the
plant. On the outside of the leaf is epidermis. This is made up of the upper
epidermis on the leaf surface and exposed to the sun, and the lower epidermis on
the bottom of the leaf. The epidermis is covered by a waxy cuticle to reduce
water loss. Within the lower epidermis are numerous openings called stomata.
These are composed of specialised cells called guard cells and are vital in
regulating gas exchange and water loss in the plant. Between the epidermis
layers is the mesophyll tissue. Most plants have two types of mesophyll. Beneath
the upper epidermis lies the palisade mesophyll, which is made up of tightly
packed cylindrical cells rich in chloroplasts that are the primary sites of
photosynthesis. Beneath this layer is the spongy mesophyll which is made up of
randomly packed cells surrounded by large air spaces connected to the stomata.

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(a)

(b)

Figure 9.3: Differences between monocot (a) and dicot (b) leaves.
Note the parallel veins on the monocot leaf and the network veination and the dicot leaf

Packed into the mesophyll are veins made up of vascular tissue. Plant leaves
show a variety of adaptations and can be modified into spines, reproductive
leaves and even carnivorous leaves (see Figure 9.4).

Figure 9.4: Cross section of a plant leaf. Note the photosynthetically vital mesophyll
tissue, the position of the vascular bundles and the presence of a stomata

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Monocots and Eudicots

Though all plants share a common structure, they are divided into two groups
based on their structure and evolutionary history. These groups are called
monocots and eudicots (also known as dicots) (see Figure 9.5). Generally,
monocots are mostly herbaceous plants with long narrow leaves with parallel
veins. Monocot flowers are usually arranged in multiples of three and monocot
seeds have single cotyledon (embryonic leaf). Examples include all grasses like
rice, as well as orchids, lilies and palm trees. Dicots, on the other hand, may be
herbaceous or woody, and have leaves that are variable in shape and often quite
complex. Their flowers occur in fours or fives or multiples of these and they have
two cotyledons in their seed. Examples include all trees, beans, sunflowers and
shrubs. Monocots are considered the more advanced plants. Table 9.2
summarises the main differences between these two groups. Figure 9.6 shows the
distribution of vascular bundles between monocot and dicot stems.
Table 9.2: Differences between Monocots and Dicots
Characteristic
Roots
Leaves
Flowers
Seeds
Secondary growth (wood and
bark)
Vascular bundles in stem
Pollen grains

Monocots
Fibrous
Parallel venation
Usually in threes
Embryo has one cotyledon
Absent

Dicots
Taproot
Netted venation
Usually in fours or fives
Embryo has two cotyledons
Usually present

Usually scattered or in complex

arrangements
One pore

Arranged in a circle or ring

Three pores

Figure 9.5: Comparison between monocot and dicot seeds showing

the single cotyledon of the monocot and the double in the dicot

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Figure 9.6: Differences in the distribution of vascular bundles between monocot

(A) and dicot (B) stems (refer to table 9.2). Note the detailed description of vascular
bundle structure

9.2

ESSENTIAL ELEMENTS FOR PLANTS

Sixteen minerals were designated as essential chemical elements for plant

growth. These elements present in plants were necessary for normal growth. By
using hydroponic system, Arnon & Stout (1939) identified three criteria to
establish the essentiality of a chemical element.
1.

In the absence of any one of these essential elements, plants displayed

characteristic abnormalities of growth or deficiency symptoms, and often
such plants did not reproduce normally (for example, a plant cannot
complete its growth cycle under a severe deficiency of nitrogen).

2.

An essential element cannot be replaced by other elements (for example,

potassium ion (K+) cannot be replaced by sodium ion (Na+) although they
have similarities in structure and valency).

3.

Essential elements must be involved directly in plant metabolism. For

example, cobalt is essential for rhizobia - bacteria that form symbiotic union
in nodules of legumes; and it benefits the host plant only indirectly. On this
basis, cobalt is not considered an essential element.

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On the basis of the normal concentrations in plants, the essential elements can be
divided into two groups, macronutrients and micronutrients. Tables 9.3
summarises the roles of macronutrients and micronutrients.
Table 9.3: The Roles of Macro and Micro Nutrients
Macro
nutrients

Elements
Nitrogen

Potassium

Phosphorus

Sulphur

Micro
nutrients

Iron

Zinc

Molybdenum

Role of the elements

Components of amino acids,
proteins, nucleotides, nucleic
acids,
chlorophyll
and
coenzymes
Exists as K+, involved in
osmotic and ionic balance,
opening
and
closing
of
stomata, as activator for many
enzymes
Component of ATP and ADP,
nucleic
acids,
several
essential
coenzymes,
phospholipids, essential in
plant metabolism and energy
transfer
Component of some amino
acids

cysteine
and
methionine, coenzyme-A and
certain nucleotides.
Involves in oxido-reductase
reactions;
required
for
chlorophyll
synthesis;
component of cytochromes
and nitrogenase; essential
component of ferrodoxin; and
the site of NAD and NADP
reductions
Activator of many enzymes
especially those involve in
oxido-reductase reactions
Nitrogen fixation; component
of
the
active
site
of
nitrogenase

Deficiency Symptoms
Occurs in mature tissues;
growth retardation; leaves turn
to yellowish brown.
Occur in young tissue; drying
of the tips of root and leaf;
twisted
leaf
morphology;
retardation of root growth;
decrease of plant growth rate.
Old leaves become dark green
colour,
appearance
of
anthocyanin
(dark
purple
pigment); delayed maturity

Rarely happens

Symptom first occur in young

tissues.

Spots on necrosis.

Chlorosis and retardation of

plant growth.

In general, deficiency of certain element usually results in growth

inhibition/retardation because it almost always causes chlorosis and necrosis
which reduces the effective area for photosynthesis. The response of growth rate
to fertiliser is positive within the insufficient zone but not within the sufficient
zone; whilst the response is negative within the toxic zone. The second and
the last tasks, obviously, are waste of resource.

SELF-CHECK 9.1
What does N:P:K of 1:2:1 mean? Discuss your findings in your
online forum.

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9.3

TRANSPORT IN PLANTS

9.3.1

PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

Soil-Plant-Atmosphere Continuum (SPAC)

Roots absorb the ground water into the plant xylem which travels upwards
against gravity and is lost to the atmosphere by a process called transpiration.
Thus, there is a continuous column of water from the ground through a plant to
the atmosphere (SPAC). This column needs specific mechanisms to be sustained.
It is now known that these mechanisms include root pressure, cohesive and
adhesive properties of water, and transpiration pull. This last terminology seems
empty unless one fills it with an energy term, in this case water potential.

9.3.2

Concepts of Water Potential

Water potential is defined as chemical potential of a solution compared to

chemical potential of pure water at a given temperature and atmospheric
pressure. When a water potential gradient is established between two areas,
water will spontaneously diffuse from the high end to the low end. This water
potential gradient can easily explain SPAC or transpiration pull.
Water potential can be measured by using Scholander Bomb and using a
psychrometer. It has two major components: osmotic and turgor pressure, related
in the following manner:
Water Potential = Osmotic Potential + Turgor pressure

Turgor Pressure
The presence of water in a closed system such as a cell creates a pressure in an
outward direction. Turgor pressure is defined as the pressure that develops in a
plant cell as a result of water molecules hitting against the plasma membrane.
Turgor to a plant is especially important in the support of non-woody plant
parts. Turgor is maintained constant (and sufficiently high) in most plant cells
because they generally exist in an environment where water loss is balanced by
water uptake. When an environment does not allow this, both anatomical and
physiological adjustments occur to strike a balance.

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9.4
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PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

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WATER UPTAKE
Root Pressure

Root pressure is the capacity of the roots (devoid of all leaves) of a plant to
absorb water. Plants absorb water from soil during the daytime when stomata
are open and water loss by transcription is inevitable. However, transpiration is
not a total loss. It confers advantages to plants, directly and indirectly, in various
ways including the following:
(i)

as a result of transpiration and subsequent water uptake into the plant,

water, together with all dissolved solutes are siphoned towards the root
into the plant from adjacent areas. There is, in other words, a sphere of
water movement whose magnitude depends on the rate of transpiration.
This movement constantly replenishes nutrients in the sphere which have
been taken up by the plant.

(ii)

evaporation of water from leaf surfaces cools down the leaf temperature
and maintains this important organ of a plant at optimal temperature. In
addition to this, a population of cooled leaves can have profound effects on
the atmospheric temperature, very much like cooling fins cool down the air
that flow through them in an air-conditioner.

Apoplast and Symplast Concept

In this concept, the root of a plant is regarded as two components, the apoplast
and symplast, which are separated by plasma membrane. The apoplast
comprises of intercellular spaces, the cell wall and xylem while symplast is the
cytoplasm of all root cells. The cytoplasm of all root cells is considered as one
because of the presence of astronomically numerous plasmodesmata (interconnecting bridges) that enable rapid mass transport within the symplast.
Apoplast is considered as a free space. In this respect, water in the soil (along
with all the dissolved minerals and other solutes) is free to move in and out of
the apoplast. Soil water thus surrounds symplast (up to the endodermis). The
endodermis, with the water-impermeable casparian strip, limits movement of
soil water to within the outer apoplast only.
The Mechanism of Root Pressure
With soil water (and dissolved minerals) surrounding it, there is no difficulty to
imagine how some of the selected minerals in ionic form, are actively transported
into the symplast. Movements of these ions occurs by active transport all the way
to xylem. Xylem at this stage act like a dumping ground accepting all the
(selected) ions from symplast with the obvious result of increased osmotic

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potential, facilitating passive water entry. Figure 9.7 shows the mechanism of
root pressure.

Figure 9.7: A flow chart to describe the mechanism of root pressure

9.4.2

Adhesive and Cohesive Theory

Adhesion is the attractive force between water molecules and other substances.
Both water and cellulose are polar molecules, so there is a strong attraction for
water within the hollow capillaries of the xylem. Cohesion is the attractive force
between molecules of the same substance. Water has an unusually high cohesive
force due to the four hydrogen bonds each water molecule potentially has with
any other molecule. It is estimated that the water s cohesive force within xylem
give it a tensile strength equivalent to that of a steel wire of similar diameter.

9.4.3

Transpiration Pull

The evaporation of water from the walls of mesophyll cells of leaves results in a
decrease in the diffusion pressure of water in these walls. This disturbance in the
water balance of the cells causes a series of changes that set in motion the entire
train of water through the plant. The living leaf cells next to the tracheids in
veinlets eventually lose water to neighbouring cells. Water moves from the
veinlets into the adjacent cells. This results in the upward movement of water in
the continuous liquid columns in the xylem. Continuity of water is maintained
even under conditions of considerable strain because of the strong cohesive
forces between water molecules.

9.4.4

Transpiration: Stomatal and Epidermal

While most of the water taken up by the plant is lost through the stomata, a small
fraction is lost through the epidermis, presumably through the apoplastic
water movement, through the cell wall. Loss through the epidermis is minimised
in most plant species by the presence of waxy cuticular layer on the epidermal
surface.

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EXERCISE 9.1

9.5

1.

Apoplast is a free space. Explain.

2.

Is phloem a part of apoplast? Explain.

PLANT ADAPTATIONS TO THEIR

ENVIRONMENTS

Because plants are non-motile (they cannot move), they cannot actively select the
best conditions to be in at any time, and they cannot move in search of water or
minerals (food). As such, they develop specific adaptations to maximise their
suitability to their growing environments. Plant adaptations are best considered
in terms of limiting factors that plants face that reduce their growth from
optimum. The two most common limiting factors for plants are sunlight and
water.
Plants growing in the hot and humid environment of a tropical rain forest
generally have sufficient water and minerals but because of the crowding and
large numbers of competitors, sunlight becomes a limiting factor. To enhance
their ability to reach more sunlight, many plants in tropical rain forest strive to
grow upwards. Trees can easily reach optimum levels of sunlight by growing
upwards through the canopy. Smaller plants, on the other hand, reach for the sun
by either climbing onto the stems of trees (like lianas do) or by growing on the
top branches of trees as epiphytes (as many species of orchids do).
Plants growing in water like lilies and lotus or in water logged soils like rice have
to be adapted to the absence of oxygen and the high concentration of methane
gases associated with anaerobic decaying matter on the bottom of ponds and
lakes. They have specialised tissue that help cope with these problems (such as
the development of aerenchyma, a loosely spaced tissue with large air spaces).
Plants growing in arid areas are commonly known as xerophytes (and are said to
be growing in xeric conditions), and they probably face the greatest challenge of
all. Water is absolutely vital for plant survival and its absence in arid deserts is
the main factor limiting the amount of vegetation that can grow. Most plants
have to balance water loss with gaseous exchange by regulating the opening and
closing of the stomata according to temperature and relative humidity, but plants
growing in arid areas have to develop other unique and interesting adaptations
to survive their harsh environment. Some of these include:

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Conversion of leaves into spines to reduce water loss from the stomata. In the
absence of green leaves, the stem turns green and takes over as the
photosynthetic tissue.

Large succulent leaves of stems that act as water storage organs.

Hidden and sunken stomata. Many plants in arid areas have their stomata in
sunken groves where dry air does not penetrate. This greatly reduces water
loss without getting rid of leaves.

Rolling of leaves. Many plants, especially grasses, can roll their leaves when
water loss is excessive. This hides the stomata and reduces the surface area
across which water loss can occur.

Thickened waxy cuticle on leaves. This further reduces water loss by

evaporation.

Evolution of the CAM photosynthetic pathway. This is a modified

photosynthetic pathway that reduces stomata opening times and thereby
reduces water loss.

Modified root systems. These include very deep roots that grow into the
ground until they reach the water table (as in Acacia trees); very widespread
superficial roots that grow in a thick network just under the soil surface to
catch as much water from rare rainfall as possible (as in many cacti); or
modified roots that act as water storage organs (as in many Euphorbia).

9.6

PHOTOSYNTHESIS

Photosynthesis is a fundamental biochemical process that occurs in green plants.

The importance of photosynthesis cannot be overstated. Most metazoan life
(animal life) is dependent on photosynthesis. The basis of most food webs on
earth is the energy trapped by plants through photosynthesis, and the oxygen we
breathe is a by product of the photosynthetic process. Photosynthesis occurs in
many kinds of bacteria and algae as well as green plants, and these organisms are
termed photoautotrophs (make their own food using light). In green plants, all
photosynthetic reactions occur with organelles called chloroplasts.
Photosynthesis takes place in three stages:
1.

Trapping energy from sunlight

2.

Using this energy to make ATP and a reducing compound known as

NADPH; and

3.

Using the energy from ATP and NADPH to manufacture organic molecules
from CO2 from the air

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The first two stages require light to proceed and are commonly called the lightdependent reactions of photosynthesis. The third stage does not require light to
proceed and can occur in the absence of light and is often called the lightindependent stage of photosynthesis. The entire process of photosynthesis occurs
within a specialised organelle found in plant cells called the chloroplast. Thus
photosynthesis can be considered a process in which cells use light energy
trapped by chloroplasts to power the synthesis of carbohydrates, and can be
summarised in the following equation:
6CO2
+
(carbon dioxide)

12H2O
(water)

C6H12O6
(glucose)

6H2O +
(water)

6O2
(oxygen)

Oxygen is vital by-product of photosynthesis and scientists believe that virtually

all the oxygen in the atmosphere has been generated by plant photosynthesis. As
such, we owe our existence and evolution, as aerobic organisms, directly to the
presence on earth of photosynthetic autotrophs.

9.6.1

The Photosynthetic Pigments

The light energy used in photosynthesis is trapped in the chloroplasts by

molecules called photosynthetic pigments. A pigment is a molecule that absorbs
light. Different pigments absorb light at different wavelengths. There are several
types of pigments used by a variety of organisms, though green plants use two
main pigments called chlorophylls and carotenoids. Chlorophylls occur in two
forms, chlorophyll a and chlorophyll b, and preferentially absorb violet-blue and
red light and reflect green light. This is the reason we perceive plants as being
green in colour. Carotenoids preferentially absorb blue light and reflect red,
orange and yellow light. This is why many plants rich in carotenoids (such as
carrots) appear to be orange in colour to our eyes. The efficiency and range of a
pigments absorption of visible light is termed its absorption spectrum.
Pigments are divided into two groups according to their function in
photosynthesis. These groups are primary pigments that play a direct role in
photosynthesis, and accessory pigments that transfer trapped light energy to
primary pigments. Chlorophyll is a primary pigment while carotenoids are
accessory pigments. Pigments are grouped together in the chloroplast into
photosystems. A photosystem is a network of chlorophyll molecules, accessory
pigments, and binding proteins forming a reaction centre held together on the
chloroplast membrane.

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9.6.2

The Chloroplast

PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

When plant cells are examined under a microscope, it is clear that the green
pigment is not uniformly distributed across the entire cell. Rather it is
concentrated in specialised organelles called chloroplasts (see Figure 9.8). The
chloroplast is a membrane bound organelle that is found in photosynthetic
eukaryotes. It has an inner and outer membrane. The inner membrane encloses a
fluid-filled space called the stroma rich in enzymes responsible for carbohydrate
synthesis. The light-independent carbon-fixing reactions occur in the stroma.
Suspended in the stroma is another membrane system called thylakoids. In
several places, the thylakoids membrane is stacked into a structure called a
granum (plural grana). Chlorophyll is found in the thylakoid membrane, and the
light-dependent reactions of photosynthesis occur in the thylakoids.

Figure 9.8: Structure of a chloroplast

9.6.3

The Process of Photosynthesis

As mentioned earlier, the process of photosynthesis is divided into the lightdependent and the light-independent stages.
(a)

The Light-Dependent Reactions of Photosynthesis

The light-dependent reactions of photosynthesis (also called the light
reactions) convert light energy into chemical energy. It involves a process
called photolysis where light energy is used to split water, freeing electrons,
removing a H+ ion and generating oxygen as a by-product. The reactions
proceed through two photosystems called Photosystem I and Photosystem
II. Electrons flow from water (freed by light energy) split at Photosystem II
to Photosystem I, and as such the process is termed electron transport
chain. These electrons are used by Photosystem II to synthesise ATP in a

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process called photophosphorylation. They are then passed to Photosystem

I where they are used to drive the synthesis of NADPH (see Figure 9.9).

Figure 9.9: The light reaction of photosynthesis. Note the two photosystems,
the splitting of water (photolysis) to yield electrons and the production of ATP

There are two forms of electron transport chains. One is called non-cyclic
electron transport and the other is called cyclic electron transport. Both
processes are used to manufacture ATP, but there are a few small
differences between the two. Non-cyclic transfer is the process described
above. Electrons are supplied to the process by the splitting of water by
Photosystem II when it is activated by light energy (with oxygen as a byproduct). These electrons pass along the electron transport chain until they
reach Photosystem I, where they are re-energised by absorption of
additional light by Photosystem I, where they are passed onto NADP+ to
form NADPH. In cyclic electron transport, only one photosystem,
Photosystem I, is involved. In addition, there is no splitting of water and
the electrons are not accepted by NADP+. Rather, they circulate through the
system, constantly returning to Photosystem I which uses their energy to
manufacture ATP.
(b)

The Light-Independent Reactions of Photosynthesis

In the light independent reactions of photosynthesis (also called the dark
reactions), cells use the energy and reducing power generated by the light
reactions to make organic molecules through a series of complex enzyme
driven reactions in the stroma of the chloroplast. This is achieved through a

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PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

process called the Calvin Cycle (the dark reactions are often referred to
simply as the Calvin Cycle). There are three main phases of the Calvin
Cycle (see Figure 9.10):

Figure 9.10: The dark reactions (Calvin Cycle) of PS. Note the entry of CO2
the release of 1G3P for sugar synthesis and the return of 5G3P into the cycle

1.

The CO2 uptake phase: The cycle begins with a molecule of CO2 reacting
with a five-carbon compound called RuBP (ribulose biphosphate) to
produce an unstable six-carbon molecule that breaks down into two
molecules of a three-carbon compound called PGA (phosphoglycerate).

2.

The carbon reduction phase: In this phase, the energy from ATP and
hydrogens from NADPH (both produced in the light reactions) are used to
convert the three-carbon PGA to a another three-carbon compound called
G3P (glyceraldehydes-3-phosphate). The G3P can leave the Calvin cycle
and be used to manufacture glucose (recall that glucose is a six-carbon
compound, and the three-carbon G3P is an intermediate of glucose
formation). Two molecules of G3P can react to form either glucose or
fructose. In many plants, glucose and fructose are then joined to produce
sucrose (sugar, like the sugar we get from sugar cane). Glucose can also be
used to manufacture starch or cellulose.

3.

The RuBP regeneration phase: In the Calvin Cycle, RuBP (the five-carbon
molecule that starts the cycle) is constantly regenerated to continue the
cycle. For every six CO2 molecules that enter the cycle, six carbon atoms
leave the cycle as two molecules of G3P. All the other carbon molecules
remain within the cycle and are used to regenerate RuBP. This can best be
understood as follows:

The cycle begins with 6 CO2 (6 1C = 6C) combining with 6 RuBP (6

5C = 30C) to give 12 PGA (36C).

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12PGA (36C) break down into 12 G3P (36C).

Of these, 2 G3P (6C) leave the cycle for carbohydrate formation, while
10 G3P (30C) are used to regenerate 5 RuBP (30C).

The Calvin Cycle is also known as C3 photosynthesis because the initial carbon
compound formed at the CO2 uptake phase is a three-carbon molecule. Some
plants also show C4 photosynthesis which will be discussed briefly later.

9.6.4

Factors Affecting the Rate of Photosynthesis

Photosynthesis is a biochemical pathway that uses light, CO2 and water in a

complex series of reactions to produce carbohydrates and energy in plants. Being
a complex series of reactions, the process of photosynthesis is highly dependent
on enzymatic control, and there are several enzymes involved which you do not
have to learn at this stage. One would expect if any of the vital components
involved in the reactions was missing, photosynthesis would not proceed at an
optimal rate.
Water in general is always available to photosynthesising cells as it is actively
provided by the plant. There are also several micro-nutrients that are important
(such as phosphorus and iron) but these are needed only in small quantities and
are generally not limiting under normal circumstances. It has been shown,
though, that there are three vital factors that affect the rate of photosynthesis.
These are light intensity, temperature and the availability of CO2 (see Figure
9.11).
1.

Light intensity: Light intensity is an indicator of the amount of energy

available for the light dependent reactions of photosynthesis. The more
light is available, the faster the light reactions can proceed. When light is
scarce, the light reactions slow down or are unable to proceed and
photosynthesis can shut down. Generally, as long as the temperature
remains high and CO2 is not limiting, the rate of photosynthesis increases
with the increase in light intensity. In nature, light can be limiting due to
strong shade, cloudy conditions, or length of daylight (in temperate and
polar regions). Often, in intensive greenhouse farming, light intensity is
increased by fixing powerful lights in the greenhouse.

2.

Temperature: Because photosynthesis is dependent on enzymatic control,

changes in temperature have a direct effect on the rate of photosynthesis
through the effect of temperature on enzyme function. Generally, as long as
other factors are not limiting, the rate of photosynthesis increases with the
increase in temperature until an optimum temperature is reached.
Thereafter, enzymes begin to denature and the rate slows down until it

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PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

completely stops at excessively hot temperatures. In nature, seasonal and

daily temperature changes in polar or high altitude zones have a direct
effect on the rate of photosynthesis in temperate and polar regions.

Figure 9.11: Changes in the rate of photosynthesis with increases in limiting factors
(light, temperature and carbon dioxide). Note point (A) where graph reaches a
constant level

3.

CO2 availability: The availability of CO2 is a natural limiting factor for all
plants because the amount of CO2 in the atmosphere is fixed (about 0.03%)
and is constant virtually everywhere in the terrestrial environment. CO2 is
vital for the dark reactions of photosynthesis where carbon fixation occurs.
Generally, the increase in CO2 availability results in an increased rate of
photosynthesis as long as other factors are not limiting. As such, for plants
growing in hot tropical areas like rain forests where sunlight and
temperature are not limiting, the rate of photosynthesis operates at a suboptimal rate that is limited by the scarcity of CO2 in the atmosphere. Often
in commercial greenhouse agriculture, plant productivity is increased by
increasing the concentration of CO2.

It is important to understand the importance of the above three factors for plants
in general. Outside the tropics, where seasonal changes in sunlight intensity and
length of daylight are very variable, and where temperatures range from warm
to freezing, the plant growth and biomass productivity is greatly limited by the
first two factors. In the tropics, where plants have ample sunlight and high
temperatures, the rate of productivity is limited by the CO2 concentration
ceiling. This is why many scientists speculate that with increased global

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temperatures from global warming, and increased CO2 concentrations from our
pollution, plant productivity might increase in the near future.

9.6.5

C4 and CAM Photosynthesis

In warm, sunlight-rich environments, plants face a critical compromise. Either

their rate of photosynthesis is limited by the fixed availability of CO2 (see
above), or they risk loosing too much moisture if they open their stomata for
lengthy periods of time as they try to increase the availability of CO2 to
photosynthesising leaf tissue. Plants have evolved two strategies to cope with
these problems, namely the C4 Pathway and CAM Pathway.
The C4 Pathway: Look back and recall that the Calvin Cycle is also called the C3
Pathway because the starting molecule is 3-carbon molecule of PGA. In the C4
pathway, plants fix CO2 into a four-carbon compound, oxaloacetate, prior to the
C3 pathway. This is done in specialised bundle-sheath cells separate from the
mesophyll cells. This is a way of accumulating extra carbon and is achieved by a
unique enzyme that has an extremely high affinity for CO2, binding to CO2 even
at very low concentrations. Due to the extra carbon accumulated, the plant does
not need to open its stomata for long periods of time to increase CO2 uptake, and
can minimise water loss by closing its stomata. Alternatively, they can increase
the rate of photosynthesis due to the extra carbon available. Plants that utilise
this strategy are called C4 plants. Many plants have C4 pathway in addition to the
standard C3 pathway (note that C4 does not replace C3 as a pathway, but is an
additional mechanism of carbon fixation). Such plants mentioned are found in
the tropics and good examples include sugar cane and corn.
The CAM Pathway: Many succulent plants growing in xeric (extremely arid)
conditions have developed a special carbon fixation pathway called CAM
(crassulacean acid metabolism). CAM plants open their stomata during the night
and fix CO2 into organic compounds through the C4 pathway. These organic
compounds accumulate overnight and are broken down (decarboxylated) during
the day to yield high levels of CO2 that is used to drive the Calvin Cycle. This
way, CAM plants solve the problem of water loss by keeping their stomata
closed during the day when high temperature and bright sunlight mean water
loss is at a maximum, and open their stomata at night when low temperatures
and lack of sunlight means water loss is minimal. Like C4 plants, CAM plants use
both the C4 and C3 pathways. However, they differ from C4 plants in that C4
occurs at night and C3 during the day. Both occur in the mesophyll cells.

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PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

EXERCISE 9.1
1.

What are three tissue types that make up a plant?

2.

Phloem and xylem are tissues vital for transport in plants. What
are their functions?

3.

Epidermis and root hairs are examples of tissue on the roots. Give
three examples of tissue or cell types found in leaves and give a
single function for each.

4.

The RuBP regeneration phase is the third stage of the Calvin

Cycle. What are the two first stages?

5.

It has been shown, though, that there are three vital factors that
affect the rate of photosynthesis. These are
,
and
the
availability
of
.

Plants are made up of three basic tissue types, namely dermal tissue, ground
tissue and vascular tissue.

The plant body is essentially made up of the root and the stem on which
leaves are formed. Tissue extend throughout the plant from the root to the
stem.

Leaves are specialised organs of photosynthesis in plants. Their structure is

characterised by tissue layers with specific functions.

Xerophytic plants are plants that are adapted to growing in xeric (arid)
conditions. Adaptations in such plants centre around reduction of water loss,
water storage and access to water.

Photosynthesis is a biochemical process that occurs in photoautotrophs

where light, energy, water and carbon dioxide is used to provide energy to
build carbohydrates. CO2 is used up in photosynthesis and oxygen is
released as a by product.

Photosynthesis occurs in the chloroplast and is divided into two stages,

namely the light-dependent reactions and the light-independent reactions

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(also called the Calvin Cycle). Photosynthetic pigments are vital in the light
reactions to trap the energy of sunlight.

The rate of photosynthesis is affected by a number of factors. The most

important of these are temperature, light intensity and CO2 availability.

The C4 pathway and the CAM pathway are alternative methods used by
plants in addition to the regular C3 pathway to increase the amount of CO2
available for photosynthesis or to decrease the need to open leaf stomata for
long periods.

1.

Which of the following statement is FALSE?

A. Xylem allows water and nutrients from soil to travel to the leaves.
B.
Xylem cells are living cells.
C. Phloem transports food generated from photosynthesis to the roots.
D. Plant vascular tissue consists of phloem and xylem.

2.

Which of the following plants is a xerophyte?

A. Hibiscus
B.
Oil palm
C. Paddy
D. Cactus

3.

How does xerophyte plants overcome extreme environment?

I
Thick waxy cuticle
II
Thick network of roots
III Modified photosynthetic pathway.
A. I only
B.
I and II only
C. II and III only
D. I, II and III.

4.

Which of the following plants contain more carotenoids?

A. Papaya
B.
Apple
C. Durian
D. Watermelon

5.

What is the factor(s) that governs the rate of photosynthesis?

I.
Light intensity
II.
Concentration of CO2

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III.
A.
B.
C.
D.

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PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS

Temperature
I only
I and II only
II and III only
I, II and III.

1.

Describe three adaptations in xerophytic plants that help to reduce water

loss from stomatal opening in the arid conditions of their habitats.

2.

Briefly describe the light-dependent reactions of photosynthesis. No

chemical names are expected in your description but the process must be
described in correct order.

Raven, P. H., Johnson, G. B., Losos, J. B., & Singer S. R. (2005). Biology. (7th ed.).
Boston: McGraw Hill.
Solomon, E. P., Berg, L. R., & Martin, D. W. (2002). Biology. (6th ed.). Toronto:
Thomas Learning Inc.