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Plant Physiology:

Transport and Photosynthesis


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.


Plants are multicellular autotrophs that have come to cover most of the earthÊs 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




explain the mechanisms of transport in plants and the fundamentals of the process of photosynthesis.



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 ÂskinÊ of the plant and protects the plant from damage and from excessive water loss.




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


Cell Types

Main Functions



Parenchyma cells

Storage, secretion,




Collenchyma cells Sclerenchyma cells (or fibres)



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


Vessel elements


Fibres Xylem parenchyma Sieve tube elements Companion cells Fibres Phloem parenchyma



Epidermal cells Guard cells Trichomes Cork cells Cork cambium cells Cork parenchyma

Protective cover Regulate stomata opening Variety of functions Protective covering Meristematic Storage






9.1.2 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.

the phloem filling the spaces between the arms of the X. Figure 9.1 (a) : Cross

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




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).

more adaptable in the rainforest soils. See Figure 9.1 (b). Figure 9.1 (b) : Shallow root

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:




94 TOPIC 9 PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS Figure 9.2 : Examples of root structure in

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 rootÊs 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




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.






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).

leaves and even carnivorou s leaves (see Figure 9.4). Figure 9.4 : Cross section of a

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




9.1.5 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








Parallel venation

Netted venation


Usually in threes

Usually in fours or fives


Embryo has one cotyledon

Embryo has two cotyledons

Secondary growth (wood and bark)


Usually present

Vascular bundles in stem

Usually scattered or in complex arrangements

Arranged in a circle or ring

Pollen grains

One pore

Three pores

circle or ring Pollen grains One pore Three pores Figure 9.5 : Comparison between monocot and

Figure 9.5: Comparison between monocot and dicot seeds showing the single cotyledon of the monocot and the double in the dicot




98 TOPIC 9 PLANT PHYSIOLOGY: TRANSPORT AND PHOTOSYNTHESIS Figure 9.6 : Differences in the distribution of

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



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.




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


Role of the elements


Deficiency Symptoms




Components of amino acids, proteins, nucleotides, nucleic acids, chlorophyll and coenzymes

Occurs in mature tissues; growth retardation; leaves turn to yellowish brown.





K + ,

involved in

Occur in young tissue; drying of the tips of root and leaf; twisted leaf morphology; retardation of root growth; decrease of plant growth rate.

osmotic and







stomata, as activator for many




Component of ATP and ADP, nucleic acids, several essential coenzymes, phospholipids, essential in plant metabolism and energy transfer

Old leaves become dark green colour, appearance of anthocyanin (dark purple pigment); delayed maturity


Component of some amino acids – cysteine and methionine, coenzyme-A and certain nucleotides.

Rarely happens




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



Symptom first occur in young tissues.






Spots on necrosis.







oxido-reductase reactions


Nitrogen fixation; component










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

9.3.1 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.






9.4.1 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:


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.


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 (inter- connecting 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




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

entry. Figure 9.7 shows the mechanism of root pressure. Figure 9.7 : A flow chart to

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 Ês 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.




EXERCISE 9.1 1. Apoplast is a „free space‰. Explain. 2. Is phloem a part of
1. Apoplast is a „free space‰. Explain.
2. Is phloem a part of apoplast? Explain.


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:




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).



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




The first two stages require light to proceed and are commonly called the light- dependent 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 light- independent 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:

6CO 2


(carbon dioxide)

12H 2 O (water)

C 6 H 12 O 6 (glucose)


6H 2 O (water)


6O 2


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 pigmentÊs 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.




9.6.2 The Chloroplast

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.

reactions of photosynthesis occur in the thylakoids. Figure 9.8 : Structure of a chloroplast 9.6.3 The

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 light- dependent 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




process called photophosphorylation. They are then passed to Photosystem I where they are used to drive the synthesis of NADPH (see Figure 9.9).

are used to drive the synthesis of NADPH (see Figure 9.9). Figure 9.9 : The light

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 by- product). 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




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):

three main phases of the Calvin Cycle (see Figure 9.10): Figure 9.10 : The dark reactions

Figure 9.10: The dark reactions (Calvin Cycle) of PS. Note the entry of CO 2 the release of 1G3P for sugar synthesis and the return of 5G3P into the cycle

1. The CO 2 uptake phase: The cycle begins with a molecule of CO 2 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 CO 2 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 CO 2 (6 × 1C = 6C) combining with 6 RuBP (6 × 5C = 30C) to give 12 PGA (36C).




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 C 3 photosynthesis because the initial carbon compound formed at the CO 2 uptake phase is a three-carbon molecule. Some plants also show C 4 photosynthesis which will be discussed briefly later.

9.6.4 Factors Affecting the Rate of Photosynthesis

Photosynthesis is a biochemical pathway that uses light, CO 2 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 CO 2 (see Figure



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 CO 2 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.


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




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.

the rate of photosynthes is in temperate and polar regions. Figure 9.11 : Changes in the

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. CO 2 availability: The availability of CO 2 is a natural limiting factor for all plants because the amount of CO 2 in the atmosphere is fixed (about 0.03%) and is constant virtually everywhere in the terrestrial environment. CO 2 is vital for the dark reactions of photosynthesis where carbon fixation occurs. Generally, the increase in CO 2 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 sub- optimal rate that is limited by the scarcity of CO 2 in the atmosphere. Often in commercial greenhouse agriculture, plant productivity is increased by increasing the concentration of CO 2 .

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 CO 2 concentration ceiling. This is why many scientists speculate that with increased global




temperatures from global warming, and increased CO 2 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 CO 2 (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 CO 2 to photosynthesising leaf tissue. Plants have evolved two strategies to cope with these problems, namely the C 4 Pathway and CAM Pathway.

The C 4 Pathway: Look back and recall that the Calvin Cycle is also called the C 3 Pathway because the starting molecule is 3-carbon molecule of PGA. In the C 4 pathway, plants fix CO 2 into a four-carbon compound, oxaloacetate, prior to the C 3 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 CO 2 , binding to CO 2 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 CO 2 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 C 4 plants. Many plants have C 4 pathway in addition to the standard C 3 pathway (note that C 4 does not replace C 3 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 CO 2 into organic compounds through the C 4 pathway. These organic compounds accumulate overnight and are broken down (decarboxylated) during the day to yield high levels of CO 2 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 C 4 plants, CAM plants use both the C 4 and C 3 pathways. However, they differ from C 4 plants in that C 4 occurs at night and C 3 during the day. Both occur in the mesophyll cells.




EXERCISE 9.1 1. What are three tissue types that make up a plant? 2. Phloem
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
photosynthesis. These are , and the availability of . • Plants are made up of three

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. CO 2 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




(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 CO 2 availability.

The C 4 pathway and the CAM pathway are alternative methods used by plants in addition to the regular C 3 pathway to increase the amount of CO 2 available for photosynthesis or to decrease the need to open leaf stomata for long periods.

to decrease the need to open leaf stomata for long periods. 1. Which of the following

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?




Oil palm





3. does xerophyte plants overcome extreme environment?



Thick waxy cuticle


Thick network of roots


Modified photosynthetic pathway.


I only


I and II only


II and III only


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




III. Temperature

A. I only

B. I and II only

C. II and III only

D. I, II and III.

I only B. I and II only C. II and III only D. I, II and

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.

but the process must be described in correct order. Raven, P. H., Johnson, G. B., Losos,

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.