BOTANY COURSE MATERIAL FOR SECOND SEMESTER 100LEVEL

STUDENTS.[2020/2021] ACADEMIC SESSION.

ROOT SYSTEM.

typical root is shown above. This is a taproot system made up of main axis which
is the taproot. From the taproot arises secondary branch roots and rootlets arise
from the branch roots.

Functions: the functions of the roots are -

1. To anchor the plants.

2. To provide support for the part of the plants above the soil surface.

3. Absorb water and mineral nutrients from the soil using the principles of
osmosis. The sap in the root tips is stronger than the solution of water and
nutrients around the root in the soil.
LEAF.

The leaves of Dicot and Monocot are green in color because they contain
chlorophyll. The veins in dicot are network and monocot parallel.

Functions:

Both types of leaves i.e dicot and monocot have same single function which is
photosynthesis.

The plant by the activity of photosynthesis are said to be auto trophic. The leaves
are generally called the source since they are the "industries" of food
manufacture. Where the photosyntate is used in the plant are called sink e.g
Growth regions

SHOOT SYSTEM

This is made of the following:

1. Bole or trunk or stem and bark, leaves, branches, flowers, fruits, buds nodes,
internodes.

Functions:

The bole or trunk or stem and bark bears the crown. The bole provide basis for
raw materials for saw milling industry, furniture industry etc.

Bark: protects the underlying tissue. the bud grows into branches or flora parts.

Nodes: point is attachment of leaf to the stem.

Inter nodes: the distance between two nodes.

Fruits/seeds: fruits are matured ovaries and provide food for man and animals.
The seeds are mature ovules and used mainly for new plant production. The seed
is covered with seed coat which protects the cotyledons. The food is used by the
embryo during early growth till the seedling starts to photosynthesize.

MOVEMENT OF MATERIALS IN PLANT.

Anatomy of the plant reveals two path ways involved in the movement of
materials in the plant .
the materials moved in the plant are_

1: water and nutrients grow the soil. These materials go through the xylem
vessels.

2. The product of photosynthesis. This moves from the source (leaf) through the
phloem vessels is where needed for growth, repairs etc.

These internal structures are known as vascular bundles. They occur in dicot and
monocot. The arrangement of vascular bundle may differ but they perform same
function.

PLANT PRODUCTION

INTRODUCTION

This deals with the techniques of providing new plants species. The plants to be
produced depends on the object of management. The production could be for
aesthetic values, regeneration, reforestation etc. The methods vary. Methods are
discussed below.
Young plant grows while in the mother plant.

This occurs naturally. Young plants grow from the seeds in the fruits attached to
parent plants. This is noticed mainly in mangrove areas e.g Avecinia resimosa
species. At a time in its development the young plant drops from the mother tree
or plant species to start a full life of plant species.

2: Apomixis.

This is a situation where the ovules are not fertilized and from the tissue of the
ovules grows new plants.
Vegetative reproduction. This deals with the production of new plants without
the fusion of male and female gametes from a zygote after fertilization.

(a) cuttings from roots, branches, stems etc are rooted and planted in the Field or
desired areas.

(b) other methods include - layering , grafting , tissue culture etc.

SEED GERMINATION

This is purely sexual reproduction which involves ovules (egg) and male gametes.
After fertilization and maturity the ovules form the seeds and the ovary form the
fruit . the seeds when sown under favourable condition grow into new plants.

Stages in seed germination:

1. Viability This is the ability of a given seed to germinate. This is usually
experimentally determined . two methods can be used - chemical and floatation .

CHEMICAL METHOD. Seeds are soaked in the appropriate chemical for 24hours.
There after seeds are removed and a cut is made through the embryo. If the
embryo shows red or pink color, this seed is viable if colourless the seed is not
viable.

FLOATATION METHOD. In this method some seeds are steeped into a container
of water and stirred. Leave it settle. Thereafter observe. Some seeds would have
sank to the bottom of the container while others float on top of the water. The
ones that sank are non viable .

DORMANCY: Seed dormancy is the inability of viable seeds to germinate even

when they are given good conditions for seed germination includes - oxygen
moisture, good substrate etc.

SEED INHIBITION. This is the ability of the seed to take in moisture to initiate the
germination process. If the viable seed is unable to imbibe water it cannot
germinate. The restricting factor is usually in the seed coat or taste. This limitation
can be over come by pre-sowing treatments .

PRE - SOWING TREATMENTS. The treatments aims at removing /to modify the
seed coat to allow imbibing of moisture. Some of the methods are, soaking in
water for some days, soaking in acids for some minutes, soaking in growth shows
etc.

There are two main types of seed germination - where the cotyledon remains in
the soil -- hypogeal and where cotyledons are carried up -- epigeal seed
germination.
INTRODUCTION.

Plant metabolism is the complex of physical and chemical events of

photosynthesis, respiration, and the synthesis and degradation of organic
compounds. Photosynthesis produces the substrates for respiration and the
starting organic compounds used as building blocks for subsequent biosynthesis
of nucleic acids, amino acids, and proteins, carbohydrates and organic acids,
lipids, and natural products.

All living organisms respire in order to release energy from glucose and make it
available in the form of ATP for chemical, osmotic and other work. Plants are no
exception. They need to respire virtually all the time in order to supply their
energy needs. They are not able to use the ATP generated in photosynthesis for
these purposes. Plants respire in the normal way using glycolysis, Krebs cycle,
oxidative phosphorylation etc. Often, the respiration is masked by the fact that
photosynthesis produces oxygen faster than respiration takes it up and
photosynthesis uses carbon dioxide faster than respiration produces it. It is only
in the dark that the full effects of respiration become apparent when
photosynthesis is brought to a halt (there is however the light and dark reactions
that will considered later in the course).

Plants need energy to take in mineral salts from the soil where they are present in
very low concentration. This needs energy to concentrate the mineral inside the
plant. Plants growing in waterlogged soils (which are short of oxygen) cannot
respire in their roots and soon show the symptoms of shortage of minerals (like
yellow leaves). Rice is interesting because it has a pithy stem through which it
enables oxygen from above the water to get down to the roots and therefore rice
thrives in "paddy fields".

Water is taken into the plant partly with the help of energy, but most of the
energy for water uptake is a result of the evaporation from the leaves "sucking"
the water up. However, moving sugars around the plant seems to require energy
as dead phloem cells do not transport sugars. Complex chemicals (like proteins)
need energy to make them from simple chemicals and require a supply of energy
to achieve this.

PLANT RESPIRATION

Plant respiration is a biochemical process whereby specific substrates are

oxidized with a subsequent release of carbon dioxide (CO 2). There is usually
conservation of energy accompanying the oxidation which is coupled to the
synthesis of energy-rich compounds, such as adenosine triphosphate (ATP),
whose free energy is then used to drive otherwise unfavourable reactions that are
essential for physiological processes such as growth.

Respiration is carried out by specific proteins, called enzymes, and it is necessary

for the synthesis of essential metabolites, including carbohydrates, amino acids,
and fatty acids, and for the transport of minerals and other solutes between cells
making respiration an essential characteristic of life itself in plants as well as in
other organisms.
Overall aerobic respiration is the end result of a sequence of many biochemical
reactions that ultimately lead to O2 uptake and CO2 evolution. In the absence of
O2, as may occur in bulky plant tissues such as the potato tuber and carrot root
and in submerged plants such as germinating rice seedlings, the breakdown of
hexose does not go to completion. The end products are either lactic acid or
ethanol, which are produced by anaerobic glycolysis or fermentation.

The sequence of reactions of anaerobic glycolysis or fermentation is shown in the

illustration. The enzymes associated with anaerobic glycolysis have been isolated
from many plant tissues, but more often ethanol and not lactic acid is the final
product. In aerobic tissues, pyruvic acid produced during glycolysis is completely
oxidized with the accompanying synthesis of much more ATP than in anaerobic
glycolysis.

Pyruvic acid oxidation takes place in the mitochondria by means of a cyclic

sequence of reactions, the Krebs cycle (also known as the citric acid cycle) which
begins when the first product of pyruvate oxidation, acetyl coenzyme A, reacts
with oxaloacetic acid to produce citric acid. Oxaloacetic acid is eventually
regenerated. Thus, the cycle can be repeated. In terms of the conservation of
chemical energy, the Krebs cycle is about 12 times more efficient than anaerobic
glycolysis per mole of glucose oxidized.
 In addition to anaerobic glycolysis and the Krebs cycle, there are two other
sequences of biochemical reactions related to respiration that are important in
plant tissues: (1) The pentose phosphate pathway permits an alternate
mechanism for converting hexose phosphate to pyruvate, and (2) in germinating
fatty seeds the reactions of the Krebs cycle are modified so that acetyl coenzyme
A is converted to succinic acid and then to hexose by a pathway called the
glyoxylate cycle.

AEROBIC RESPIRATION

Aerobic respiration (red arrows) is the main means by which both fungi and plants
utilize energy in the form of organic compounds that were previously created
through photosynthesis (green arrow).
Aerobic respiration requires oxygen in order to generate ATP. Although
carbohydrates, fats, and proteins can all be processed and consumed as
reactants, it is the preferred method of pyruvate breakdown in glycolysis and
requires that pyruvate enter the mitochondrion in order to be fully oxidized by
the Krebs cycle. The product of this process is carbon dioxide and water but the
energy transferred is used to break strong bonds in ADP as the third phosphate
group is added to form ATP (adenosine triphosphate), by substrate-level
phosphorylation, NADH and FADH2.

Simplified reaction:

C6H12O6 (s) + 6 O2 (g) → 6 CO2 (g) + 6 H2O (l) + heat

ΔG = -2880 kJ per mole of C6H12O6

The negative ΔG indicates that the reaction can occur spontaneously.

The reducing potential of NADH and FADH2 is converted to more ATP through an
electron transport chain with oxygen as the "terminal electron acceptor". Most of
the ATP produced by aerobic cellular respiration is made by oxidative
phosphorylation. This works by the energy released in the consumption of
pyruvate being used to create a chemiosmotic potential by pumping protons
across a membrane. This potential is then used to drive ATP synthase and
produce ATP from ADP and a phosphate group. Biology textbooks often state that
38 ATP molecules can be made per oxidised glucose molecule during cellular
respiration (2 from glycolysis, 2 from the Krebs cycle, and about 34 from the
electron transport system). However, this maximum yield is never quite reached
due to losses (leaky membranes) as well as the cost of moving pyruvate and ADP
into the mitochondria's matrix and current estimates range around 29 to 30 ATP
per glucose.

Aerobic metabolism is up to 15 times more efficient than anaerobic metabolism

(which yields 2 molecules ATP per 1 molecule glucose). However, some
anaerobic organisms, such as Methanogen are able to continue with anaerobic
respiration, yielding more ATP by using other inorganic molecules (not oxygen) as
a final electron acceptors in the electron transport chain. They share the initial
pathway of glycolysis but aerobic metabolism continues with the Krebs cycle and
oxidative phosphorylation. The post glycolytic reactions take place in the
mitochondria in eukaryotic cells, and in the cytoplasm in prokaryotic cells.

PHOTORESPIRATION

Photorespiration is the process occurring in plant cells under the action of light by
which oxygen is absorbed and carbon dioxide is given off. Although the
mechanism of photorespiration and the enzymes participating in the process have
not yet been thoroughly studied, it is speculated that in photorespiration the
reduced substances formed during electron transfer in photosynthesis are
oxidized in reactions of reciprocal conversions of glycolic and glyoxylic acids. In
some plants, photorespiration is intense, with as much as 50 percent of the
reduced nicotinamide adenine dinucleotide phosphate (NADP-N) formed during
photosynthesis being expended. In a number of tropical plants photorespiration is
not observed. It is believed that selective suppression of photorespiration by
means of specific inhibitors might increase the productivity of a number of
agricultural plants.

It can also be referred to the process in which green plants, algae, and
cyanobacteria utilize the energy of sunlight to manufacture carbohydrates from
carbon dioxide and water in the presence of chlorophyll. Some of the plants that
lack chlorophyll, e.g., the Indian pipe, secure their nutrients from organic
material, as do animals, and a few bacteria manufacture their own carbohydrates
with hydrogen and energy obtained from inorganic compounds (e.g., hydrogen
sulphide) in a process called chemosynthesis.

However, the vast majority of plants contain chlorophyll concentrated, in the

higher land plants, in the leaves. In these plants, water is absorbed by the roots
and carried to the leaves by the xylem, and carbon dioxide is obtained from the
air that enters the leaves through the stomata and diffuses to the cells containing
chlorophyll. The green pigment chlorophyll is uniquely capable of converting the
active energy of light into a latent form that can be stored (in food) and used
when needed.
When Light-dependent carbon dioxide is released and there is oxygen uptake in
photosynthetic organisms caused by the fixation of oxygen instead of carbon
dioxide during photosynthesis, the (oxygenation) reaction forms
phosphoglycolate. This represents carbon lost from the photosynthetic pathway.
Phosphoglycolate also inhibits photosynthesis if it is allowed to accumulate in the
plant. The reactions of photorespiration break down phosphoglycolate and
recover 75% of the carbon to the photosynthetic reaction sequence. The
remaining 25% of the carbon is released as carbon dioxide. Photorespiration
reduces the rate of photosynthesis in plants in three ways: carbon dioxide is
released; energy is diverted from photosynthetic reactions to photorespiratory
reactions; and competition between oxygen and carbon dioxide reduces the
efficiency of the important photosynthetic enzyme ribulose-bisphosphate (RuBP)
carboxylase. There is no known function of the oxygenation reaction but most
scientists believe it is an unavoidable side reaction of photosynthesis.

The rate of photosynthesis can be stimulated as much as 50% by reducing

photorespiration. Since photosynthesis provides the material necessary for plant
growth, photorespiration inhibits plant growth by reducing the net rate of carbon
dioxide assimilation (photosynthesis). Plants grow faster and larger under non-
photorespiratory conditions, in either low oxygen or high carbon dioxide
atmospheres. Most of the beneficial effects on plant growth achieved by
increasing CO2 may result from the reduced rate of photorespiration.

There are some plants that avoid photorespiration under certain conditions by
actively accumulating carbon dioxide inside the cells that have ribulose-
bisphosphate carboxylase/oxygenase. Many Cacti plants do this by taking up
carbon dioxide at night and then releasing it during the day to allow normal
photosynthesis. These plants are said to have crassulacean acid metabolism
(CAM). Another group of plants, including corn (Zea mays), take up carbon
dioxide by a special accumulating mechanism in one part of the leaf, then
transport it to another part of the leaf for release and fixation by normal
photosynthesis. The compound used to transport the carbon dioxide has four
carbon atoms, and so these plants are called C4 plants. Plants that have no
mechanism for accumulating carbon dioxide produce the three-carbon compound
phosphoglycerate directly and are therefore called C3 plants. Most species of
plants are C3 plants.

ANAEROBIC RESPIRATION

Cellular respiration is the process by which biological fuels are oxidised in the
presence of an inorganic electron acceptor (such as oxygen) to produce large
amounts of energy, to drive the bulk production of ATP. Anaerobic respiration is
used by some microorganisms in which neither oxygen (aerobic respiration) nor
pyruvate derivatives (fermentation) is the final electron acceptor. Rather, an
inorganic acceptor such as sulphate or nitrate is used.

PHOTOSYNTHESIS.

Photosynthesis is the manufacture in light of organic compounds (primarily

certain carbohydrates) from inorganic materials by chlorophyll- or
bacteriochlorophyll-containing cells. It is a process that requires a supply of
energy in the form of light. In chlorophyll-containing plant cells and in
cyanobacteria, photosynthesis involves oxidation of water (H2O) to oxygen
molecules, which are released into the environment. In contrast, bacterial
photosynthesis does not involve O2 evolution instead of H2O, other electron
donors, such as H2S, are used.

The light energy absorbed by the pigments of photosynthesizing cells, especially

by the pigment chlorophyll or bacteriochlorophyll, is efficiently converted into
stored chemical energy. Together, the two aspects of photosynthesis, the
conversion of inorganic into organic matter, and the conversion of light energy
into chemical energy make it the fundamental process of life on Earth: it is the
ultimate source of all living matter and of all life energy.

The net overall chemical reaction of plant photosynthesis is shown in the

equation below,

where {CH2O} stands for a carbohydrate (sugar).

The photochemical reaction in photosynthesis belongs to the type known as
oxidation-reduction, with CO2 acting as the oxidant (hydrogen or electron
acceptor) and water as the reductant (hydrogen or electron donor). The unique
characteristic of this particular oxidation-reduction is that it goes “in the wrong
direction” energetically; that is, it converts chemically stable materials into
chemically unstable products. Light energy is used to make this “uphill” reaction
possible.

Photosynthesis is a complex, multistage process. Its main parts are (1) the primary
photochemical process in which light energy absorbed by chlorophyll is converted
into chemical energy, in the form of some energy-rich intermediate products; and
(2) the enzyme-catalyzed “dark” (that is, not photochemical) reactions by which
these intermediates are converted into the final products carbohydrates and free
oxygen.

Experiments suggest that plants contain two pigment systems. One (called
photosystem I, or PS I, sensitizing reaction I) contains the major part of
chlorophyll a; the other (called photosystem II, or PS II, sensitizing reaction II)
contains some chlorophyll a and the major part of chlorophyll b or other auxiliary
pigments (for example, the red and blue pigments, called hycobilins, in red and
blue-green algae, and the brown pigment fucoxanthol in brown algae and
diatoms). It appears That efficient photosynthesis Requires the absorption of an
equal number of light quanta in PS I and in PS II; and that within both systems
excitation energy undergoes resonance migration from one pigment to another
until it ends in special molecules of chlorophyll a called the reaction centers. The
latter molecules then enter into a series of chemical reactions that result in the
oxidation of water to produce O2 and the reduction of nicotinamide adenine
dinucleotide phosphate (NADP+).

Chromatophores from photosynthetic bacteria and chloroplasts from green

plants, when illuminated in the presence of adenosine diphosphate (ADP) and
inorganic phosphate, also use light energy to synthesize adenosine triphosphate
(ATP); this photophosphorylation could be associated with some energy-releasing
step in photosynthesis.
Schematic outline of the Calvin (C3) carbondioxide assimilation cycle

The light-dependent conversion of radiant energy into chemical energy as ATP

and reduced nicotinamide adenine dinucleotide phosphate (NADPH) serves as a
prelude to the utilization of these compounds for the reductive fixation of CO2
into organic molecules. Such molecules, broadly designated as photosynthates,
are usually but not invariably in the form of carbohydrates such as glucose
polymers or sucrose, and form the base for the nutrition of all living things.
Collectively, the biochemical processes by which CO2 is assimilated into organic
molecules are known as the photosynthetic dark reactions, not because they
must occur in darkness, but because light in contrast to the photosynthetic light
reactions is not required.

The initial process in photosynthesis is the decomposition of water (H 2O) into

oxygen, which is released, and hydrogen. Direct light is required for this process.
The hydrogen and the carbon and oxygen of carbon dioxide (CO2) are then
converted into a series of increasingly complex compounds that result finally in a
stable organic compound, glucose (C6H12O6), and water. This phase of
photosynthesis utilizes stored energy and therefore can proceed in the dark. The
simplified equation used to represent this overall process is
6CO2+12H2O+energy=C6H12O6+6O2+6H2O. In general, the results of this process
are the reverse of those in respiration, in which carbohydrates are oxidized to
release energy, with the production of carbon dioxide and water.

The intermediary reactions before glucose is formed involve several enzymes,

which react with the coenzyme ATP to produce various molecules. Studies using
radioactive carbon have indicated that among the intermediate products are
three-carbon molecules from which acids and amino acids, as well as glucose, are
derived. This suggests that fats and proteins are also products of photosynthesis.
The main product, glucose, is the fundamental building block of carbohydrates
(e.g., sugars, starches, and cellulose). The water-soluble sugars (e.g., sucrose and
maltose) are used for immediate energy. The insoluble starches are stored as tiny
granules in various parts of the plant (chiefly the leaves, roots including tubers),
and fruits and can be broken down again when energy is needed. Cellulose is used
to build the rigid cell walls that are the principal supporting structure of plants.

C3 photosynthesis

The essential details of C3 photosynthesis can be seen in Fig. 3. Three molecules

of CO2 combine with three molecules of the five-carbon compound ribulose
bisphosphate (RuBP) in a reaction catalyzed by RuBP carboxylase to form three
molecules of an enzyme-bound six-carbon compound. These are hydrolyzed into
six molecules of the three-carbon compound phosphoglyceric acid (PGA), which
are phosphorylated by the conversion of six molecules of ATP (releasing ADP for
photophosphorylation via the light reactions). The resulting compounds are
reduced by the NADPH formed in photosynthetic light reactions to form six
molecules of the three-carbon compound phosphoglyceraldehyde (PGAL). One
molecule of PGAL is made available for combination with another three-carbon
compound, dihydroxyacetone phosphate, which is isomerized from a second
PGAL (requiring a second “turn” of the Calvin-cycle wheel) to form a six-carbon
sugar. The other five PGAL molecules, through a complex series of enzymatic
reactions, are rearranged into three molecules of RuBP, which can again be
carboxylated with CO2 to start the cycle turning again. The net product of two
“turns” of the cycle, a six-carbon sugar (glucose-6-phosphate) is formed either
within the chloroplast in a pathway leading to starch (a polymer of many glucose
molecules), or externally in the cytoplasm in a pathway leading to sucrose
(condensed from two six-carbon sugars, glucose and fructose).
Fig 4: Schematic outline of the Hatch-Slack (C4) carbon dioxide assimilation route
in two cell types of a NADP-ME-type plant.

C4 photosynthesis

Initially, the C3 cycle was thought to be the only route for CO 2 assimilation,
although it was recognized by plant anatomists that some rapidly growing plants
(such as maize, sugarcane, and sorghum) possessed an unusual organization of
the photosynthetic tissues in their leaves (Kranz morphology). It was then
demonstrated that plants having the Kranz anatomy utilized an additional CO2
assimilation route now known as the C4 dicarboxylic acid pathway (Fig. 4). Carbon
dioxide enters a mesophyll cell, where it combines with the three-carbon
compound phosphoenolpyruvate (PEP) to form a four-carbon acid, four-carbon
acid moves into bundle sheath cells, where the acid is decarboxylated, the CO2
assimilated via the C3 cycle, and the resulting three-carbon compound, pyruvic
acid, moves back into the mesophyll cell and is transformed into PEP, which can
be carboxylated again. The two cell types, mesophyll and bundle sheath, are not
necessarily adjacent, but in all documented cases of C4 photosynthesis, the
organism had two distinct types of green cells. C4 metabolism is classified into
three types, depending on the decarboxylation reaction used with the four-
carbon acid in the bundle sheath cells:

1. NADP-ME type (sorghum):

2. NAD-ME type (Atriplex species):

3. PCK type (Panicum species):

CAM photosynthesis

Under arid and desert conditions, where soil water is in short supply,
transpiration during the day when temperatures are high and humidity is low may
rapidly deplete the plant of water, leading to desiccation and death. By keeping
stomata closed during the day, water can be conserved, but the uptake of CO2,
which occurs entirely through the stomata, is prevented. Desert plants in the
Crassulaceae, Cactaceae, Euphorbiaceae, and 15 other families evolved,
apparently independently of C4 plants, an almost identical strategy of assimilating
CO2 by which the CO2 is taken in at night when the stomata open; water loss is
low because of the reduced temperatures and correspondingly higher humidities.
First studied in plants of the Crassulaceae, the process has been called
crassulacean acid metabolism (CAM).

In contrast to C4, where two cell types cooperate, the entire process occurs
within an individual cell; the separation of C4 and C3 is thus temporal rather than
spatial. At night, CO2 combines with PEP through the action of PEP carboxylase,
resulting in the formation of oxaloacetic acid and its conversion into malic acid.
The PEP is formed from starch or sugar via the glycolytic route of respiration.
Thus, there is a daily reciprocal relationship between starch (a storage product of
C3photosynthesis) and the accumulation of malic acid (the terminal product of
night time CO2 assimilation).

PROCESS OF PHOTOSYNTHESIS

Photosynthesis in plants occurs in two stages. These stages are known as the
light-dependent reactions and the Calvin Cycle.

Light-dependent Reactions

The first stage of photosynthesis is the light dependent reactions. These reactions
take place on the thylakoid membrane inside the chloroplast. During this stage
light energy is converted to ATP (chemical energy) and NADPH (reducing power).
Fig. 5: Light-dependent Reactions

Light is absorbed by two Photosystems called Photosystem I (PSI) and

Photosystem II (PSII). These protein complexes contain light harvesting
chlorophyll molecules and accessory pigments called antenna complexes. The
photosystems are also equipped with reactions centres (RC). These are
complexes of proteins and pigments which are responsible for energy conversion.
The chlorophyll molecules of PSI absorb light with a peak wavelength of 700nm
and are called P700 molecules. The chlorophyll molecules of PSII absorb light
with a peak wavelength of 68Onm and are called P68O molecules.

The light dependent reactions begin in PSII.

A photon of light is absorbed by a P680 chlorophyll molecule in the light

harvesting complex of PSII.

· The energy that is generated from the light is passed from one P680 chlorophyll
molecule to another until it reaches the reaction center (RC) of PSII.
· At the RC is a pair of P680 chlorophyll molecules. An electron in the chlorophyll
molecules becomes excited as a result of a higher level of energy. The excited
electron becomes unstable and is released. Another electron is released following
the capture of another photon of light by the light harvesting complex and the
transfer of energy to the reaction center.

· The electrons are transported in a chain of protein complexes and mobile

carriers called an electron transport chain (ETC). Plastoquinone is the mobile
carrier that transports the electrons from the reaction center of PSII to the
Cytochrome b6f Complex as shown in the diagram above.

· The electrons lost from PSII are replaced by splitting water with light in a process
called Photolysis. Water is used as the electron donor in oxygenic photosynthesis
and is split into electrons (e-), hydrogen ions (H+, protons) and oxygen (O2). The
hydrogen ions and oxygen are released into the thylakoid lumen. Oxygen is later
released into the atmosphere as a by-product of photosynthesis.

· While the electrons pass through the ETC via Plastoquinone, hydrogen ions
(protons) from the stroma are also transferred and released into the thylakoid
lumen. This results in a higher concentration of hydrogen ions (proton gradient)
in the lumen.

· As a result of the proton gradient in the lumen, hydrogen ions are transferred to
ATP synthase and provide the energy needed for combining ADP and Pi to
produce ATP.

· Cytochrome b6f transfers the electrons to Plastocyanin which then transports

them to Photosystem I.

The electrons have now arrived at PSI.

· They again receive energy, but this time from light absorbed by P700 chlorophyll
molecules.

· The electrons are transferred to mobile carrier, ferredoxin.

· They are then transported to ferredixin NADP reductase (FNR), which is the final
electron acceptor. At this point the electrons and a hydrogen ion are combined
with NADP+ to produce NADPH.

· The lost electrons from PSI are replaced by electrons from PSII via the electron
transport chain.

Summary of Light-dependent Reactions

Flow of Electrons

Photosystem II —–> b6-f complex —–> Photosystem I —-> NADP reductase

Role of Photolysis

Utilizes light to split water into the following:

· Electrons – donated to PSII to replace lost electrons

· Hydrogen ions – carried to ATP synthase to provide energy for the production of
ATP

· Oxygen – released into the atmosphere as a by-product

Products

· ATP – chemical energy

· NADPH – reducing power/electron donor

Light-dependent Reactions Animation

THE CALVIN CYCLE

The second stage of photosynthesis is the Calvin Cycle. These reactions occur in
the stroma of the chloroplast. Energy from ATP and electrons from NADPH are
used to convert carbon dioxide into glucose and other products.
Fig. 6: Overview of the Calvin Cycle pathway

· One molecule of carbon dioxide is combined with one molecule of Ribulose

Bisphosphate (RuBP). It is important to note that RuBP is a 5-carbon molecule.
When it is combined with CO2 the reaction produces an unstable 6-carbon
intermediate.

· The unstable 6-carbon intermediate quickly breaks down to form two 3-carbon
molecules known as 3-phosphoglycerate (PGA).

· The two 3-phosphoglycerate molecules receive energy from ATP and produce
two molecules of 1,3-bisphosphoglycerate (BPGA).

· An electron from NADPH is combined with each 1,3-bisphosphoglycerate

molecule to produce two molecules of Glyceraldehyde 3-phosphate (G3P).
Two Glyceraldehyde 3-phosphate molecules are needed to make one molecule of
glucose.

The next important step in the cycle is to regenerate RuBP. The problem is there
is not enough G3P. We only ran the cycle once with one molecule of CO2 and one
molecule of RuBP. Only two molecules of G3P were produced. We still need an
additional ten molecules of G3P for the cycle to continue.

If you take another look at the photosynthesis equation you will notice that six
molecules of carbon dioxide (6CO2) are needed for the process of photosynthesis.

These six molecules of CO2 must be used to produce twelve G3Ps. This means
that the steps above would have to be repeated five more times to produce ten
additional molecules of G3P.

Two molecules of G3P will be used to produce glucose and the other ten will be
used for the regeneration of RuBP.

IMPORTANCE OF PHOTOSYNTHESIS

Animals and plants both synthesize fats and proteins from carbohydrates; thus,
glucose is a basic energy source for all living organisms. The oxygen released (with
water vapor, in transpiration) as a photosynthetic by-product, principally of
phytoplankton, provides most of the atmospheric oxygen vital to respiration in
plants and animals, and animals in turn produce carbon dioxide necessary to
plants. Photosynthesis can therefore be considered the ultimate source of life for
nearly all plants and animals by providing the source of energy that drives all their
metabolic processes.

Fig. 7: Glycolysis and Citric Acid Cycle in Typical eukaryotic cell

 CELLULAR RESPIRATION

Cellular respiration is the set of the metabolic reactions and processes that take
place in the cells of organisms to convert biochemical energy from nutrients into
adenosine triphosphate (ATP), and then release waste products. The reactions
involved in respiration are catabolic reactions, which break large molecules into
smaller ones, releasing energy in the process as weak so-called "high-energy"
bonds are replaced by stronger bonds in the products. Respiration is one of the
keyways a cell gains useful energy to fuel cellular activity. Cellular respiration is
considered an exothermic redox reaction. The overall reaction is broken into
many smaller ones when it occurs in the body, most of which are redox reactions
themselves. Although technically, cellular respiration is a combustion reaction, it
clearly does not resemble one when it occurs in a living cell. This difference is
because it occurs in many separate steps. While the overall reaction is a
combustion reaction, no single reaction that comprises it is a combustion
reaction.

Nutrients that are commonly used by animal and plant cells in respiration include
sugar, amino acids and fatty acids, and a common oxidizing agent (electron
acceptor) is molecular oxygen (O2). The energy stored in ATP (its third phosphate
group is weakly bonded to the rest of the molecule and is cheaply broken allowing
stronger bonds to form, thereby transferring energy for use by the cell) can then
be used to drive processes requiring energy, including biosynthesis, locomotion or
transportation of molecules across cell membranes.
GLYCOLYSIS

Glycolysis is a metabolic pathway that takes place in the cytosol of cells in all
living organisms. This pathway can function with or without the presence of
oxygen. Aerobic conditions produce pyruvate and anaerobic conditions produce
lactate. In aerobic conditions, the process converts one molecule of glucose into
two molecules of pyruvate (pyruvic acid), generating energy in the form of two
net molecules of ATP. Four molecules of ATP per glucose are actually produced;
however, two are consumed as part of the preparatory phase. The initial
phosphorylation of glucose is required to increase the reactivity (decrease its
stability) in order for the molecule to be cleaved into two pyruvate molecules by
the enzyme Aldolase. During the pay-off phase of glycolysis, four phosphate
groups are transferred to ADP by substrate-level phosphorylation to make four
ATP, and two NADH are produced when the pyruvate are oxidized. The overall
reaction can be expressed as:

Glucose + 2 NAD+ + 2 Pi + 2 ADP → 2 pyruvate + 2 NADH + 2 ATP +2 H+ + 2 H2O +

heat

Starting with glucose, 1 ATP is used to donate a phosphate to glucose to produce

glucose 6-phosphate. Glycogen can change into glucose 6- phosphate as well with
the help of glycogen phosphorylase. During Energy metabolism, glucose 6-
phosphate turns into fructose 6-phosphate. An additional ATP is used to
phosphorylate fructose 6-phosphate into fructose 1,6-disphosphate by the help of
phosphofructokinase. Fructose 1,6-diphosphate then splits into two
phosphorylated molecules with three carbon chains that later degrades into
pyruvate.

Oxidative Decarboxylation of pyruvate

Pyruvate is oxidized to acetyl-CoA and CO2 by the pyruvate dehydrogenase

complex (PDC). The PDC contains multiple copies of three enzymes and is located
in the mitochondria of eukaryotic cells and in the cytosol of prokaryotes. In the
conversion of pyruvate to acetyl-CoA, one molecule of NADH and one molecule of
CO2 is formed. This step is also known as the link reaction or transition step, as it
links glycolysis and the Krebs cycle.

CITRIC ACID CYCLE

This is also called the Krebs cycle or the tricarboxylic acid cycle. When oxygen is
present, acetyl-CoA is produced from the pyruvate molecules created from
glycolysis. When oxygen is present, the mitochondria will undergo aerobic
respiration which leads to the Krebs cycle. However, if oxygen is not present,
fermentation of the pyruvate molecule will occur. In the presence of oxygen,
when acetyl-CoA is produced, the molecule then enters the citric acid cycle (Krebs
cycle) inside the mitochondrial matrix, and gets oxidized to CO2 while at the same
time reducing NAD to NADH. NADH can be used by the electron transport chain to
create further ATP as part of oxidative phosphorylation. To fully oxidize the
equivalent of one glucose molecule, two acetyl-CoA must be metabolized by the
Krebs cycle. Two waste products, H2O and CO2, are created during this cycle.

The citric acid cycle is an 8-step process involving different enzymes and co-
enzymes. Throughout the entire cycle, acetyl-CoA (2 carbons) + Oxaloacetate (4
carbons), Citrate (6 carbons) is rearranged to a more reactive form called
Isocitrate(6carbons). Isocitrate (6 carbons) modifies to become α-Ketoglutarate (5
carbons), Succinyl-CoA, Succinate, Fumarate, Malate, and finally, Oxaloacetate.
The net energy gain from one cycle is 3 NADH, 1 FADH2, and 1 GTP; the GTP may
subsequently be used to produce ATP. Thus, the total energy yield from one
whole glucose molecule (2 pyruvate molecules) is 6 NADH, 2FADH2, and 2 ATP.

FERMENTATION

Without oxygen, pyruvate (pyruvic acid) is not metabolized by cellular respiration

but undergoes a process of fermentation. The pyruvate is not transported into
the mitochondrion, but remains in the cytoplasm, where it is converted to waste
products that may be removed from the cell. This serves the purpose of oxidizing
the electron carriers so that they can perform glycolysis again and removing the
excess pyruvate. Fermentation oxidizes NADH to NAD+ so it can be re-used in
glycolysis. In the absence of oxygen, fermentation prevents the build-up of NADH
in the cytoplasm and provides NAD+ for glycolysis. This waste product varies
depending on the organism.

In skeletal muscles, the waste product is lactic acid. This type of fermentation is
called lactic acid fermentation. In strenuous exercise, when energy demands
exceed energy supply, the respiratory chain cannot process all of the hydrogen
atoms joined by NADH. During anaerobic glycolysis, NAD+ regenerates when pairs
of hydrogen combine with pyruvate to form lactate. Lactate formation is
catalyzed by lactate dehydrogenase in a reversible reaction. Lactate can also be
used as an indirect precursor for liver glycogen. During recovery, when oxygen
becomes available, NAD+ attaches to hydrogen from lactate to form ATP. In yeast,
the waste products are ethanol and carbon dioxide. This type of fermentation is
known as alcoholic or ethanol fermentation. The ATP generated in this process is
made by substrate-level phosphorylation, which does not require oxygen.

Fermentation is less efficient at using the energy from glucose since only 2 ATP
are produced per glucose, compared to the 38 ATP per glucose produced by
aerobic respiration. This is because the waste products of fermentation still
contain plenty of energy. Ethanol, for example, can be used in gasoline (petrol)
solutions. Glycolytic ATP, however, is created more quickly. For prokaryotes to
continue a rapid growth rate when they are shifted from an aerobic environment
to an anaerobic environment, they must increase the rate of the glycolytic
reactions. For multicellular organisms, during short bursts of strenuous activity,
muscle cells use fermentation to supplement the ATP production from the slower
aerobic respiration, so fermentation may be used by a cell even before the
oxygen levels are depleted, as is the case in sports that do not require athletes to
pace themselves, such as sprinting.

STRUCTURE OF LEAVES
Fig. 9: Leaf Tissue Structure
The typical plant leaf includes the following

· Upper and lower epidermis – the upper epidermis is the outer layer of the cells
that controls the amount of water that is lost through transpiration.

· Stomata – these are pores (holes) in the leaves that are responsible for the
exchange of gases between the plant leaves and the atmosphere. Carbon dioxide
is absorbed from the atmosphere and oxygen is released.

· Mesophyll – these are photosynthetic (parenchyma) cells that are located

between the upper and lower epidermis. These cells contain the chloroplasts.

· Vascular bundle – these are tissues that form part of the transport system of the
plant. Vascular bundles consist of xylem and phloem vessels which transport
water, dissolved minerals and food to and from the leaves.

MINERAL NUTRITION OF PLANTS

Mineral Nutrition is the assimilation by plants from the environment of the ions of
mineral salts needed for normal development. Mineral nutrients include the
elements nitrogen, phosphorus, sulphur, potassium, calcium, and magnesium, as
well as such trace elements as iron, boron, copper, zinc, and magnesium. Mineral
nutrition of plants involves the absorption of minerals in the form of ions, their
transport within plants, and their inclusion in metabolism. Unicellular organisms
and aquatic plants absorb ions over their entire surface, whereas higher
terrestrial plants absorb the ions on the surface cells of the roots, predominantly
on root hairs. After absorption by the cell membranes, the ions penetrate the
cytoplasm through its surrounding lipoprotein membrane (plasmalemma).
Cations (except K+) penetrate the membrane passively, by diffusion; anions and
K+ (at low concentrations) do so actively, by means of molecular “ion pumps,”
which transport ions with the expenditure of energy. The rate of active ion
transport depends on the quantity of carbohydrates available to the cell and on
the respiration rate, whereas the rate of passive absorption depends on the
permeability of biological membranes and the difference in concentrations and
electric potentials between the solution and the cell. Membrane permeability is
not the same for the different ions: for the cation K+ it is 100 times higher than
for Na+ and 500 times higher than for anions. Absorbed ions travel from cell to
cell by way of the connecting cytoplasmic bridges (plasmodesmata). The roots
and stems of higher plants have a special vascular system for transporting
minerals and their organic compounds (which are also partly synthesized in the
roots) to the leaves. As the lower leaves age, some mineral substances flow from
them into the growing organs, where they can be used again.

Every element of mineral nutrition plays a specific role in metabolism and cannot
be completely replaced by another element. Nitrogen forms part of the proteins,
the main substances of the cytoplasm, and of the amides, nucleic acids,
hormones, alkaloids, vitamins (B1, B2, B6, and PP) and chlorophyll. Nitrogen is
absorbed in the form of the anion NO3-(nitrate) and cation NH4+ (ammonium),
which are created by the decomposition of humus by soil microorganisms.
Molecular nitrogen (N2), the main constituent of air (79 percent), can be
assimilated only by certain species of lower plants. Nitrates are reduced to
ammonium by means of the enzyme nitroreductase. Ammonium combines with
organic acids to form amino acids, which are then incorporated into proteins.
Phosphorus forms part of the nucleoproteids of the cell nucleus, phospholipids of
cell membranes, phosphatides, and phosphoric esters of sugars. The participation
of phosphorus in photophosphorylation is particularly important. In this process,
solar energy accumulated in the form of energy-rich adenosine triphosphate
(ATP) bonds is used to take up CO2 from the air and to form organic matter.
Energy released during respiration by the oxidation of organic matter formed in
the course of photosynthesis is also stored in the form of macroergic ATP bonds.
Phosphorus is absorbed as an anion of orthophosphoric acid (PO43-, or
phosphate) and inalterably incorporated into organic compounds in hundredths
of a second. In addition, plants always contain a large quantity of inorganic
phosphate, whose physiological function is unknown.

Sulphur, like nitrogen, is a constituent of all proteins, as well as peptides

(glutathione) and some amino acids (cystine, cysteine, and methionine) and
essential oils. Sulphur is taken up by plants in the form of the anion (SO4 2-, or
sulphate), which is reduced in the cells to form disulphide (—S—S—) and
sulfhydryl (—SH) groups (the latter form bonds that strengthen the configuration
of the protein macromolecule).

Potassium is absorbed in the form of the K+ cation and remains in the same form
in the cell without forming stable organic compounds. It participates only in weak
adsorption interactions with proteins and in exchange reactions with inorganic
acids. Unlike nitrogen, phosphorus, and sulphur, which participate directly in the
creation of organic material in the plant cell, potassium is not a nutrient in the full
sense of the word. It increases the water-retention capacity of the cytoplasm and
the rate of photosynthesis and outflow of assimilated material and participates in
the functioning of the stomata.

Calcium and magnesium are absorbed in the form of bivalent cations, Ca2+ and
Mg2+. The main function of calcium is stabilization of the cell structures. The Ca2+
ions (“calcium bridges”) connect the lipid molecules to one another and ensure
their orderly arrangement in cell membranes. Compounds of calcium with
pectin’s cement the membranes of adjacent cells. Unlike other elements of
mineral nutrition, calcium is almost immobile in plants. It is virtually not reused,
and it accumulates in aging organs. It is essential to maintain the structure of the
ribosomes, where protein is synthesized.

Manganese is a constituent of chlorophyll, and it activates the enzymes that

transport phosphate from ATP to the sugar molecule.

Iron is part of several enzymes, including the respiratory enzymes (cytochromes).

It is involved in the synthesis of chlorophyll, although it does not become part of
it. Plants can also receive mineral nutrients through the leaves.

Together with aerial nutrition (photosynthesis), the mineral nutrition of plants

constitutes a single process of exchange between the plants and the
environment. It affects all physiological functions (respiration, growth,
development, photosynthesis, water conditions, and so on) and, in turn, is
dependent on them. Therefore, one of the most effective means of controlling
crop productivity is regulation of mineral nutrition by means of fertilizers.

TRANSLOCATION is the movement of materials from leaves to other tissues

throughout the plant. Plants produce carbohydrates (sugars) in their leaves by
photosynthesis, but non-photosynthetic parts of the plant also require
carbohydrates and other organic and nonorganic materials. For this reason,
nutrients are translocated from sources (regions of excess carbohydrates,
primarily mature leaves) to sinks (regions where the carbohydrate is needed).
Some important sinks are roots, flowers, fruits, stems, and developing leaves.
Leaves are particularly interesting in this regard because they are sinks when they
are young and become sources later, when they are about half grown.

Phloem Structure and Function

The tissue in which nutrients move is the phloem. The phloem is arranged in long,
continuous strands called vascular bundles that extend through the roots and
stem and reach into the leaves as veins. Vascular bundles also contain the xylem,
the tissue that carries water and dissolved minerals from the roots to the shoots.
When plants increase in diameter (secondary growth) they do so by divisions of a
layer of cells just under the bark; this cell layer makes new xylem to the inside
(forming the wood of the tree trunk) and a thin, continuous cylinder of new
phloem to the outside.

The contents of the phloem can be analyzed by cutting off the stylets (mouth
parts) of phloem-feeding insects such as aphids and collecting the drops of sap
that exude. Phloem sap is composed largely of sugar dissolved in water. All plants
translocate sucrose (table sugar) and some also transport other sugars such as
stachyose, or sugar alcohols such as sorbitol. Many other organic compounds are
found, including amino acids, proteins, and hormones . Glucose, the sugar found
in the circulatory system of animals, is not translocated.

In order to accommodate the flow of sap, the internal structure of the conducting
cells of the phloem, the sieve elements, is drastically altered. As the sieve
elements mature, they lose many of the organelles commonly found in living cells
and they modify others. The nucleus disappears, as do the vacuoles,
microfilaments, microtubules, ribosomes, and Golgi bodies. Therefore, the inside
(lumen) of the cell is left essentially open. The sieve elements are greatly
elongated in the direction of transport and are connected to one another to form
long sieve tubes. Large pores perforate the end walls of the sieve elements to
facilitate flow through the tube. The connecting walls thus look like a sieve, giving
the cell type its name.

Some sieve elements can live for a long time, as many as one hundred years in
palm trees, even though they have no nucleus or any of the machinery needed for
protein synthesis. Cells closely associated with them, called companion cells,
apparently keep them alive. The association of sieve elements and companion
cells is one of the most intimate and complex in nature, and one of the least
understood. It now appears that both small and large molecules can move from
companion cells to sieve elements through the plasmodesmata that connect
them. Plasmodesmata are minute pores that traverse the common walls between
plant cells. They have an intricate internal structure. Interest in plasmodesmata is
high because viruses move through them to cause infections. If a virus enters the
phloem this way it will travel with the sap, spread widely around the plant, and
infect sink organs. Since viruses are much larger than plasmodesmata, they must
be disassembled in one cell and reassembled when they get to their destination.

Sugars synthesized in the chloroplasts are actively pumped into the sieve tubes.
Water follows by osmosis, creating high pressure. Sugar is then removed by active
transport, and water again by osmosis, lowering the pressure in the sieve tube.

THE PRESSURE -FLOW MECHANISM

The rate of translocation in angiosperms (flowering plants) is approximately 1

meter per hour. In conifers, it is generally much slower, but even so this is far too
fast to be accounted for by diffusion. Instead, the sap flows, like a river of dilute
syrup water. What is the force that drives the flow of material in the phloem? It is
pressure, generated in the sieve elements and companion cells in source tissues.
In leaves, sugar is synthesized in mesophyll cells (the middle layer of the leaf), and
is then actively pumped into the phloem, using metabolic energy. By using
energy, the sugar is not only transferred to the phloem but is also concentrated.
When a solute such as sugar is concentrated inside cells, water enters the cells by
osmosis. Since the plant cells have a rigid cell wall, this influx of water creates a
great deal of internal pressure, over ten times the pressure in an automobile tire.
The pressure causes sap to move out through the pores of the sieve element,
down the tube.

At the other end of the transport stream, in the sinks, sugar is constantly leaving
the phloem and being used by surrounding cells. Some is consumed as an energy
source, some is stored as sugar or starch, and some is used to make new cells if
the sink tissue is growing. Since sugar leaves the phloem in the sink, water exits
too (again by osmosis) and the pressure goes down. Therefore, there is a
difference in pressure between source and sink phloem. This causes the solution
to flow, just as water flows along a pressure gradient in a garden hose. This
process is known as the pressure-flow mechanism.

Sugar Loading and Unloading

How is sugar actively pumped (loaded) into the phloem? There are two known
mechanisms, operating in different species. In one, sucrose enters the cell walls
near the phloem in the smallest (minor) veins of the leaf. It then enters the
phloem by attaching to sucrose transporter proteins embedded in the plasma
membranes of the sieve elements and companions cells. In the second
mechanism, sucrose enters the companion cells of the minor vein through small
plasmodesmata, and is converted to larger sugars, raffinose, and stachyose. These
larger sugars are unable to diffuse back through these plasmodesmata due to
their size. Therefore, they are trapped in the phloem of the leaf and build up to
high concentration. They enter the sieve elements through larger plasmodesmata
and are carried away toward the sinks.

When sugars and other nutrients arrive in sink tissues they unload from the
phloem and enter surrounding cells, either through plasmodesmata or by crossing
from one cell to another across the cell walls. The size and metabolic activity of
the different sinks determines the amount of material that is delivered to them.
Thus, the use of sugar in the sinks determines how much sugar flows to them.

Water Movement in Plants

Long-distance water movement is crucial to the survival of land plants. Although

plants vary considerably in their tolerance of water deficits, they all have their
limits, beyond which survival is no longer possible. About 85 percent of the fresh
weight of leaves can be water. On a dry, warm, sunny day, a leaf can evaporate
100 percent of its water weight in just an hour. Water loss from the leaves must
be compensated for by the uptake of water from the soil. Water transport is also
important for the uptake of essential mineral nutrients from the soil. Shortages of
mineral nutrients such as nitrogen, potassium, and phosphorus are often limiting
to plant growth, which is why fertilizers are often added to the soil to improve
plant productivity and appearance.

THE COHENSION-TENSION THEORY

The major mechanism for long-distance water transport is described by the

cohesion-tension theory, whereby the driving force of transport is transpiration,
i.e., the evaporation of water from the leaf surfaces. Water molecules cohere
(stick together), and are pulled up the plant by the tension, or pulling force,
exerted by evaporation at the leaf surface.

Water will always move toward a site with lower water potential, which is a
measure of the chemical free energy of water. By definition, pure water has a
water potential of 0 MegaPascals (MPa). In contrast, at 20 percent relative
humidity, the water potential of the atmosphere is -500 MPa. This difference
signifies that water will tend to evaporate into the atmosphere. The water within
plants also has a negative potential, indicating water will tend to evaporate into
the air from the leaf. The leaves of crop plants often function at -1 MPa, and some
desert plants can tolerate leaf water potentials as low as -10 MPa. The water in
plants can exist at such low water potentials due to the cohesive forces of water
molecules. The chemical structure of water molecules is such that they cohere
very strongly. By the cohesion-tension theory, when sunlight strikes a leaf, the
resultant evaporation first causes a drop in leaf water potential. This causes water
to move from stem to leaf, lowering the water potential in the stem, which in turn
causes water to move from root to stem, and soil to root. This serves to pull water
up through the xylem tissue of the plant.

From Root to Leaf

Plants have root hairs and often mycorrhizal fungi at the root surface, both of
which serve to filter the soil water as it enters the plant. Mycorrhizae are
symbiotic associations between plant roots and fungi. The root cells and
mycorrhizal fungi both actively uptake certain mineral nutrients. Mycorrhizae can
be particularly important for the uptake of phosphate. The active uptake of
minerals by living cells of the root and the subsequent transfer of minerals to the
xylem can result in positive root pressures, with water potentials above 0 MPa.
This occurs only under certain conditions, such as at night or during rainstorms,
when water loss from the leaves is minimal. Such positive root pressures
disappear with the onset of leaf transpiration.

Water molecules move from the soil into living cells of the root, and eventually
into the transport cells of the xylem, known as tracheids and vessels. These xylem
cells are dead and hollow, allowing rapid water transport. They also have
hardened cell walls to help them resist the tendency to collapse as water is
sucked through them. Both tracheids and vessels have pits on the sides of their
walls, which include porous areas for side-to-side transport. Unlike tracheids, a
vessel is composed of many cells stacked end to end, with perforations between
cells, allowing for more efficient transport.

The long-distance transport of the water molecule occurs first within the xylem
cells of the root, then the xylem of the stem and branch, and then into the xylem
of a leaf midrib and vein. Driven by transpiration, the water molecule is pulled
from the non-living tracheids and vessels of the xylem in the living cells of the leaf
mesophyll (middle layer) and to the surface of mesophyll cell walls. The water
molecule then evaporates into a leaf inter-cellular air space and finally out of a
stomatal pore and into the atmosphere. Though photosynthetic action consumes
some water, only a small fraction of the water that travels through the plant is
used directly for the photo-synthetic reaction, which occurs in leaf mesophyll
cells. Instead, most water is lost by transpiration through the stomates.

THE ROLE OF STOMATES

Leaves of land plants are covered with a waxy cuticle that prevents water loss and
gas exchange. The stomates at the leaf surface have guard cells that open and
close the stomate to regulate the uptake of carbon dioxide and release of oxygen,
as required for photosynthesis. They also serve to regulate water loss from
transpiration. During the day, the stomates normally open up in response to
sunlight, allowing for photosynthetic gas exchange, but also allowing for
transpiration. At night, the stomates normally close, preventing unnecessary
water loss. When excessive water loss occurs during the day, drops in leaf water
potential can cause stomates to close. Were it not for stomate closure in
response to water stress, the leaves would suffer excessive water loss, the leaf
cell membranes and photosynthetic apparatus would be destroyed, and
"cavitation" would occur in the xylem cells. Cavitation, which is a break in the
water column, occurs when air is pulled into the xylem vessel or tracheid. This can
make the xylem cell unable to conduct water. Plants vary considerably in their
vulnerability to cavitation, but for most plants, stomate closure can prevent
cavitation from occurring.

The transpirational water loss allows for uptake of mineral nutrients from the soil.
However, much of the water loss that land plants exhibit can be viewed as a
"necessary evil." The stomates must open up to allow for photosynthesis to occur,
and during the process of letting carbon dioxide into the leaf, water vapor is lost
to the atmosphere. When the stomates close to prevent excess water loss,
photosynthesis is compromised.