Def Physio

Download as doc, pdf, or txt
Download as doc, pdf, or txt
You are on page 1of 34

TREE PHYSIOLOGY

TERMINOLOGY:
Absorption – the intake of water, and nutrients in the roots and leaves or any part of the
plant.
Accumulation – concentration of ions or substances inside the cell or organ.
Accumulation ratio – ratio of the amount of ions or substances inside the cell to the
amount of ions outside the cell (external solution)
Acid – a substance that can donate a proton.
Active absorption – movement of nutrients, substance or dissolved materials into the
cell or organ against an electrochemical potential gradient
Active mechanism – process that requires expenditure of energy
Adsorption – the concentration and accumulation of ions or molecules of a substance
at a surface or interface.
Allocation – metabolic fate of carbon either newly assimilated in the source leaf or
delivered to a sink.
Amino acids – organic compounds containing amino and carboxylic groups; building
blocks of protein
Amphoteric compound – a compound which can either donate or accept a proton and
has the properties of both acid and base
Anabolism – synthesis of complex molecules from simpler substances
Antagonism – presence or excess of one ion impairs the absorption of another ion or
induces a deficiency of another.
Antiport – linked movement of two entities in opposite directions at the same site in a
protoplasmic membrane.
Apical dominance – inhibition of lateral buds by the apex or apical meristem
Assimilation – metabolic transformation of simple nutrients and substances to organic
plant constituents.
Base – a substance that can accept a proton (H+).
Buffer solutions – solutions which can resist abrupt changes in pH
Catabolism – breakdown of complex molecules into simpler substances.
Cell – basic unit of life.
Cell wall – layer of material chiefly elongated polymers laid down outside the plasma
membrane to protect the protoplast and limit its expansion.
Chloroplast – a plastid that contains chlorophyll pigment that absorb light energy during
photosynthesis.
Cohesion – strong mutual attraction between water molecules resulting from hydrogen
bonding.

FRE - TREE PHYSIOLOGY 1


Colloids – particles small enough to remain in suspension but too large to go into
solution.
Critical nutrient concentration – level of nutrient element in the tissue below which
deficiency symptoms occur and above which an optimum growth is obtained
Cytosol – non-particulate (liquid) portion of the cytoplasm.
Denaturation – collapse of the enzyme structure caused by high temperature or other
factors.
Development – ordered change in the complexity of an organism; a combination of both
growth and differentiation
Diffusion – movement of atoms, ions, molecules or particles from an area of higher
concentration to an area of lower concentration.
Diffusion pressure deficit – difference in diffusion pressure between a solution and a
pure solvent when both are subjected to the same pressure
Electron transport chain – a membrane bound system that controls the flow of electron
from reduced to oxidized compound so that some of the energy carried by the
electron is used to form ATP; found in mitochondria and chloroplast.
Emulsion – a colloidal system consisting of one liquid dispersed in another liquid.
Endoplasmic Reticulum (ER) – single, highly convoluted membrane continuous with
the nuclear envelope and serves as the channel of communication and transport
between the nucleus and cytoplasm.
Enzymes – a protein that acts as a catalyst to speed up chemical reaction.
Field capacity – amount of water retained in the soil after large capillary space have
been drained by gravity
Function – natural activity of a thing whether a cell, organ or molecule.
Geotropism – movement in response to gravity
Golgi bodies – stacks of flattened membranous sac called cisternae which assemble
and process carbohydrate chain of glycoproteins.
Growth – an irreversible increase in size or volume; a quantitative aspect of
development
Growth correlation – relationship of the growth of one part to the growth of another part
Growth movement – irreversible movement that occurs due to the growth and
elongation of cells
Gutation – water loss in liquid form through the hydathodes.
Hydathodes – small opening at the leaf tip
Hypertonic solution – external solution has more negative osmotic potential than the
cell or water concentration inside the cell is higher than the external solution
leading to plasmolysis.
Hypotonic solution – external solution has higher concentration of water than the cell
or external solution has less negative water potential than the cell

FRE - TREE PHYSIOLOGY 2


Imbibition (in plants) – absorption and adsorption of water by insoluble solid, hydrophilic
protoplasmic and cell wall constituents
Incipient plasmolysis – point at which 50% of the cell is plasmolyzed
Ion – an atom or group of atoms that are electrically charged. Positively charged ions
are called cations and the negatively charged are anions.
Isotonic solution – external solution has equal osmotic potential with the cell.
Metabolite – a chemical or substance that is a normal cell constituent capable of
entering into the biochemical transformations within living cells.
Microbody – a single membrane cellular organelle frequently spherical, 20 to 60 nm in
diameter and containing a variety of enzyme.
Mitochondrion – a double membrane organelle where aerobic respiration take place;
powerhouse of the cell.
Mycorrhiza – composite organs formed by specific association of particular specialized
fungi with the roots; association between fungus and roots of plants
Nitrogen fixation – conversion of dinitrogen (N2) to ammonium either chemically or
biologically in a reaction catalyzed by nitrogenase
Nucleic acid – large polymeric molecules composed of repeating units called
nucleotides.
Nucleus – a double membrane; information center of the cell.
Oleosome – lipid storage bodies
Organelles – discrete, membrane-limited compartments in the cytoplasm.
Osmosis – movement of water across a differentially permeable membrane from a
region of higher concentration to a region of lower concentration.
Osmotic pressure – force to be applied to stop the osmotic movement of water across
membrane
Outer space – space in the cell, tissue or organ open for free diffusion of ions
Partitioning – distribution of assimilate between sinks.
Phloem loading – translocation of assimilates from the photosynthetic mesophyll cells
into the sieve elements at the source end.
Phloem unwading – translocation of assimilates from the sieve elements into the target
cells at the sink end.
Photoperiodism – response of plants to the relative length of light and dark period
Photorespiration – a light-dependent oxidation of organic compound (glycolate) to CO 2
occurring in chlorophyllous tissues
Photosynthesis – absorption of light energy and its conversion to useful chemical
energy
Physiology
Physis – nature
Logos – discourse or study

FRE - TREE PHYSIOLOGY 3


Plant Physiology – a science concerned with functions and processes, the response of
plants to changes in the environment, and the growth and development that
result from such responses.
Plagiogeotropism – lateral branches become oriented at angles around 90o
Plant - a machine that takes in energy and simple inorganic molecules from the physical
environment and uses them to assemble chemical structure; a biochemical
factory that literally built itself.
Plant growth regulators – organic compounds other than nutrients which in small
amount promote, inhibit or otherwise modify any physiological processes in
plants
Plant hormones (phytohormones) – organic substances produced naturally in plants
controlling growth and other physiological functions at sites remote from the
place of production and eliciting response at minimal amount.
Plasmodesma (plural- plasmodesmata) – fine protoplasmic thread or strands that
penetrates the cell wall through the pit.
Plasmolysis – shrinking away of protoplasm from the cell wall due to coming out of
water from cell.
Polar transport – directed movement within plants of compounds (usually hormones),
predominantly in one direction
P-protein – proteinaceous contents of phloem sieve tube member; sometimes called
slime.
Process – natural continuing sequence of events. i.e. Photosynthesis
Quantasomes – photosynthetic unit composed of 300 chlorophyll molecules involved in
processing a quantum of light
Respiration – oxidative breakdown of organic compounds accompanied by the release
of energy.
Ribosome- complex aggregates of RNA and protein and site of protein synthesis.
Rough Endoplasmic Reticulum – endoplasmic reticulum with associated ribosome on
its surface.
Senescence – deterioration that ends the functional life of an organ or organism
Sieve tubes – long, vertical arrays of sieve tube members joined end to end.
Sink strength – a measure of capacity of the sink to accumulate metabolites.
Smooth Endoplasmic Reticulum – major site for lipid biosynthesis and membrane
formation and devoid of ribosome.
Solute – a dissolved substance
Solution – a homogenous mixture of two or more substances.
Solvent – a substance usually a liquid having the property of dissolving other
substances.
Suberin – a waxy material found in the cell wall of cork cells and endodermis.

FRE - TREE PHYSIOLOGY 4


Symport – linked movement of two entities across a protoplasmic membrane at the site
in the same direction (i.e. sucrose – proton translocators).
Synergism – presence or excess of one ion in the solution promotes the absorption of
another.
Tensile strength – maximum tension that an uninterrupted column of any material can
withstand without breaking.
Tonoplast – cytoplasmic membrane bordering the vacuole or vacuolar membrane.
Tracheid – an elongated tapering xylem cells with liquified pitted walls adapted for
conduction and support.
Translocation – movement of materials from cell to cell, tissue to tissue and organ to
organ within the plant body.
Transpiration – water loss in vapor form through the aerial part of the plant.
Tropism – an orientation of the direction of growth in an organ of the plant guided by an
external stimulus such as light or gravity.
Turgor pressure – outwardly directed pressure exerted by the contents of the cell on
the cell wall.
Uniport – movement of one entity across a protoplasmic membrane at one site.
Vacuole – depository organ of the cell.
Vernalization – acceleration of the ability to flower by cold treatment (does not induce
flowering but merely prepares the plant to flower)
Vesicles – tiny vacuole-like structures
Vessel – long tracheary elements made up of individual units known as vessel
members, arranged end to end in longitudinal series.

FRE - TREE PHYSIOLOGY 5


Growth - end result of the interaction between numerous physiological processes.

Hereditary
Potentialities Environment

TREE GROWTH

Photosynthesis
Respiration
Absorption
Translocation
Transpiration
etc.

Kleb’s Concept - emphasizes that the only way in which heredity and environment
can affect growth of an organism is by affecting its internal
processes and conditions.

The physiological processes of a plant constitute the machinery through which


heredity and environment operate to control growth.

The ultimate shape, size, form and degree of complexion


of a plant are the result of the interaction between its genetic composition and the
environment in which it grows.

FRE - TREE PHYSIOLOGY 6


CELL
- basic unit of life

Parts of a plant cell:


1. cell wall – non-living component which provides rigidity and maintains cell shape and
protects the protoplast
a) primary wall – thin-walled and consists mainly of cellulose, hemicellulose and
pectic substances.
b) secondary wall – thicker and more rigid that primary wall; laid down on the inside
of the primary wall which contains 45% cellulose, less hemicelluloses and little
pectic substances.
c) middle lamella – holds together the primary wall of adjacent cells; principal
constituent is pectic substance
2. cytoplasmic membrane or plasmalemma
3. protoplasm – living component
 cell organelles – discrete, membrane limited compartments.
a) Nucleus – information center of the cell which contains DNA and RNA.
 Nucleulos – site of ribosome synthesis
 Nuclear envelope
 Nuclear pores – channel for the export of RNA from the nucleus to the
cytoplasm.
b) Mitochondrion – site of cellular respiration
c) Plastid
 Chloroplast –contains green pigment
 Leucoplast – colorless plastid
 Chromoplast – colored plastid (orange, yellow, red, brown)
d) endoplasmic reticulum (ER) – single, highly convoluted membrane network
continuous with the outer membrane of the nucleus.
 rough ER – with ribosome
 smooth ER – major site of lipid biosynthesis and membrane formation;
serves as channel of communication and transport between the nucleus
and cytoplasm
e) Golgi bodies – stack of flattened, membranous sacs called cisternae;
assemble and process glycoproteins; synthesize complex polysaccharides
f) vacuole – depository organ of the cell.
g) ribosomes – complex aggregates of protein and RNA; site of protein
synthesis.
h) Microbodies
 peroxisomes – contain large amount of catalase used to remove harmful
metabolic by-products.
 Glyoxysomes – contain enzymes of glyoxylate cycle used to convert fatty
acids to glucose.
 spherosome

FRE - TREE PHYSIOLOGY 7


Types of cell according to cell wall thickness:
1) parenchyma – isodiametric cells surrounded only by primary wall; found throughout
the plant body.
2) Collenchyma – cells with thickened primary wall; support for young tissue.
3) Sclerenchyma – cells with thick secondary walls that are heavily lignified; for strength
and support.

Vascular tissues:
1) xylem – water conducting tissue
a. xylem fibers
b. xylem parenchyma
c. tracheary elements
 tracheid
 vessel elements
2) phloem – food conducting tissue
a. sieve tube members
b. companion cells
c. phloem parenchyma

CARBOHYDRATES
 Carbohydrates are a group of organic compounds containing the elements carbon,
hydrogen and oxygen in the ratio of 1:2:1 or approaching the empirical formula of
(CH2O) n.
 Carbohydrates that participate in metabolic reaction are generally present as
monosaccharide such as glucose, whereas the storage and structural carbohydrates
are found as polysaccharides.

Importance of Carbohydrates:
1. It represents an important food reserve or as a means for storage of the energy
trapped from the sun through photosynthesis (food storage function).
2. Forms the supporting tissue of the plant enabling it to achieve erect growth
(structural function).
3. Provides the carbon skeleton for most if not all of the organic compounds that make
up a plant.

Classification:
a) Monosaccharide – carbohydrate monomer
1. Triose sugars – 3 carbon sugar; the simplest monosaccharide; serve as
important intermediates in photosynthesis and respiration
2. Tetrose – 4 carbon sugar (erythrose); respiratory intermediates; use in the
synthesis of phenylalamine, tyrosine, lignin and anthocyanin pigment; exists in
phosphorylated form.

FRE - TREE PHYSIOLOGY 8


3. Pentose – 5 carbon sugar (ribose, deoxyribose)
 Divided into aldopentoses (ribose, xylose, deoxyribose) and ketopentose
(ribulose, xylulose)
 Form part of polymers in cell walls, arabans and xylans
 Ribose and deoxyribose occur as structural parts of RNA and DNA,
respectively
 Part of nucleotide coenzymes such as NAD+NADP, FAD, CoA
 Important intermediates in photosynthesis and respiration.
Hexoses – 6 carbon sugar (glucose, galactose, mannose, and fructose)
 D-glucose is also known as dextrose, corn sugar and grape sugar
 Considered as starting materials and end products of respiration and
photosynthesis, respectively.
 Involved in the synthesis of more complex carbohydrates
 Galactose is a constituent of agar; part of secondary walls of wood, straw and
seed.
4. Heptose – 7 carbon sugars which exists in phosphorylated forms (redoheptalose
7-phosphate); respiratory and photosynthetic intermediate
b) Oligosaccharide – 2-4 monosaccharides joined together
1. Disaccharide – two monosaccharides joined together
a) Sucrose – principal form of carbohydrate transported within the plants. A
condensation product of glucose and fructose joined together in  (12) link
b) Maltose – composed of two molecules of glucose joined together in an (1
4) link; product of starch hydrolysis.
c) Cellobiose – composed of two glucose molecules joined together by  (1
4) link; product of cellulose hydrolysis.
2. Trisaccharides – 3 monosaccharides joined together
a) Raffinose – galactose-glucose-fructose; found in storage organs and
consumed during germination; a non-reducing sugar.
b) Gentiose – 2 molecules of glucose+ fructose; a non-reducing sugar.
3. Tetrasaccharides – 4 monosaccharides joined together (Stachyose).
c) Polysaccharides – large, high molecular weight polymers of monosaccharides.
Storage polysaccharides
 Starch
 Fructosans –inulin –abundant in monocots
levan – abundant in grasses

Structural polysaccharides:
 Cellulose
 Hemicellulose; mannans and xylans
 Pectin
 Gums and muscilages

FRE - TREE PHYSIOLOGY 9


Starch
 a complex mixture of straight and branch chain glucans (glucose molecules)
 Components: a) amylose b) amylopectin
Cellulose - is a fundamental component of cell wall.
Pectic Substances – polymers containing primarily galacturonic acid.

Types:
a) Pectic acid – unbranched molecules consisting of about 100 D-galacturonic acid
residues bound together by (1-4) linkages
b) Pectins – closely resemble pectic acid, except that many of the carboxyl groups are
methylated.
c) Protopectins – insoluble pectic substances
Hemicellulose - cellulose-like and can be extracted from cell walls with a strong base.
Gums and muscilages - hydrophillic (not hygroscopic) polymers of rhamnose arabinose,
xylose, galactose, glucuronic acid

Importance of gums and muscilages:


 Have large water-holding capacity hence may function in seed germination by
absorbing water during imbibition.
 Function as water-holding molecules in plants adapted to arid conditions
Fructosans – reserve food of compositae and consist of 35 fructose units joined by  (2-
1) link.

LIPIDS

 Lipids (Gp. Lipos; fat) are a rather mixed group of compounds that have the
characteristic of being hydrophobic.
 They are soluble in organic solvents such as acetone, chloroform and others, but
sparingly in water.

Group of Lipids:
1) neutral lipids
2) phospholipids – component of cellular membrane
3) glycolipids – component of chloroplast membrane
4) terpenoids
5) waxes

Importance of lipids:
 Neutral lipids including fats and oils function primarily as storage molecules.
 Phospholipids and glycolipids function as structural lipids because they are
components of membranes.

FRE - TREE PHYSIOLOGY 1


 The terpenoids which include the sterols and a variety of secondary products such
as rubber, chicle, and the essential oils are important commercially.
 Waxes are protective lipids which prevent excessive loss of water from leaves and
stems of plants. Plant waxes function largely for protection.

PROTEINS
Proteins - are large and complex molecules with high molecular weight and considered
as the ultimate translational product of RNA.

Functions of Protein
 Regulate cellular metabolism by acting as catalyst in various chemical reactions
taking place in living cell (regulatory function).
 Serve as structural components of membranes and cell walls. Most structural
proteins of plants have regulatory functions as well (structural function).
 Storage molecules especially in seeds i.e. zein (storage function)

Types/Classification of Protein:
1. Simple
a) Protamines - small, water-soluble proteins with high number of basic amino
acids, soluble in ammonia, uncoagulable by heat.
b) Albumins - common proteins of plants and animals that are soluble in water and
dilute salt; readily precipitated by saturated (NH4)2SO4 coagulable by heat (i.e. -
amylase)
c) Histones - large, water soluble proteins with high number of basic amino acids;
arginine and lysine rich protein.
d) Globulins - common proteins of plant and animals that are insoluble in water but
soluble in dilute salt solution; precipitated by ½ saturated (NH4)2 SO4 solutions
(i.e. -amylase).
e) Glutelins - plant proteins that are insoluble in water and dilute salt solution (i.e.
Oryzenin from rice).
f) Prolamins - plant proteins found in seeds that are soluble in 70-80% alcohol
solution; insoluble in water and salt solution (i.e. zein from corn).
2. Conjugated protein – protein with prosthetic grp.
a) Mucoprotein or glycoprotein – protein with carbohydrate moiety; plays an
important role in cell recognition system and cell defense against fungi and
microorganisms.
b) Lipoprotein - with lipid group
c) Nucleoprotein - with nucleic acid group; involved in transmission of information
during nuclear division and protein synthesis.
d) Chromoprotein - with chromatophore (pigment) grp.
e) Metalloprotein - with metal group (i.e. calcium, potassium, etc.)

FRE - TREE PHYSIOLOGY 1


Structure of Proteins
1. Primary - number, kind and sequence of amino acids linked together by peptide
bond
2. Secondary - coiling of polypeptides in the form of a helix stabilized by hydrogen
bonding
3. Tertiary - further folding or coiling of the peptide chains into a sphere-like shape or
three dimensional structure stabilized by weak and strong forces. The hydrogen
bonds between amino and carboxyl groups and other kinds of bonds cross-linked
one chain to another.
4. Quarternary - aggregate of polypeptides. Each individual chain has its own primary,
secondary and tertiary structure.

Groups/Classes of enzymes based on catalytic functions:


1. Oxidoreductases - catalyze oxidation-reduction reactions
2. Transferases - transfer of groups such as amino, phosphate, etc.
3. Hydrolases - catalyze hydrolysis by addition of water i.e. cellulases
4. Lyases - catalyze removal or addition of groups not involving water
5. Ligases - condensing enzymes

Factors Affecting Enzyme Activity


1. Substrate concentration
2. Enzyme concentration
3. Temperature - 0oC–rate of enzyme-catalyzed reaction is 0; reaction rate increases
2.5 times for every 10oC increase in temperature up to 25oC. As 30oC is
approached, factors leading to denaturation of enzymes become apparent. Further
increases in T increase the thermal activity causing the hydrogen bonds to be
stretched and finally broken.
4. Hydrogen concentration (pH) - Different enzymes have different pH. Change of pH
can cause denaturation of enzymes.
5. Inhibitors - interaction of enzymes with substances other than the normal substrates
lead to an alteration of structure essential in catalytic activity resulting to loss in
catalytic efficiency or complete inactivation of the enzyme.

Type of inhibition:
1. Competitive inhibition - structural analogs compete with the normal substrate for
active sites on the enzymes
2. Non-competitive inhibition - non-competitive inhibitors do not compete with the active
sites on the enzyme, but reacts either with parts of the enzyme not involve in
catalytic activity or with the enzyme-substrate complex.

FRE - TREE PHYSIOLOGY 1


PHOTOSYNTHESIS
1. Photosynthesis – absorption of light energy and its conversion to useful chemical
energy.
2. Overall equation:
light
6CO2 + 12 H2O C6H12O6 + 6H2O + 6O2
chlorophyll

3. Only 5.8% of the total light energy which strikes the leaves is converted to chemical
free energy of organic compounds.
4. Quantum requirement: 4 to 8 quanta of light is needed to evolve one volume of
oxygen or reduce one volume of CO2
5. Portion of the spectrum between 400nm and 700nm is called photosynthetically
active radiation (PAR).
6. Blue and red light are absorbed by the chlorophyll.
7. Photosynthetic pigments
a. chlorophyll – a magnesium porphyrin with a long hydrocarbon phytol tail.
b. Carotenoids – linear tetraterpenoids which are usually red, orange, yellow or
brown in color; includes carotenes and xantophyll
b.1. Function
a. prevent photo-oxidation of chlorophyll
b. serves as accessory pigment for gathering light and transferring energy
to chlorophyll.
Chlorophyll a – universally present in all photosynthetic plants and serves as the
primary energy trap during photosynthesis

Properties of Pigment
Properties Chlorophyll a Chlorophyll b Carotenoids
Structure of side Methyl group Aldehyde group ---
chain at ring II (CH3) (HC=O)

Color Blue-green Yellow green Yellow, red, brown,


orange

Solubility Insoluble in water, Soluble in polar Soluble in benzene,


soluble in acetone, solvent petroleum ether,
ether ethyl alcohol,
chloroform,
acetone, insoluble
in water

Fluorescence Deep blood red Brownish ----

Absorption peak 430nm (blue 430-490nm 500-570nm


violet) (violet to blue) (green to yellow)
640nm (orange
red)

FRE - TREE PHYSIOLOGY 1


8. Phases/stages of photosynthesis
a. light phase
b. dark phase
A. Light phase (photochemical phase)
a. Basic features
1. photolysis of water (Hill reaction)
2. generation of reduced pyrimidine nucleotide (NADH, NADPH)
 Overall reaction:
Chlo. + light + H2O + NADP + ADP + Pi ½O2 + NADPH + H+ +
ATP + Chlo
 Photophosphorylation (photosynthetic electron transport system) –
light – driven production of ATP by chloroplast

Importance:
1. Produces ATP and reductants (NADPH) for the reduction of CO 2 to
carbohydrate level.
2. Provides continuous supply of ATP to support a variety of other metabolic
activities in the chloroplast.

Types:
1. Cyclic photophosphorylation – involves production of ATP; electrons
cycled back to P700; associated with photosystem I (PSI)
2. Non-cyclic – electrons from water are transported in a unidirectional
manner to ferredoxin (Fd) and reduce NADP; involves production of ATP
and reduction of NADP.
3. Pseudocyclic – electrons are transferred from Fd back to oxygen reducing
it to water and in the process ATP may be produced.
 Illumination of chloroplast with long wavelength stops non-cyclic
photophosphorylation, thus, NADPH is no longer formed and CO2
assimilation is retarded.
 Emerson Enhancement Effect – states that the effect of two
superimposed beams of light on the rate of photosynthesis exceeds
the sum effect of both beams used separately.
 Red drop – decrease in quantum yield at wavelength greater than 680
nm, an area of the spectrum occupied by the red wavelength.

Two types of Photosystem:


1) Photosystem I (PS I)
 long wavelength system
 chlorophyll A 683
 carotenoids
 P 700 as reaction center (energy sink)
 more chlorophyll A than other pigments
 the reaction center is an energy sink; longest wavelength and lowest
energy absorbing chlorophyll in the complex
2) Photosystem II (PS II)
 short wavelength system

FRE - TREE PHYSIOLOGY 1


 chlorophyll A 673
 chlorophyll B
 P 680 as reaction center
 More chlorophyll B
 Accessory pigment
B. Dark phase (Carbon reductions)
1. Reductive Pentose Cycle (Calvin-Benson Cycle or C3 cycle)
Net assimilation of 3CO2 produces 1 net mole of glyceraldehyde – 3 –
phosphate (GAP). During the process, 3 moles ribulose -1, 5-diphosphate
(RuBP) are consumed along with 9 ATPs and 6 NADPH. The cycle is
completed with the regeneration of 3 moles of RuBP
Balance sheet: 3CO2 + 9ATP + 6 NADPH 1GAP + 9ADP + 6NADP
ATP and NADPH come from the photochemical phase of photosynthesis
2. Hatch – Slack Pathway (C4 cycle)
CO2 is fixed by phosphoenolpyruvate (PEP) to form oxaloacetate (OAA)
which is immediately reduced to malic acid through reductive amination or
transamination in the mesophyll cell. Malate diffuses into the bundle sheath
cells where it is decarboxylated to form pyruvate. CO 2 is refixed by the C3
cycle while the pyruvate diffuses into the mesophyll cells where it is again
converted back to PEP.
2. Crassulacean acid metabolism (CAM)
Metabolic pathway, is similar to C 4 cycle except that the carboxylation and
decarboxylation reactions are separated by time

Features of CAM plants:


 generally succulent
 nocturnal stomatal opening
 nighttime transpiration
 nighttime CO2 uptake
 increase in acidity of cell sap during the night
 starch depletion during the night

Factors Affecting Photosynthesis (PS):


1. Plant factors
a. Age of leaves – low photosynthesis in young leaves; PS increase with light
intensity and physiological age of leaves up to full expansion then decreases as
the leaf ages.
b. Crown shape – tall conical form will intercept more light than with foliage oriented
in horizontal plane.

c. Structure and arrangement of leaves


 leaves near the apex have more chlorophyll per unit area of leaf surface than
the basal leaves.

FRE - TREE PHYSIOLOGY 1


 In trees with dense foliage, the most shaded oldest foliage can be below the
compensation point for most of the time and when the rate of respiration is
higher, can constitute a drain in the carbohydrate reserve.
 Exposed sun leaves are smaller, thicker and more leathery compared to
shade leaves. Shade leaves have high photosynthesis under conditions of
low light intensity, and light saturation is reached at lower light intensity. They
are also darker green than light – adapted ones and absorb light more
efficiently.
d. Carbohydrate accumulation
Accumulation of carbohydrates reduces photosynthesis. Girdling which
prevents the transport of carbohydrates to the lower parts of tree, reduces
photosynthesis by 100%
2. Environmental Factors
a. Light - In a hardwood forest, the light that reaches the understorey is low in blue,
high in yellow and green, similar to sunlight in orange, red and exceptionally high
in far red. Trees accumulate more photosynthate if expose to a long day than at
short day. Photosynthesis of shade tolerant plants is always regulated by light
intensity, while photosynthesis of sun plants is affected by light only on overcast
days.
b. Temperature – high temperature causes stomatal closure due to water stress.
Photosynthesis of both woody and herbaceous plants increases from 6 to 25°C
but only if neither CO2 nor light intensity is limiting.
c. Carbon dioxide concentration – photosynthesis is commonly limited by CO 2
availability within the dense foliage; increase in CO 2 level increases
photosynthesis by 2 to 3 times.
 At low CO2 concentration, photosynthesis decreases until it is equal to the
rate of photorespiration.
 The point where CO2 uptake and output are equal is known as CO 2
compensation point
 Large amount of CO2 is evolved from the forest soil especially after the
summer rain which is ecologically important to the understorey plants.
d. Water
1. Low water supply restricts Hill reactions, electron transport system and NADP
reduction.
2. Low water content causes stomatal closure by restricting CO2 uptake.
3. Water stress alters hydration of enzymes involved in photosynthesis.
e. Air pollutants/chemical sprays
1. Damage the stomatal apparatus restricting the entrance of CO2 into the leaf.
2. At high concentration, certain gases react with cellular constituent and cause
the death of the tissue.
3. Many fungicidal and insecticidal sprays reduced photosynthetic efficiency.
The reduction is greater when only the lower leaf surface is sprayed
compared to the upper surface. Photosynthesis is reduced due to the
clogging of stomates and interference with inward diffusion of CO2.
f. Diseases/Pest Infestation

FRE - TREE PHYSIOLOGY 1


1. Leaf diseases/pests often reduce photosynthesis by reducing the
photosynthetic area and the efficiency of the remaining photosynthetic
tissues.
2. Root diseases which interfere with water and mineral chloride, manganese
and phosphorous will decrease photosynthesis.

Mineral Nutrients
 Magnesium is a component of chlorophyll
 Iron (Fe) and Copper (Cu) are cofactors in the electron flow
(photophosporylation)
 Chlorine (Cl) and Manganese (Mn) are involved in Hill reaction.
 Phosphorous (P) is needed in ATP production
 Chlorotic leaves have lower concentration of chlorophyll.

PHOTORESPIRATION
 A light-dependent oxidation of organic compound (glycolate) to CO 2 occurring in
chlorophyllous tissues.
 Represents a potentially serious loss of energy from a plant.
 Over 30% of the carbon reduced during photosynthesis is reoxidized to CO 2 during
Pr. Thus, the process contributes to the reduction in dry matter accumulation and
ultimately yield.
 For every mole of CO2 lost in Pr, 6 ATPs and 3 NADPH are consumed.
 One ATP and 2 NADPH are still used for reassimilation of NH3
 An increase in O2 concentration in the external environment will increase the amount
of glycolic acid synthesis in the illuminated leaves.
 Organelles involved:
 Peroxisomes
 Mitochondria
 Chloroplast
 To sum up, 2 molecules of phosphoglycolic acid produced by oxygenase reactions
are converted to one molecule each of O2 and PGA.
Peroxisomes in C4 plants are present in the bundle sheath cells while in C3, numerous
peroxisomes occur in the mesophyll cells

Reasons why Photorespiration occurs:


1. Oxygenase reaction is inevitable Energy-consuming photosynthetic carbon
oxidation (PCO) cycle is the most effective way of salvaging 75% of the carbon and
putting it back into the profitable stream of the reductive pentose cycle (RPC)
2. Serves as a mechanism for dissipation of energy without which the photosynthesis
apparatus would be totally unstable.
3. Consuming O2 during Pr helps to prevent the build up of hyperoxidative state in
chloroplast of C3 plants.

FRE - TREE PHYSIOLOGY 1


RESPIRATION
 defined as the oxidative breakdown of organic compounds accompanied by the
release of energy .
 usually starts with glucose or a hexose sugar.
C6H12O6+6O2 6 CO2 + 6H2O + energy (ATP)
 Oxygen does not directly react with glucose. Instead, water molecules are added to
intermediate products of glucose degradation (one molecule of H 2O for each carbon
atom in the glucose molecule) and hydrogen atoms in the intermediate products are
transferred to O2 which is reduced to water.
 The overall summary reaction based on the biochemical considerations is
C6H12O6+6H2O + 6O2 6 CO2 + 12H2O + E
 Energy released per mole of hexose oxidized:
2,900 kilojoules or 686,000 calories
 The energy released is stored as chemical energy
1. Part of the energy is used to maintain the structure of the protoplasm –
Maintenance respiration
2. A part is used in synthetic processes – growth respiration
3. Some is released as mechanical energy such as that exhibited in protoplasmic
streaming, transport of materials.
4. Some occurs as electrical energy
5. A part is dissipated as heat
6. Part of the energy is stored as chemical energy of high energy phosphate bonds
which can be used to drive energy-requiring reactions.
 Minimal rate of respiration is essential for survival and a higher rate is necessary for
growth, however, the rate often rises for above the essential level which results in the
unproductive consumption of food which might have been used to produce new
tissues or accumulated in storage organs.
 A large fraction of the food produced by trees is used in respiration of non-
photosynthetic tissues.
 Productivity of forest trees may be increased more by treatments which reduce
respiration such as removal of parasitic branches than by treatments to increase
photosynthesis
 Respiration occurs in all living tissues all the time while photosynthesis occurs only in
the light and chiefly in the leaves. Therefore, the rate of photosynthesis must be
several times as rapid as the rate of respiration per unit of tissue, if any net increase
in dry weight is to occur.

Factors Affecting Respiration:


1. Age of tissue – the younger and the more active the tissue, the greater the oxygen
consumption per unit of tissue

FRE - TREE PHYSIOLOGY 1


2. Oxygen – increase in oxygen aerobic respiration
 At very low oxygen, both aerobic and anaerobic respiration occurs.
 As O2 concentration approaches O, Co2 released is due to fermentation.
 High level of O2 inhibits glycolysis
3. Carbon dioxide – increases CO2 level represses respiration
4. Light – stimulates respiration
 Exposure to light increases glycolylic acid formation in chloroplast which is
eventually oxidized to O2
5. Soil atmosphere
 Respiration of roots and soil organisms tends to deplete the oxygen and
increases the CO2 concentration
 Flooded or compact soil greatly reduces gas exchange
 Filing in with soil over an established root system injures or kills trees by cutting
off oxygen supply to the roots
6. Soil moisture
 excess soil moisture interferes with oxygen availability to the roots
 Leaf respiration increases in soil that is drying down from field capacity toward
wilting point.
7. Disease infections, injuries, wounding and mechanical disturbance
 Injury to respiring tissues increases respiration due to the increased in oxygen
and substrate availability
 Infection by diseases increases respiration due to more rapid utilization of ATP
for growth processes. Pentose phosphate pathway is more active in diseased
plants
 Handling, rubbing, stroking or bending increases respiration by as much as 20
to 180%
 Increased in respiration after injury to tissues is due to the initiation of
meristematic activity in the wounded area resulting to the development of wood
callus.
8. Chemical sprays and inhibitors
 Cyanide, axide, flouride, malonate, iodoacetate, dietheyldithiocarbornate and
carbon monoxide reduce respiration
 Hydrocarbon oils used as insecticide affect respiration of twigs and leaves in a
complex manner. The effect varies with age and conditions of tissues and type
of oil used
 Biocides or chemical sprays affect the respiration of younger tissue more than
the older tissue
 In general, respiration is a good index of physiological activity and measurement
of respiration serves as an indicator of the relative toxicity of various chemical
sprays to the tree.
9. Starvation – starved plants have lower respiration due to the low sugar or starch
reserve. Severe starvation results to protein and lipid breakdown.

FRE - TREE PHYSIOLOGY 1


Water
Importance of Water
 Solvent for various biological compounds, substances and mineral nutrients
 Maintain the structure of polymeric compounds such as protein and amino acids,
nucleic acids
 Substrate for biochemical reactions
 Imparts turgidity to cell
 Constituent of protoplasm
 Important in the elongation phase of cell growth
 Responsible for a variety of plant movement

Physical Properties
1. High heat of vaporization
a. Provides maximum cooling per unit water lost. Each gram of water lost by a leaf
requires 2258 joules hence, this much heat will be lost for each gram of water
transpired.
b. Minimize the lost of water.
2. High heat of fusion – heat required to change a solid to a liquid is high for water (ice)
compared to other substances

3. Extremely high surface tension- the tendency of the surface to contract and resemble
an elastic membrane is very high for water. Water droplets tend to form spheres, a
condition resulting in less energy per unit of surface relative to volume.
The energy associated with the surface is surface tension. The greater the surface
tension, the greater the rise of water in the capillary.
4. High specific heat (great heat capacity) – water has the highest specific heat among
the common liquids.

Importance:
a. Great heat capacity of water causes it to be a temperature buffer which protects
living things against violent changes in temperature
b. Water has a better storage capacity for heat than air. Living organism containing
large quantities of water tend to be protected from significant changes in
temperature due to small changes in heat load.
5. Viscosity – resistance to flow is higher in water than in most liquids due to hydrogen
bonding. Viscosity decreases with increasing temperatures.
6. Volume and density
a. Water is more dense at 4 C and less dense above and below 4 C. As
temperature increases or decreases from 4 C, the volume occupied by 1 g of
water increases.
b. Water is less dense as a solid than as a liquid.
c. Water expands when it freezes while all other substances contract upon cooling
and freezing

FRE - TREE PHYSIOLOGY 2


d. Water molecules in ice have an open structure and when ice melts, there is
partial collapse of its lattice, thus water molecules occupy the spaces that are
void or empty
7. Scatters short-wavelength light (blue) and transmit the visible long wavelength (red)
 Absorbs long-wave heat radiation (infra-red) while ultra violet light will penetrate
substantially.
9. Universal solvent – most substance of biological significance are extremely soluble in
water. It is also a carrier of mineral elements.
10. Water is liquid at the temperature prevailing over most of the earth surfaces.

WATER POTENTIAL
1. Principle involved in water transport: Water moves from a region of higher free
energy (high water potential) to a region of low free energy (low water potential) or
simply stated “Water runs downhill”.
Free energy – energy available for work
2. Difference in water potential between the source and sink indicates the direction of
water flow.
3. Free energy of water in the sink must be less than the free energy in the source for
spontaneous water flow.
4. Basic unit for water potential is bar
a) 1 bar = 106 dynes/cm2; b) 0.987 bar = 1 atm
5. Two most important components of water potential
a) Osmotic potential (solute potential)
 The decrease in water potential brought about by substances either in ionic
or non-ionic form in the solution is referred to as solute potential
 Solute potential takes the place of osmotic pressure expressed in bars with
positive sign while osmotic potential has negative sign. When solute potential
decrease, the solution becomes more negative
Pressure Potential () or turgor pressure – In an osmotic system, water will
flow from higher concentration to a lower concentration on a water
potential gradient.
 The pressure normally exerted in the contents of a plant cell is mainly due to
the tensile strength and elastic stretch of the cell wall. The hydrostatic
pressure that develops in the vacuole as a result of inwardly directed wall
pressure is called turgor pressure.
 Turgor pressure is the outwardly directed pressure exerted by the contents of
the cell on the cell wall. It is equal in magnitude and opposite in direction to
the pressure of the cell wall.

FRE - TREE PHYSIOLOGY 2


 Pressure potential in plant cell is positive. i. e. leaf cells of herbaceous plants
have turgor pressure of +3 to +5 bars in the afternoon of a warm sunny day to
about +15 bars or more during the night hours.
 Pressure potential may become zero or negative in some cases. Plasmolyze
cells have zero or almost zero turgor pressure. Negative hydrostatic
pressure exists in the water in the xylem during periods of rapid transpiration.

Importance of turgor pressure:


 Ensures maintenance of normal rigidity in plant cell.
 Aids in cell enlargement
b) Osmotic pressure is basically a quantitative index of the lowering of water
potential in a solution due to the presence of solute. Addition of solute to water
will reduce the free energy of water and then, reduce the water potential.
Osmotic potential can be estimated numerically from osmotic pressure by
changing the sign from + to -.
 Osmotic potential depends on the number of solute particles (molecules or
ions) in a solution rather than the kind of charges.
 Variations in solute potential occur during day and night. At daylight, leaf
water content falls and the concentration of solutes in the leaf cell increases,
hence  becomes more negative. During night time, leaf water content
rises and concentration of solutes in leaf cells decreases, so  - becomes
less negative.

Factors affecting the Osmotic potential of solution


1) Concentration – an increase in the concentration of a solution results in its osmotic
potential becoming more negative. If a solute is a non-electrolyte and its molecule
does not acquire water of hydration, the osmotic potential is almost strictly proportion
to the molal concentration that is to the proportion of solute to solvent molecule.
2) Ionization of the solute molecules – Osmotic potential is a colligative property, that is,
one which depends in the number of solute particles regardless of kind, in proportion
to the number of solute particles in a solution. If the solutes dissociate, either in part
or entirely, the osmotic potential will be more negative. If an electrolyte such as NaCl
is present at 1 molal concentration completely in the form of free ions, the theoretical
osmotic potential of the solution would be –45.5 (2X-22.7). The measured  is –
43.2 bars which indicate that a condition of maximum mobility is closely approached
in such a solution.
3) Hydration of the solute molecules - micelles of hydrophobic solutes is highly
hydrated. Water molecules adhere to many kinds of solutes molecules and ions.
Water associated with the particles of a solute is called water of hydration. Water
molecules which are bound to solute particle as water of hydration are no longer
effective as part of the solvent. In effect, a solution containing hydrated solute
particles is more concentrated than its molality would indicate and its  is
correspondingly more negative.
4) Temperature – The  of a solution becomes more negative with an increase in
temperature. For an ideal solution, the osmotic potential is proportional to the
absolute temperature.

FRE - TREE PHYSIOLOGY 2


Water Uptake and Transport
Processes by which water absorption is accomplished:
a) imbibition – process of adsorption of water to nearly dry surfaces such as seeds or
wood accompanied by the release of heat.

Conditions necessary for imbibition:


 Water potential gradient must exist between the imbibant and the liquid imbibed.
 A certain affinity must exist between components of the imbibant and the imbibed
substance

Factors Affecting Imbibition:


 Temperature – increase in temperature increases the rate of imbibition
 Osmotic potential of the substance to be imbibed – addition of solute to pure
water causes a more negative water potential. A decrease in water potential
gradient will decrease the rate of which water is imbibed.
b) Osmosis – a special type of diffusion which involves movement of water through a
differentially permeable or semi-permeable membrane from the region of high
concentration to the region of lower concentration. It involves the movement of a
solvent not a solution across the membrane. In living organisms, water is the only
important solvent which moves by osmosis.
c) Diffusion – movement of water from region of higher concentration to lower
concentration.

Factors Affecting Absorption of Water:


 Temperature of the soil – low soil temperature reduces water absorption because
water is less viscous at low temperature, a factor which reduces its mobility, and
protoplasm is less permeable at low temperature.
 Concentration of the soil solution – increase in salt concentration in the soil solution
decreases water absorption. If the osmotic potential of the soil solution is more
negative than the cell sap of the root cells, water will be drawn out of the plant
instead of being absorbed.
 Aeration of soil – root growth and metabolism are retarded under conditions of low
oxygen. Accumulation of CO2 increases the viscosity of protoplasm and decreases
its permeability, thus retarding water absorption.
 Availability of soil water – not all of the water in the soil is available to the plant. As
the soil in the immediate viscosity of the root system is depleted of its water supply,
absorption of water by the plant becomes more and more difficult.

Mechanism involved in the translocation of water (ascent of sap):


 Root pressure – a pressure that develops in the tracheary elements of the xylem as
a result of the metabolic activities of the root. It is referred to as an active process,
however, the movements of water up to the stem as a result of root pressure is due
to osmotic mechanisms (passive), which are created as a result of the active
absorption of salts by the roots.

FRE - TREE PHYSIOLOGY 2


 Cohesion-tension theory – water molecules cohere to each other and adhere to the
side or wall of the xylem. Evaporation of water from the mesophyll cells created a
tension or negative pressure which is transmitted through the water column from the
leaves down to the roots forcing water from the soil to enter the roots.
 Vital theory – water is moved in response to the living function of the cells in the stem
and metabolic activities of the tracheary elements.

TRANSPIRATION
Transpiration – is the water loss in vapor form through the aerial parts of the plant

Type of Transpiration:
1) Stomatal – accounts for 90% or more water loss
2) Lenticular
3) Cuticular

Importance:
 Lowers the internal temperature of leaves during hot days, since evaporation of liquid
exerts cooling effect. On bright days, transpiration may lower the leaf temperature
by 20 o.
 Furnishes a pull in the leaves that is principally responsible for the rise of sap in the
plants.
 Rapid transpiration favors a corresponding rapid absorption of nutrient ions from the
soil.

Disadvantages:
 Causes excessive loss of water resulting in wilting and oftentimes death of the
plants. If transpiration outbalances absorption, wilting is inevitable.

Stomatal Movement
 The degree of stomatal opening is influenced by changes in the turgor pressure of
the guard cells and epidermal cells, especially those adjacent to the guard cells. In
general, an increase in the turgor pressure of the guard cells relative to epidermal
cells leads to the widening of stomatal aperture.
 The usual diurnal pattern in the movement of the stomates indicates that guard cells
are more turgid during daylight than during nighttime. Osmotic potential of guard
cells is usually more negative during daytime than nighttime.

Causes of decrease water potential:


1) Increase in the organic content of the cell sap due to synthesis of sugars and other
soluble compounds.
2) Conversion of insoluble carbohydrates such as starch into soluble ones.
3) Transfer of solutes from adjacent cell to guard cell.
4) Movement or influx of K+ into the guard cells upon exposure to light.

FRE - TREE PHYSIOLOGY 2


Factors Affecting Stomatal Movement
1. Light
a. Osmotic theory of stomatal movement in response to light:
Light photosynthesis reduction of CO2 concentration throughout the leaf
increase in pH of guard cells activation of enzymes which convert insoluble
polysaccharides into soluble carbohydrates increase in the negativity of the
osmotic potential of the guard cell sap increase in negativity of water potential
movement of water into the guard cells from adjoining cells  increase in
pressure potential of the guard cells opening of stomates.
b. Role of Potassium in stomatal movement:
Light potassium ions from nearby epidermal cells move into the guard cells
decrease osmotic potential of guard cell sap inward osmosis increase
turgidity of guard cells opening of stomates.
2. Water – development of internal water deficit may lead to stomatal closure due to the
reduction in the turgor or pressure potential of the guard cells.
3. Temperature – stomatal opening increases with increase in temperature up to 25 to
30oC.

Factors Affecting Transpiration:


a) environmental factors
 Solar radiation – principal effect of light is on the opening and closing of
stomates. In most species, stomates are closed in the absence of light, thus,
causing a complete cessation of stomatal transpiration. The second important
effect of solar radiation upon transpiration is its influence on leaf temperatures.
 Humidity – for any given temperature of an atmosphere, there is a maximum
vapor pressure which can prevail called the saturation vapor pressure. In
general, the greater the vapor pressure of the atmosphere, the lower the
rate of transpiration. Whenever, the stomates are substantially open, the
rate of diffusion of water vapor out of the leaf is largely conditioned by the
difference between the vapor pressure in the intercellular spaces and the
vapor pressure in the outside atmosphere.
 Temperature – effect is on the difference in vapor pressure between the
intercellular spaces and the outside atmosphere. In general, the higher the
temperature for a given vapor pressure gradient, the higher the rate of diffusion.
 Wind – exposure of leaves to wind results in a lower leaf temperature than that
would prevail under conditions of intense solar radiation. Overall effect of wind
on transpiration depends on its relative influence on the steepness of the vapor
pressure gradient through the stomates and on the leaf temperature.
 Availability of water – in general, if soil condition retards water absorption, the
rate of transpiration is also retarded.
b) Plant factors
 Total leaf surface
 Distribution and gross morphology of the root system

FRE - TREE PHYSIOLOGY 2


 Thickness of cutin layers
 Presence of epidermal hairs – slows down air movement
 Size, spacing, distribution and structural peculiarities of the stomates
 Ratio of internal exposed surface area of mesophyll cells to the external surface area of
the leaf.

Properties of colloidal systems:


1) Tyndall effect – a beam of light flashed through a glass of colloidal system can
easily be discerned or path of the light beam becomes visible.
2) Brownian movement – random motion of small particles in suspension caused by the
random bombardment of molecules of the dispersing agent.
3) Stable suspension – difficult to separate or separation of colloidal particles from
water is very difficult.
4) Adsorption – tendency of colloidal particles to adhere to the surface of certain solids
or liquids.
5) Electrical property – colloidal particles carry electrical charges.
6) Two phase system – consists of a solid phase, the colloidal unicells, suspended in a
liquid phase.

Ways of measuring transpiration:


1) Weighing method – rate of transpiration is measured as the weight of a plant in a
definite period of time.
2) Photometer method – based on the principle that the rate of water transpired is equal
to the rate at which water is absorbed by the same plant. The rate of transpiration
measured by the distance traveled by the bubble from photometer attached to the
plant divided by the time it takes the air bubble to travel such a distance.
Disadvantage: the method assumes that the rate of transpiration is equivalent to the
rate of water absorption which is not the case because usually, the rate of water loss
is much greater than the rate of water absorption.
3) Cobalt-chloride method – filter papers are impregnated with 3% solution of cobalt-
chloride and dried in the oven. The amount of water loss can be measured by
measuring the time needed for the color of the filter paper to change from blue to
pink.
4) Collection and weighing water vapor lost in transpiration – the method makes use of
a box. Water vapor is accumulated by calcium chloride deposited in a U-tube
connected to the box. A common source of air through a pump pushes the water
vapor along the tube towards the U-tube containing CaCl 2. Rate of transpiration is
computed as :

Final Weight of CaCl2 - Initial Weight of CaCl2


TIME

FRE - TREE PHYSIOLOGY 2


MINERAL NUTRITION
Sources of Mineral Nutrients
a) Air (atmosphere)
b) Water
c) Soil

Classification of mineral elements:


1. According to essentiality
a. essential elements – elements that satisfy the three criteria for essentiality
formulated by Arhon and Stout (1939)
 The element must be needed to complete the life cycle of the plant.
 Action of the element must be specific and cannot be replaced or substituted
by another element.
 The element must be directly involved in the nutrition of the plant or acting
directly inside the plant and not simply causing some other element to be
more readily available or simply antagonizing a toxic effect of another
element.
b. non-essential
2. According to mobility
 Highly mobile – elements which in an appreciable amount remain in the cell sap
and do not enter into an essential constituents of some metabolically
indispensable component. Deficiency symptoms appear in the older leaves and
becomes less marked in the upper (new) leaves i.e. K+, Mg++.
 Slightly mobile – deficiency symptoms appear with equal severity on both old and
new leaves i.e. N, S, P
 Immobile – deficiency symptoms appear only or mainly on the upper i.e. Ca++,
Bo, Fe, Mn, S.
3. According to plant needs
 macronutrient - elements needed in large amounts i.e. N, P, K, Ca, Mg, S
 micronutrient - needed in small amounts i.e. Zn, Cu, B, Mo, Mn, Fe

Role of Mineral Elements in Plants


A. General role or functions
 Building materials for protoplasm, cell walls, enzymes, metabolites and other
organic constituent of the plant. e.g. S – as constituent of proteins, P of ADP,
ATP, Mg of chlorophyll and C, H, O for carbohydrates etc.
 Regulator of membrane permeability e.g. K, Ca
 Maintenance of a desirable degree of hydration of cell colloids – monovalent
cations increases hydration divalent particularly the polyvalent cations decrease
it.

FRE - TREE PHYSIOLOGY 2


 Buffering action – PO4, HCO3 and CO3 may act as buffers and prevent marked
changes in H+ concentration.
 Catalytic roles – catalyze reactions by activating certain enzyme systems or as
constituent of certain enzymes.
 Maintenance of electrostatic neutrality. Within the cells, electrostatic neutrality
exists, positive and negative charges balancing each other. e.g. Na+, K+, Ca++
and Mn++ balance the negative charges of the organic acid and inorganic anions.

Specific Functions/Roles and Deficiency symptoms of Essential Elements

Elements Role(s) Deficiency Symptom(s)

N Constituent of amino acids, amides, Leaf changes color from green to pale
proteins, nucleic acids, nucleotides and green; in severe deficiency, the color
coenzymes, hexoseamines, purines, changes to lemon yellow and finally to
purimidines. Proteins consist of 18%N; yellow; stunted growth.
70% leaf N is in chloroplast.
P Component of sugar phosphates, nucleic Leaf color changes to slightly pale then
acids, nucleotides, co-enzymes, to dark green; red or purple coloration at
phospholipids, phytic acid, etc. Has key the base of the stem or at the leaf midrib.
role in reactions in which ATP is involved. Gradually, the leaf color changes to dull
grayish green. Sign of burning or
necrosis appears along the leaf margin.
K Required as a cofactor for 40 or more Leaf color changes to slightly ash-gray at
enzymes. Has a role in stomatal the tips, gradually spreading over the leaf
movements. Maintains electro-neutrality in margin. Necrosis lesions beginning at
plant cells. Promotes translocation of the older leaf tips gradually progressing
photosynthates from leaves. Involves in along the margins to the younger cells
osmotic and ionic balance. near the leaf base.
S Component of cysteine, cystine, Generalized chlorosis
methionine, and thus of proteins.
Constituent of lipoic acid, coenzyme A,
thiaminepyrophosphate, gluthionin, biotin,
adenosine-5’-phosphosulfate and 3’-
phosphoadenosine, 5’-phos-phosulfate and
other compounds. Volatile compounds
containing contri-bute to the odors of
onions, mustard, etc.
Ca A constituent of the middle lamella of cell Initial necrosis will appear on top of the
walls as Ca pectate. Required as a new leaves or the upper portion of the
cofactor by some enzymes involved in the outer edges and does not develop over
hydrolysis of ATP and phos-pholipids. Ca the entire leaf. Most severe in new
is a metal in the metalloenzyme, amylase. leaves; formation of black spot at the
Needed for germination and growth of center of the new plant because of tissue
pollen tube. Acts as counter-ion for necrosis.
negative charges. Maintains membrane
per-meability.

FRE - TREE PHYSIOLOGY 2


Elements Role(s) Deficiency Symptom(s)

Mg Required nonspecifically by large number Interveinal chlorosis; grayish areas


of enzymes involved in phosphate transfer. appearing on the leaf tip and then above
A constituent of the chlorophyll molecule. the outer edges. Gradually, color
Represents 2.7% of the molecular weight changes to yellow and then brown. Color
of Mg porphyrins. Mg in chlorophyll takes shape of con-centric circles.
represents only about 10% of the total Mg
in the leaf.
Fe Essential for chlorophyll synthesis. A Outer margin of the leaf becomes lemon
constituent of cytochromes, nonheme iron yellow. This slowly expands over the
proteins, which are involved in whole area of the leaf surface, producing
photosynthesis, N2 fixation, and a mosaic chlorosis with only the areas
respiratory-linked dehydrogenases. around the leaf veins remaining green.
Suddenly, color changes to yellow with
dirty brown coloration and necrotic spots;
in severe cases, leaves may turn white.
Mn Required for activity of some Symptoms are very similar to those of
dehydrogenases, decarboxylases, kinases, iron deficiency; mosaic pattern is less
oxidases, peroxidases, and nonspecifically clearly marked than with iron.
by other vibalent, cation-activated
enzymes. Required for photosynthetic
evolution of O 2. Manganin, a
manganoprotein was isolated in higher
plants. Function as a counter-ion for
unionic groups.
B Indirect evidence for involvement of B in Show corky patches on the leaf stalk,
carbohydrate transport, Borate forms newly formed leaf changes to brownish
complexes with certain carbohydrates, but and then black.
natural borate complexes in plants have
not been identified.
Cu An essential component of ascorbic acid Leaf color changes to dark green, leaves
oxidase, tyrosinase, lacease, monoamine small and growth is retarded and stunted.
oxidase, cytochrome oxidase, and
galactose oxidase. Component of
pastocyanin compund forming part of the
photosynthetic electron transport chain,
polyphenol oxidase. Deficiency in algae
causes decreased activity of cytochrome
photo-oxidase.
Zn Essential constituent of alcohol Main vein becomes silver white and a
dehydrogenase, glutamic dehydro-genase, marked strip along the middle of the leaf;
lactic dehydrogenase, car-bonic necrosis appears on the vein and
anhydrase, alkaline phos-phatase, expands to half of the leaf; short
carboxypeptidase B, and other enzymes. internode and smaller leaves.
Needed in the production of IAA (trytophan
pre-cursors of IAA).
Mo Involved in nitrogen fixation and nitrate Leaves pale green to yellow green in
reducing enzyme system. It is a color, growth is stunted; poor seed
constituent of nitrate reductase of fungi, production.
bacteria, and higher plants and of xanthine
oxidase, aldehyde oxidase from animal

FRE - TREE PHYSIOLOGY 2


sources and essential for N2 fixation.
Technique in the study of Plant Nutrition:
1. Water culture (hydroponics, soiless culture, tray agriculture, nutriculture) – the only
technique that has enabled scientists to determine which elements are essential for
higher plants. Also required for precise studies involving variation in plant, variation
in properties of nutrients and for many studies concerning growth requirement of
plants.
2. Solid growth medium – soil, sand, perlite and vermiculite
3. Aeroponics – confining the root system in humidity-rich and air-tight container and
supplying the culture solution to the roots in spray or mist form at regular interval.

MECHANISM OF NUTRIENT UPTAKE AND TRANSLOCATION


PASSIVE ABSORPTION – physical, non-metabolic absorption

Diffusible ions
Outer space =
External concentration
Where:
Diffusible ion (DI) – umole/g. fresh weight
External concentration (EC) – umole/ml
Outer space (OP) – ml/g. f.w.

Ways by which passive absorption is accomplished:

1. Ion exchange – ions are adsorbed on the surface of the cell walls and may exchange
with ions from the external solution.
2. Donnan equilibrium – takes into account the effect of fixed or indiffusible ions. The
membrane is permeable to some ions and the inner side of the membrane has
anions to which the membrane is impermeable (fixed anion).
3. Mass flow – ions move through the roots along with water. Increase transpiration
increases ion absorption.
4. Adsorption – accumulation of ions at interfaces. In many ways, roots act as a
sponge in the absorption of ions. Most substances in contact with water assume a
negative charge, including roots. The site of the charge.
5. Diffusion – the simplest process by which ions enter a cell or root. In diffusion, there
is net movement of ions from the region of higher concentration to a region of lower
concentration.

ACTIVE ABSORPTION – requires expenditure of energy. It is a process by which ions


are moved against an electrochemical potential gradient and, is therefore
dependent on a decrease in free energy in some metabolic processes.

Features of active absorption:


 linear with time, no equilibrium being reached

FRE - TREE PHYSIOLOGY 3


 selective with respect to various ions and groups of ions
 involves ions essentially non-exchangeable
 requires expenditure of energy

Mechanisms by which active absorption are accomplished:


a. Carrier concept
Sub units of the membrane called carriers bind the ions at the external face of
the membrane forming a carrier-ion complex. Carrier is also known as porter,
transporter and translocator.
b. Cytochrome pump (Lundegardh hypothesis)
c. Via Y-glutamyl peptide
d. Dual mechanism of ion absorption – involves mechanism I which operates at very low ion
concentrations (may be as low as 0.002 mM for K) and mechanism II, which operates at
higher concentration usually at the range of 1m 50 nM.
e. APOPLAST AND SYMPLAST CONCEPT (Craft-Boyer Hypothesis)
 The symplast consists of the total protoplasts of the cells, which connected from
cell to cell via small strands of cytoplasm called plasmodesmata that penetrates
the cell wall through the pit.
 Apoplast – consists of all space in the roots that roughly equate with free spaces
plus the tissue in the stele which gives free access to water, mainly the xylem
vessels.
Soil solution with ions  external portion of the apoplast outside the endodermis
 symplast  apoplast of the stele  xylem vessels
f. ATPase – dependent ion pump
g. Transport of ions via phosphatidic acid cycle

Factors Affecting Ion Absorption:


 Temperature
 Light
 Oxygen
 PH
 Nutrient status of the medium
 Growth stages of the plant
 Interaction among elements
a. antagonism
b. synergism

Forms of nitrogen available to the plants


1. Nitrate nitrogen
2. Ammonium nitrogen
3. Organic nitrogen
4. Molecular nitrogen – confine only to few groups of plants

FRE - TREE PHYSIOLOGY 3


Requirements for nitrogen-fixation
1. Continuous supply of ATP
2. Reduced substrate capable of donating protons and electrons
3. Nitrogenase enzyme
4. Magnesium
5. Anaerobic condition (for nitrogenase)

Plant hormones
1. Auxins
2. Gibberellins
3. Cytokinins
4. Abscissic acid
5. Ethylene – gaseous hormones

Causes of senescence
1. Competition for nutrients
2. Production of inhibitors
3. Change in enzyme pattern – more catabolic than anabolic reactions

Processes that take place during senescence


1. Decrease in starch, chlorophyll, protein and RNA in the leaves
2. Transport of hydrological products to younger regions of the plant
3. Synthesis of anthocyanins
4. Unmasking of the yellow and orange pigments
5. Increase respiration

Physiological changes during abscission


1. Loss of dry weight, chlorophyll, proteins, phosphate compounds and RNA from the
distal tissue
2. Documentation of the substances in the region proximal to the abscission Layer

Translocation - movement of materials from cell to cell, tissue to tissue and organ to
organ within the plants.

Pathways Used in Translocation

Xylem – upward movement of water, minerals and other substances


Phloem – downward movement of all kinds of solute and substances
Both phloem and xylem are continuous through the plants from the tips of the root to the
tips of the stem and the smallest vein in the leaves.

Salt movement outside the vascular system can occur in three ways:
1. Active transport in the cytoplasm
2. Diffusion
3. Mass movement in transpiration stream

FRE - TREE PHYSIOLOGY 3


Root grafts - connections between the root systems of adjacent trees by root grafts
occur frequently.
A group of trees may be connected so effectively through root grafts that if all but
one tree is cut the root system will survive on carbohydrates supplied by the
remaining tree.

Factors Affecting Translocation


1. Temperature – increase in temperature increase the rate of translocation
2. Aeration – supply of oxygen is necessary for translocation of food - poor soil
aerations hinder the movement of food to the roots. Internal factors – all growing
regions of the plant (stem and root tips, cambiums, fruits and seeds) compete for the
product of photosynthesis. The success in competition varies with age and stage of
growth.
i.e. shoot of conifers have an advantage over the side branches, young
leaves get food and minerals at the expense of roots and other vegetative structures.

ACCUMULATION
Accumulation of reserve food is very important for growth and survival of a tree.
This is particularly true for deciduous trees where the maintenance of life during summer
and resumption of growth during the wet season are entirely dependent on food
accumulated during the previous season.

Food economy of a tree can be expressed as follows:


Income = food manufactures by photosynthesis
Expenditures = food used in assimilation and respiration
Balance = food accumulated

Uses of Reserved Foods


1. Maintaining respiration and other processes at times when food is not supplied
directly from photosynthesis. All metabolic activities at night are dependent on food
accumulated during the day.
2. Use in growth and production of new branches, flowers and leaves.
3. Production of stump sprouts depends on the stored food in the roots and stumps.
Number and six of sprouts are dependent or related to the amount of stored food.

Forms and Sites in which Food Accumulates


Forms
1. Carbohydrates chiefly starch – hardwoods
2. Fats – chief storage from in conifers
3. Proteins – stored in seeds of some trees species
* - Fats and proteins are important reserved foods in seeds than in vegetative
structures.

FRE - TREE PHYSIOLOGY 3


Sites of Accumulation
1. In vegetative structures, food accumulate in the parenchyma cells especially in those
of the xylem and phloem and the rays (xylem parenchyma, phloem parenchyma, ray
parenchyma)
2. Accumulation in leaves is usually temporary and much of the food in leaves move
out before they fall by natural condition.

Factors Affecting Food Accumulation


1. Factors Affecting Photosynthesis – specifically the leaf area. Defoliation by insects,
fungi, fires and storms cause reduction in growth and even deaths of trees because
of the considerable decrease in the photosynthesis surface.
2. Factors Affecting Use of Food
a. Assimilation – excessive stimulation of vegetative growth uses so much
carbohydrates preventing flower bud formation.
b. Flowering and Fruiting – production of flowers, fruits and seeds uses large
amount of carbohydrates. Fruit production reduces vegetative growth.
c. Respiration – photosynthesis occurs only in chlorophyll – containing tissues
when exposed to light while respiration occurs in every living cell all the time.
Anything that increases respiration above that necessary for growth represents a
waste of food.

Role of mycorrhiza
1. absorption of nutrients – phosphorous is the main nutrient involved
2. increase absorption of water
3. helps in biological control of pathogenic organisms
4. promotes nodulation and nitrogen fixation
5. improves soil aggregation
6. promotes nutrient cycling
7. production of growth promoting hormones which improve plant growth

Effect of mycorrhizal infection


1. increases in total potential absorbing surface of the plant which significantly
increases nutrient absorption
2. prolongs the life of the short rootlets and stimulate them to branch
3. Mycorrhizal infection causes a considerable drain on photosynthetic products of the
plant since the fungus is dependent on the plant for carbohydrate supply.

FRE - TREE PHYSIOLOGY 3

You might also like