English Rule

GOVERNMENT OF TAMIL NADU
BOTANY
HIGHER SECONDARY FIRST YEAR
VOLUME - 2
Untouchability is Inhuman and a Crime

A publication under Free Textbook Programme of Government of Tamil Nadu
Department of School Education
TN_GOVT_BOTANY_LV_XI_Page I-IV Front.indd 1 27-04-2018 15:21:40

Government of Tamil Nadu
First Edition - 2018
NOT FOR SALE
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The wise
possess all
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© SCERT 2018
Printing & Publishing
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Services Corporation
www.textbooksonline.tn.nic.in
II

Learning objectives are brief statements that describe what
students will be expected to learn by the end of school year,
Learning Objectives: course, unit, lesson or class period.
Chapter Outline Illustrate the complete overview of chapter
Amazing facts, Rhetorical questions to lead students

to biological inquiry
Activity Directions are provided to students to conduct activities

in order to explore, enrich the concept.
Infographics Visual representation of the lesson to enrich learning .
Evaluation Assess students to pause, think and check their understanding
HOW TO USE
THE BOOK To motivate the students to further explore the content
digitally and take them in to virtual world
ICT To enhance digital Science skills among students
Conceptual diagram that depicts relationships between

Concept Map concepts to enable students to learn the content schematically
References List of related books for further details of the topic
Web links Digital resources
Glossary Explanation of scientific terms
Tamil terminology for Botanical terms given for easy

List of Botanical terms understanding
Competitive Model questions to face various competitive exams

Exam questions
III

Scope of Botany
Higher Studies and Career Opportunities
TN_GOVT_BOTANY_LV_XI_Page I-IV Front.indd 4

TNAU TNMGRMU
TNMG
GRM
RMU AIIMS SCIENCE
MEDICAL Indian Medicine and Undergraduate Courses (UG) Courses in Arts & Science Colleges
B.Sc. Agriculture,
Homoeopathy Courses and Universities
B.Sc. Horticulture
MBBS MBBS
B.Sc. Forestry, B.Sc. Botany
M.D/M.S/M.D.S B.A.M.S. - Ayurvedic Medicine B.Sc Nursing (post Certificate)
B.Sc Sericulture B.Sc. Plant Biology & Plant Biotechnology
M.Ch. (5 year course) B.H.M.S. - Homoeopathic Medicine B.Sc. (Hons.) Nursing
B.Tech Biotechnology B.Sc Biochemisty
B.D.S B.N.Y.S. - Naturopathy and Yogic Paramedical Courses (PM)
B.Tech Agricultural Engineering B.Sc Bio-computing
M.D.S B.S.M.S. - Siddha Medicine B.Sc. (Hons.) Opthalmic Techniques
B.Tech Horticulture B.Sc. Plant Pathology
B.U.M.S. - Unani Medicine B.Sc. (Hons.) Medical Technology
B.Tech Food process Engineering M.Sc. Botany
B.Tech Energy and Allied Health Sciences M.Sc Biotechnology
Environmental Engineering Postgraduate Courses (PG)
M.Sc. Bio-chemistry
B.Sc.(N)- Bachelor of Science in Nursing
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B.P.T.- Bachelor of Physiotherapy M.D/M.S/M.D.S
B.Sc Agribusiness Management M.Sc Immunology and Microbiology
M.P.T. - Master of Physiotherapy M.Ch. (5 year course)
B.Tech Agricultural IT M.Sc. Applied Medical Biotechnology & clinical
B.O.T. - Bachelor of Occupational Therapy M.Sc. / M. Biotechnology
M. Tech. Environmental Engineering Research
IV
M. Sc in Agriculture M.O.T. - Master of Occupational Therapy
M.Sc. Genetic Engineering & Plant Breeding
M. Sc in Agricultural Extension B.Sc. - Accident & Emergency Care Technology
M.Sc. Applied Plant Science
M. Sc in Agronomy B.Sc. - Audiology & speech Language Pathology
M.Sc. Plant Biology & Plant Biotechnology
M. Sc in Soil Science B.Sc. - Cardiac Technology
M.Sc. Plant molecular Biology
V
M. Sc in Agricultural Biotechnology B.Sc. - Cardio Pulmonary Perfusion Care Technology
M.Sc. Mycology & Plant pathology
M. Sc in Agricultural Marketing B.Sc. - Critical Care Technology
M.Sc. Plant science
M. Sc in Agricultural Microbiology B.Sc. - Dialysis Technology
M. Tech in Agricultural Engineering B.Sc. - Neuro Electrophysiology
M. E in Agricultural Engineering B.Sc. - Medical Sociology
Master of Agriculture in Entomology B.Sc. - Nuclear Medicine Technology
Master of Agriculture in Horticulture B.Sc. - Operation Theatre & Anaesthesia Technology
Master of Agriculture in Animal Sciences B.Sc. - Physician Assistant
Master of Agriculture in Entomology B.Sc. - Radiology Imaging Technology Integrated
tegrat
atted
d cours
courses
Master of Agriculture in Plant Pathology B.Sc. - Radiotherapy Technology
Master of Agriculture in Agricultural B.Sc. - Fitness and Lifestyle Modifications Mode of selection: Entrance conducted by
Economics and Rural Sociology B.Sc. - Clinical Nutrition concern institution or NEET
Master In Agriculture And M.Sc in Life sciences- 5 year Integrated
Rural Development course
Diploma Course
Indian institute of Science, Bengaluru
Accident & Emergency Care Technology ANNA UNIVERSITY
TANUVAS Website: http://www.iisc.ac.in/
Critical Care Technology
B.V.Sc and Animal Husbandry National Institute of Science
Health Care Aide (as per 245th GC)
B.Tech Food Technology Education and Research (NISER) , B.E. Bio Medical Engineering
Operation Theatre & Anaesthesia Technology
B.Tech Poultry processing
Bhubaneswar, Kolkata , Pune , B.Tech. Industrial Bio technology
Ophthalmic Nursing Assistant Mohali, Bhopal ,Thiruvananthapuram , B.Tech. Food technology
B.Tech Dairy Technology Scope Support Technology Tirupati and Berhampur
B.Tech. Bio technology
M.V.Sc. Medical Record Science Website: http://www.niser.ac.in
M.Tech. Food Techology Optometry Technology B.Sc.,B.Ed -5 year Integrated course
M.Sc., Bioinformatics/BioStatistics Radiology & Imaging Technology Regional Institute of Education
M.B.A. Medical Lab Technology Ajmer, Bhopal, Bhubaneswar, Mysuru
Cardiac Non Invasive Technology and Shilillong
Post Graduate Diploma
Dialysis Technology Website: www.riemysore.ac.in
27-04-2018 15:21:41
Research Institutions in various areas of Botany
Name of the Institution Research Areas Website
International Centre for Genetic Engineering Mammalian Biology; Plant Biology; Synthetic Biology and Biofuels. www.icgen.org
and Biotechnology (ICGEB), New Delhi
National Institute of Virology, Pune Epidemology, Basic virology; Diagnostics. www.niv.co.in

Center for DNA Fingerprinting and Diagnos- Computational Biology, Bioinformatics; Protein structure, Dynamic www.cdfd.org.in
tics, Hyderabad and Interactions Epigenetic
Institute of Life Sciences, Bhubaneswar Infectious disease;Immune biology; Cancer biology; Nanotechnology www.ils.res.in
Centre for Cellular and Molecular Biology, Genetics & evolution, Genomics; Cell Biology & Development. www.ccmb.res.in
Hyderabad.
Central food Technological Research Institute, Food science and Technology
Mysore.
Central Institute of medicinal and Aromatic Agronomy & soil sciences; Biotechnology, Crop protection; Genetics www.cimap.res.in
Plants, Lucknow. and plant breeding;
National Botanical Research Institute, Genetics and molecular biology; Plant microbe interaction & Phar- www.nbri.res.in
V
Lucknow. macogonosy.
Institute of Genomics and Integrative Biology Genomics and Molecular medicine, Chemical and systems biology. www.igib.res.in
Bose Institute, Kolkatta Molecular and cellular biology www.boseinst.ernet.in
National Centre for Biological Sciecnes, Biochemistry, Biophysics, Bioinformatics, Genetics and develop- www.ncbs.res.in
Bengaluru ment;Cellular organization & signelling neurobiology etc.
Birbal Sahni Institute od Palaeobotany (BSIP) Palynology in fossil fuel exploration; Dendrochronology; Ethnobota- www.bsip.res.in
Lucknow. ny; Micropaleontology; Carbon 14Dating
School of Medical Science and Technology, Indian Tissue Engineering; Biomaterials; Herbal medicine & Bio-Engineering. www.smstweb.iitkgp.
Institute of Technology, Kharagpur, West Bengal. ernet.in
Institute of Wood Science and Technology, Tree improvement and Genetics; Chemistry of Forest Products. iwst.icfre.gov.in
Bengaluru.
Centre for Ecological Sciences, Indian Institute Behaviour Ecology; Evolution; climate change & conservation. www.ces.iisc.ernet.in
of Science. Bengaluru.
Botanical Survey of India(BSI), Kolkatta www.bsi.gov.in
endangered species.
27-04-2018 15:21:41
Indian Agricultural Research Institute (IARI) Genetics & Plant Breeding; Plant Pathology; Microbiology; Post Har- www.iari.res.in
New Delhi vest Technology
Indian Institute of Horticultural Research, Horticultural Research; Biotechnology; Entomology; Pathology www.iihr.res.in

Bengaluru
Agharkar Research Institute, Pune Biodiversity & Palaeobiology, Bioenergy, BioprospectingNanobiosci- www.aripune.org
cence
National Bureau of Plant Genetic Resources Plant genetic resources management and use. www.nbpgr.ernet.in
(NBPGR) New Delhi
Institute of Forest Genetics and Tree Breeding, Tree improvement; Bio-prospecting of Forest Natural Resources www .ifgtb.icfre.gov.in
Coimbatore.
Central Soil Salinity Research Institute, Karnal, www .cssri.nic.in
Haryana of waste waters. Carbon Sequestration
Central Institute of Post Harvest Engineering & Rapid Evaluation of Food Quality and Safety; Packaging and storage www.ciphet.in
VI
Technology, Ludiana of agricultural produce and products.
Central Plantation crops Research Institute, Crop improvement; Production; Protection; Plant physiology and www.cpcri.gov.in
Kerala Biochemistry.
Indian Institute of Crop Processing Technology, Agricultural Process Engineering Renewable energy for food process- www.iicpt.edu.in
ing .
Central Tuber Crops Research Institute, Development of Agro techniques for tuber crops www.ctcri.org
National Centre for Integrated Pest Pest Management www.ncipm.org.in

Management (ICAR) New Delhi
Indian Institute of Spices Research, Kozhikode. Collection, conservation, evaluation and cataloging of germplasm. www.spices.res.in
Central Institute for Cotton Research, Nagpur, Crop improvement, Crop Production and Crop Protection. www.cicr.org.in
(Regional station: Coimbatore & Sirsa)
Central Institute for Research on Cotton Tech- Improvement in Ginning of cotton; Improvement and quality evalua- www.circot.res.in
nology, (CIRCOT) Mumbai
Directorate of Cashewnut & Cocoa, Agri, Kerala Cocoa production and processing www.dccd.gov.in
27-04-2018 15:21:42
National Research Center on Plant Biotechnol- Genetic engineering for biotic resistance. www.nrcpb.org
ogy, New Delhi

Indian Institute of Soil Sciences (IISS), Bhopal www.iiss.nic.in
activity.
National Institute of Plant Genome Research Structural and Functional Genomics in Plants; Computational biolo- www.nipgr.res.in
(NIPGR), New Delhi gy; Genome analysis and molecular mapping.
Sugarcane Breeding Institute, ICAR, Breeding of superior sugarcane varieties/ genotypes; www.sugarcane.res.in
Coimbatore.
National Centre for Agricultural Economics Agricultural technology policy. www.ncap.res.in
and Policy Research (NCAP), New Delhi
National Institute of Abiotic Stress Basic and strategic research on management of abiotic stresses of crop www.niam.res.in
VII
Management., Pune plants.
Central Research Institute for Dryland Dryland, Agrometerology and Crop sciences crida.in
Agriculture, Hyderabad
Central Research Institute for Jute & Allied Crop improvement, Crop production, Crop protection, Agricultural www.crijaf.org.in
Fibres, Kolkata, West Bengal research.
Indian Institute of Pulses Research (IIPR), Genetics & Plant Breeding and Seed Science www.iipr.res.in
Kanpur
National Research Centre for Groundnut(N- Productivity and quality of groundnut; repository of groundnut ger- www.nrcg.res.in
RCG) Junagarh, Gujarat mplasm and information on groundnut researches
Indian Institutes of Science Education and Re- Microbial Ecology; Marine Molecular Ecology; Marine Biology. www.iiserkol.ac.in
search(IISER) - Berhampur, Bhopal, www.issertvm.ac.in
and Tirupati.
27-04-2018 15:21:42
CONTENTS
BOTANY
UNIT IV: Plant Anatomy (Structural Organisation)

Chapter 9 Tissue and Tissue System 1
Chapter 10 Secondary Growth 38
UNIT V: Plant Physiology (Functional Organisation)

Chapter 11 Transport in plants 62
Chapter 12 Mineral Nutrition 95
Chapter 13 Photosynthesis 112
Chapter 14 Respiration 144
Chapter 15 Plant Growth and Development 168
Annexure
References 206
Glossary 207
English-Tamil Terminology 210
Competitive Exam Questions 214
E-book Assessment DIGI links
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VIII

Unit IV: Plant Anatomy
(Structural organisation)
Chapter
9 Tissue and Tissue System
Learning Objectives
The learner will be able to,
Nehemiah Grew
• Study major types of plant cells and Father of Plant
their function. Anatomy
• Differentiate the various types of
1641–1712
cells.
• Study the relationship between the Katherine Esau (1898–1997)
distribution of tissues in the various A legendary Role model for women in
parts of plants. science. She was a scintillating Botany
• Describes the ground tissue system teacher and pioneering researcher for
[cortex and pith] and vascular six decades. Her classic book Anatomy
systems of Seed Plants is the best literature in
• Interpret cross sections and Plant Anatomy. In
longitudinal sections of dicot and recognition of her
monocot root, stem and leaf. distinguished service
to science, she was
• Compare the internal organization awarded National
of dicot root and monocot root. Medal of Science
(1989) by USA.
This chapter introduces the internal

structure of higher Plants. The study of
Chapter Outline
internal structure and organisation of
9.1 Meristematic tissue plant is called plant Anatomy (Gk: Ana =
9.2 Permanent tissues as under; temnein = to cut). Plants have
9.3 The tissue system cells as the basic unit. The cells are
organised into tissues. The tissues in turn
9.4 Epidermal tissue system
are organised into organs. The different
9.5 Fundamental tissue system
organs in a plant have different internal
9.6 Vascular tissue system structures. It is studied by means of
9.7 Comparision of primary structure dissection and microscopic examination.
TN_GOVT_BOTANY_XI_CH09_001-037.indd 1 27-04-2018 11:15:30

Milestones in Anatomy of tissue is called Histology. A plant is
• 1837 Hartig: Coined the term Sieve made up of different types of tissues.
tubes There are two principal groups:
• 1839 Schleiden: Coined the term 1. Meristematic tissues
Collenchyma 2. Permanent tissues
• 1857 Hofmeister: Proposed Apical cell
9.1 Meristematic Tissue
theory
• 1858 Nageli. C: Coined the term Xylem 9.1.1 Characteristics and classification
and Phloem, Meristem and supporter The characters of meristematic tissues:
of Apical cell theory (Gr. Meristos-Divisible)
• 1865 Mettenius: Coined the term The term meristem is coined by
Sclerenchyma C. Nageli 1858.
• 1868 Hanstein: Proposed Histogen • The meristematic cells are isodiametric
theory and they may be, oval, spherical or
• 1885 Tschirch: Coined the term polygonal in shape.
Sclereids Named Four types of • They have generally dense cytoplasm
Sclereids (Brachy, Macro, Osteo & with prominent nucleus.
Astro) in 1889
• Generally the vacuoles in them are
• 1914 Haberlandt: Coined the term
either small or absent.
xylem as Hadrome and Phloem
as Leptome and Classification of • Their cell wall is thin, elastic and
meristem. essentially made up of cellulose.
• 1924 Schmidt A: Proposed Tunica – • These are most actively dividing cells.
Corpus theory
• Meristematic cells are self-perpetuating.
• 1926 Schűepp: Mass, rib, & plate
meristem Classification of Meristem
• 1946 Bloch: Discovered the Meristem has been classified into several
Trichosclereids types on the basis of position, origin,
• 1952 Popham: Explained the function and division.
organization of Shoot apex of
Apical meristem
Angiosperms
• 1955 Duchaigne: Discovered the
Annular collenchyma
Intercalary meristem
• 1961 Clowes: Proposed Quiescent
centre concept
• 1963 Sanio: Coined the term Tracheids Lateral meristem
The Tissues
A Tissue is a group of cells that are alike in Figure 9.1: Different types of meristems
origin, structure and function. The study on the basis of position in plant body
2

Classification of Meristem
Position Origin Function Plane of division
Apical meristem Protoderm

Primary Mass meristem
Present in apices of root It gives rise to
Meristem It divides in all
and shoot. It is responsible epidermal tissue
It is derived planes. Example:
for increase in the length system and
from endosperm,young
of the plant, it is called as develops into
embryonic embryo and
primary growth. epidermis,stomata
stages and sporangium
and hairs.
differentiated
into primary
Intercalary meristem
permanent Rib meristem or
Occurs between the tissues. Procambium File meristem
mature tissues. It is
It gives rise to It divides
responsible for elongation
primary vascular anticlinally in one
of internodes.
Secondary tissues. Example: plane. Example:
meristem It is xylem and phloem . development of
derived during cortex and pith
Lateral meristem
later stage of
Occurs along the
development
longitudinal axis of stem Plate meristem
of the plant Ground Meristem
and root. It is responsible
body. It It divides
for secondary tissues and It gives rise to
produces cork anticlinally in two
thickening of stem and all tissues except
cambium and planes. Example:
root. Example: vascular epidermis and
interfascicular development of
cambium and cork vascular strands.
epidermis
cambium. cambium.
Theories of Meristem Organization and Shoot Apical Meristem

Function Apical Cell Theory
Many anatomists illustrated the root Apical cell theory is proposed by
and shoot apical meristems on the basis Hofmeister (1852) and supported by
of number and arrangement and accordingly Nageli (1859). A single apical cell is the
proposed the following theories – An structural and functional unit.
extract of which are discussed below.
Apical cell
Tunica
Leaf primodium Leaf primordia

Dermatogen
Periblem Histogen Corpus
Plerome
a. b. c.
Figure 9.2: Shoot apical meristem a) Apical cell theory, b) Histogen theory,
c) Shoot Tunica corpus theory
3

This apical cell governs the growth at their apices and the apical meristem
and development of whole plant body. It is present below the root cap. The
is applicable in Algae, Bryophytes and in different theories proposed to explain
some Pteridophytes. root apical meristem organization is
given below.
Histogen Theory
Apical Cell Theory
Histogen theory is proposed by Hanstein
(1868) and supported by Strassburgur. The Apical cell theory is proposed by Nageli.
shoot apex comprises three distinct zones. The single apical cell or apical initial
composes the root meristem. The apical
1. Dermatogen: It is a outermost layer.
initial is tetrahedral in shape and produces
It gives rise to epidermis.
root cap from one side. The remaining
2. Periblem: It is a middle layer. It gives three sides produce epidermis, cortex and
rise to cortex. vascular tissues. It is found in vascular
3. Plerome: It is innermost layer. It gives cryptogams.
rise to stele
Histogen Theory
Tunica Corpus Theory Histogen theory is proposed by Hanstein
Tunica corpus theory is proposed by (1868) and supported by Strassburgur.
A. Schmidt (1924). The histogen theory as appilied to the
Two zones of tissues are found in apical root apical meristem speaks of four
meristem. histogen in the meristem. They are
respectively
1. The tunica: It is the peripheral zone of
shoot apex, that forms epidermis. i. Dermatogen: It is a outermost layer. It
2. The corpus: It is the inner zone of gives rise to root epidermis.
shoot apex,that forms cortex and stele ii. Periblem: It is a middle layer. It gives
of shoot. rise to cortex.
Root Apical Meristem iii. Plerome: It is innermost layer. It gives
Root apex is present opposite to the rise to stele
shoot apex. The roots contain root cap iv. Calyptrogen: It gives rise to root cap.
Epidermis Stele Cortex Cortex Stele Cortex

Protoderm
T
Ground tissue
Quiescent
Vascular cambium Root centre
cap Inverted ‘T’
division
(Y division) Cap
Plerome c.
b. Calyptrogen
Periblem
Dermatogen / Calyptrogen
Figure 9.3: Root apical meristem
a) Histogen Theory, b) Korper kappe theory,
Root cap
a. c) Quiescent Centre Concept
4

Korper Kappe Theory Parenchyma (Gk: Para-beside;
Korper kappe theory is proposed by enehein- to pour)
Schuepp. There are two zones in root Parenchyma is generally present in all organs
apex – Korper and Kappe of the plant. It forms the ground tissue in a
1. Korper zone forms the body. plant. Parenchyma is a living tissue and made
2. Kappe zone forms the cap. This up of thin walled cells. The cell wall is made
theory is equivalent to tunica corpus up of cellulose. Parenchyma cells may be oval,
theory of shoot apex.The two divisions polyhedral, cylindrical, irregular, elongated
are distinguished by the type of T or armed. Parenchyma tissue normally has
(also called Y divisions). Korper is prominent intercellular spaces. Parenchyma
characterised by inverted T divisions may store various types of materials like,
and kappe by straight T divisions. water, air, ergastic substances. It is usually
colourless. The turgid parenchyma cells help
Quiescent Centre Concept in giving rigidity to the plant body. Partial
Quiescent centre concept was proposed conduction of water is also maintained
by Clowes (1961) to explain root apical through parenchymatous cells.
meristem activity. These centre is located
Intercellular spaces
between root cap and differentiating
cells of the roots. The apparently inactive
region of cells in root promeristem is
called quiescent centre. It is the site of
hormone synthesis and also the ultimate
source of all meristematic cells of the
meristem.
9.2 Permanent Tissues Figure 9.4: Parenchyma

The Permanent tissues develop from apical
meristem. They lose the power of cell Occsionally Parenchyma cells which
division either permanently or temporarily. store resin, tannins, crystals of calcium
They are classified into two types: carbonate, calcium oxalate are called
idioblasts. Parenchyma is of different types
1. Simple permanent tissues.
and some of them are discussed as follows.
2. Complex permanent tissues.
Types of Parenchyma
Simple Permanent Tissues
Starch
Simple tissues are composed of one type g
grains
Intercellular
of cells only. The cells are structurally and spaces
functionally similar. It is of three types.
1. Parenchyma
a. b.
2. Collenchyma
Figure 9.5: Types of Parenchyma
3. Sclerenchyma
a) Aerenchyma, b) Storage parenchyma
5

1. Aerenchyma:
Parenchyma which contains air in its intercellular spaces. It helps
in aeration and buoyancy. Example: Nymphae and Hydrilla.
5. Prosenchyma: 2. Storage Parenchyma:

Parenchyma cells became Parenchyma stores food
elongated, pointed and slightly Parenchyma materials. Example: Root and
thick walled. It provides stem tubers.
mechanical support.
4. Chlorenchyma 3. How?.... Stellate

Parenchyma cells with Parenchyma
chlorophyll. Function is Star shaped parenchyma.
photosynthesis. Example: Example: Petioles of Banana
Mesophyll of leaves. and Canna.
Intercellular Intercellular small chloroplast or none. Tannin maybe

Space Spaces
Chloroplasts present in collenchyma.Based on pattern
Palisade of pectinisation of the cell wall, there are
Parenchyma
three types of collenchyma
Spongy
Parenchyma
Types of Collenchyma
a. b. c. 1. Angular collenchyma
Figure 9.5: c) Stellate parenchyma, It is the most common type of collenchyma
with irregular arrangement and
d) Chlorenchyma, e) Prosenchyma thickening at the angles where cells meets.
Example:Hypodermis of Datura and
Collenchyma (Gk. Colla-glue; Nicotiana
enchyma – an infusion)
Collenchyma is a simple, living mechanical 2. Lacunar collenchyma
tissue. Collenchyma generally occurs in The collenchyma cells are irregularly
hypodermis of dicot stem. It is absent arranged. Cell wall is thickening on the
in the roots and also occurs in petioles walls bordering intercellular spaces.
and pedicels. The cells are elongated Example:Hypodermis of Ipomoea
and appear polygonal in cross section.
The cell wall is unevenly thickened. 3. Lamellar collenchyma
It contains more of hemicellulose and The collenchyma cells are arranged
pectin besides cellulose. It provides compactly in layers(rows). The Cell wall
mechanical support and elasticity to the is thickening is at tangential walls.These
thickening appear as successsive tangential
growing parts of the plant. Collenchyma layers. Example:Hypodermis of Helianthus
consists of narrow cells. It has only a few
6

Diagramatic structures
Nucleus
Thickened Intercellular
corners thickenings
Protoplasm
Vacuole Lamellar
Cell wall thickenings
a. b. c.
Figure 9.6: Types of Collenchyma a) Angular collenchyma, b) Lacunar collenchyma,

c) Lamellar collenchyma
1. Sclereids
Annular Collenchyma: Duchaigne (1955)
2. Fibres
reported another type called Annular
collenchyma in petiole of Nerium. The Sclereids (Stone Cells)
lumen is more or less circular in shape.
Sclereids are dead cells, usually these
are isodiametric but some are elongated
Sclerenchyma (Gk. Sclerous- hard:
too. The cell wall is very thick due
enchyma-an infusion)
to lignification. Lumen is very much
The sclerenchyma is dead cell and reduced. The pits may simple or branched.
lacks protoplasm. The cells are long or Sclereids are mechanical in function.
short, narrow thick walled and lignified They give hard texture to the seed coats,
secondary walls. The cell walls of these cells endosperms etc., Sclereids are classified
are uniformly and strongly thickened. The into the following types.
sclerenchymatous cells are of two types:
Types of Sclereids
1. Branchysclereids or Stone cells: 2. Macrosclereids:

Isodiametric sclereids, with hard cell Elongated and rod shaped cells, found in
wall. It is found in bark, pith cortex, hard the outer seed coat of leguminous plants.
endosperm and fleshy portion of some Example: Crotalaria and Pisum sativum.
fruits. Example: - Pulp of Pyrus.
3. Osteosclereids (Bone cells):

Rod shaped with dilated ends. They occur in leaves and seed coats. Example: seed coat of Pisum
and Hakea
4. Astrosclereids: 5. Trichosclereids:
Star cells with lobes or arms diverging Hair like thin walled sclereids. Numerous
form a central body. They occur in petioles small angular crystals are embedded in the
and leaves. Example: Tea, Nymphae and wall of these sclereids, present in stems and
Trochodendron. leaves of hydrophytes. Example: Nymphaea
leaf and Aerial roots of Monstera.

Diagramatic Structures
Macro
Sclereid Lumen
Lumen cell Thick

cell wall
Lumen
Thick cell wall
Pith
a. b. c.
Thick Tricho
cell wall Sclereids
Lumen
d. e.
Figure 9.7: Types of Sclereids a) Brachysclereids, b) MacroSclereids, c) Osteosclereids,

d) Astrosclereids, e) Trichosclereids
Pointed
pits. They provide end
Filiform Sclereids: The sclereids mechanical
are present in the leaf lamina of strength and
Olea europaea. They are very much protect them
elongated fibre like and about 1m.m from the strong
length. wind. It is also
called supporting
Sclerenchyma Found in Some Fruits tissues. Fibres
have a great
commercial value
Lumen
in cottage and
textile industries.
Fibres are of five
Figure 9.8: a) Pear fruit,
types
b) Strawberry, c) Guava Figure 9.9 T.S of fibre
Wood Fibres or Xylary Fibres
Fibres
These fibres are associated with the
Fibres are very much elongated secondary xylem tissue. They are also
sclerenchyma cells with pointed tips. called xylary fibres. These fibres are
Fibres are dead cells and have lignified derived from the vascular cambium.
walls with narrow lumen. They have simple These are of four types. a. Libriform fibres
8

b. Fibre tracheids c. Septate fibres Mesocarp Fibres
d. Gelatinous fibres. Fibres obtained from the mesocarp of
drupes like Coconut.
Fibres are the longest Leaf Fibres
plant cells. Longest
Fibres obtained from the leaf of Musa,
Fibres occur in
Agave and Sensciveria.
Boehmeria (Ramie
fibre) 55 cm long Fibres in Our Daily Life
Economically fibres may be grouped as
a. Libriform fibres: These fibres have follows
slightly lignified secondary walls with
1. Textile Fibres: Fibres utilized for the
simple pits. These fibres are long and
manufacture of fabrics, netting and
narrow.
cordage etc.
b. Fibre tracheids: These are shorter
a. Surface Fibres: Example: Cotton.
than the libriform fibres with moderate
secondary thickenings in the cell walls. b. Soft Fibres: Example: Flax, Jute and
Pits are simple or bordered. Ramie
c. Septate fibres: Fibres that have thin c. Hard fibres: Example: Sisal,
septa separating the lumen into distinct Coconut, Pineapple, Abaca etc.
chambers. Eg. Teak 2. Brush fibre: Fibres utilized for the
d. Gelatinous fibres: Fibres in which lignin manufacture of brushes and brooms.
is less in amount and cellulose is more in 3. Rough weaving fibres: Fibres utilized
this cell walls. in making baskets, chairs, mats etc.
These fibres are characteristic of tension 4. Paper making fibres: Wood fibres
wood which is formed in the underside of utilized for paper making.
leaning stems and branches. 5. Filling fibres: Fibres used for stuffing
cushions, mattresses, pillows, furniture
Bastfibres or Extra Xylary Fibres etc. Example: Bombax and Silk cotton.
These fibres are present in the phloem.
Natural Bast fibres are strong and Complex Tissues
cellulosic. Fibres obtaining from the A complex tissue is a tissue
phloem or outer bark of jute, kenaf, with several types of cells
flax and hemp plants. The so called but all of them function
pericyclic fibres are actually phloem together as a single unit. It
fibres. is of two types – xylem and
Surface Fibres phloem.
These fibres are produced from the surface Xylem
of the plant organs. Cotton and silk cotton The xylem is the principal water conducting
are the examples.They occur in the testa tissue in a vascular plant. The term xylem
of seeds. was introduced by Nageli(1858) and is
9

derived from the Gk. Xylos – wood. The Tracheids
xylem which is derived from Procambium is Tracheids are dead, lignified and
called primary xylem and the xylem which elongated cells with tapering ends. Its
is derived from vascular cambium is called lumen is broader than that of fibres. In
secondary xylem. Early formed primary cross section, the tracheids are polygonal.
xylem elements are called protoxylem,
There are different types of cell wall
whereas the later formed primary xylem
thickenings due to the deposition of
elements are called metaxylem.
secondary wall substances. They are
Protoxylem lies towards the periphery annular (ring like), spiral (spring like),
and metaxylem that lies towards the centre scalariform (ladder like) reticulate (net
is called Exarch. It is common in roots. like) and pitted (uniformly thick except
Protoxylem lies towards the centre and at pits). Tracheids are imperforated cells
meta xylem towards the periphery this with bordered pits on their side walls.
condition is called Endarch. It is seen in Only through this conduction takes place
stems. in Gymnosperms. They are arranged one
Protoxylem is located in the centre above the other. Tracheids are chief water
surrounded by the metaxylem is called conducting elements in Gymnosperms
Centrarch. In this type only one vascular and Pteridophytes. They also offer
strand is developed. Example: Selaginella mechanical support to the plants.
sp.
Protoxylem is located in the centre
surrounded by the metaxylem is
called Mesarch.In this type several
vascular strands are developed.
Example: Ophioglossum sp.
Student Activity Annular Spiral Reticulate Scalariform Pitted thickening
Cell lab: students prepare the slide Figure 9.10: Types of secondary wall
and identify the different types tissues.
thickenings in tracheids and vessels
Xylem Consists of Four Types of Cells

Vessels or Trachea
1. Tracheids
Vessels are elongated tube like structure.
2. Vessels or Trachea
They are dead cells formed from a row of
3. Xylem Parenchyma vessel elements placed end to end. They
4. Xylem Fibres are perforated at the end walls. Their
lumen is wider than Tracheids. Due to
Xylem is called hadrome phloem the dissolution of entire cell wall, a single
is called leptome. These terms are pore is formed at the perforation plate.
coined by haberlandt (1914) It is called simple perforation plate,
Example: Mangifera. If the perforation
10

plate has many pores, it is called multiple Xylem Parernchyma
perforation plate. Example Liriodendron. The parenchyma cells associated with the
The secondary wall thickening of xylem are known as xylem parenchyma.
vessels are annular, spiral, scalariform, These are the only living cells in xylem
reticulate, or pitted as in tracheids, Vessels tissue. The cell wall is thin and made
are chief water conducting elements in up of cellulose. Parenchyma arranged
Angiosperms and absent in Pteridophytes longitudinally along the long axis is called
and Gymnosperms. In Gnetum of axial parenchyma. Ray parenchyma is
Gymnosperm, vessels occur. The main arranged in radial rows. Secondary xylem
function is conduction of water, minerals consists of both axial and ray parenchyma,
and also offers mechanical strength. Parenchyma stores food materials and
also helps in conduction of water.
Xylem Fibre
The fibres of sclerenchyma associated Phloem
with the xylem are known as xylem fibres. Phloem is the food conducting complex
Xylem fibres are dead cells and have tissues of vascular plants. The term
lignified walls with narrow lumen. They phloem was coined by C. Nageli (1858)
cannot conduct water but being stronger The Phloem which is derived from
provide mechanical strength. They are procambium is called primary phloem and
present in both primary and secondary the phloem which is derived from vascular
xylem. Xylem fibres are also called cambium is called secondary phloem.
libriform fibres. Early formed primary phloem elements
The fibres are abundantly found in many are called protophloem whereas the later
plants. They occur in patches, in continuous formed primary phloem elements are
bands and sometimes singly among other called metaphloem. Protophloem is short
cells. Between fibres and normal tracheids, lived. It gets crushed by the developing
there are many transitional forms which are metaphloem.
neither typical fibres nor typical tracheids.
The transitional types are designated as Phloem Consists of Four Types of Cells
fibre-tracheids. The pits of fibre-tracheids 1. Sieve elements
are smaller than those of vessels and typical 2. Companion cells
tracheids. 3. Phloem parenchyma
Vessels are found in 4. Phloem fibres

Gymnosperms like Sieve Elements
Ephedra, Gnetum and
Welwitschia Sieve elements are the conducting
elements of the phloem. They are of two
Vesselless angiospermic families
types, namely sieve cells and sieve tubes.
Winteraceae, Tetracentraceae and
Trochodendracae. Sieve Cells
These are primitive type of conducting
11

elements found in Pteridophytes and Companion Cells
Gymnosperms. Sieve cells have sieve areas The thin walled, elongated, specialized
on their lateral walls only. They are not parenchyma cells, which are associated with
associated with companion cells. the sieve elements, are called companion
Sieve Tubes cells. These cells are living and they have
Sieve tubes are long tube like conducting cytoplasm and a prominent nucleus. They
elements in the phloem. These are formed are connected to the sieve tubes through
from a series of cells called sieve tube pits found in the lateral walls. Through
elements. The sieve tube elements are these pits cytoplasmic connections are
arranged one above the other and form maintained between these elements.
vertical sieve tube. The end wall contains These cells are helpful in maintaining
a number of pores and it looks like a the pressure gradient in the sieve tubes.
sieve. So it is called as sieve plate. The Usually the nuclei of the companion
sieve elements show nacreous thickenings cells serve for the nuclei of sieve tubes as
on their lateral walls. They may possess they lack them. The companion cells are
simple or compound sieve plates The present only in Angiosperms and absent
function of sieve tubes are believed to be in Gymnosperms and Pteridophytes. They
controlled by campanion cells.
assist the sieve tubes in the conduction of
In mature sieve tube, Nucleus is absent. food materials.
It contains a lining layer of cytoplasm.
A special protein (P. Protein = Phloem Phloem Parenchyma
Protein) called slime body is seen in it. In
mature sieve tubes, the pores in the sieve The parenchyma cells associated with the
plate are blocked by a substance called phloem are called phloem parenchyma.
callose (callose plug).The conduction These are living cells. They store starch
of food material takes place through and fats. They also contain resins and
cytoplasmic strands. Sieve tubes occur tannins in some plants. Primary phloem
only in Angiosperms. consists of axial parenchyma and
Sieve plate
secondary phloem consists of both axial
and ray parenchyma. They are present
Sieve tube
in Pteridophytes,Gymnosperms and
Dicots.
Phloem
parenchyma
Phloem Fibres (or) Bast Fibres
The fibres of sclerenchyma associated
Companian
with phloem are called phloem fibres or
Cells bast fibres. They are narrow, vertically
elongated cells with very thick walls and
a small lumen. Among the four phloem
elements, phloem fibres are the only dead
Figure 9.11: Different types of tissue. These are the strengthening as well
phloem elements as supporting cells.
12

Concept Map
Plant tissues
Meristematic tissue: Permanent tissues:

Capable of active cell division. Lose the power of cell division.
Thin walled and living. Have definite shape, size and form.
Compactly arranged.
Found in root and shoot apex.
Simple tissues: Complex tissues:
One type of cells. More than one type
Based on position: of cells.
1. Apical
2. Intercalary Parenchyma:
3. Lateral Thin walled,isodiametric,
found in all the parts. Xylem: Water
Types: conducting tissue.
Based on origin: 1. Aerenchyma. 1. Tracheids: Dead,
2. Storage parenchyma. elongated with
1. Primary
3. Stellate parenchyma. tapering end
2. Secondary
4. Chlorenchyma. 2. Vessels: Made of
5. Prosenchyma. row of dead cells
3. Xylem fibres:
Based on function: Lignified and
1. Periderm (Epidermis ) Collenchyma: sclerenchymatous.
2. Procambium (Primary Hypodermal position. Provide 4. Xylem
vascular tissues) mechanical strength. parenchyma:
3. Ground meristem (Cortex Types: Living and
and Pith) 1. Angular collenchyma. cellulosic
2. Lacunar collenchyma.
3. Lamellar collenchyma.
Based on division:
1. Mass meristem: Divides in all Phloem: Food
Sclerenchyma: Dead cells and conducting tissue
planes lignified walls.
2. Rib meristem: Anticlinal 1. Sieve elements: Sieve
Types: cells & sieve tubes
division in one plane 1. Sclereids
3. Plate meristem: Anticlinal 2. Companion cells:
2. Fibres Only in Angiosperms.
division in two planes.
3. Phloem
Sclereids: parenchyma:
1. Brachysclereids: Stone cells Living & absent in
2. Macro sclereids: Rod shaped Monocots.
3. Osteosclereids: Bone shaped 4. Phloem fibres:
Syncyte: Cell Thick walled &
4. Astrosclereids: Star shaped
which is formed sclerenchymatous,
5. Trichosclereids: Hair cells
by fusion of giving mechanical
cell is called strength.
Syncyte. Fibres:
1. Wood fibres xylary fibres
Example: Vessels (Dead 2. Bastfibres: Extra xylary fibres
syncyte), sieve tube (living 3. Surface fibres: Cottan
syncyte) 4. Mesocarp fibres: Ccoconut
5. Leaf fibres: Musa, Agave
13

Table 9.1: Different types of tissues
Distribution Main functions Nature Cell shape Wall materials
Parenchyma Cortex, Pith Packing tissue, Living Usually Mainly
medullary rays support, gaseous Isodiametric Cellulose and
and Packing exchange, food Pectinase
tissues in storage
vascular system
Collenchyma Outer region Mechanical Living Elongated, Mainly
of cortex as in Polygonal Cellulose,
angles of stems, Pectin and
mid-rib of leaves Hemi-cellulose
Sclerenchyma Outer region of Mechanical Dead Elongated Mainly Lignin
(a) Fibre cortex, pericycle and
of stems, vascular Polygonal
bundles with tapering
ends
(b) Sclereids Cortex, Pith, Mechanical Dead Roughly Mainly lignin
Phloem shells Protection Isodiametric
and stones of with much
fruits and seed variation
coats
Tracheids and Vascular System Translocation Dead Elongated Mainly lignin
Vessels of water and and Tubular
mineral salts
Phloem Sieve Vascular System Translocation of Living Elongated Cellulose,
tubes organic solutes and Tubular Pectin and
Hemicellulose
Companion Vascular System Work in Living Elongated Cellulose,
Cells association with and narrow Pectin and
sieve tubes Hemicellulose
Difference Between Meristematic Tissue and Permanent Tissue

Meristematic tissue Permanent tissue
• Cells divide repeatedly • Do not divide
• Cells are undifferentiated • Cells are fully differentiated
• Cells are small and Isodiametric • Cells are variable in shape and size
• Intercellular spaces are absent • Intercellular spaces are present
• Vacuoles are absent • Vacuoles are present
• Cell walls are thin • Cell walls maybe thick or thin
• Inorganic inclusions are absent • Inorganic inclusions are present
14

Difference Between Collenchyma and Sclerenchyma
Collenchyma Sclerenchyma
• Living Cells • Dead cells
• Contains Protoplasm • Cells are empty
• Cell walls are cellulosic • Cell walls are lignified
• Thickening of cell wall is not uniform • Thickening of cell wall is uniform
• Keeps the plant body soft • Keeps plant body stiff and hard
• Sometimes it has chloroplast • Do not have chloroplast
Difference between Fibre and Sclereids

Fibre Sclereids
• Long cells • Short cells
• Narrow, Elongated pointed ends • Usually short and broad
• Occurs in bundles • Occurs individually or in small groups
• Commonly unbranched • Maybe branched
• Derived directly from meristematic • Develops from secondary sclerosis
tissue parenchyma cells
Difference between Tracheids and Fibres

Tracheids Fibres
• Not much elongated • Very long cells
• Possess oblique end walls • Possess tapering end walls
• Cell walls are not as thick as Fibres • Cell wall are thick and lignified
• Possess various types of thickenings • Possess only pitted thickenings
• Responsible for the conduction and also • Provide only mechanical support
mechanical support
Difference Between Sieve Cells and Sieve Tubes

Sieve cells Sieve tubes
• Have no companion cells • Have companion cells
• The sieve areas do not form sieve plates • The sieve areas are confined to sieve
• The sieve areas are not well plates
differentiated • The sieve areas are well differentiated
• They are elongated cells and are quite • They consist of vertical cells placed
long with tapering end walls one above the other forming long tubes
• The sieve are smaller and numerous connected at the walls by sieve pores
• Found in Pteridophytes and • The sieve pores are longer and fewer
Gymnosperms • Found in Angiosperms
15

9.3 The Tissue System recognized three tissue systems in the
plants. They are:
Introduction to Tissue System, Types
and Characteristics of tissue System 1. Epidermal tissue system (derived from
protoderm)
As you have learnt, the plant cells are
organised into tissues, in turn the tissues 2. Ground tissue system (derived from
are organised into organs. Different ground meristem)
organs in a plant show differences in their 3. Vascular tissue system (derived from
internal structure. This part of chapter procambium)
deals with the different type of internal
structure of various plant organs and its
adaptations to diverse environments. Histology
A group of tissues (Greek. histos – web,
performing a similar logos – science) It is
function, irrespective of the study of tissues,
its position in the plant their composition, and structure
body, is called a tissue as observed with the help of
system. In 1875, German Figure 9.12: microscope.
Scientist Julius von Sachs Julius von Sachs
Figure 9.13: Tissue system

16

Table 9.2: Types and characteristics of tissue systems
S.No. Types/ Epidermal tissue Ground or Vascular or
Characters system fundamental tissue conduction tissue
system system
1. Formation Forms the outermost Forms the ground Forms the
covering protoderm meristem procambial bundles
2. Components epidermal Simple permanent Xylem and Phloem
cells, stomata tissues –
and epidermal Parenchyma and
outgrowths Collenchyma
3. Functions Protection of plant Gives mechanical Conducts water
body; absorption support to the and food; gives
of water in roots; organs; prepares and mechanical strength
gas exchange for stores food in leaf
photosynthesis and stem
and respiration;
transpiration in
shoots
9.4 Epidermal Tissue System Stem Epidermis
Introduction It is protective in function and forms the

outermost layer of the stem. It is a single
Epidermal tissue system is the outer most
layer of parenchymatous rectangular cells.
covering of plants. It is in direct contact
The cells are compactly arranged without
with external environment. It consists
intercellular cells. The outer walls of
of epidermis derived from protoderm.
epidermal cells have a layer called cuticle.
Epidermis is derived from two Greek words,
The cuticle checks transpiration. The
namely ‘Epi’ and ‘Derma’. ‘Epi’ means
cuticle is made up of cutin. In many plants
upon and ‘Derma’ means skin. Although
it is also mixed wax to form epicuticular
epidermis is a continuous outer layer, it is
wax. Epidermal pores may be present
interrupted by stomata in many plants.
here and there. Epidermal cells are living.
Root Epidermis Chloroplasts are usually absent except in
guard cells of stomata. In many plants a
The outer layer of the root is known as
large number of epidermal hairs occur on
piliferous layer or epiblema. It is made up
the epidermis.
of single layer of parenchyma cells which are
arranged compactly without intercellular Leaf Epidermis
spaces. It is devoid of epidermal pores and
The leaf is generally dorsiventral. It has
cuticle. Root hair is always single celled, it
upper and lower epidermis. The epidermis
absorbs water and mineral salts from the
is usually made up of a single layer of cells
soil. The another important function of
that are closely packed. Generally the
piliferous layer is protection.
17

cuticle on the upper epidermis is thicker epidermal cells called guard cells. In most
than that of lower epidermis. The minute dicots and monocots the guard cells are
openings found on the epidermis are bean-shaped. While in grasses and sedges,
called stomata (singular: stoma). Usually, the guard cells are dumbbell- shaped. The
stomata are more in number on the lower guard cells contain chloroplasts, whereas
epidermis than on the upper epidermis. A the other epidermal cells normally do not
stoma is surrounded by a pair of specialised have them.
Subsidiary cell
Figure 9.14: (a) Stoma with bean-shaped guard cells.

(b) Stoma with dumb-bell shaped guard cells
Some cells of upper epidermis mainly in the epidermis of leaves. In

(Example: Grasses) are larger and thin some plants addition to guard cells,
walled. They are called bulliform cells specialised epidermal cells are present
or motor cells. These cells are helpful which are distinct from other epidermal
for the rolling and unrolling of the leaf cells. They are called Subsidiary cells.
according to the weather change. Some Based on the number and arrangement
of the epidermal cells of the grasses are of subsidiary cells around the guard
filled with silica. They are called silica cells, the various types of stomata
cells. are recognised. The guard cells and
subsidiary cells help in opening and
Check Your Grasp! closing of stomata during gaseous
In which group of plants the guard exchange and transpiration.
cells are dumb-bell shaped?
Sunken Stomata
Grasses and s edges In some Xerophytic plants (Examples:
Cycas, Nerium), stomata is sunken beneath
the abaxial leaf surface within stomatal
Subsidiary Cells
crypts. The sunken stomata reduce water
Stomata are minute pores surrounded loss by transpiration.
by two guard cells. The stomata occur
18

Multilayered or Multiseriate Epidermis In Nerium, in the multilayered
Generally, epidermis is single layered, epidermis the outer layer alone is
but in certain leaves, multilayered upper cutinized.
epidermis is present, Example: Ficus, Epidermal Outgrowths
Nerium, and Peperomea.
There are many types of epidermal
In Ficus upper epidermal layer contains outgrowths in stems. The unicellular or
cystoliths made up of calcium carbonate multicellular appendages that originate
crystals.
Cuticle
Upper epidermis
(Multilayered)
Palisade parenchyma
Bundle sheath
Xylem
Phloem
Spongy parenchyma
Sunken stoma
Hair
Lower epidermis
(Multilayered)
Figure 9.15: T.S. of Nerium Leaf
from the epidermal cells are called Trichoblasts are elongate into root hairs.
trichomes. Trichomes may be branched or Epidermal hairs can also be in the form
unbranched and are one or more one celled of stellate hairs (star shaped) present in
thick. They assume many shapes and sizes. plants. Example: styrax, many members
They may also be glandular (Example: of Malvaceae and Solanaceae.
Rose, Ocimum) or non-glandular.
Long Short cell
cell (Trichoblast)
The trichomes on the
leaves of insectivorous
plants secrete muco-
polysaccharides that Unicellular Uniseriate Branched Rhizodermis
hair unbranched multiseriate and root hairs
trap insects. hair hair
Piliferous layer of the root has two types of

epidermal cells, long cells and short cells. Stellate hairs
The short cells are called trichoblasts. Figure 9.16: Types of Trichomes
19

Prickles main zones – cortex, pericycle and pith.
Prickles, are one type of It is classified into extrastelar ground
epidermal emergences tissue (Examples: cortex and endodermis)
with no vascular supply. and intrastelar ground tissue (Examples:
They are stiff and sharp pericycle, medullary ray and pith)
in appearance. (Example: Extrastelar Ground Tissue
Rose).
The ground tissues present outside the
Figure 9.17:
Functions of Epidermal stele is called extrastelar ground tissue.
Prickles (Cortex)
Tissue System
1. This system in the shoot checks Intrastelar Ground Tissue
excessive loss of water due to the
The ground tissues present within the
presence of cuticle.
stele are called intrastelar ground tissues.
2. Epidermis protects the underlying (pericycle, medullary rays and pith).
tissues.
Different Components of Ground
3. Stomata is involved in transpiration
Tissue Systems are as follows
and gaseous exchange.
4. Trichomes are also helpful in the Hypodermis
dispersal of seeds and fruits, and One or two layers of continuous or
provide protection against animals. discontinuous tissue present below the
5. Prickles also provide protection against epidermis, is called hypodermis. It is
animals and they also check excessive protective in function.
transpiration In dicot stem, hypodermis is generally
6. In some rose plants they also help in collenchymatous, whereas in monocot
climbing. stem, it is generally sclerenchymatous.
7. Glandular hairs repel herbivorous In many plants collenchyma form the
animals. hypodermis.
General Cortex
9.5 Fundamental Tissue System The Cortex occurs between the epidermis
The ground or fundamental tissue system and pericycle. Cortex is a few to many
constitutes the main body of the plants. It layers in thickness, In most cases, it is
includes all the tissues except epidermis made up of parenchymatous tissues.
and vascular tissues. In monocot stem, Intercellular spaces may or may not be
ground tissue system is a continuous present.
mass of parenchymatous tissue in which The cortical cells may contain non
vascular bundles are found scattered. living inclusions of starch grains, oil,
Hence ground tissue is not differentiated tannins and crystals.
into cortex, endodermis, pericycle and Sometimes in young stem, chloroplasts
pith. Generally in dicot stem, ground develop in peripheral cortical cells, which
tissue system is differentiated into three is called chlorenchyma.
20

In the leaves, the ground tissue consists Pericycle
of chlorenchyma tissues. This region is Pericycle is single or few layered parenchymatous
called mesophyll. In hydrophytes, cortex is found inner to the endodermis. It is the
Aerenchymatous (with air cavities). outermost layer of the stele. Rarely thick walled
Its general function is storage of food sclerenchymatous. In angiosperms, pericycle
as well as providing mechanical support gives rise to lateral roots.
to organs.
Pith or Medulla
Endodermis
The central part of the ground tissue is
The cells of this layer are barrel shaped and known as pith or medulla. Generally this
arranged compactly without intercellular is made up of thin walled parenchyma
spaces. cells with intercellular spaces. The cells
Endodermis is the innermost cortical in the pith generally stores starch, fatty
layer that separates cortex from the stele. substances, tannins, phenols, calcium
This layer may be a true endodermis as in oxalate crystals, etc.
root or it is an endodermis like layer in stems.
This layer is morphologically homologous Albuminous Cells: The cytoplasmic
to the endodermis found in the root. nucleated parenchyma, is associated
The cells of endodermis like layer with the sieve cells of Gymnosperms.
had living cells containing starch grains. Albuminous cells in Conifers are
Hence it is known as starch sheath. In analogous to companion cells of
true root endodermis, radial and inner Angiosperms. It also called as
tangential walls of endodermal cells strasburger cells.
possess thickenings of lignin, suberin and
some other carbohydrates in the form of 9.6 Vascular Tissue System
strips they are called casparian strips.
This section deals with the vascular tissue
The endodermal cells, which are
opposite to the protoxylem elements, system of gymnosperms and angiosperms
are thin walled without casparian strips. stems and roots.The vascular tissue
These cells are called passage cells. system consists of xylem and phloem. The
Their function is to transport water and elements of xylem and phloem are always
dissolved salts from the cortex to the organized in groups. They are called
protoxylem. vascular bundles.
Water cannot pass through other The stems of both groups have an
endodermal cells due to casparian strips. eustele while roots are protostele. In
The main function of casparian strips eustelic organization, the stele contains
in the endodermal cells is to prevent the usually a ring of vascular bundles separated
re-entry of water into the cortex once by interfascicular region or medullary ray
water entered the xylem tissue. The structural and organizational
The other suberized cells acts as variation in vascular bundles is shown
water-tight layer between vascular and non- below.
vascular regions to check the loss of water.
21

Outer Phloem
Phloem
Outer Cambium
Cambium
Xylem
Xylem Inner Cambium

Inner Phloem
a c e
Outer Phloem
Phloem Outer Cambium
Cambium Xylem
Xylem
Inner Cambium
Inner Phloem
b d f
Xylem Xylem
Phloem Phloem
g i k
Xylem
Xylem
Phloem
Phloem
h j l
Figure 9.18: Types of vascular bundles
(a) and (b) - Conjoint, collateral and open; (c) and (d) - Conjoint, collateral and closed
(e) and (f ) - Conjoint, bicollateral and open; (g) and (h) - Concentric and amphicribral;
(i) and (j) - Concentric and amphivasal; (k) and (l) - Radial
22

Types of vascular Bundles
Radial Conjoint Concentric
Xylem and phloem are present on Xylem and phloem Xylem and phloem are
different radii alternating with each are present on the present in concentric
other. The bundles are separated same radius in one circles one around the
by parenchymatous tissue. bundle. ( All stems ) other in some stems.
(Monocot and Dicot roots)
Collateral Bicollateral
Xylem placed towards inside Phloem occurs on both the

and phloem towards outside outer and inner sides of xylem
Example: Cucurbitaceae
Open Closed
Cambium is Amphicribral/Hadrocentric Amphivasal/Leptocentric

Cambium is present
between xylem and absent
between
phloem. (Stems of Xylem lies in the centre Phloem lies in the centre
Dicots and xylem and
phloem. with phloem surrounding
Gymnosperms) with xylem surrounding it.
Stems of it. Example: Ferns
Example: Dragon plant-
Monocots) (Polypodium) dicots
and aquatic Dracena and Yucca
angiosperms
Table 9.3: Comparison of vascular tissues

Proto xylem Meta xylem
• First formed primary xylem • Later formed primary xylem
• Found in developing organs • Found in developed primary organs
• Elements relatively smaller in size • Elements relatively larger in size
Proto phloem Meta phloem
• First formed primary phloem • Later formed primary phloem
• Found in developing organs • Found in developed primary organs
• Elements relatively smaller in size • Elements relatively larger in size
Primary xylem Secondary xylem
• The primary xylem is derived from the • The secondary xylem is derived from
procambium of the apical meristem the vascular cambium which is a lateral
meristem
Primary phloem Secondary phloem
• The primary phloem is derived from the • The secondary phloem is derived from
procambium of the apical meristem the vascular cambium, which is a lateral
meristem
23

9.7 Comparison of Primary of the cortex is endodermis. Endodermis
Structure – Dicot and Monocot is made up of single layer of barrel shaped
Root, Stem and Leaf parenchymatous cells. Stele is completely
surrounded by endodermis. The radial and
Anatomy of Dicot and Monocot Roots the inner tangential walls of endodermal
In different parts of the plants, the various cells are thickened with suberin and lignin.
tissues are distributed in characteristic This thickening was first noted by Robert
patterns. This is best understood by studying Casparay in 1965. So these thickenings are
their internal structure by cutting sections called casparian strips. But these casparian
(transverse or longitudinal or both) of the strips are absent in the endodermis cells
part to be studied. which are located opposite the protoxylem
elements. These thin-walled cells without
Primary Structure of Dicot Root – casparian strips are called passage cells
Bean Root through which water and mineral salts are
The transverse section of the dicot root conducted from the cortex to the xylem
(Bean) shows the following plan of elements. Water cannot pass through other
arrangement of tissues from the periphery endodermal cells due to the presence of
to the centre. casparian thickenings.
Piliferous Layer or Epiblema Check Your Grasp!

The outermost layer of the root is called Give the exact location and function
piliferous layer or epiblema. It is made up of passage cells?
of single layer of parenchyma cells which are In roots some cells of the
arranged compactly without intercellular endodermis usually the ones opposite
spaces. It is devoid of epidermal pores and to protoxylem, remain thin walled.
cuticle. It possesses root hairs which are These cells are called passage cells.
single celled. It absorbs water and mineral They help in radial diffusion of water.
salts from the soil. The chief function of
piliferous layer is protection. Stele
Cortex All the tissues present inside endodermis
Cortex consists of only parenchyma comprise the stele. It includes pericycle
cells. These cells are loosely arranged and vascular system.
with intercellular spaces to make gaseous Pericycle
exchange easier. These cells may store food
Pericycle is generally a single layer of
reserves. The cells are oval or rounded in
parenchymatous cells found inner to the
shape. Sometimes they are polygonal due
endodermis. It is the outermost layer
to mutual pressure. Though chloroplasts
of the stele. Lateral roots originate from
are absent in the cortical cells, starch
the pericycle. Thus, the lateral roots are
grain are stored in them. The cells also
endogenous in origin.
possess leucoplasts. The innermost layer
24

Vascular System arrangement of tissues from the periphery
Vascular tissues are in radial arrangement. to the centre.
The tissue by which xylem and phloem
are separated is called conjunctive tissue. Root hair
Piliferous layer
In bean, the conjuctive tissue is composed Cortex
of parenchyma tissue. Xylem is in exarch Endodermis
condition. The number of protoxylem Pith
points is four and so the xylem is called

tetrach. Each phloem patch consists
of sieve tubes, companion cells and Ground plan
phloem parenchyma. Metaxylem vessels

are generally polygonal in shape. But in Root hair
Piliferous layer
monocot roots they are circular.
Root hair
Piliferous layer
Cortex Cortex
Phloem Passage cell
Metaxylem Casparian strip
Conjunctive tissue
Endodermis
Pericycle
Ground plan Passage cell

Phloem
Root hair Protoxylem
Piliferous layer Metaxylem

Pith
Cortex A sector enlarged
Figure 9.20: T.S of Monocot root

(Maize root)
Endodermis
Phloem
Piliferous Layer or Epiblema
Pericycle
Protoxylem The outermost layer of the root is known as
Conjunctive tissue
Metaxylem
piliferous layer. It consists of a single row of
Casparian strip thin-walled parenchymatous cells without
Passage cell
any intercellular space. Epidermal pores
A sector enlarged and cuticle are absent in the piliferous layer.
Root hairs that are found in the piliferous
Figure 9.19: T.S. of Dicot root (Bean root)
layers are always unicellular. They absorb
Primary Structure of Monocot water and mineral salts from the soil. Root
Root-maize Root hairs are generally short lived. The main
function of piliferous layer is protection of
The transverse section of the monocot the inner tissues.
root (maize) shows the following plan of
25

Cortex transport water and dissolved salts from
The cortex is homogenous. i.e. the cortex the cortex to the xylem. Water cannot
is made up of only one type of tissue called pass through other endodermal cells due
parenchyma. It consists of many layers to casparian strips. The main function of
of thin-walled parenchyma cells with casparian strips in the endodermal cells is
lot of intercellular spaces. The function to prevent the re-entry of water into the
of cortical cells is storage. Cortical cells cortex once water entered the xylem tissue.
are generally oval or rounded in shape.
Stele
Chloroplasts are absent in the cortical
cells, but they store starch. The cells All the tissues inside the endodermis
are living and possess leucoplasts. The comprise the stele. This includes pericycle,
inner layer of the cortex is endodermis. vascular system and pith.
It is composed of single layer of barrel
Pericycle
shaped parenchymatous cells. This forms
a complete ring around the stele. There Pericycle is the outermost layer of the
is a band like structure made of suberin stele and lies inner to the endodermis. It
and lignin present in the radial and inner consists of single layer of parenchymatous
tangential walls of the endodermal cells. cells.
They are called casparian strips named
after casparay who first noted the strips. Vascular System
The endodermal cells, which are opposite Vascular tissues are seen in radial
the protoxylem elements, are thin walled arrangement. The number of protoxylem
without casparian strips. These cells are groups is many. This arrangement of
called passage cells. Their function is to xylem is called polyarch. Xylem is in
Anatomical differences between dicot root and monocot root

S.No. Characters Dicot root Monocot root
1. Pericyle Gives rise to lateral roots, Gives rise to lateral roots
phellogen and a part of only.
vascular cambium.
2. Vascular tissue Usually limited number of Usually more number of
xylem and phloem strips. xylem and phloem strips,
3. Conjunctive Parenchymatous; Its cells are Mostly sclerenchymatous
tissue differentiated into vascular but sometimes
cambium. parenchymatous. It is never
differentiated in to vascular
cambium.
4. Cambium It appears as a secondary It is altogether absent.
meristem at the time of
secondary growth.
5. xylem Usually tetrach Usually polyarch
26

exarch condition, the tissue which is Inner to the hypodermis, a few layers
present between the xylem and the of collenchyma cells are present. This zone
phloem, is called conjunctive tissue. In is called hypodermis. It gives mechanical
maize, the conjunctive tissue is made up strength to the stem. These cells are living
of sclerenchymatous tissue. and thickened at the corners. Inner to the
hypodermis, a few layers of chlorenchyma
Pith cells are present with conspicuous
The central portion is occupied by a large intercellular spaces. This region performs
pith. It consists of thin-walled parenchyma photosynthesis. Some resin ducts also
cells with intercellular spaces. These cells occur here. The third zone is made up of
are filled with abundant starch grains. parenchyma cells. These cells store food
materials. The innermost layer of the cortex
Anatomy of Dicot and Monocot Stems is called endodermis. The cells of this layer
The transverse section of the dicot stem are barrel shaped and arrange compactly
[sunflower] shows the following plan of without intercellular spaces. Since starch
arrangement of tissues from the periphery grains are abundant in these cells, this
to the centre. layer is also known a starch sheath. This
layer is morphologically homologous
Epidermis to the endodermis found in the root. In
It is protective in function and forms the most of the dicot stems, endodermis with
outermost layer of the stem. It is a single casparian strips is not developed.
layer of parenchymatous rectangular cells.
The cells are compactly arranged without Check Your Grasp!
intercellular spaces. The outer walls of Why the endodermis in dicot stem is
epidermal cells have a layer called cuticle. also referred to as the starch sheath?
The cuticle checks the transpiration. The The cells of the endodermis are
cuticle is made up of waxy substance rich in starch grains and thus this layer
known as cutin. Stomata may be present is also referred to as the starch sheath.
here and there. Epidermal cells are living.
Chloroplasts are usually absent. A large
number of multicellular hairs occur on Stele
the epidermis. The central part of the stem inner to the
endodermis is known as stele. It consists
Cortex of pericyle, vascular bundles and pith. In
Cortex lies below the epidermis. The dicot stem, vascular bundles are arranged
cortex is differentiated into three zones. in a ring around the pith. This type of stele
Below the epidermis, there are few layers is called eustele.
of collenchyma cells. This zone is called
hypodermis. It gives mechanical strength Pericycle
of the Stem. These cells are living and Pericycle is the layers of cells that occur
thickened at the corners. between the endodermis and vascular
bundles. In the stem of sunflower
27

(Helianthus),a few layers of sclerenchyma Pith
cell occur in patches outside the phloem The large central portion of the stem is
in each vascular bundle. This patch of called pith. It is composed of parenchyma
sclerenchyma cell is called Bundle cap cells with intercellular spaces. The pith
or Hardbast. The bundle caps and the is also known as medulla. The pith
parenchyma cells between them constitute extends between the vascular bundles.
the pericycle in the stem of sunflower. These extensions of the pith between the
Vascular Bundles vascular bundles are called primary pith
rays or primary medullary rays. Function
The vascular bundles consist of xylem, of the pith is storage of food.
phloem and cambium. Xylem and phloem
in the stem occur together and form the Epidermal hair
Epidermis
vascular bundles. These vascular bundles
Hypodermis
are Wedge shaped. They are arranged in Cortex
the form of a ring. Each vascular bundle is Endodermis
Pericycle
conjoint, collateral, open and endarch. Vascular bundle
Pith
Phloem Ground plan
Primary phloem lies towards the Epidermal hair
periphery. It consists of protophloem Cuticle
Epidermis
and metaphloem. Phloem consists of Collenchyma
sieve tubes, companion cells and phloem Chlorenchyma
Resin duct
parenchyma. Phloem fibres are absent in
Parenchyma
the primary phloem. Phloem conducts Starch sheath
organic food materials from the leaves to Bundle cap
Primary phloem
other parts of the plant body. Cambium
Metaxylem
Cambium Protoxylem
Cambium consists of brick shaped and Pith

thin walled meristematic cells. It is one Primary
medullary ray
to four layers in thickness. These cells A sector enlarged
are capable of forming new cells during Figure 9.21: T. S of Dicot Stem
secondary growth.
(Sunflower stem)
Xylem
Xylem consists of xylem fibres, xylem Primary Structure of Monocot
parrenchyma vessels and tracheids. Stem-maize Stem
Vessels are thick walled and arranged in The outline of the maize in transverse section
a few rows. is more or less circular. The transverse
Xylem conducts water and minerals section of the monocot stem [maize] shows
from the root to the other parts of the the following plan of arrangement of tissues
plant body. from the periphery to the centre.
28

Epidermis mass of parenchyma cells lying inner to
Hypodermis the hypodermis forms the ground tissue.
Ground tissue
Vascular The cell wall is made up of cellulose.
bundles The cells contain reserve food material
like starch. The cells of the ground tissue
Ground plan next to the hypodermis are smaller in size,
polygonal in shape and compactly arranged.
Cuticle
Epidermis Towards the centre, the cells are loosely
Hypodermis arranged, rounded in shape and bigger in
Chlorenchyma
size. The vascular bundles lie embedded
in this tissue. The ground tissue stores
Vascular bundle food and performs gaseous exchange.
Ground tissue
Vascular Bundles
Crushed protophloem Vascular bundles are scattered
Metaphloem
Metaxylem (atactostele) in the parenchymatous
Protoxylem ground tissue. Each vascular bundle is
Protoxylem lacuna
surrounded by a sheath of sclerenchy-
Sclerenchymatous
bundle sheath matous fibres called bundle sheath. The
A sector enlarged vascular bundles are conjoint, collateral,
endarch and closed.Vascular bundles are
Figure 9.22: T.S. Monocot stem
numerous, small and closely arranged in
(Maize stem)
the peripheral portion. Towards the cen-
Epidermis tre, the bundles are comparatively large in
size and loosely a rranged. Vascular bun-
It is the outermost layer of the stem. It is
dles are skull or oval shaped.
made up of single layer of tightly packed
parenchymatous cells. Their outer walls are Phloem
covered with thick cuticle. The continuity
The phloem in the monocot stem consists
of this layer may be broken here and there
of sieve tubes and companion cells. Phloem
by the presence of a few stomata. There
parenchyma and phloem fibres are absent. It
are no epidermal outgrowths.
can be distinguished into an outer crushed
Hypodermis protophloem and an inner metaphloem.
A few layer of sclerenchymatous cells Xylem
lying below the epidermis constitute the
hypodermis. This layer gives mechanical Xylem vessels are arranged in the form
strength to the plant. It is interrupted here of ’Y’ the two metaxylem vessels are
and there by chlorenchyma cells. located at the upper two arms and one
or two protoxylem vessels at the base. In
Ground Tissue a mature bundle, the lowest protoxylem
There is no distinction into cortex, disintegrates and forms a cavity known as
endodermis, pericycle and pith. The entire protoxylem lacuna.
29

Table 9.4: Anatomical differences between dicot stem and monocot stem
S.No. Characters Dicot Stem Monocot Stem
1. Hypodermis Collenchymatous Sclerenchymatous
2. Ground tissue Differentiated into cortex, Not differentiated, but it
endodermis and pericycle is a continuous mass of
and pith parenchyma.
3. Starch Sheath Present Absent
4. Medullary rays Present Absent
5. Vascular (a) Collateral and open (a) Collateral and closed
bundles (b) Arranged in a ring (b) Scattered in ground tissue
(c) Secondary growth occurs (c) Secondary growth usually
does not occur.
Table 9.5: Anatomical differences between root and stem

S.No. Characters Root Stem
1. Epidermis Absence of cuticle and Presence of cuticle and
epidermal pores. epidermal pores.
Presence of unicellular root Presence of unicellular and
hairs. multicellular trichomes
2. Outer Cortical Chlorenchyma absent Chlorenchyma present
cells
3. Endodermis Well defined ill-defined or absent.
4. Vascular Radial arrangement Conjoint arrangement
bundles
5. Xylem Exarch Endarch
Anatomy of a Dicot and Monocot Leaves In dorsiventral leaves the mesophyll

Leaves are very important vegetative is differentiated into palisade and spongy
organs. They are mainly concerned with parenchyma, the former occurring on the
photosynthesis and transpiration. Like upper side and the later on the lower side
stem and roots, leaves also have the Example: Sunflower. In isobilateral leaf
three tissue system – dermal, ground palisade is present on both sides of the leaf
and vascular. The dermal tissue system and inbetween them spongy parenchyma
consists of an upper and lower epidermis. is present. Example: Nerium. In some
The ground tissue system that lies between plants Example: Ficus calcium crystals
the epidermal layers of leaf is known as are present. There are also leaves where
mesophyll tissue. Often it is differentiated spongy tissue alone is present in some
into palisade parenchyma on the adaxial epidermal cells Example: Grasses.
(upper) side and spongy parenchyma on The presence of air spaces is a special
the abaxial (lower) side. feature of spongy cells. They facilitate the
30

gaseous exchange between the internal surrounded by a pair of bean shaped
photosynthetic tissue (mesophyll) and the cells called guard cells.
external atmosphere through the stomata. Each stoma internally opens into an
The vascular tissue system is composed air chamber. These guard cells contain
of vascular bundles. They are collateral chloroplasts, whereas other epidermal
and closed. The vascular tissues forms cells do not contain chloroplasts. The
the skeleton of the leaf and are known as main function of the epidermis is to
veins. The veins supply water and minerals give protection to the inner tissue called
to the photosynthetic tissue. Thus the mesosphyll. The cuticle helps to check
morphological and anatomical features of transpiration. Stomata are used for
the leaf help in its physiological functions. transpiration and gas exchange.
Mesophyll
The entire tissue between the upper and
lower epidermis is called the mesophyll
(GK meso = in the middle, phyllome
= leaf). There are two regions in the
mesophyll. They are palisade parenchyma
and spongy parenchyma. Palisade
parenchyma cells are seen beneath the upper
epidermis. It consists of vertically elongated
cylindrical cells in one or more layers.
These cells are compactly arranged and
Figure 9.23: Anatomy of Leaf
are generally without intercellular spaces.
Palisade parenchyma cells contain more
Anatomy of a Dicot Leaf-sunflower Leaf
chloroplasts than the spongy parenchyma
Internal structure of dictoyledonous cells. The function of palisade parenchyma
leaves reveal epidermis, Mesophyll and is photosynthesis. Spongy parenchyma lies
vascular tissues. below the palisade parenchyma. Spongy cells
are irregularly shaped. These cells are very
Epidermis
loosely arranged with numerous airspaces.
This leaf is generally dorsiventral. It As compared to palisade cells, the spongy
has upper and lower epidermis. The cells contain lesser number of chloroplasts.
epidermis is usually made up of a single Spongy cells facilitate the exchange of gases
layer of cells that are closely packed. with the help of air spaces. The air space
The cuticle on the upper epidermis is that is found next to the stomata is called
thicker than that of lower epidermis. The respiratory cavity or substomatal cavity.
minute openings found on the epidermis
are called stomata. Stomata are more Vascular Tissues
in number on the lower epidermis than Vascular tissues are present in the veins
on the upper epidermis. A stomata is of leaf. Vascular bundles are conjoint,
31

Collateral and closed. Xylem is present Xylem consists of metaxylem and
towards the upper epidermis, while the protoxylem elements. Protoxylem is
phloem towards the lower epidermis. present towards the upper epidermis,while
Vascular bundles are surrounded by the phloem consists of sieve tubes,
a compact layer of parenchymatous companion cells and phloem parenchyma.
cells called bundle sheath or border Phloem fibres are absent. Xylem consists of
parenchyma. vessels and xylem parenchyma. Tracheids
and xylem fibres are absent.
Cuticle
Upper epidermis
Palisade parenchyma
Protoxylem
Metaxylem
Spongy parenchyma
Phloem
Bundle sheath
Stoma
Epidermal hair
Lower epidermis
Respiratory cavity
Figure 9.24: T.S. of Dicot Leaf (Sunflower)
Anatomy of a Monocot Leaf – Grass Leaf These cells are called subsidiary cells.
A transverse section of a grass leaf reveals Some cells of upper epidermis are large
the following internal structures. and thin walled. They are called bulliform
cells or motor cells. These cells are helpful
Epidermis for the rolling and unrolling of the leaf
The leaf has upper and lower epidermis. according to the weather change.
They are made up of a single layer of thin Some of the epidermal cells of the
walled cells. The outer walls are covered grass are filled with silica. They are called
by thick cuticle. silica cells.
The number of stomata is more or less
Mesophyll
equal on both the epidermis. The stomata
is surrounded by dumb – bell shaped The ground tissue that is present between
guard cells. The guard cells-contain the upper and lower epidermis of the leaf
chloroplasts, whereas the other epidermal is called mesophyll. Here, the mesophyll
cells do not have them. is not differentiated into palisade and
Some special cells surround the spongy parenchyma. All the mesophyll
guard cells. They are distinct from other cells are nearly isodiametric and thin
epidermal cells. walled. These cells are compactly arranged
32

with limited intercellular spaces. They have such sclerenchymatous patches.
contain numerous chloroplasts. Vascular bundles are conjoint, collateral
and closed. Each vascular bundle is
Vascular Bundles surrounded by a parenchymatous bundle
Vascular bundles differ in size. Most sheath. The cells of the bundle sheath
of the vascular bundles are smaller in generally contain starch grains. The
size. Large bundles occur at regular xylem of the vascular bundle is located
intervals.Two patches of sclerenchyma towards the upper epidermis and the
are present above and below the large phloem towards the lower epidermis.In
vascular bundles. These sclerenchyma C 4 grasses, the bundle sheath cells are
patches give mechanical support to the living and involve in C 4 photosynthesis.
leaf. The small vascular bundles do not This sheath is called Kranz sheath.
Cuticle
Upper epidermis
Sub-stomatal chamber
MesophyII
Bundle sheath
Xylem
Phloem
Lower epidermis
Stoma
Figure 9.25: T.S. of monocot leaf (Grass)
Water Stomata (or) Hydathodes chloroplasts. These cells open out into one
A hydathode is a type of epidermal pore, or more sub-epidermal chambers. These,
commonly found in higher plants. in turn, communicate with the exterior
through an open pore. The water stoma
Structurally, hydathodes are modified
structurally resembles an ordinary stoma,
stomata, usually located at leaf tips or
but is usually larger and has lost the power
margins, especially at the teeth.
of movement.They are connected to the
Hydathodes occur in the leaves plant vascular system by a tracheid or
of submerged aquatic plants such as vessel element.
Ranunculus fluitans as well as in many
Hydathodes discharge liquid water
herbaceous land plants.
with various dissolved substances from
Hydathodes are made of a group of the interior of the leaf to its surface. This
living cells with numerous intercellular process is called guttation. Example many
spaces filled with water, but few or no grasses.
33

Differences Between Stomata and Halophiles
Hydathodes • Plants that grow in salty environment
are called Halophiles.
Stomata Hydathodes
• Plant growth in saline habitat developed
Occur in epidermis Occur at the tip or numerous adaptations to salt stress.
of leaves, young margin of leaves The secretion of ions by salt glands is the
stems. that are grown in best known mechanism for regulating
moist shady place. the salt content of plant shoots.
• Salt glands typically are found in
Stomatal aperture Aperture of
halophytes. (Plants that grow in saline
is guarded by two hydathodes are
environments)
guard cells. surrounded
by a ring of
cuticularized cells.
The two guard Subsidiary cells
cells are generally are absent.
surrounded by
subsidiary cell.
Opening and closing Hydathode pores
of the stomatal remain always
aperture is regulated open.
by guard cells.
These are involved These are involved
in transpiration and in guttation.
exchange of gases. Figure 9.26: Halophytes
Can mangroove trees

grow in salt water?
These amazing trees
and shrubs cope with
salt. Salt water can kill Plants, so
mangroves must extract fresh water
from the sea water that surrounds
them. Many mangrove species survive
by filtering out as much as 90 percent
of the salt found in seawater as it
enters their roots.
Mangrove excrete salt through
glands in their leaves. Figure 9.27: Removes excess salts
through special salt glands on leaves
34

Summary root, xylem is tetrach. Its phloem patch
consists of sieve tubes, companion cells
A Tissue is a group of cells that are alike in
and phloem parenchyma. In monocot
origin, structure and function.There are two
(Example: maize) root, xylem is polyarch.
principal groups: (1) Meristematic tissues
and (2) Permanent tissues. Meristematic In dicot (Example: sunflower) stem,
tissues comprise of self-perpetuating cells. stele is eustele type and its vascular
Meristems are classified into several types bundles are wedge shaped, conjoint,
on the basis of position, origin, function collateral, open and endarch. In monocot
and activity. Many anatomists illustrated stem (Example: maize) vascular bundles
the root and shoot apical meristems on are scattered and skull shaped, conjoint,
the basis of the type and arrangement and collateral, closed and endarch.
accordingly proposed many theories. The In dicot (Example: sunflower) and
permanent tissues normally develop from monocot (Example: grass) leaves vascular
apical meristem. They are classified into bundles are conjoint, collateral and closed.
two types: 1)Simple permanent tissues Hydathodes discharge liquid water
and 2)Complex permanent tissues. Simple with various dissolved substances from
tissues are composed of a single type of cells the interior of the leaf to its surface. Plants
only. It is of three types: (1) Parenchyma that grow in salty environment are called
(2) Collenchyma and (3) Sclerenchyma. A halophiles. Salt glands typically are found
complex tissue is a tissue with several types in halophytes.
of cells but all of them function together as
a single unit. It is of two types – xylem and
Evaluation
phloem. Secretory tissues produce different
types of chemicals. Some are in the form of 1. Refer to the given figure and select
enzymes, hormones, rubber, gum etc. the correct statement
The tissues can be classified on the basis
of their function, structure and location
A
into epidermal tissue system, ground B
tissue system and vascular tissue system. C
Epidermal tissue system develops as the
outermost covering of the entire plant
body. It consists of epidermal cells and i. A, B, and C are histogen of shoot
associated structures. All tissues except apex
epidermis and vascular tissues constitute
ii. A Gives rise to medullary rays.
the ground tissue. The vascular tissue
system is formed of vascular bundles. iii. B Gives rise to cortex
iv. C Gives rise to epidermis
In the primary structure, the outermost
layer of the root is called piliferous layer. a. i and ii only
Cortex consists of only parenchyma cells. b. ii and iii only
All the tissues present inside endodermis c. i and iii only
comprise the stele. In dicot (Example: bean) d. iii and iv only
35

2. Read the following sentences and 5. Grafting is successful in dicots but
identify the correctly matched not in monocots because the dicots
sentences. have
i. In exarch condition, the a. Vascular bundles arranged in a ring
protoxylem lies outside of b. Cambium for secondary growth
metaxylem. c. Vessels with elements arranged end
ii. In endarch condition, the to end
protoxylem lie towords the centre. d. Cork cambium
iii. In centarch condition, metaxylem 6. Why the cells of sclerenchyma and
lies in the middle of the tracheids become dead?
protoxylem.
7. Explain sclereids with their types.
iv. In mesarch condition, protoxylem
8. What are sieve tubes ?explain.
lies in the middle of the metaxylem.
9. Distinguish the anatomy of dicot root
a. i, ii and iii only
from monocot root.
b. ii, iii and iv only
10. Distinguish the anatomy of dicot stem
c. i, ii and iv only from monocot stem.
d. All of these
3. In Gymnosperms, the activity of sieve
tubes are controlled by
a. Nearby sieve tube members.
b. Phloem parenchyma cells
c. Nucleus of companion cells.
d. Nucleus of albuminous cells.
4. When a leaf trace extends from a
vascular bundle in a dicot stem, what
would be the arrangement of vascular
tissues in the veins of the leaf?
a. Xylem would be on top and the
phloem on the bottom
b. Phloem would be on top and the
xylem on the bottom
c. Xylem would encircle the phloem
d. Phloem would encircle the xylem
36

t ICT Corner
Plant and Animal Tissues
Let’s explore Plant tissues.
Steps
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• Type online labs and install it.
• Select biology and select plant and animal tissues
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• Choose theory tab to know the basic about anatomical structure
• Choose animation to view the sectioning process
Activity
• Choose simulation tab and view the section of plant parts under microscope
Step 2
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37

Chapter
10 Secondary Growth
We have studied in the previous chapters

Learning Objectives the primary internal structure of monocots
The students should be able to, and dicots. If you look at the stem of grass
(monocot), it is soft, whereas in the neem
• Analyze primary and secondary
(dicot), the stem is very hard and woody,
growth.
why? It is the secondary growth which
• Discuss the increase confers the hardness to wood of dicot stems
in length and width and roots. In monocots, usually there is no
of the plant.
secondary growth and so they are soft.
• Explain secondary
The increase in girth is called
growth in dicot stems.
secondary growth or growth in girth and
• Understand the use of wood we shall discuss the details of secondary
products to lead comfortable life. growth in this chapter.
• Explain secondary growth in dicot The plant organs originating from the
roots.
apical meristems pass through a period
• Discuss anomalous secondary of expansion in length and width. The
growth in dicots and monocots. roots and stems grow in length with the
• Explain the seasoning, grain, help of apical meristems. This is called
texture and figure of wood. primary growth or longitudinal growth.
The gymnosperms and most angiosperms,
Chapter Outline including some monocots, show an increase
10.1 Secondary Growth in Dicot Stem in thickness of stems and roots by means of
secondary growth or latitudinal growth.
10.2 Secondary Growth in Dicot Root
The secondary growth in dicots and
10.3 Anomalous Secondary Growth
gymnosperms is brought about by two
10.4 Timber lateral meristems.
How do the trees increase their girth? • Vascular Cambium and
• Cork Cambium
Activity
Generally monocots do not have
secondary growth, but palms and
bamboos have woody stems. Find the
Figure 10.1: Taxus wood reason.
38

10.1 Secondary Growth in Dicot Stem The cells of vascular cambium do not fit
into the usual description of meristems
Vascular Cambium
which have isodiametric cells, with a dense
The vascular cambium is the lateral cytoplasm and large nuclei. While the active
meristem that produces the secondary vascular cambium possesses cells with large
vascular tissues. i.e., secondary xylem and central vacuole (or vacuoles) surrounded
secondary phloem. by a thin, layers of dense cytoplasm.
Origin and Formation of Vascular Further, the most important character
Cambium of the vascular cambium is the presence
of two kinds of initials, namely, fusiform
A strip of vascular cambium that is
initials and ray initials.
believed to originate from the procambium
is present between xylem and phloem of Fusiform Initials
the vascular bundle. This cambial strip is These are vertically elongated cells. They
known as intrafascicular or fascicular give rise to the longitudinal or axial
cambium. In between the vascular system of the secondary xylem (treachery
bundles, a few parenchymatous cells of elements, fibers, and axial parenchyma)
the medullary rays that are in line with the and phloem (sieve elements, fibers, and
fascicular cambium become meristematic axial parenchyma).
and form strips of vascular cambium. It is Based on the arrangement of the
called interfascicular cambium. fusiform initials, two types of vascular
This interfascicular cambium joins cambium are recognized.
with the intrafascicular cambium on both Storied (Stratified cambium) and
sides to form a continuous ring. It is called Non-Storied (Non-stratified cambium)
a vascular cambial ring. The differences
between interfascicular and intrafascicular
cambia are summarised below: Ray initials
Intrafascicular Interfascicular Fusiform initials
cambium cambium
Present inside the Present in between a
vascular bundles the vascular

Ray initials
bundles.
Originates from Originates from
Fusiform initials
the procambium. the medullary rays.
Initially it forms a From the
part of the primary beginning it b
meristem. forms a part of Figure 10.2: Tangential longitudinal

the secondary
section (TLS) of cambium (a) Storied
meristem.
cambium (b) Non-storied cambium
Organization of Vascular Cambium If the fusiform initials are arranged in
39

horizontal tiers, with the end of the cells phloem and inward secondary xylem.
of one tier appearing at approximately At places, cambium forms some
the same level, as seen in tangential narrow horizontal bands of parenchyma
longitudinal section (TLS), it is which passes through secondary phloem
called storied (stratified) cambium. and xylem. These are the rays.
It is the characteristic of the plants with Due to the continued formation of
short fusiform initials. Whereas in plants secondary xylem and phloem through
with long fusiform initials, they strongly vascular cambial activity, both the primary
overlap at the ends, and this type of xylem and phloem get gradually crushed.
cambium is called non-storied (non-
startified) cambium. Secondary Xylem
Ray Initials The secondary xylem, also called wood, is
These are horizontally elongated cells. formed by a relatively complex meristem,
They give rise to the ray cells and form the the vascular cambium, consisting of
elements of the radial system of secondary vertically (axial) elongated fusiform initials
xylem and phloem. and horizontally (radially) elongated ray
Activity of Vascular Cambium initials.
The vascular cambial ring, when active,

Xylotomy
cuts off new cells both towards the
inner and outer side. The cells which The study of wood
are produced outward form secondary by preparing sections
for microscopic
a A Portion of cambium
observation.
Cambium
b
First formed xylem
The axial system consists of vertical files
Cambium of treachery elements, fibers, and wood
c Second formed xylem parenchyma. Whereas the radial system
First formed xylem consists of rows of parenchymatous cells
Cambium oriented at right angles to the longitudinal
d
Third formed xylem
Second formed xylem
axis of xylem elements.
First formed xylem The secondary xylem varies very greatly
First formed phloem from species to species with reference to
Cambium relative distribution of the different cell
e Third formed xylem
Second formed xylem types, density and other properties. It is
First formed xylem
of two types.
First formed phloem
Second formed phloem
Cambium
Porous Wood or Hard Wood
Fourth formed xylem
f Third formed xylem Generally, the dicotyledonous wood,
Second formed xylem
First formed xylem
which has vessels is called porous wood
or hard wood. Example: Morus rubra.
Figure 10.3: Diagrammatic representa-
tion of vascular cambial activity (a–f) Non- Porous Wood or Soft Wood
40

Generally, the gymnosperm wood, which Differences between Porous Wood and
lacks vessels is known as non- porous Non-porous Wood
wood or soft wood. Example: Pinus.
Porous wood or Non porous wood or
Hard wood, Soft wood,
Example: Morus Example: Pinus
Common in Common in
a b angiosperms gymnosperms
Figure 10.4: Structure of porous (a) Porous because it Non-porous because
contains vessels it does not contain
and non-porous wood (b)
vessels
Epidermis
Cortex
Bundle cap
Primary Phloem
Fascicular cambium
Primary Xylem
Inter fascicular cambium
Pith
a b
Phellem (Cork)
Phellogen (Cork cambium)
Phelloderm
(Secondary cortex)
Secondary Phloem
Vascular
cambial ring
Secondary xylem
Medullary ray
c d
Figure 10.5: Secondary growth in dicot stem (diagrammatic) -

stages in transverse section (a–d)
41

Lenticel
Complementary cells
Epidermis
Cork (phellem)
Cork cambium (Phellogen)
Secondary cortex (Phelloderm)
Collenchyma
Parenchyma
Secondary phloem
Cambium
Autumn wood (secondary xylem)
Spring wood (secondary xylem)
Secondary medullary ray
Primary xylem
Pith
Figure 10.6: Secondary growth in two year old dicot stem – A portion enlarged
42

Primary and Secondary Growth
Primary growth in stem
Growth of plant derived from shoot apical
meristem increases the length of stem
Ground meristem
Pro
ocam
oca
am
Procambium
Primary
P rim
xylem
xxy
y le
Apical Prima
Primary
growth phloem
p h lo
oee
Secondary growth
th
h iin stem
t
In gymnosperm and most
angiosperm plants, the lateral meristem
of stem produces an increase in the girth
Vascular Cork cambium

cambium Secondary
xylem
y m
Primary
Prim
maarry
phloem
phlooeeem
m
Primary
Prim
maar
Late
Lateral xylem
xylem
grow
growth Secondary
Seconnd
phloem
phloe
phlo
hloe
oe
Secondary growth in root

In gymnosperm and most angiosperm
plants the lateral meristem of root
produces an increase in the girth
Vascular
V
Va
ascularr
ascular Primary
P
Prim a
ccambium
am
ambium
bi m xyle
y eem
xylemm
Secondary
Seco
SSe on
onndd
xylem
xxy
ylemm
Primary
Pr
rm
ri
APICAL AND phloem
ph
hlo
LATERAL MERISTEMS. Lateral
growth Secondary
SSe
ecoon
onnd
d
Apical meristems phloem
p
ph
hloeeem
hlo
hl
h m
produce the primary plant
body. In some plants, the
lateral meristem Primary growth in root
increase the girth of a plant. Growth of plant derived from root apical
This type of growth is secondary meristem increases the length of root
because the lateral meristem are Primary
not directly produced by apical xylem
meristems. Woody plants have two
Primary
types of lateral meristems: a vascular phloem
cambium that produces xylem,
Apical
phloem tissues and cork cambium that Ground meristem
growth
produces the bark of a tree.
Procambium
43

Annual Rings Sometimes annual rings are called growth
The activity of vascular cambium is rings but it should be remembered all the
under the control of many physiological growth rings are not annual. In some trees
and environmental factors. In temperate more than one growth ring is formed with
regions, the climatic conditions are not in a year due to climatic changes.
uniform throughout the year. In the Additional growth rings are
spring season, cambium is very active developed within a year due to adverse
and produces a large number of xylary natural calamities like drought, frost,
elements having vessels/tracheids with defoliation, flood, mechanical injury
wide lumen. The wood formed during and biotic factors during the middle
this season is called spring wood or early of a growing season,which results in
wood. The tracheary elements are fairly the formation of more than one annual
thin walled. In winter, the cambium is less ring. Such rings are called pseudo- or
active and forms fewer xylary elements false- annual rings.
that have narrow vessels/ tracheids and Each annual ring corresponds to one
this wood is called autumn wood or late year’s growth and on the basis of these
wood.The treachery elements are with rings, the age of a particular plant can
narrow lumen, very thick walled. easily be calculated. The determination of
the age of a tree by counting the annual
• Usually more
rings is called dendrochronology.
distinct annual
rings are formed in
Importance of Studying Growth Rings
the regions where
climatic variations are sharp. • Age of wood can be calculated.
• Usually more distinct annual rings • The quality of timber can be
are formed in temperate plants ascertained.
and not in tropical plants. • Radio-Carbon dating can be
• Usually least distinct annual rings verified.
are formed in seashore region
• Past climate and archaeological
because the climatic conditions
dating can be made.
remain same throughout the year.
• Generally annual rings are also • Provides evidence in forensic
less distinct in desert plants. investigation.
The spring wood is lighter in colour and Dendroclimatology

has a lower density whereas the autumn It is a branch of dendrochronology
wood is darker and has a higher density. concerned with constructing records
The annual ring denotes the of past climates and climatic events by
combination of early wood and late wood analysis of tree growth characteristics,
and the ring becomes evident to our eye especially growth rings.
due to the high density of late wood.
44

Heart wood Another feature of wood related to
Sap wood seasonal changes is the diffuse porous and
Early wood ring porous condition. On the basis of
Late wood
diameter of xylem vessels, two main types
of angiosperm woods are recognized.
 Diffuse porous woods
Diffuse porous woods are woods
in which the vessels or pores are
rather uniform in size and distribution
throughout an annual ring.
Figure 10.7: Structure of wood – Image Example: Acer
shows early wood and late wood  Ring porous woods
Differences Between Spring Wood and The pores of the early wood are
distinctly larger than those of the late
Autumn Wood
wood. Thus rings of wide and narrow
Spring wood or Autumn wood or vessels occur.
Early wood Late wood Example: Quercus
The activity of Activity of
cambium is faster. cambium is Late wood
slower
Early wood
Produces large Produces fewer
Rays
number of xylem xylem elements.
elements. Tyloses
Xylem vessels/ Xylem vessels/

trachieds have trachieds have
wider lumen. narrow lumen.
a b
Wood is lighter Wood is darker in
Figure 10.8: Transverse section of wood
in colour and has colour and has a showing. a. Diffuse porous. b. Ring porous
lower density higher density.
The word “Porous”

is used by the wood
anatomists to refer
to the appearance
of the vessels as pores in transverse
The age of American,
section.
Sequoiadendron tree
is about 3500 years.
45

Differences Between Diffuse Porous Usually, these structures are formed
Wood and Ring Porous Wood in secondary xylem vessels that have last
Diffuse porous Ring porous their function i.e., in heart wood.
wood wood In fully developed tyloses, starchy
This type of This type of wood crystals, resins, gums, oils, tannins or
wood is formed is formed where the coloured substances are found.
where the climatic climatic conditions
conditions are are not uniform. There are tylosoids
uniform. in gymnosperms and
The vessels are The vessels are wide angiosperms
more or less equal and narrow within In gymnosperms,
in diameter in any any annual ring. the resin ducts are blocked by tylose-
annual ring. like ingrowths from the neighbouring
resin producing parenchyamatous
The vessels The vessels are cells. Example: Pinus.
are uniformly not uniformly In angiosperms, the sieve tubes are
distributed distributed blocked by tylose- like ingrowths from
throughout the throughout the the neighbouring parenchyamatous
wood. wood. cells. Example: Bombox.
These are called tylosoids
Tyloses
In many dicot plants, the lumen of the Wood is also classified into sap wood and
xylem vessels is blocked by many balloon- heart wood.
like ingrowths from the neighbouring
parenchymatous cells. These balloon-like Sap Wood and Heart Wood
structures are called tyloses. Sap wood and heart wood can be
distinguished in the secondary xylem.
In any tree the outer part of the wood,
which is paler in colour, is called sap
wood or alburnum. The centre part of the
Parenchyma cell
wood, which is darker in colour is called
heart wood or duramen. The sap wood
Tyloses conducts water while the heart wood
stops conducting water. As vessels of the
heart wood are blocked by tyloses, water
is not conducted through them. Due to
the presence of tyloses and their contents
Vessel wall the heartwood becomes coloured, dead
and the hardest part of the wood.
Vessel lumen From the economic point of view,
generally the heartwood is more useful
Figure 10.9: Structure of tyloses
46

than the sapwood. The timber from the
heartwood is more durable and more When, the heart wood
resistant to the attack of microorganisms of a tree is destroyed,
and insects than the timber from no vital function of
sapwood. the plant is affected.
When, the sap wood is destroyed,
the plant will die because conduction
of water will be blocked.
Differences Between Sap Wood

(alburnum) and Heart Wood (duramen)
Sap Wood Heart Wood
(Alburnum) (Duramen)
Living part of the Dead part of the
Figure 10.10: Cross section of wood. wood.
natural wood It is situated on It is situated in
the outer side of the centre part of
Bark
wood wood
Sap wood It is less in It is dark in
coloured coloured
Heart wood Very soft in nature Hard in nature
Annual rings Tyloses are absent Tyloses are
present
It is not durable It is more durable
and not resistant and resists
to microorganisms microorganisms
Figure 10.11: Cross - section of wood
showing annual ring
Abies balsamea is a
The dye, haematoxylin is
gymnospermic plant. It
obtained from the heart
produces canada balsam,
wood of Haematoxylum
from its resin ducts.
campechianum used to stain
It is used as mounting
plant materials for observation
medium for microscopic
under microscope, especially
slide preparation.
the nucleus of the cell.
a. Haematoxylin b. Canada balsam
47

Fossil resins-Amber
Plants secrete resins for their
protective benefits.Amber
is a fossilized tree resin
especially from the wood,
which has been appreciated
c. Microscopic slide
for its colour and natural
A slide of 60-years-old holotype specimen of beauty since neolithic times.
a flatworm (Lethacotyle fijiensis) permanently d. Ant inside Much valued from antiquity
mounted in canada balsam. to the present as a gemstone,
blastic amber
amber is made into a variety
of decorative objects. Amber
is used in jewellery. It has
also been used as a healing
agent in folk medicine.
Figure 10.12: Economic importance of wood (a–d)

Secondary Phloem iii. Sun hemp-Crotalaria juncea
The vascular cambial ring produces iv. Jute-Corchorus capsularis
secondary phloem or bast on the outer
side of the vascular bundle. Be friendly with your environment
Just as the secondary xylem, the (Eco friendly)
secondary phloem also has two tissue Why should not we use the natural
systems – the axial (vertical) and the products which are made by plant
radial (horizontal) systems derived fibres like rope, fancy bags, mobile
respectively from the vertically elongated pouch, mat and gunny bags etc.,
fusiform initials and horizontally instead of using plastics or nylon?
elongated ray initials of vascular
cambium. While sieve elements, phloem
fibre, and phloem parenchyma represent Periderm
the axial system, phloem rays represent Whenever stems and roots increase
the radial system. Life span of secondary in thickness by secondary growth, the
phloem is less compared to secondary periderm, a protective tissue of secondary
xylem. Secondary phloem is a living origin replaces the epidermis and often
tissue that transports soluble organic primary cortex. The periderm consists of
compounds made during photosynthesis phellem, phellogen, and phelloderm.
to various parts of plant.
Some commercially important phloem Phellem (Cork)
or bast fibres are obtained from the It is the protective tissue composed of
following plants. non-living cells with suberized walls and
i. Flax-Linum ustitaissimum formed centrifugally (outward) by the
phellogen (cork cambium) as part of the
ii. Hemp-Cannabis sativa
periderm. It replaces the epidermis in
48

older stems and roots of many seed plants. Differences Between Phellem and
It is characterized by regularly arranged Phelloderm
tiers and rows of cells. It is broken here Phellem (Cork) Phelloderm
and there by the presence of lenticels. (Secondary
cortex)
Cuticle
Epidermis It is formed on It is formed on
First cork cell the outer side of the inner side of
Phellogen
(Cork cambium) phellogen. phellogen.
Cortex
a
Cells are Cells are loosely
Cuticle compactly arranged with
Epidermis
Phellem(Cork)
arranged in intercellular
Phellogen regular tires and spaces.
(Cork cambium)
Phelloderm rows without
(Secondary cortex)
Cortex
intercellular
b spaces.
Figure 10.13: The cross section of Protective in As it contains
function. chloroplast, it
periderm (a–b)
synthesises and
stores food.
Phelloids Consists of non- Consists of
Phellem (Cork) like cells which lack living cells with living cells,
suberin in their walls. suberized walls. parenchymatous
in nature and does
not have suberin.
Phellogen (Cork Cambium) Lenticels are Lenticels are
It is a secondary lateral meristem. It present. absent.
comprises homogenous meristematic cells
unlike vascular cambium. It arises from
epidermis, cortex, phloem or pericycle Rhytidome is a
technical term used
(extrastelar in origin). Its cells divide
for the outer dead
periclinally and produce radially arranged bark which consists of
files of cells. The cells towards the outer periderm and isolated
side differentiate into phellem (cork) and cortical or phloem tissues formed
those towards the inside as phelloderm during successive secondary growth.
(secondary cortex). Example: Quercus.
Polyderm is found in the roots
Phelloderm (Secondary cortex) and underground stems.eg. Rosaceae.
It is a tissue resembling cortical living It refers to a special type of protective
parenchyma produced centripetally tissues consisting of uniseriate
(inward) from the phellogen as a part of the suberized layer alternating with
periderm of stems and roots in seed plants. multiseriatenonsuberized cells in
periderm.
49

Differences Between Vascular barks normally do not peeled off, scale barks
Cambium and Cork Cambium peeled off.
Vascular cambium Cork cambium
Also called Also called
cambium phellogen
It arises from It arises from
procambium and epidermis, cortex,
interfascicular phloem, or
parenchyma in pericyle in both
stems and from stems and roots
conjunctive
parenchyma in
roots Figure 10.14: Quercus Tree-showing
It comprises long It comprises of ring bark
fusiform and homogenous cells.
short ray initials.
It produces It produces
secondary phloem phellem(cork)
towards the outer towards outer side
side and secondary and phelloderm
xylem towards (secondary cortex)
inner side. towards inner side.
Bark
Figure 10.15: Guava tree showing scale
The term ‘bark’ is commonly applied to all
bark
the tissues outside the vascular cambium
of stem (i.e., periderm, cortex, primary Lenticel
phloem and secondary phloem). Bark
Lenticel is raised opening or pore on the
protects the plant from parasitic fungi and
epidermis or bark of stems and roots.
insects, prevents water loss by evaporation
It is formed during secondary growth
and guards against variations of external
in stems. When phellogen is more active
temperature. It is an insect repellent, decay
in the region of lenticels, a mass of loosely
proof, fireproof and is used in obtaining
arranged thin-walled parenchyma cells
drugs or spices. The phloem cells of the bark
are formed. It is called complementary
are involved in conduction of food while
tissue or filling tissue.
secondary cortical cells involved in storage.
Lenticel is helpful in exchange of
If the phellogen forms a complete cylinder
gases and transpiration called lenticular
around the stem, it gives rise to ring barks.
transpiration.
Example: Quercus. When the bark is formed
in overlapping scale like layers, it is known
as scale bark. Example: Guava. While ring
50

Lenticel
Complementary cell
Epidermis
Phellem(Cork)
Phellogen
(Cork cambium)
Phelloderm
(Secondary cortex)
Figure 10.16: Structure of Lenticel
Know your Commercial Barks
Cinchona bark is Turpentine (Resin)

medicinally active, – obtained from bark
containing a variety of of Confiers, is used as
alkaloids including the thinner for oil based
antimalarial compound paints and organic
a. Quinine quinine. e. Turpentine solvents.
Cork is an impermeable Example: Pinus
buoyant material, the
phellem layer of bark
Cinnamon (Oldest
tissue that is harvested for Spice) – Its bark is
commercial use primarily used as ingredients
from Quercus suber. Cork of curry powder,
is composed of suberin, a
medicine for cardiac
hydrophobic substance and,
b. Cork because of its impermeable, f. Cinnamomum stimulant, diarrhoea
buoyant, elastic, and fire bark and vomiting. Example:
retardant properties, used as Cinnamomum
a bottle stoppers. zeylanicum
Cork is also used as an Gum Arabic
essential element in the Transverse incisions
production of badminton
are made with a small
shuttle cocks. Example:
Quercus suber
axe and thin strip of
c. Shuttle cocks g. Tree shows
the outer bark are torn
gum exudes
off. From that, gum
slowly exudes as a
Rubber is obtained from viscous liquid, collects
latex vessels of inner bark.
in a drop and hardens.
Example: Hevea brasiliensis
Example: Acacia
d. Rubber tree h. Gum senegal.
51

10.2 Secondary Growth in Dicot root combination of conjunctive tissue located
Secondary growth in dicot roots is essential just below the phloem bundles, and as a
to provide strength to the growing aerial portion of pericycle tissue present above
parts of the plants. It is similar to that of the the protoxylem to form a complete and
secondary growth in dicot stem. However, continuous wavy ring. This wavy ring later
there is marked difference in the manner of becomes circular and produces secondary
the formation of vascular cambium. xylem and secondary phloem similar to
the secondary growth in stems.
The vascular cambium is completely
secondary in origin. It originates from a
Epidermis
Endodermis
Pericycle
Primary phloem
Cambium
Primary xylem
a b
Epidermis
Cortex
Endodermis
Pericycle
Primary phloem
Secondary phloem
Cambial ring
Primary xylem
Secondary xylem
c d
Epidermis
Phellogen (Cork cambium)
Pericycle
Primary phloem
Secondary phloem
Phloem ray
Cambial ring
Primary xylem
e Secondary xylem
Xylem ray
Figure 10.17: Different stages of the secondary growth (diagrammatic)

in a typical dicot root (a–e)
52

Differences Between Secondary Growth in Dicot Stem and Root
Secondary growth in dicot stem Secondary growth in dicot root
The cambial ring formed is circular in The cambial ring formed is wavy in the
cross section from the beginning. beginning and later becomes circular.
The cambial ring is partially primary The cambial ring is completely secondary
(fascicular cambium)and partially in origin.
secondary (Interfascicular cambium) in
origin.
Generally, periderm originates from the Generally, periderm originates from the
cortical cells (extrastelar in origin). pericyle.
(intrastealar in origin)
More amount of cork is produced as stem Generally, less amount of cork is produced
is aboveground as root is underground.
Lenticels of periderm are prominent. Lenticels of periderm are not very
prominent.
Pre-structure of Secondary
Primary Structure
Primary Structure secondary growth Secondary Structure
Structure
Procambium Fascicular Axial Phloem

cambium Fusiform
initials
Vascular
Vascular Axial Xylem
cambium
cambium
DICOT Inter fascicular Phloem rays

DICOT STEM
STEM Medullary rays Ray initials
cambium
Xylem rays
Epidermis
Cortex Phellem (cork)
Cork cambium
Phloem Phelloderm
(Phellogen)
(Secondary cortex)
Lenticels
Axial Phloem
Fusiform initials
Vascular
Vascular
Conjunctive cambium Axial Xylem
cambium
tissue
Ray initials Phloem rays
DICOT
DICOT ROOT
ROOT
Xylem rays
Pericycle Cork cambium Phellem (cork)

(Phellogen) Phelloderm
(Secondary cortex)
Tissue Lineages During Secondary Growth in Dicot Stem and Root
53

10.3 Cambial variants (Anomalous 10.3.2 Abnormal behavior of normal
Secondary Growth) cambium
Cambial variants (previously known Certain segments of cambia cease to
as anomalous secondary growth) is a produce secondary xylem. But at the
deviation from normal secondary growth same time, it produces secondary phloem
and production of secondary vascular and only on the outer side. The remaining
non-vascular tissues. A normal cambium cambial segments do the normal activity;
with abnormal activity, accessory As a result, ridged and furrowed steles are
(additional) cambia or abnormally formed. Example: Bignonia.
situated cambia with normal activity can Epidermis
Parenchyma
produce anomalous secondary growth.
Sclerenchyma
The anomalies may be listed as follows,
10.3.1 Anomalous position of vascular

Starch sheath
cambium Secondary phloem
Cambial strips
Unusual shapes in stems are formed by the
unusual position of cambium. Example: Secondary xylem
Thinonia scandens, Serjania ichthyoctona
Pith
and Bauhinia langsdorffiana.
Periderm
Figure 10.19: Transverse section of
Secondary phloem Bignonia stem (diagrammatic) showing
Cambium
Secondary xylem anomalous secondary growth
Main vascular cylinder
Pith
Peripheral vascular cylinder 10.3.3 Successive cambium
a
In plants like Aviccenia,
Periderm
Cycas, Gnetum a number
Secondary phloem
of cambial rings arise in
Cambium
succession and produce
Secondary xylem
concentric rings of
Periderm
Periderm
secondary tissues.
Secondary
S econdary p
phloem
hloem
Successive
vascular ring
Secondary xylem
Secondary phloem
Cambium
Figure 10.18: Transverse section (T.S)
of stems showing anomalous scondary
growth (diagrammatic) (a) Thinouia
scandens (b) Sarjania ichthyoctona Figure 10.20: Transverse section (T.S) of
(c)Bauhinia langsdorffiana Gnetum stem (diagrammatic) showing
anomalous secondary growth
54

10.3.4 Interxylary or Included Phloem These are called medullary vascular
The secondary phloem found in the form bundles. Example: Boerhaavia.
of strands (islands) embedded in the 10.3.6 Presence of cortical bundles
secondary xylem is called interxylary or
along with normal cambial activity
included phloem.
Example: Strychnos, Combretum and In this type, a ring of vascular bundles
Salvadora. are found, from which a normal cambial
ring is formed. In addition, in the cortical
Epidermis
region additional vascular bundles are
Cortex
Endodermis
found. Example: Nyctanthes.
Pericycle Epidermal hair
Secondary phloem Epidermis
Cambium Parenchyma
Secondary xylem
Cortex
Interxylary phloem
Endodermis
Secondary phloem
Pith Cambium
Intra xylary phloem Secondary xylem
(Primary phloem) Cortical vascular
bundle (Inverted)
Pith
Figure 10.21: Transverse section (T.S) of
Strychnos stem (diagrammatic) showing
10.3.5 Presence of medullary bundles
Nyctanthes stem (diagrammatic) showing
along with normal cambial activity
Epidermis
Hypodermis
(Collenchyma) 10.3.7 Intraxylary or Internal Phloem
Parenchyma Epidermis
Starch sheath Collenchyma
Secondary phloem
Parenchyma
Cambium
Endodermis
Secondary xylem
Conjunctive tissue External phloem
Cambium
Medullary bundles
(Primary) Xylem
Ground tissue Internal (or)

Intraxylary phloem
Pith

Boerhaavia stem (diagrammatic) showing
In this type, the normal vascular bundles Solanum tuberosum stem (diagrammatic)
are arranged in a ring from which a normal showing anomalous secondary growth
cambial ring is formed. In addition,
vascular bundles are scattered or found The cambium usually forms primary
in a ring in the pith or medullary region. phloem on the outer side. In some plants,it
55

forms the primary phloem on the inner II) Scattered Vascular bundles along
side along with xylem towards pith. This with cambial activity in dicots.
is called intraxylary or internal Phloem. In dicot stem, normally the vascular
Example: Solanum tuberosum bundles are arranged in a ring, whereas
some dicot stems possess scattered vascular
10.3.8 Secondary Growth in Monocot
bundles in the cortex like monocot stems.
In some monocotyledons, the stems Example: Piper, Peperomia.
exhibit secondary growth. Example:
Epidermis
Dracaena. Here, a secondary thickening
meristems (STM) originate from the Scattered
ground tissue outside of the vascular vascular
bundles
bundles. STM cuts off cells on the inside Phloem
from additional vascular bundles are
Cambium
formed along with parenchyma cells in
Xylem
between the vascular bundles. The STM
consists of only one type of cells unlike
normal vascular cambium. STM does not
produce continuous xylem on the inside Figure 10.26: Transverse section (T.S) of
but only vascular bundle with xylem and Peperomia stem (diagrammatic) showing
phloem. Also no phloem is formed on the anomalous primary growth
outer side.
Epidermis
10.4 Timber
Cork cells
Cortex Timber is derived from wood logs
Cambium (STM)
through the process of sawing. There are
Conjunctive tissue
Secondary vascular many methods of sawing. Among them,
bundle
Secondary xylem
Secondary phloem
Ground tissue
Primary vascular
bundle
Primary phloem
Primary xylem
Figure 10.25: Transverse section of

Dracaena stem (diagrammatic) showing a.
10.3.9 Anomalous primary growth

I) Absence of vessels in the xylem.
Usually, vessels are found in the xylem
of angiosperms, whereas some plants are
lacking vessels in the xylem. Example:
Hydrilla, Winteraceae family members. b.
Figure 10.27: a. Saw mill b. Timber
56

rib sawing is the most common method fans, ensuring the removal of moisture
in making timber. Timber is mainly uniformly, rapidly and completely.
used for carpentry and building houses.
In order to enrich the quality of timber,
seasoning of wood is done. Timber is the
most important tissue that sequestrates
atmospheric carbon and this reduces
global warming.
10.4.1 Seasoning of wood

It is the process in which moisture content
from the wood is removed. There are two
types of seasoning. Figure 10.29: Kiln Seasoning
1. Air Seasoning is the process in
which the moisture can be removed Activity
without resorting to artificial heat. In this List out the uses of saw dust, shavings
method of artificial seasoning, the cut and wood flour.
timber pieces are left exposed in the open
air and the moisture is removed naturally
10.4.2 Grain, Texture, and Figure of
and gradually by the heat of the sun. It
Wood
increases the strength, the combustibility
and renders the wood less subject to decay. Grain refers to the structural arrangement
of wood, while texture shows the relative
size and quality of the various types of
wood. Figure of wood refers to the pattern
formed by grains in wood when the wood
is cut in the longitudinal direction. It
depends on the grain and texture and
their exposure by direction of sawing.
Ply wood
Figure 10.28: Air Seasoning It is manufactured by
2. Kiln Seasoning is the process in gluing together 3 to 9
which the moisture can be removed thin layers or piles of
by artificial method in an enclosed wood veneers. It is used in flooring,
condition. The timber pieces are enclosed walls, false roof and vehicle interiors.
in a steam-heater chamber into which
air is introduced and circulated by
57

fusiform and ray initials. Fusiform initials
give rise to the axial tissue system whereas
ray initials give rise to radial tissue system
of stems and roots.
Wood is a very important product
of secondary growth. It represents
secondary xylem. It is classified in various
ways. Based respectively on the presence
or absence of vessels, wood is classified
a. into two types. i.e., porous and non-
porous wood. Based on the wood formed
during seasons, it is classified into spring
wood and autumn wood. The spring and
autumn wood, together is called annual
ring. The wood is also classified into sap
wood (pale in colour) and heart wood
(dark in colour). The lumen of the xylem
vessels of heart wood are blocked by many
b. balloon like ingrowths from neighbouring
Figure 10.30: Shows grain, texure and parenchymatous cells called tyloses.
figure of wood (a–b). The periderm, a secondary protective
tissue consists of phellem, phellogen and
phelloderm. Secondary growth produces
a corky bark around the tree trunk that
Activity protects the interior parts from heat, cold,
infection etc. Secondary growth of root
Collect some pieces of plywood,
is different from stem in the method of
analyse the layers and discuss yourself
formation of vascular cambium.
how it is made.
Anomalous secondary growth is now
referred as cambial variants. They are
Summary abnormal types of secondary growth
that occur in some dicots and monocots.
Secondary growth deals with the formation Timber is derived from wood logs. In
of additional vascular tissue by the order to enrich the quality of timber,
activities of vascular and cork cambia and seasoning of wood is done through air
secondary thickening meristem (STM). and kiln drying. Wood is characterized by
It increases the girth of stem and roots colour, grain, texture and figure.
of gymnosperms, most angiosperms, and
some monocot plants. Vascular cambium
possesses two kinds of initials they are,
58

Evaluation b. A. Complementary tissue,
B. Phellem, C. Phellogen,
1. Consider the following
D. Phelloderm.
statements
c. A. Phellogen, B. Phellem,
In spring season
C. Phelloderm, D. complementary
vascular cambium
tissue
i. is less active
d. A. Phelloderm, B. Phellem,
ii. produces a large number of xylary
C. Complementary tissue,
elements
D. Phellogen
iii. forms vessels with wide cavities of
4. Inner, darker & harder portion of
these,
secondary xylem that cannot conduct
a. (i) is correct but (ii) and (iii) are not water in an older dicot stem is called
correct
a. Alburnum b. Bast
b. (i) is not correct but (ii) and (iii) are
c. Wood d. Duramen
correct
5. The common bottle cork is a product of
c. (i) and (ii) are correct but (iii) is not
a. Dermatogen
correct
b. Phellogen
d. (i) and (ii) are not correct but (iii) is
correct. c. Xylem
2. Usually, the monocotyledons do not d. Vascular cambium
increase their girth, because 6. What is the fate of primary xylem in a
a. They possess actively dividing dicot root showing extensive secondary
cambium growth?
b. They do not possess actively a. It is retained in the center of the axis
dividing cambium b. It gets crushed
c. Ceases activity of cambium c. May or may not get crushed
d. All are correct d. It gets surrounded by primary
3. In the diagram of lenticel identify the phloem
parts marked as A,B,C,D
Assertion and Reason
7. These questions consist of two
B
statements each printed as Assertion
and Reason. While answering
A these questions you are required to
choose any one of the following four
D
responses.
C A. If both Assertion and Reason are
true but the Reason is a correct
a. A. phellem, B. Complementary
explanation of the Assertion.
tissue, C. Phelloderm, D. Phellogen.
59

B. If both Assertion and Reason are 15. A timber merchant bought 2 logs of
true but Reason is not a correct wood from a forest & named them
explanation of the Assertion A & B, The log A was 50 year old & B
C. If Assertion is true but the Reason is was 20 years old. Which log of wood
false. will last longer for the merchant?
D. If both Assertion and Reason are Why?
false. 16. A cross section of tree trunk contains
1. Assertion: In woody stems the 60 lighter and 60 darker rings.
amount of heart wood continue to Determine the age of the tree and
increase year after Year justify
Reason: The activity of cambial ring 17. A transverse section of the trunk of
continues uninterrupted a tree shows concentric rings which
are known as growth rings. How are
a. A b. B
these rings formed? What are the
c. C d. D significance of these rings?
2. Assertion: Secondary growth in 18. There are many tissues produced
dicot roots occurs with the help of outside the vascular cambium in the
vascular cambium and phellogen. stem. Explain them
Reason: Vascular cambium is 19. When you go to a timber mart to
completely primary in origin. collect timber for your construction
a. A b. B of a new house, how will you select
c. C d. D good timber?
20. Explain artificial seasoning.
Answer the Following
8. When the plants shed their leaves
how do they respire?
9. What is wood botanically?
10. In a forest, if the bark of a tree is
damaged by the horn of a deer, How
will the plant overcome the damage?
11. Differentiate the wood formed in
Pinus from that of Morus
12. In which season the vessels of
angiosperms are larger in size, why?
13. Central part of the wood is always
dark. Why?
14. Continuous state of dividing tissue
is called meristem. In connection
to this, what is the role of lateral
meristem?
60

ICT Corner
Characteristics of Dicot and Monocot Stem and Root
Let’s explore inside

Stem and Root
Steps
• Scan the QR code or go to Google play store.

• Select biology and select Characteristics of dicot and monocot stem and root.
• Click free sign up and provide your basic information with valid mail-Id.
• Login with your registered mail id and password.
• Choose theory tab to know the basic about anatomical structure of plant parts.
• Choose animation to view the sectioning process.
• Choose simulation tab and view the section of plant parts under microscope.
Activity
• Do the section through simulation and record your observations.
Step 2 Step 3
Step 1 Step 4 Step 5
URL:
h�ps://play.google.com/store/apps/details?id=in.edu.olabs.olabs&hl=en
* Pictures are indica�ve only
61

Unit V: Plant Physiology
(Functional Organisation)
Chapter
11 Transport in Plants
Learning Objectives
The learner will be able to, Over 450 million years ago (the Ordovician
• Recall knowledge of basic physical period in Paleozoic era) plants migrated
and biological processes studied in from their own sophisticated water world
previous classes. to newly formed land. The land had harsh
• Classify, differentiate and compare environment; water availability was deeper
the process of active and passive and so plants struggled for getting water for
transport. their very existence. Some of them failed
to survive and rest adopted themselves to
• Understand the mechanism of
the new world. The biggest adaptations
absorption of water.
followed for their survival was building
• Analyse the various theories in their own water absorbing systems to
ascent of sap. draw water from deep inside the land. The
• Understand the process of creation and updating of water absorbing
transpiration and Compare the system (vascular tissues) led to the diversity
various types of transpiration. of the plant kingdom. The gregarious
• Discuss the mechanism of phloem growth of prehistoric pteridophytes,
translocation. gymnosperms and present-day flowering
• Understand the process behind plants led to the biggest challenge in the
mineral absorption. transport of water from root to several
meters high trees against gravity. In this
chapter, we will study the events taking
Chapter Outline place between the gain of water in roots
and loss in leaves and the mechanisms
11.1 Types of transport
behind the basic physical and biological
11.2 Cell to Cell transport
processes in the movement of water, gases
11.3 Plant water relations and minerals in plants. Further, we study
11.4 Absorption of water how food material synthesized in the leaf
11.5 Ascent of Sap can be transported to various utilizing
11.6 Transpiration and storage areas against struggles and
challenges.
11.7 Translocation of organic solutes
11.8 Mineral absorption
62

11.1 Types of Transport
The Plumbing system of Plants and
Humans Transport is the process of moving water,
minerals and food to all parts of the plant
body. Conducting tissues such as xylem
and phloem play an important role in this.
What is the need for transport? Water
absorbed from roots must travel up to
leaves by xylem for food preparation by
photosynthesis. Likewise, food prepared
from leaves has to travel to all parts of the
plant including roots. Both the processes
are interconnected and depend on
each other.
 Based on the distance travelled by
water (sap) or food (solute) they are
classified as
(a) Short distance (Cell to cell
transport) and (b) Long distance
transport.
i. Short-distance (Cell to cell transport):
Involvement of few cells, mostly in the
Plants and animals evolved separately
lateral direction. They are the connecting
but developed comparable structures to
link to xylem and phloem from root hairs
control transport of water and dissolved
or leaf tissues respectively. Examples:
chemicals. But whose transport system
Diffusion, Imbibition, and Osmosis.
is optimally designed to offer selective
advantage? In plants, transport through ii. Long-distance transport: Transport
xylem has allowed growth in height within the network of xylem or phloem
and colonization of diverse habitats is an example for long-distance
and the system has to be extensive transport. Examples: Ascent of Sap and
as Photosynthesis requires water. Translocation of Solutes.
Murray’s law predicts the thickness  Based on energy expenditure during
of branches in transport networks, transport, they are classified as
such that the cost for transport and (a) passive transport and (b) active
maintenance of the transport medium transport.
is minimized. This law is observed in i. Passive transport: It is a downhill
the vascular and respiratory systems of process which utilizes physical forces
animals, xylem in plants, and the like gravity and concentration. No
respiratory system of insects. Further energy expenditure is required. It
research in this area will improve our includes diffusion, facilitated diffusion,
understanding of natural world. imbibition, and osmosis.
63

ii. Active transport: It is a b
iological
process and it runs based on the Diffusion: The net movement of
energy obtained from respiration. It molecules from a region of their
is an uphill process. higher concentration to a region
of their lower concentration along
a concentration gradient until an
11.2 Cell to Cell Transport
equilibrium is attained.
Cell to cell or short distance transport
covers the limited area and consists of few
Characteristics of diffusion
cells. They are the facilitators or tributaries
to the long-distance transport. The i. It is a passive process, hence no energy
driving force for the cell to cell transport expenditure involved.
can be passive or active (Figure 11.1). The ii. It is independent of the living system.
following chart illustrate the various types iii. Diffusion is obvious in gases and
of cell to cell transport: liquids.
Cell to cell transport
Passive Transport Active Transport
Diffusion Facilitated Diffusion

Channel Protein
Carrier Protein
Channel Protein Carrier Protein Pumps
Figure 11.1: Cell to cell transport
A
11.2.1 Passive Transport
1. Diffusion
When we expose a lightened incense stick
or mosquito coil or open a perfume bottle
HIGH CONCENTRATION LOW CONCENTRATION
in a closed room, we can smell the odour Lorem ipsum dolor sit amet, consectetuer adipiscing elit, sed diam nonummy nibh
B
euismod tincidunt ut laoreet dolore magna aliquam erat volutpat. Ut wisi enim ad
everywhere in the room. This is due to the minim veniam, quis nostrud exerci tation ullamcorper suscipit lobortis nisl ut
aliquip ex ea commodo consequat. Duis autem vel eum iriure dolor in hendrerit in
even distribution of perfume molecules vulputate velit esse molestie consequat, vel illum dolore eu feugiat nulla facilisis at
vero eros et accumsan et iusto odio dignissim qui blandit praesent luptatum zzril
throughout the room. This process is

delenit augue duis dolore te feugait nulla facilisi.
Lorem ipsum dolor sit amet, cons ectetuer adipiscing elit, sed diam nonummy nibh
euismod tincidunt ut laoreet dolore magna aliquam erat volutpat. Ut wisi enim ad
called diffusion. minim veniam, quis nostrud exerci tation ullamcorper suscipit lobortis nisl ut
aliquip ex ea commodo consequat.
Lorem ipsum dolor sit amet, consectetuer adipiscing elit, sed diam nonummy nibh
In diffusion, the movement of euismod tincidunt ut laoreet dolore magna aliquam erat volutpat. Ut wisi enim ad
molecules is continuous and random in Figure 11.2: Distribution of molecules in

order in all directions (Figure 11.2). diffusion (A) Initial stage (B) Final stage
64

iv. Diffusion is rapid over a shorter i. Size of molecule: Smaller molecules
distance but extremely slow over a diffuse faster.
longer distance. ii. Solubility of the molecule: Lipid
v. The rate of diffusion is determined by soluble substances easily and rapidly
temperature, concentration gradient pass through the membrane. But water
and relative density. soluble substances are difficult to pass
through the membrane. They must be
Significance of diffusion in Plants facilitated to pass the membrane.
i. Gaseous exchange of O2 and CO2
between the atmosphere and stomata Types of Membrane Permeability
of leaves takes place by the process A solution is made up of solute
of diffusion. O2 is absorbed during particles dissolved in a solvent and the
respiration and CO2 is absorbed during permeability of the above components
photosynthesis. depends on the nature of cell
ii. In transpiration, water vapour from membranes, which is given below:
intercellular spaces diffuses into Impermeable: Inhibit the movement
atmosphere through stomata by the
of both solvent and solute molecules.
process of diffusion.
Example: Suberised, cutinesed or
iii. The transport of ions in mineral salts liginifid cell walls.
during passive absorption also takes
Permeable: They allow diffusion
place by this process.
of both solvent and solute molecules
through them. Example: Cellulosic cell
Diffusion for
wall.
sterilization in
surgical theatres Semi permeable: Semi permeable
allow diffusion of solvent molecules
Surgical theatres must
but do not allow the passage of solute
be free from germs to prevent infection
molecule. Example: Parchment paper.
during surgeries. A mixture of
Selectively permeable: All bio
Formalin and Potassium permanganate
membranes allow some solutes to pass
produces enormous fumes which will
in addition to the solvent molecules.
kill all pathogens in an enclosed area.
Example: Plasmalemma, tonoplast, and
This method is known as fumigation
membranes of cell organelles.
and operates by diffusion.
In facilitated diffusion, molecules cross

2. Facilitated Diffusion
the cell membrane with the help of special
Cell membranes allow water and nonpolar membrane proteins called transport
molecules to permeate by simple diffusion. proteins, without the expenditure of ATP.
For transporting polar molecules such as
There are two types of transport
ions, sugars, amino acids, nucleotides and
proteins present in the cell membrane.
many cell metabolites is not merely based
They are channel protein and a carrier
on concentration gradient. It depends on,
protein.
65

I. Channel Protein
Discovery of Aquaporin
Channel protein forms a channel or
Peter Agre,
tunnel in the cell membrane for the easy
discovered
passage of molecules to enter the cell. The
the “Water
channels are either open or remain closed.
Pore” Aquaporin in RBC
They may open up for specific molecules.
and received Nobel Prize
Some channel proteins create larger pores
for chemistry in 2003.
in the outer membrane. Examples: Porin
and Aquaporin.
i. Porin II. Carrier Protein
Porin is a large transporter protein Carrier protein acts as a vehicle to
found in the outer membrane of plastids, carry molecules from outside of the
mitochondria and bacteria which membrane to inside the cell and vice versa
facilitates smaller molecules to pass (Figure 11.4). Due to association with
through the membrane. molecules to be transported, the structure
ii. Aquaporin of carrier protein gets modified until the
Aquaporin is a water channel protein dissociation of the molecules.
embedded in the plasma membrane.
It regulates the massive amount of
water transport across the membrane
(Figure 11.3). Plants contain a variety of OUTSIDE
MEMBRANE
OUTSIDE
INSIDE
MEMBRANE
Carrier Protein
Figure 11.4: Carrier Protein
There are 3 types of carrier proteins

INSIDE
classified on the basis of handling of
OPEN CHANNEL CLOSED CHANNEL molecules and direction of transport
Figure 11.3: Aquaporin (Figure 11.5). They are, i) Uniport
ii) Symport iii) Antiport.
aquaporins. Over 30 types of aquaporins
are known from maize. Currently, they are i. Uniport: In this molecule of a single
also recognized to transport substrates like type move across a membrane
glycerol, urea, CO2, NH3, metalloids, and independent of other molecules in one
Reactive Oxygen Species (ROS) in addition direction.
to water. They increase the permeability ii. Symport or co-transport: The term
of the membrane to water. They confer symport is used to denote an integral
drought and salt stress tolerance. membrane protein that simultaneously
66

of ions or molecules. The pump action is
OUTSIDE
an example of active transport. Example:
MEMBRANE
Na+-K+-ATPase pump (Table 11.1).
INSIDE
Table 11.1 Comparison of different
transport mechanisms
UNIPORT SYMPORT ANTIPORT
Passive transport
Figure 11.5: Direction of transport Active
Property Simple Facilitated transport
transports two types of molecules diffusion diffusion
across the membrane in the same Nature of
Physical Biological Biological
direction. process
iii. Antiport or Counter Transport: An Requirement
antiport is an integral membrane for presence
No Yes Yes
transport protein that simultaneously of membrane
transports two different molecules, protein
in opposite directions, across the Selectivity of
No Yes Yes
membrane. molecule
Saturation of
11.2.2 Active Transport No Yes Yes
transport
The main disadvantage of passive transport Uphill
No No Yes
processes like diffusion is the lack of control transport
over the transport of selective molecules. Energy
There is a possibility of harmful substances requirement No No Yes
entering the cell by a concentration gradient (ATP)
in the diffusion process. But selective Sensitivity to
No Yes Yes
permeability of cell membrane has a great inhibitors
control over entry and exit of molecules.

Active transport is the entry of molecules Check your grasp!
against a concentration gradient and an What are the similarities and
uphill process and it needs energy which differences between co- transport and
comes from ATP. Passive transport uses counter transport?
kinetic energy of molecules moving down Solution:
a gradient whereas, active transport uses
Similarity: In both system two molecules
cellular energy to move them against a
are involved for the unidirectional
gradient. The transport proteins discussed in
transport.
facilitated diffusion can also transport ions or
molecules against a concentration gradient Difference: In co-transport, two
with the expenditure of cellular energy as molecules are transported together
an active process. Pumps use a source of whereas, in counter transport two
free energy such as ATP or light to drive molecules are transported in opposite
the thermodynamically uphill transport direction to each other.
67

11.3 Plant Water Relations Significance of imbibition
Water plays an essential role in the life i. During germination of seeds, imbibition
of the plant. The availability of water increases the volume of seed enormously
and leads to bursting of the seed coat.
influences the external and internal
structures of plants as protoplasm is made ii. It helps in the absorption of water by
of 60-80% water. Water is a universal roots at the initial level.
solvent since most of the substances
get dissolved in it and the high tensile Activity
strength of water molecule is helpful in
Imbibition experiment
the ascent of sap. Water maintains the
internal temperature of the plant as well Collect 5 gm of gum from Drumstick
as the turgidity of the cell. tree or Babool tree or Almond tree.
Immerse in 100ml of water. After 24
11.3.1 Imbibition hours observe the changes and discuss
the results with your teacher.
Colloidal systems such as gum, starch,
proteins, cellulose, agar, gelatin when
placed in water, will absorb a large volume
of water and swell up. These substances are
called imbibants and the phenomenon is
imbibition.
Examples: 1. The swelling of dry

seeds 2.The swelling of wooden windows,
tables, doors due to high humidity during
the rainy season.
The Power of 11.3.2 Water Potential (Ψ)

Imbibition
The concept of water potential was
In olden days, small
introduced in 1960 by Slatyer and Taylor.
wooden pegs were
Water potential is potential energy of
inserted into crevices of rocks
water in a system compared to pure water
followed by continuous hydration.
when both temperature and pressure are
Due to imbibition the volume of
kept the same. It is also a measure of how
wooden peg increases and cuts off
freely water molecules
rocks precisely.
can move in a particular
environment or system.
The gluten from wheat can take as Water potential is denoted
much as 300% of its own weight by the Greek symbol Ψ
68

(psi) and measured in Pascal (Pa). At withdrawal of water from the cell decreases
standard temperature, the water potential the water potential and the cell becomes
of pure water is zero. Addition of solute flaccid.
to pure water decreases the kinetic energy
3. Matric Potential (ΨM)
thereby decreasing the water potential.
Comparatively a solution always has Matric potential represents the attraction
low water potential than pure water. between water and the hydrating colloid
In a group of cells with different water or gel-like organic molecules in the cell
potential, a water potential gradient is wall which is collectively termed as matric
generated. Water will move from higher potential. Matric potential is also known
water potential to lower water potential. as imbibition pressure. The matric
Water potential (Ψ) can be determined by, potential is maximum (most negative
value) in a dry material. Example: The
1. Solute concentration or Solute
swelling of soaked seeds in water.
potential (ΨS)
2. Pressure potential (ΨP) 11.3.3 Osmotic Pressure and Osmotic
By correlating two factors, water potential Potential
is written as,
When a solution and its solvent (pure
ΨW = ΨS + ΨP water) are separated by a semipermeable
membrane, a pressure is developed in the
Water Potential = S olute potential +
solution, due to the presence of dissolved
Pressure potential
solutes. This is called osmotic pressure
(OP). Osmotic pressure is increased
1. Solute Potential (ΨS)
with the increase of dissolved solutes in
Solute potential, otherwise known as
the solution. More concentrated solution
osmotic potential denotes the effect of
(low Ψ or Hypertonic) has high osmotic
dissolved solute on water potential. In pure
pressure. Similarly, less concentrated
water, the addition of solute reduces its
solution (high Ψ or Hypotonic) has low
free energy and lowers the water potential
osmotic pressure. The osmotic pressure
value from zero to negative. Thus the value
of pure water is always zero and it
of solute potential is always negative. In a
increases with the increase of solute
solution at standard atmospheric pressure,
concentration. Thus osmotic pressure
water potential is always equal to solute
always has a positive value and it is
potential (ΨW= ΨS).
represented as π.
2. Pressure Potential (ΨP) Osmotic potential is defined as
Pressure potential is a mechanical force the ratio between the number of solute
working against the effect of solute particles and the number of solvent
potential. Increased pressure potential particles in a solution. Osmotic potential
will increase water potential and water and osmotic pressure are numerically
enters cell and cells become turgid. This equal. Osmotic potential has a negative
positive hydrostatic pressure within the value whereas on the other hand osmotic
cell is called Turgor pressure. Likewise, pressure has a positive value.
69

11.3.4 Turgor Pressure and Wall It is equal to the difference of osmotic
Pressure pressure and turgor pressure of a cell.
The following three situations are seen in
When a plant cell is placed in pure water
plants:
(hypotonic solution) the diffusion of water
into the cell takes place by endosmosis. It • DPD in normal cell: DPD = OP – TP.
creates a positive hydrostatic pressure on • DPD in fully turgid cell: Osmotic
the rigid cell wall by the cell membrane. pressure is always equal to turgor
Henceforth the pressure exerted by the cell pressure in a fully turgid cell.
membrane towards the cell wall is Turgor • OP = TP or OP-TP =0. Hence DPD of
Pressure (TP). fully turgid cell is zero.
The cell wall reacts to this turgor • DPD in flaccid cell: If the cell is in
pressure with equal and opposite force, flaccid condition there is no turgor
and the counter-pressure exerted by the pressure or TP=0. Hence DPD = OP.
cell wall towards cell membrane is wall
pressure (WP). 11.3.6 Osmosis
Turgor pressure and wall pressure
Osmosis (Latin: Osmos-impulse, urge) is a
make the cell fully turgid.
special type of diffusion. It represents the
TP + WP = Turgid. movement of water or solvent molecules
through a selectively permeable
Activity membrane from the place of its higher
Find the role of turgor pressure in concentration (high water potential) to
sudden closing of leaves when we the place of its lower concentration (low
touch the ‘touch me not’ plant. water potential).
Types of Solutions based on concentration
11.3.5 Diffusion Pressure Deficit (DPD) i. Hypertonic (Hyper = High; tonic =
or Suction Pressure (SP) solute): This is a strong solution (low
solvent/ high solute / low Ψ) which
Pure solvent (hypotonic) has higher
attracts solvent from other solutions.
diffusion pressure. Addition of solute in
pure solvent lowers its diffusion pressure. ii. Hypotonic (Hypo = low; tonic = s olute):
The difference between the diffusion This is a weak solution (high solvent
pressure of the solution and its solvent at /low or zero solute / high Ψ) and it
a particular temperature and atmospheric diffuses water out to other solutions
pressure is called as Diffusion Pressure (Figure 11.7).
Deficit (DPD) termed by Meyer (1938).
iii. Isotonic (Iso = identical; tonic = soute):
DPD is increased by the addition of solute
It refers to two solutions having same
into a solvent system. Increased DPD
concentration. In this condition the net
favours endosmosis or it sucks the water
movement of water molecule will be zero.
from hypotonic solution; hence Renner
(1935) called it as Suction pressure. The term hyper, hypo and isotonic are
relative terms which can be used only
70

in comparison with another solution.
Thistle funnel experiment
1. Types of osmosis
Based on the direction of movement of
water or solvent in an osmotic system,
two types of osmosis can occur, they are
Endosmosis and Exosmosis.
i. Endosmosis: Endosmosis is defined as
the osmotic entry of solvent into a cell
or a system when it is placed in a pure
water or hypotonic solution.
Figure 11.6: Thistle Funnel Experiment For example, dry raisins (high solute
and low solvent) placed in the water,
Mouth of a thistle funnel is tied with it swells up due to turgidity.
goat bladder. It acts as a semipermeable
membrane. Pour concentrated sugar ii. Exosmosis: Exosmosis is defined as
solution in the thistle funnel and the osmotic withdrawal of water from
mark the level of solution. Place a cell or system when it is placed in a
this in a beaker of water. After some hypertonic solution. Exosmosis in a
time, water level in the funnel rises plant cell leads to plasmolysis.
up steadily. This is due to the inward
diffusion of water molecules through 2. Plasmolysis (Plasma = cytoplasm;
the semipermeable membrane lysis = breakdown)
(Figure 11.6). When a plant cell is kept in a hypertonic
Conversely, if water in the beaker is solution, water leaves the cell due to
replaced by a sugar solution and sugar exosmosis. As a result of water loss,
solution in the thistle funnel replaced protoplasm shrinks and the cell membrane
by water, what will be happen? is pulled away from the cell wall and finally,
the cell becomes flaccid. This process is
named as plasmolysis.
Wilting of plants noticed under the
condition of water scarcity is an indication
of plasmolysis. Three types of plasmolysis
occur in plants: i) Incipient plasmolysis
ii) Evident plasmolysis and iii) Final
plasmolysis. Differences among them are
given in table 11.2.
Significance
Plasmolysis is exhibited only by living
Figure 11.7: Types of solution based on cells and so it is used to test whether the
concentration cell is living or dead.
71

3. Deplasmolysis 4. Reverse Osmosis
The effect of plasmolysis can be reversed, Reverse Osmosis follows the same
by transferring them back into water or principles of osmosis, but in the reverse
hypotonic solution. Due to endosmosis, direction. In this process movement of
the cell becomes turgid again. It regains its water is reversed by applying pressure to
original shape and size. This phenomenon force the water against a concentration
of the revival of the plasmolysed cell gradient of the solution. In regular
is called deplasmolysis. Example: osmosis, the water molecules move from
Immersion of dry raisin in water. the higher concentration (pure water
= hypotonic) to lower concentration
Potato Osmoscope (salt water = hypertonic). But in reverse
osmosis, the water molecules move from
the lower concentration (salt water =
hypertonic) to higher concentration (pure
water = hypotonic) through a selectively
permeable membrane (Figure 11.9).
Uses: Reverse osmosis is used for

purification of drinking water and
desalination of seawater.
Figure 11.8: Demonstration of Pressure

Endosmosis by Potato Osmoscope
Pure Water
i. Take a peeled potato tuber and
make a cavity inside with the help Salt Water
of a knife.
ii. Fill the cavity with concentrated Membrane
sugar solution and mark the initial Movement of Water
level.
Figure 11.9: Reverse Osmosis
iii. Place this setup in a beaker of pure
water.
iv. After 10 minutes observe the sugar
solution level and record your Check your grasp!
findings (Figure 11.8).
If a cell in the cortex with DPD of 5atm
v. With the help of your teacher is surrounded by hypodermal cells with
discuss the results. DPD of 2atm, what will be direction of
Instead of potato use beetroot or bottle- movement of water?
guard and repeat the above experiment. Solution: Water will move from low
Compare and discuss the results. DPD to high DPD (hypodermis 2 atm
to cortex 5 atm).
72

Table 11.2: Difference between plasmolysis types.
Incipient plasmolysis Evident plasmolysis Final plasmolysis
No morphological Wilting of leaves appear. Severe wilting and
symptoms appear in plants. drooping of leaves appear.
The plasma membrane Plasma membrane completely Plasma membrane
separates only at the corner detaches from the cell wall. completely detaches from
from the cell wall of cells. cell wall with maximum
shrinkage of volume.
It is reversible. It is reversible. It is irreversible.
11.4 Absorption of Water

Terrestrial plants have to absorb water from
the soil to maintain turgidity, metabolic
activities and growth. Absorption of water
from soil takes place in two steps:
1. From soil to root hairs – either
actively or passively.
Figure 11.10: Structure of Root Hair
2. From root hairs further transport in
the lateral direction to reach xylem, the 11.4.2 Path of Water Across Root Cells
superhighway of water transport. Water is first absorbed by root hair
and other epidermal cells through
11.4.1 Water Absorbing Organs
imbibition from soil and moves radially
Usually, absorption of water occurs in and centripetally across the cortex,
plants through young roots. The zone endodermis, pericycle and finally reaches
of rapid water absorption is root hairs. xylem elements osmotically.
They are delicate structures which get
There are three possible routes of
continuously replaced by new ones.
water (Figure 11.11). They are i) Apoplast
Root hairs are unicellular extensions of
ii) Symplast iii) Transmembrane route.
epidermal cells without cuticle. Root hairs
are extremely thin and numerous and they 1. Apoplast
provide a large surface area for absorption The apoplast (Greek: apo = away;
(Figure 11.10). plast = cell) consists of everything external
73

Figure 11.11: Path of water across root cells
to the plasma membrane of the living 11.4.3 Mechanism of Water Absorption
cell. The apoplast includes cell walls,
Kramer (1949) recognized two distinct
extra cellular spaces and the interior of
mechanisms which independently operate
dead cells such as vessel elements and
in the absorption of water in plants.
tracheids. In the apoplast pathway, water
They are, i) active absorption ii) passive
moves exclusively through the cell wall or
the non-living part of the plant without absorption.
crossing any membrane. The apoplast is a
continuous system. 1. Active Absorption
The mechanism of water absorption due to
2. Symplast
forces generated in the root itself is called
The symplast (Greek: sym = within; plast = active absorption. Active absorption may
cell) consists of the entire mass of cytosol be osmotic or non-osmotic.
of all the living cells in a plant, as well as the
i. Osmotic active absorption
plasmodesmata, the cytoplasmic channel
that interconnects them. The theory of osmotic active
absorption was postulated by Atkins
In the symplastic route, water has to cross
(1916) and Preistley (1923). According to
plasma membrane to enter the cytoplasm
this theory, the first step in the absorption
of outer root cell; then it will move within
is soil water imbibed by cell wall of the root
adjoining cytoplasm through plasmodesmata
hair followed by osmosis. The soil water
around the vacuoles without the necessity to
is hypotonic and cell sap is hypertonic.
cross more membrane, till it reaches xylem.
Therefore, soil water diffuses into root
3. Transmembrane route hair along the concentration gradient
(endosmosis). When the root hair becomes
In transmembrane pathway water
fully turgid, it becomes hypotonic and
sequentially enters a cell on one side and
water moves osmotically to the outer most
exits from the cell on the other side. In
cortical cell. In the same way, water enters
this pathway, water crosses at least two
into inner cortex, endodermis, pericycle
membranes for each cell. Transport across
and finally reaches protoxylem. As the
the tonoplast is also involved.
74

sap reaches the protoxylem a pressure is in the rate of respiration and also the rate
developed known as root pressure. This of absorption of water.
theory involves the symplastic movement
of water. 2. Passive Absorption
Objections to osmotic theory: 1.The In passive absorption, roots do not play
cell sap concentration in xylem is not any role in the absorption of water and
always high. 2. Root pressure is not is regulated by transpiration only. Due to
universal in all plants especially in trees. transpiration, water is lost from leaf cells
ii. Non-Osmotic active absorption along with a drop in turgor pressure. It
increases DPD in leaf cells and leads to
Bennet-Clark (1936), Thimann (1951)
withdrawal of water from adjacent xylem
and Kramer (1959) observed absorption
cells. In xylem, a tension is developed
of water even if the concentration of cell
and is transmitted downward up to root
sap in the root hair is lower than that of
resulting in the absorption of water from
the soil water. Such a movement requires
the soil.
an expenditure of energy released by
respiration (ATP). Thus, there is a In passive absorption (Table 11.3),
link between water absorption and the path of water may be symplastic or
respiration. It is evident from the fact that apoplastic. It accounts for about 98% of
when respiratory inhibitors like KCN, the total water uptake by plants.
Chloroform are applied there is a decrease
Concept Map - Movement of water in an osmotic system

based on various parameters
High Water Low Water

Potential Potential
Low / Zero solute (Negative value) High solute
(Zero)
concentration concentration
Low High
High solvent Low solvent
DPD DPD
Concentration Concentration
Low Osmotic High Osmotic Hypertonic

Hypotonic Pressure
Pressure
(Zero) (Positive value)
High osmotic Low osmotic

potential potential
(Zero) PURE SALT (Negative Value)
WATER WATER
75

Table: 11.3 Differences between Active
Absorption and Passive Absorption
Active Passive
absorption absorption
Active absorption The pressure for
takes place by the absorption is not
activity of root and developed in roots
root hairs and hence roots
play passive role
Transpiration has Absorption
no effect on active regulated by
absorption transpiration
The root hairs The absorption
have high DPD occurs due to
as compared to tension created
soil solution and in xylem sap by
therefore water is transpiration pull,
taken by tension thus water is sucked
in by the tension
Respiratory energy Respiratory energy
needed not required
It involves Both symplast
Figure 11.12: Balsam plant and eosin
symplastic and apoplast dye experiment
movement of water movement of water
involved 11.5.1 The Path of Ascent of Sap
11.5 Ascent of Sap There is no doubt; water travels up along the

vascular tissue. But vascular tissue has two
In the last chapter, we studied about water
components namely Xylem and Phloem.
absorption from roots to xylem in a lateral
Of these two, which is responsible for the
direction and here we will learn about
ascent of sap? The following experiment
the mechanism of distribution of water
will prove that xylem is the only element
inside the plant. Like tributaries join
through which water moves up.
together to form a river, millions of root
hairs conduct a small amount of water and Cut a branch of balsam plant and
confluence in xylem, the superhighway of place it in a beaker containing eosin
water conduction. Xylem handles a large (red colour dye) water. After some
amount of water to conduct to many parts time, a red streak appears on the stem
in an upward direction. indicating the ascent of water. Remove
The water within the xylem along with the plant from water and cut a transverse
dissolved minerals from roots is called sap section of the stem and observe it under
and its upward transport is called ascent the microscope. Only xylem element is
of sap. coloured red, which indicates the path
76

of water is xylem. Phloem is not colored into the inner cortex of the stem, the
indicating that it has no role in the ascent galvanometer showed high electrical
of sap (Figure 11.12). activity. Bose believed a rhythmic
Mechanism of Ascent of Sap pulsating movement of inner cortex like a
pump (similar to the beating of the heart)
In ascent of sap, the biggest challenge is the
is responsible for the ascent of sap. He
force required to lift the water to the top
concluded that cells associated with xylem
of the tallest trees. A number of theories
exhibit pumping action and pumps the
have been put forward to explain the
sap laterally into xylem cells.
mechanism of the ascent of sap. They are,
A. Vital force theories, B. Root pressure Objections to vital force theories
theory, and C. Physical force theory. i. Strasburger (1889) and Overton
(1911) experimentally proved that living
11.5.2 Vital Force Theories
cells are not mandatory for the ascent of
According to vital force theories, living cells sap. For this, he selected an old oak tree
are mandatory for the ascent of sap. Based trunk which when immersed in picric
on this the following two theories derived: acid and subjected to excessive heat killed
1. Relay pump theory of Godlewski all the living cells of the trunk. The trunk
(1884) when dipped in water, the ascent of sap
Periodic changes in osmotic pressure took place.
of living cells of the xylem parenchyma ii. Pumping action of living cells should
and medullary ray act as a pump for the be in between two xylem elements
movement of water. (vertically) and not on lateral sides.
2. Pulsation theory of J.C.Bose (1923)
11.5.3 Root Pressure Theory
Bose invented an instrument called
Crescograph, which consists of an electric If a plant which is watered well is cut a few
probe connected to a galvanometer inches above the ground level, sap exudes
(Figure 11.13). When a probe is inserted out with some force. This is called sap
exudation or bleeding. Stephen Hales,
father of plant physiology observed this
phenomenon and coined the term ‘Root
Pressure’. Stoking (1956) defined root
pressure as “a pressure developing in the
tracheary elements of the xylem as a result
of metabolic activities of the root”. But the
following objections have been raised
against root pressure theory:
i. Root pressure is totally absent in
gymnosperms, which includes some of
the tallest plants.
ii. There is no relationship between
Figure 11.13: J.C. Bose the ascent of sap and root pressure. For
77

example, in summer, the rate of the ascent 3. Cohesion-tension or Cohesion and
of sap is more due to transpiration in spite transpiration pull theory
of the fact that root pressure is very low. Cohesion-tension theory was originally
On the other hand, in winter when the proposed by Dixon and Jolly (1894) and
rate of ascent of sap is low, a high root again put forward by Dixon (1914, 1924).
pressure is found. This theory is based on the following
iii. Ascent of sap continues even in the features:
absence of roots i. Strong cohesive force or tensile
iv. The magnitude of root pressure strength of water
is about 2atm, which can raise the water Water molecules have the strong
level up to few feet only, whereas the tallest mutual force of attraction called cohesive
trees are more than 100m high. force due to which they cannot be easily
separated from one another. Further, the
11.5.4 Physical Force Theory attraction between a water molecule and
Physical force theories suggest that ascent the wall of the xylem element is called
of sap takes place through the dead xylem adhesion. These cohesive and adhesive
vessel and the mechanism is entirely force works together to form an unbroken
physical and living cells are not involved. continuous water column in the xylem.
The magnitude of the cohesive force is
1. Capillary theory
much high (350 atm) and is more than
Boehm (1809) suggested that the xylem enough to ascent sap in the tallest trees.
vessels work like a capillary tube. This
ii. Continuity of the water column in
capillarity of the vessels under normal
the plant
atmospheric pressure is responsible for
An important factor which can break
the ascent of sap. This theory was rejected
the water column is the introduction of
because the magnitude of capillary force
air bubbles in the xylem. Gas bubbles
can raise water level only up to a certain
expanding and displacing water within
height. Further, the xylem vessels are
the xylem element is called cavitation
broader than the tracheid which actually
or embolism. However, the overall
conducts more water and against the continuity of the water column remains
capillary theory. undisturbed since water diffuses into the
2. Imbibition theory adjacent xylem elements for continuing
This theory was first proposed by Unger ascent of sap.
(1876) and supported by Sachs (1878). iii. Transpiration pull or Tension in
This theory illustrates, that water is the unbroken water column
imbibed through the cell wall materials The unbroken water column from leaf
and not by the lumen. This theory was to root is just like a rope. If the rope is
rejected based on the ringing experiment, pulled from the top, the entire rope will
which proved that water moves through move upward. In plants, such a pull is
the lumen of the cell and not by a cell wall. generated by the process of transpiration
which is known as transpiration pull.
78

79

Water vapour evaporates from
mesophyll cells to the intercellular Activity
spaces near stomata as a result of active Select a leafy twig of fully grown plant
transpiration. The water vapours are then in your school campus. Cover the
transpired through the stomatal pores. twig with a transparent polythene bag
Loss of water from mesophyll cells causes a and tie the mouth of the bag at the
decrease in water potential. So, water moves base of the twig. Observe the changes
as a pull from cell to cell along the water after two hours and discuss with your
potential gradient. This tension, generated teacher
at the top (leaf) of the unbroken water
column, is transmitted downwards from
petiole, stem and finally reaches the roots. 11.6.1 Types of Transpiration
The cohesion theory is the most accepted Transpiration is of following three types:
among the plant physiologists today. 1. Stomatal transpiration
Stomata are microscopic structures present
11.6 Transpiration in high number on the lower epidermis of
Water absorbed by roots ultimately leaves. This is the most dominant form of
reaches the leaf and gets released into transpiration and being responsible for
the atmosphere in the form of vapour. most of the water loss (90 - 95%) in plants.
Only a small fraction of water (less than 2. Lenticular transpiration
5%) is utilized in plant development and In stems of woody plants and trees, the
metabolic process. epidermis is replaced by periderm because
The loss of excess of water in the form of secondary growth. In order to provide
of vapour from various aerial parts of the gaseous exchange between the living cells
plant is called transpiration. Transpiration and outer atmosphere, some pores which
is a kind of evaporation but differs by the looks like lens-shaped raised spots are
involvement of biological system. The present on the surface of the stem called
amount of water transpired is astounding Lenticels. The loss of water from lenticels
(Table 11.4). The water may move through is very insignificant as it amounts to only
the xylem at a rate as fast as 75cm /min. 0.1% of the total.
3. Cuticular transpiration
Table: 11.4 Rate of Transpiration in The cuticle is a waxy or resinous layer
some plants of cutin, a fatty substance covering the
Plant Transpiration per day epidermis of leaves and other plant parts.
Loss of water through cuticle is relatively
Corn plant 2 Litres
small and it is only about 5 to 10 % of the
Sunflower 5 Litres
total transpiration. The thickness of cuticle
Maple tree 200 Litres
increases in xerophytes and transpiration
Date palm 450 Litres is very much reduced or totally absent.
80

11.6.2 Structure of Stomata Different theories have been proposed
The epidermis of leaves and green stems regarding opening and closing of stomata.
possess many small pores called stomata. The important theories of stomatal
The length and breadth of stomata is movement are as follows,
about 10-40µ and 3-10µ respectively. 1. Theory of Photosynthesis in guard cells
Mature leaves contain between 50 and 2. Starch – Sugar interconversion theory
500 stomata per mm2. Stomata are made 3. Active potassium transport ion concept
up of two guard cells, special semi-lunar
1. Theory of Photosynthesis in guard
or kidney-shaped living epidermal cells
cells
in the epidermis. Guard cells are attached
to surrounding epidermal cells known as Von Mohl (1856) observed that stomata
subsidiary cells or accessory cells. The open in light and close in the night.
guard cells are joined together at each According to him, chloroplasts present
end but they are free to separate to form a in the guard cells photosynthesize in
pore between them. The inner wall of the the presence of light resulting in the
guard cell is thicker than the outer wall production of carbohydrate (Sugar) which
(Figure 11.14). The stoma opens to the increases osmotic pressure in guard cells.
interior into a cavity called sub-stomatal It leads to the entry of water from other
cavity which remains connected with the cell and stomatal aperture opens. The
intercellular spaces. above process vice versa in night leads to
closure of stomata.
Demerits
1. Chloroplast of guard cells is
poorly developed and incapable of
performing photosynthesis.
2. The guard cells already possess
much amount of stored sugars.
Guard cells
2. Starch – Sugar Interconversion theory
i. According to Lloyd (1908), turgidity
Figure 11.14: Structure of Stomata of guard cell depends on interconversion,
of starch and sugar. It was supported by
11.6.3 Mechanism of Stomatal Movement Loftfield (1921) as he found guard cells
Stomatal movements are regulated by the containing sugar during the daytime when
change of turgor pressure in guard cells. they are open and starch during the night
When water enters the guard cell, it swells when they are closed.
and its unevenly thickened walls stretch ii. Sayre (1920) observed that the
up resulting in the opening of stomata. opening and closing of stomata depends
This is due to concave non-elastic nature upon change in pH of guard cells.
of inner wall pulled away from each According to him stomata open at high
other and stretching of the convex elastic pH during day time and become closed
natured outer wall of guard cell. at low pH at night. Utilization of CO2
81

by photosynthesis during light period
causes an increase in pH resulting in
the conversion of starch to sugar. Sugar
increase in cell favours endosmosis and
increases the turgor pressure which
leads to opening of stomata. Likewise,
accumulation of CO2 in cells during
night decrease the pH level resulting in
the conversion of sugar to starch. Starch
decreases the turgor pressure of guard cell
and stomata close.
iii.
The discovery of enzyme
phosphorylase in guard cells by Hanes Figure 11.15: Steward Scheme
(1940) greatly supports the starch-sugar
interconversion theory. The enzyme iii. It fails to explain the drastic change
phosphorylase hydrolyses starch into sugar in pH from 5 to 7 by change of CO2.
and high pH followed by endosmosis and 3. Theory of K+ transport
the opening of stomata during light. The This theory was proposed by Levit
vice versa takes place during the night. (1974) and elaborated by Raschke (1975).
According to this theory, the following
steps are involved in the stomatal opening:
iv. Steward (1964) proposed a

slightly modified scheme of starch-sugar
interconversion theory. According to
him, Glucose-1-phosphate is osmotically
inactive. Removal of phosphate from
Glucose-1-phosphate converts to Glucose
which is osmotically active and increases
the concentration of guard cell leading to
opening of stomata (Figure 11.15).
Objections to Starch-sugar
interconversion theory Figure 11.16: Theory of K+ transport
i. In monocots, guard cell does not Opening of stomata
have starch.
ii. There is no evidence to show the In light
presence of sugar at a time when starch i. In guard cell, starch is converted
disappears and stomata open. into organic acid (malic acid).
82

ii. Malic acid in guard cell dissociates i. In dark photosynthesis stops and
to malate anion and proton (H+). respiration continues with accumulation
iii. Protons are transported through of CO2 in the sub-stomatal cavity.
the membrane into nearby subsidiary cells ii. Accumulation of CO2 in cell lowers
with the exchange of K+ (Potassium ions) the pH level.
from subsidiary cells to guard cells. This iii. Low pH and a shortage of water in
process involves an electrical gradient and the guard cell activate the stress hormone
is called ion exchange. Abscisic acid (ABA).
iv. This ion exchange is an active iv. ABA stops further entry of K+ ions
process and consumes ATP for energy. and also induce K+ ions to leak out to
v. Increased K+ ions in the guard subsidiary cells from guard cell.
cell are balanced by Cl– ions. Increase in v. Loss of water from guard cell
solute concentration decreases the water reduces turgor pressure and causes closure
potential in the guard cell. of stomata (Figure 11.17).
vi. Guard cell becomes hypertonic
and favours the entry of water from 11.6.4 Factors Affecting Rate of
surrounding cells. Transpiration
vii. Increased turgor pressure due to The factors affecting the rate of
the entry of water opens the stomatal pore transpiration can be categorized into
(Figure 11.16). two groups. They are 1. External or
Environmental factors and 2. Internal or
In Dark plant factors.
1. External or Environmental factors
i. Atmospheric humidity: The rate
of transpiration is greatly reduced when
the atmosphere is very humid. As the air
becomes dry, the rate of transpiration is
also increased proportionately.
ii. Temperature: With the increase
in atmospheric temperature, the rate of
transpiration also increases. However, at
very high-temperatures stomata closes
because of flaccidity and transpiration stop.
iii. Light: Light intensity increases the
temperature. As in temperature, transpiration
is increased in high light intensity and
is decreased in low light intensity. Light
Figure 11.17: Theory of K+ transport also increases the permeability of the
Closing of stomata cell membrane, making it easy for water
molecules to move out of the cell.
83

iv. Wind velocity: In still air, the 2. Internal factors
surface above the stomata get saturated i. Leaf area: If the leaf area is more,
with water vapours and there is no need transpiration is faster and so xerophytes
for more water vapour to come out. If reduce their leaf size.
the wind is breezy, water vapour gets ii. Leaf structure: Some anatomical
carried away near leaf surface and DPD is features of leaves like sunken stomata, the
created to draw more vapour from the leaf presence of hairs, cuticle, the presence of
cells enhancing transpiration. However, hydrophilic substances like gum, mucilage
high wind velocity creates an extreme help to reduce the rate of transpiration. In
increase in water loss and leads to a xerophytes the structural modifications
reduced rate of transpiration and stomata are remarkable. To avoid transpiration, as
remain closed. in Opuntia the stem is flattened to look
like leaves called Phylloclade. Cladode
or cladophyll in Asparagus is a modified
Activity
stem capable of limited growth looking
What will happen if an indoor plant is like leaves. In some plants, the petioles
placed under fan and AC? are flattened and widened, to become
phyllodes example Acacia melanoxylon.
11.6.5 Plant Antitranspirants
v. Atmospheric pressure: In low
atmospheric pressure, the rate of The term antitranspirant is used to
transpiration increases. Hills favour high designate any material applied to plants
transpiration rate due to low atmospheric for the purpose of retarding transpiration.
pressure. However, it is neutralized by low An ideal antitranspirant checks the
temperature prevailing in the hills. transpiration process without disturbing
vi. Water: Adequate amount of water in the process of gaseous exchange. Plant
the soil is a pre-requisite for optimum plant antitranspirants are two types:
growth. Excessive loss of water through 1. To act as a physical barrier above the
transpiration leads to wilting. In general, stomata
there are three types of wilting as follows, Colourless plastics, Silicone oil and low
a. Incipient wilting: Water content of viscosity waxes are sprayed on leaves
plant cell decreases but the symptoms are forming a thin film to act as a physical
not visible. barrier (for transpiration) for water but
b. Temporary wilting: On hot summer permeable to CO2 and O2. The success rate
days, the freshness of herbaceous plants of a physical barrier is limited.
reduces turgor pressure at the day time 2. Induction of Stomata closure
and regains it at night. Carbon-di-oxide induces stomatal
c. Permanent wilting: The absorption closure and acts as a natural
of water virtually ceases because the plant antitranspirant. Further, the advantage
cell does not get water from any source of using CO2 as an antitranspirant is its
and the plant cell passes into a state of inhibition of photorespiration. Phenyl
permanent wilting. Mercuric Acetate (PMA), when applied
84

as a foliar spray to plants, induces partial near vein endings (xylem and Phloem).
stomatal closure for two weeks or more The liquid coming out of hydathode is
without any toxic effect. Use of abscisic not pure water but a solution containing a
acid highly induces the closing of stomata. number of dissolved substances.
Dodecenyl succinic acid also effects on 11.6.7 Measurement of Transpiration
stomatal closure.
1. Ganongs potometer
Uses:
Ganongs potometer is used to measure
• Antitranspirants reduce the the rate of transpiration indirectly. In this,
enormous loss of water by transpiration the amount of water absorbed is measured
in crop plants. and assumed that this amount is equal to
• Useful for seedling transplantations the amount of water transpired.
in nurseries. Apparatus consists of a horizontal
11.6.6 Guttation graduated tube which is bent in opposite
directions at the ends. One bent end is wide
During high humidity in the atmosphere,
and the other is narrow. A reservoir is fixed
the rate of transpiration is much reduced.
to the horizontal tube near the wider end.
When plants absorb water in such a
The reservoir has a stopcock to regulate
condition root pressure is developed due
water flow. The apparatus is filled with
to excess water within the plant. Thus
water from reservoir. A twig or a small plant
excess water exudates as liquid from the
is fixed to the wider arm through a split
edges of the leaves and is called guttation.
cock. The other bent end of the horizontal
Example: Grasses, tomato, potato, brinjal
tube is dipped into a beaker containing
and Alocasia. Guttation occurs through
coloured water. An air bubble is introduced
stomata like pores called hydathodes
into the graduated tube at the narrow end
generally present in plants that grow in
(Figure 11.19). keep this apparatus in bright
moist and shady places. Pores are present
sunlight and observe.As transpiration takes
over a mass of loosely arranged cells with
place, the air bubble will move towards
large intercellular spaces called epithem
the twig. The loss is compensated by water
(Figure 11.18). This mass of tissue lies
Figure 11.18: Structure of Hydathode Figure 11.19: Ganongs Potometer

85

absorption through the xylem portion of utilization is known as translocation of
the twig. Thus, the rate of water absorption organic solutes. The term solute denotes
is equal to the rate of transpiration. food material that moves in a solution.
2. Cobalt chloride (CoCl2) paper method 11.7.1 Path of Translocation
Select a healthy dorsiventral leaf and clean
It has now been well established that
its upper and lower surface with dry cotton.
phloem is the path of translocation of
Now place a dry Cobalt chloride (CoCl2)
solutes. Ringing or girdling experiment
strips on both surface and immediately cover
the paper with glass slides and immobilize will clearly demonstrate the translocation
them. It will be observed after some time of solute by phloem.
that the CoCl2 strip of lower epidermis turns 11.7.2 Ringing or girdling experiment
pink. This indicates that CoCl2 becomes
hydrated (CoCl2. 2H2O or CoCl2. 4H2O) The experiment involves the removal of all
due to water vapours coming out through the tissue outside to vascular cambium (bark,
stomata. The rate of transpiration is more on cortex, and phloem) in woody stems except
the lower surface than in the upper surface xylem. Xylem is the only remaining tissue in
of the dorsiventral leaf. the girdled area which connects upper and
lower part of the plant. This setup is placed in a
11.6.8 Significance of transpiration
beaker of water. After some time, it is observed
Transpiration leads to loss of water, as that a swelling on the upper part of the ring
stated earlier in this lesson 95% of absorbed appears as a result of the accumulation of food
water is lost in transpiration. It seems to be material (Figure 11.20). If the experiment
an evil process to plants. However, number continues within days, the roots die first. It
of process like absorption of water, ascent is because, the supply of food material to the
of sap and mineral absorption directly root is cut down by the removal of phloem.
relay on the transpiration. Moreover plants The roots cannot synthesize their food and so
withstand against scorching sunlight due they die first. As the roots gradually die the
to transpiration. Hence the transpiration upper part (stem), which depends on root for
is a “necessary evil” as stated by Curtis. the ascent of sap, will ultimately die.
11.7 Translocation of Organic Solutes
Leaves synthesize food material through
photosynthesis and store in the form of starch
grains. When required the starch is converted
into simple sugars. They must be transported
Ring off bark
to various parts of the plant system for removed
re ved
Swollen
Swol
tissu
tissue
further utilization. However, the site of food Xylem
production (leaves) and site of utilization are
separated far apart. Hence, the organic food
Water
has to be transported to these areas.
The phenomenon of food transportation
from the site of synthesis to the site of Figure 11.20: Ringing
Initial stage experiment
Final stage
86

11.7.3 Direction of Translocation Sink is defined as any organ in plants
Phloem translocates the products of which receives food from source.Example:
photosynthesis from leaves to the area Roots, tubers, developing fruits and
of growth and storage, in the following immature leaves (Figure 11.21).
directions,
11.7.5 Phloem Loading
Downward direction: From leaves to
stem and roots. The movement of photosynthates
(products of photosynthesis) from
Upward direction: From leaves to
mesophyll cells to phloem sieve elements
developing buds, flowers, fruits for
of mature leaves is known as phloem
consumption and storage. Germination
loading. It consists of three steps.
of seeds is also a good example of upward
translocation.
Radial direction: From cells of pith to Why plants transport
cortex and epidermis, the food materials sugars as sucrose
are radially translocated. and not as starch or
glucose or fructose?
11.7.4 Source and Sink
Glucose and Fructose are simple
Source is defined as any organ in plants
monosaccharides, whereas, Sucrose
which are capable of exporting food
is a disaccharide composed of
materials to the areas of metabolism or
glucose and fructose. Starch is a
to the areas of storage. Examples: Mature
polysaccharide of glucose. Sucrose
leaves, germinating seeds.
and starch are more efficient in
energy storage when compared to
glucose and fructose, but starch is
insoluble in water. So it cannot be
transported via phloem and the
next choice is sucrose, being water
soluble and energy efficient, sucrose
is chosen as the carrier of energy
from leaves to different parts of the
plant. Sucrose has low viscosity even
at high concentrations and has no
reducing ends which makes it inert
than glucose or fructose.During
photosynthesis, starch is synthesized
and stored in the chloroplast stroma
and sucrose is synthesized in the
leaf cytosol from which it diffuses
to the rest of the plant.
Figure 11.21: Source and Sink

87

i. Sieve tube conducts sucrose only. 2. Activated diffusion theory
But the photosynthate in chloroplast This theory was first proposed by Mason
mostly in the form of starch or and Maskell (1936). According to this
trios-phosphate which has to be
theory, the diffusion in sieve tube is
transported to the cytoplasm where it
accelerated either by activating the
will be converted into sucrose for further
diffusing molecules or by reducing the
translocation.
protoplasmic resistance to their diffusion.
ii. Sucrose moves from mesophyll to
3. Electro-Osmotic theory
nearby sieve elements by short distance
transport. The theory of electro osmosis was
proposed by Fenson (1957) and Spanner
iii. From sieve tube to sink by (1958). According to this, an electric-
long-distance transport. potential across the sieve plate causes the
11.7.6 Phloem Unloading movement of water along with solutes.
This theory fails to explain several
From sieve elements sucrose is translocated
problems concerning translocation.
into sink organs such as roots, tubers,
4. Munch Mass Flow hypothesis
flowers and fruits and this process is
termed as phloem unloading. It consists Mass flow theory was first proposed by
of three steps: Munch (1930) and elaborated by Crafts
(1938). According to this hypothesis,
1. Sieve element unloading: Sucrose
organic substances or solutes move from
leave from sieve elements.
the region of high osmotic pressure (from
2. Short distance transport: Movement
mesophyll) to the region of low osmotic
of sucrose to sink cells.
pressure along the turgor pressure
3. Storage and metabolism: The final step gradient. The principle involved in this
when sugars are stored or metabolized
hypothesis can be explained by a simple
in sink cells.
physical system as shown in figure 11.22.
11.7.7 Mechanism of Translocation Two chambers “A” and “B” made
Several hypotheses have been proposed to up of semipermeable membranes are
explain the mechanism of translocation. connected by tube “T” immersed in a
Some of them are given below: reservoir of water. Chamber “A” contains
1. Diffusion hypothesis highly concentrated sugar solution
while chamber “B” contains dilute sugar
As in diffusion process, this theory states
solution. The following changes were
the translocation of food from higher
observed in the system,
concentration (from the place of synthesis)
to lower concentration (to the place of i. The high concentration sugar
utilization) by the simple physical process. solution of chamber “A” is in a hypertonic
However, the theory was rejected because state which draws water from the reservoir
by endosmosis.
the speed of translocation is much higher
than simple diffusion and translocation is ii. Due to the continuous entry of
a biological process which any poison can water into chamber “A”, turgor pressure is
halt. increased.
88

iii. Increase in turgor pressure in the phloem to the cells of stem and roots
chamber “A” force, the mass flow of sugar along the gradient turgor pressure.
solution to chamber “B” through the tube In the cells of stem and roots, the organic
“T” along turgor pressure gradient. solutes are either consumed or converted
iv. The movement of solute will continue into insoluble form and the excess water
till the solution in both the chambers is released into xylem (by turgor pressure
attains the state of isotonic condition and gradient) through cambium.
the system becomes inactive.
Merits:
v. However, if new sugar solution is
added in chamber “A”, the system will start i. When a woody or herbaceous plant
to run again. is girdled, the sap contains high sugar
containing exudates from cut end.
A similar analogous system as given in
the experiment exists in plants: ii.
Positive concentration gradient
disappears when plants are defoliated.
Chamber “A” is analogous to mesophyll
cells of the leaves which contain a higher Objections:
concentration of food material in soluble i. This hypothesis explains the
form. In short “A” is the production point unidirectional movement of solute only.
called “source”. However, bidirectional movement of
Chamber “B” is analogous to cells of solute is commonly observed in plants.
stem and roots where the food material is ii. Osmotic pressure of mesophyll
utilized. In short “B” is consumption end cells and that of root hair do not confirm
called “sink”. the requirements.
Tube “T” is analogous to the sieve tube iii. This theory gives passive role to
of phloem. sieve tube and protoplasm, while some
Mesophyll cells draw water from the workers demonstrated the involvement
xylem (reservoir of the experiment) of of ATP.
the leaf by endosmosis leading to increase
in the turgor pressure of mesophyll cell. 11.8 Mineral Absorption
The turgor pressure in the cells of stem Minerals in soil exist in two forms, either
and the roots are comparatively low and dissolved in soil solution or adsorbed by
hence, the soluble organic solutes begin colloidal clay particle. Previously, it was
to flow en masse from mesophyll through mistakenly assumed that absorption of
mineral salts from soil took place along
with absorption of water. But absorption
of minerals and ascent of sap are identified
as two independent processes. Minerals
are absorbed not only by root hairs but
also by the cells of epiblema.
Plasma membrane of root cells are not
Figure 11.22: A model demonstrating permeable to all ions and also all ions of
the Mass flow hyphothesis same salt are not absorbed in equal rate.
89

Penetration and accumulation of ions into tightly but oscillate within a small volume
living cells or tissues from surrounding of space called oscillation volume. Due to
medium by crossing membrane is called small space, both ions overlap each other’s
mineral absorption. Movement of ions oscillation volume and exchange takes
into and out of cells or tissues is termed place (Figure 11.23).
as transport or flux. Entry of the ion ii. Carbonic Acid Exchange Theory:
into cell is called influx and exit is called According to this theory, soil solution
efflux. Various theories have been put plays an important role by acting as
forward to explain this mechanism. a medium for ion exchange. The CO2
They are categorized under passive released during respiration of root cells
mechanisms (without the involvement of combines with water to form carbonic acid
metabolic energy) and active mechanisms (H2CO3). Carbonic acid dissociates into
(involvement of metabolic energy). H+ and HCO3– in the soil solution. These
H+ ions exchange with cations adsorbed
11.8.1 Passive Absorption
on clay particles and the cations from
1. Ion-Exchange: micelles get released into soil solution and
Ions of external soil solution were gets adsorbed on root cells (Figure 11.24).
exchanged with same charged (anion for
anion or cation for cation) ions of the root
cells. There are two theories explaining
this process of ion exchange namely:
i. Contact exchange and ii. Carbonic
acid exchange.
i. Contact Exchange Theory:
According to this theory, the ions
adsorbed on the surface of root cells and
clay particles (or clay micelles) are not held
Figure 11.24: Carbonic Acid Exchange
theory
Figure 11.23: Contact Exchange theory

90

11.8.2 Active Absorption is called as anion respiration or salt
respiration. Based on this observation
Absorption of ions against the
Lundegardh (1950 and 1954) proposed
concentration gradient with the
cytochrome pump theory which is based
expenditure of metabolic energy is called on the following assumptions:
active absorption. In plants, the vacuolar
i. The mechanism of anion and cation
sap shows accumulation of anions and
absorption are different.
cations against the concentration gradient
ii.
Anions are absorbed through
which cannot be explained by theories of
cytochrome chain by an active process,
passive absorption. Mechanism of active
cations are absorbed passively.
absorption of salts can be explained
iii. An oxygen gradient responsible
through Carrier concept.
for oxidation at the outer surface of the
Carrier Concept: membrane and reduction at the inner
This concept was proposed by Van den surface.
Honert in 1937. The cell membrane is According to this theory, the enzyme
largely impermeable to free ions. However, dehydrogenase on inner surface is responsible
the presence of carrier molecules in the for the formation of protons (H+) and
membrane acts as a vehicle to pick up or electrons (e–). As electrons pass outward
bind with ions to form carrier-ion-complex, through electron transport chain there is a
which moves across the membrane. On the corresponding inward passage of anions.
Anions are picked up by oxidized cytochrome
inner surface of the membrane, this complex
oxidase and are transferred to other members
breaks apart releasing ions into cell while
of chain as they transfer the electron to the
carrier goes back to the outer surface to pick
next component (Figure 11.26).
up fresh ions (Figure 11.25).
The theory assumes that cations (C+)
move passively along the electrical gradient
created by the accumulation of anions (A–)
at the inner surface of the membrane.
Figure 11.25: Carrier Concept A-
A- Fe 2+ Fe A- A-
3+
Fe
2+
The concept can be explained using two

e-
OUTSIDE
e
-
theories:
INSIDE
Fe Fe
3+ 3+
Fe 2+
A- A-
1. Lundegardh’s Cytochrome Pump H Dehydrogenase

Theory: ¼O 2 Reactions
Lundegardh and Burstrom (1933)

½H 2O H-
observed a correlation between respiration
c+ c+
and anion absorption. When a plant is
transferred from water to a salt solution
Figure 11.26: Cytochrome Pump theory
the rate of respiration increases which
91

Main defects of the above theory are: cation concentration would be greater in the
(i) Cations also induce respiration. internal than in the external solution. This
(ii) Fails to explain the selective uptake electrical balance or equilibrium controlled
of ions. by electrical as well as diffusion phenomenon
is known as the Donnan equilibrium.
(iii) It explains absorption of anions
only.
Summary
2. Bennet-Clark’s Protein-Lecithin Theory:
In 1956, Bennet-Clark proposed that the There are two types of transports namely
carrier could be a protein associated with short and long distance in plants to
phosphatide called as lecithin. The carrier translocate sap and solutes. Based on
is amphoteric (the ability to act either as an energy requirement, the transport may
acid or a base) and hence both cations and either be passive or active. The process of
anions combine with it to form Lecithin- diffusion, facilitated diffusion, imbibition
ion complex in the membrane. Inside and osmosis are driven by concentration
the membrane, Lecithin-ion complex gradient like a ball rolling down to a slope
is broken down into phosphatidic acid and hence, no energy is needed. The water
and choline along with the liberation absorbed (either active or passive) from
of ions. Lecithin again gets regenerated the soil by root hairs must reach the xylem
from phosphatidic acid and choline in the for further transportation. There are three
presence of the enzyme choline acetylase possible routes to reach the xylem from root
and choline esterase (Figure 11.27). ATP hairs. They are i) apoplast ii) symplast and/
is required for regeneration of lecithin. or iii) transmembrane. Various theories
explain the path of sap in the xylem and
C+
Dixon’s Cohesion-tension theory is the
C+ C+ Phosphatidic
Lecithinase
_ Lecithin
Acid
_ most accepted one. Transpiration is mostly
Choline A
OUTSIDE
A
INSIDE
A
_
Choline
carried out by stomata, which has guard
Ch tera
Acetylase
Es
cells. The general mechanism of stomatal

ol se
ATP
in
e
Acetyl Choline
movement is based on entry and exit of
water molecules in guard cells. Many
theories are there to explain how water
Figure 11.27: Protein-Lecithin theory
enters and exits from guard cells. The
11.8.3 Donnan equilibrium theory of potassium transport enumerates
Within the cell, some of the ions never two different reactions separately run for
diffuse out through the membrane. They opening and closing of stomata. Contrary
are trapped within the cell and are called to ascent of sap by xylem in an upward
fixed ions. But they must be balanced by direction, the path of solute which
the ions of opposite charge. Assuming that consists of the photosynthetic products
a concentration of fixed anions is present is always in phloem and translocate
inside the membrane, more cations would be multidirectional. The point of origin of
absorbed in addition to the normal exchange translocation is photosynthetic leaves
to maintain the equilibrium. Therefore, the which are the source. On the other
92

hand, point of utilization is called 5. Munch hypothesis is based on
sink. According to Munch mass flow a. Translocation of food due to TP
hypothesis, the solutes move along the gradient and imbibition force
concentration gradient in a bulk flow. b. Translocation of food due to TP
Although minerals are dissolved in c. Translocation of food due to
soil water, they do not tend together imbibition force
with water to enter the root hairs during d. None of the above
absorption of water. Mineral absorption is 6. If the concentration of salt in the soil is
independent of water absorption. Minerals too high and the plants may wilt even
are absorbed either actively or passively. if the field is thoroughly irrigated.
Explain
Evaluation
7. How phosphorylase enzyme
1. In a fully turgid cell open the stomata in starch sugar
a. DPD = 10 atm; OP = 5 atm; interconversion theory?
TP = 10 atm 8. List out the non-photosynthetic
b. DPD = 0 atm; OP = 10 atm; parts of a plant that need a supply of
TP = 10 atm sucrose?
c. DPD = 0 atm; OP = 5 atm; 9. What are the parameters which
TP = 10 atm control water potential?
d. DPD = 20 atm; OP = 20 atm; 10. An artificial cell made of selectively
TP = 10 atm permeable membrane immersed in a
2. Which among the following is correct? beaker (in the figure). Read the values
i. apoplast is fastest and operate in and answer the following questions?
nonliving part
ii. Transmembrane route includes
vacuole
iii. symplast interconnect the nearby
cell through plasmadesmata
iv. symplast and transmembrane route a. Draw an arrow
are in living part of the cell to indicate the direction of water
a. i and ii b. ii and iii movement
c. iii and iv d. i, ii, iii, iv b. Is the solution outside the cell
isotonic, hypotonic or hypertonic?
3. What type of transpiration is possible
c. Is the cell isotonic, hypotonic or
in the xerophyte Opuntia?
hypertonic?
a. Stomatal b. Lenticular d. Will the cell become more flaccid,
c. Cuticular d. All the above more turgid or stay in original size?
4. Stomata of a plant open due to e. With reference to artificial cell
a. Influx of K+ b. Efflux of K+ state, the process is endosmosis or
c. Influx of Cl– d. Influx of OH– exosmosis? Give reasons
93

t ICT Corner
Membrane transport
Let’s play with

membrane proteins.
Steps
• Open PhET:
Method 1: By scanning the QR Code given
Method 2: Through Google – Open PhET by typing PhET
• Select play with simulation & enter
• Click Biology – select Membrane Channels & run
• Select Membrane channel in PhET
• Select round molecule and pump it by pressing red button in one column
• Select square molecule and pump it by pressing the same action
• Observe the movement of molecules across membrane
Activity
• Use leakage channel and gated channel in closed and open position and observe
the molecules movement.
Step 1 Step 2
Step 3 Step 4
URL:
https://phet.colorado.edu/
* Pictures are indicative only
94

Chapter
12 Mineral Nutrition
Learning Objectives A solution to Pollution

• Recognise the need of mineral
nutrition. Pebble Gunny bags
• Analyse the classification and
styrofoam
criteria for essential minerals.
Bamboo tray
• Learn the techniques of Hydroponics
and Aeroponics.
• Correlate different types of special Root hairs Microbes
modes of nutrition.
Bacteria
• Ability to recall and analyse
nitrogen fixation.
A new solution has come up for high
Chapter Outline nutrient pollution and eutrophication
12.1 Classification of Minerals in surface waters. Floating Treatment
12.2 Functions, mode of absorption, Wetlands (FTWs) offer promising
deficiency symptoms of solution and it is a built structure
Macronutrients which measures around 3,000 sq.ft
and comprises four layers: floatable
12.3 Functions, mode
bamboo at base, styrofoam second
of absorption and
layer, a third layer of gunny bags with
deficiency symptoms of
gravels and final layer to support
Micronutrients
cleaning agents (plants). Native plants
12.4 Deficiency Diseases and symptoms
including Vetivers, Citronella, Tulsi
12.5 Critical Concentration and Toxicity and Withania are being researched
of minerals for use as cleaning agents. FTW works
12.6 Hydroponics and Aeroponics on the principle of Hydroponics
12.7 Nitrogen Fixation which is explained in this chapter.
12.8 Nitrogen Cycle and Nitrogen Microbes grown on the roots of these
Metabolism plants break down and consume organic
matter in water and reduce pollution.
12.9 Special Modes of Nutrition
95

As a traveller you would have got 12.1 Classification of minerals
a chance to observe the plants. It is an
interesting fact that all plants are not 12.1.1 Classification of minerals based
unique. Just spend some time to listen on their quantity
to nature. You can notice plants with Essential elements are classified as
attractive leaves, flowers and fruits. Macronutrients, Micronutrients and
Can you say all plants are healthy and Unclassified minerals based on their
uniform in growth? Some plants are not requirements. Essential minerals which
healthy and show symptoms like texture are required in higher concentration are
changes, stunted growth, chlorosis, necrosis called Macronutrients. Essential minerals
and so on. Can you tell what is the reason which are required in less concentration
for all these symptoms? It may be due to called are as Micronutrients.
infection of microbial pathogens or climatic Minerals like Sodium, Silicon, Cobalt
factors or due to mineral deficiency. and Selenium are not included in the list
In this chapter we are going to learn
about classification of minerals, their Historical events in mineral
functions, deficiency diseases and
nutrition
symptoms, nitrogen metabolism and
special modes of nutrition. Further, Van Helmont (1648) – made first
how can these ideas help us to improve observation of mineral nutrition,
productivity in agriculture? noticed over a period of 5 years soil
lost only 56 g in nourishing a seedling
Plants naturally obtain nutrients from
into tree. Increase in organic substance
atmosphere, water and soil. Carbon,
comes from water alone.
hydrogen and oxygen are called as skeletal
elements and constitute about 94% of Wood word (1699) – Soil provides
mineral nutrients required for their
dry weight. These elements play an
growth.
important role in the formation of organic
compounds such as carbohydrates, fats De Saussure (1804) – plant growth
and protein. These non-mineral elements depends on nitrogen and other
are obtained from air and water. Minerals elements absorbed by roots from soil.
are classified based on essentiality. Arnon Liebig (1840) – gave the “law
and Stout (1939) gave criteria required for of minimum” which states that
essential minerals: productivity of soil depends on
1. Elements necessary for growth and amount of essential elements present
development. in minimum quantity.
Julius Von Sachs (1860) – Demon-
2. They should have direct role in the
strated growing a plant in a defined
metabolism of the plant.
nutrient solution.
3. It cannot be replaced by other elements.
William Frederick Goerick (1940) –
4. Deficiency makes the plants impossible Gave the term Hydroponics and com-
to complete their vegetative and mercial technique.
reproductive phase.
96

Table 12.1: Mineral Types
Macro nutrients Micro nutrients Unclassified minerals
Excess than 10 mmole Kg-1 Less than 10 mmole Kg-1 Required for some plants
in tissue concentration or in tissue concentration or in trace amounts and have
0.1 to 10 mg per gram of equal or less than 0.1 mg some specific functions.
dry weight. per gram of dry weight.
Example: C, H, O, N, P, K, Example: Fe, Mn, Cu, Mo, Example: Sodium, Cobalt,

Ca, Mg and S Zn, B, Cl and Ni Silicon and Selenium
of essential nutrients but are required by mobile minerals and 2. Relatively

some plants, these minerals are placed immobile minerals (Figure 12.1).
in the list of unclassified minerals. These
minerals play specific roles for example, a. Actively mobile minerals
Silicon is essential for pest resistance, Nitrogen, Phosphorus, Potassium,
prevent water lodging and aids cell wall Magnesium, Chlorine, Sodium, Zinc and
formation in Equisetaceae (Equisetum), Molybdenum.
Cyperaceae and Gramineae (Table 12. 1). Deficiency symptoms first appear on
old and senescent leaves due to active
12.1.2 Classification of minerals based
movement of minerals to younger leaves.
on mobility
b. Relatively immobile minerals
If you observe where the deficiency Calcium, Sulphur, Iron, Boron and
symptoms appear first, you can notice Copper shows deficiency symptoms first
differences in old and younger leaves. It is that appear on young leaves due to the
mainly due to mobility of minerals. Based immobile nature of minerals
on this, they are classified into 1. Actively
12.1.3 Classification of minerals based
on their functions
a.
Structural component minerals:
Minerals like Carbon, Hydrogen,
Oxygen and Nitrogen
b. Enzyme function: Molybdenum (Mo)
X
is essential for nitrogenase enzyme
during reduction of atmospheric
Mobile minerals Immobile minerals nitrogen into ammonia. Zinc (Zn)
Necrosis Minerals Chlorosis is an important activator for alcohol
dehydrogenase and carbonic
Movement of Minerals
anhydrase. Magnesium (Mg) is the
X Movement blocked
activator for RUBP carboxylase-
Figure 12.1: Mobility of Minerals oxygenase and PEP carboxylase.
97

Nickel (Ni) is a constituent of urease balance by ion-exchange. It is absorbed
and hydrogenase. as K1 ions.
c. Osmotic Potential: Potassium (K) Deficiency symptoms: Marginal
plays a key role in maintaining osmotic chlorosis, necrosis, low cambial activity,
potential of the cell. The absorption loss of apical dominance, lodging in
of water, movement of stomata and cereals and curled leaf margin.
turgidity are due to osmotic potential. 4. Calcium (Ca): It is involved in synthesis
d. Energy components: Magnesium of calcium pectate in middle lamella,
(Mg) in chlorophyll and phosphorous mitotic spindle formation, mitotic
(P) in ATP. cell division, permeability of cell
membrane, lipid metabolism, activation
12.2 Functions, mode of absorption of phospholipase, ATPase, amylase and
and deficiency symptoms of activator of adenyl kinase. It is absorbed
macronutrients as Ca21 exchangeable ions.
Macronutrients, their functions, their Deficiency symptoms: Chlorosis,

mode of absorption, deficiency symptoms necrosis, stunted growth, premature
and deficiency diseases are discussed here: fall of leaves and flowers, inhibit seed
1. Nitrogen (N): It is required by the formation, Black heart of Celery,
plants in greatest amount. It is an Hooked leaf tip in Sugar beet, Musa
essential component of proteins, and Tomato.
nucleic acids, amino acids, vitamins,
5. Magnesium (Mg): It is a constituent
hormones, alkaloids, chlorophyll and
of chlorophyll, activator of enzymes
cytochrome. It is absorbed by the
of carbohydrate metabolism (RUBP
plants as nitrates (NO3).
Carboxylase and PEP Carboxylase)
Deficiency symptoms: Chlorosis,

and involved in the synthesis of DNA
stunted growth, anthocyanin formation.
and RNA. It is essential for binding of
2. Phosphorus (P): Constituent of
ribosomal sub units. It is absorbed as
cell membrane, proteins, nucleic
Mg21 ions.
acids, ATP, NADP, phytin and sugar
phosphate. It is absorbed as H2PO41 Deficiency symptoms: Inter veinal

and HPO42 ions. chlorosis, necrosis, anthocyanin
Deficiency symptoms: Stunted (purple) formation and Sand drown of
growth, anthocyanin formation, tobacco.
necrosis, inhibition of cambial activity, 6. Sulphur (S): Essential component
affect root growth and fruit ripening. of amino acids like cystine, cysteine
3. Potassium (K): Maintains turgidity and methionine, constituent of
and osmotic potential of the cell, coenzyme A, Vitamins like biotin and
opening and closure of stomata, thiamine, constituent of proteins and
phloem translocation, stimulate ferredoxin.plants utilise sulphur as
activity of enzymes, anion and cation sulphate (SO42) ions.
98

Deficiency symptoms: Chlorosis,
in many plants. Example: Boron is essential
anthocyanin formation, stunted growth, for translocation of sugars, molybdenum
rolling of leaf tip and reduced nodulation is involved in nitrogen metabolism and
in legumes. zinc is needed for biosynthesis of auxin.
Here, we will study about the role of micro
nutrients, their functions, their mode of
NPK Fertilizers absorption, deficiency symptoms and
It consists of nitrogen, deficiency diseases.
phosphate with 1. Iron (Fe): Iron is required lesser
potassium in different than macronutrient and larger than
proportions. The number labelled on micronutrients, hence, it can be
the bags as 15-15-15 indicates N, P & placed in any one of the groups. Iron is
K in equal proportions. an essential element for the synthesis
of chlorophyll and carotenoids. It
is the component of cytochrome,
ferredoxin, flavoprotein, formation
Chelating Agents of chlorophyll, porphyrin, activation
EDTA (Chemical Chelating Agent) of catalase, peroxidase enzymes. It is
Plants which are growing in alkaline absorbed as ferrous (Fe 21) and ferric
soil when supplied with all nutrients (Fe 31) ions. Mostly fruit trees are
including iron will show iron sensitive to iron.
deficiency. To rectify this, we have to Deficiency: Interveinal Chlorosis,

make iron into a soluble complex by formation of short and slender stalk and
adding a chelating agent like EDTA inhibition of chlorophyll formation.
(Ethylene Diamine Tetra Acetic acid)
2. Manganese (Mn): Activator of
to form Fe-EDTA.
carboxylases, oxidases, dehydrogenases
Siderophores (Biological Chelating and kinases, involved in splitting of
agent)
water to liberate oxygen (photolysis). It
Siderophores (iron carriers) are is absorbed as manganous (Mn21) ions.
Iron chelating agents produced by
Deficiency: Interveinal chlorosis,
bacteria. They are used to chelate
grey spot on oats leaves and poor root
ferric Iron (Fe31) from environment
system.
and host.
3. Copper (Cu): Constituent of
plastocyanin, component of
12.3 Functions, mode of absorption phenolases, tyrosinase, enzymes
and deficiency symptoms of involved in redox reactions, synthesis of
micronutrients ascorbic acid, maintains carbohydrate
Micronutrients even though required and nitrogen balance, part of oxidase
in trace amounts are essential for the and cytochrome oxidase. It is absorbed
metabolism of plants. They play key roles as cupric (Cu21) ions.
99

Deficiency: Die back of citrus,

Reclamation disease of cereals and Calmodulin
legumes, chlorosis, necrosis and Calmodulin is a Ca21
Exanthema in Citrus. modulating protein
4. Zinc (Zn): Essential for the synthesis of in eukaryotic cells. It
Indole acetic acid (Auxin) activator of is a heat stable protein involved in
carboxylases, alcohol dehydrogenase, fine metabolic regulations.
lactic dehydrogenase, glutamic acid
dehydrogenase, carboxy peptidases 12.4 Deficiency diseases and symptoms
and tryptophan synthetase. It is
The following table (Table 12.2) gives
absorbed as Zn21 ions.
you an idea about Minerals and their
Deficiency: Little leaf and mottle leaf
Deficiency symptoms:
due to deficiency of auxin, Inter veinal
chlorosis, stunted growth, necrosis
and Khaira disease of rice. Activity
5. Boron (B): Translocation of
Collect leaves showing mineral
carbohydrates, uptake and utilisation
deficiency. Tabulate the symptoms
of Ca11, pollen germination, nitrogen
like Marginal Chlorosis, Interveinal
metabolism, fat metabolism, cell
Chlorosis, Necrotic leaves,
elongation and differentiation. It is
Anthocyanin formation in leaf, Little
absorbed as borate BO32 ions.
leaf and Hooked leaf. (Discuss with
Deficiency: Death of root and shoot

your teacher about the deficiency of
tips, premature fall of flowers and
minerals)
fruits, brown heart of beet root,
internal cork of apple and fruit cracks.
6. Molybdenum (Mo): Component of
nitrogenase, nitrate reductase, involved 12.5 Critical concentration and toxicity
in nitrogen metabolism, and nitrogen of minerals
fixation. It is absorbed as molybdate
12.5.1 Critical Concentration
(Mo21) ions.
Deficiency:
Chlorosis, necrosis, To increase the productivity and also
delayed flowering, retarded growth to avoid mineral toxicity knowledge of
and whip tail disease of cauliflower. critical concentration is essential. Mineral
nutrients lesser than critical concentration
7. Chlorine (Cl): It is involved in Anion –
cause deficiency symptoms. Increase
Cation balance, cell division, photolysis
of mineral nutrients more than the
of water. It is absorbed as Cl2 ions.
normal concentration causes toxicity. A
Deficiency: Wilting of leaf tips concentration, at which 10 % of the dry
8. Nickel (Ni): Cofactor for enzyme weight of tissue is reduced, is considered
urease and hydrogenase. as toxic. Figure 12.2 explains about Critical
Concentration.
Deficiency: Necrosis of leaf tips.
100

Table 12.2: Deficiency diseases and Symptoms
Name of the deficiency
Deficiency minerals
disease and symptoms
1. Chlorosis (Overall) Nitrogen, Potassium, Magnesium, Sulphur,
Iron, Manganese, Zinc and Molybdenum.
a. Interveinal chlorosis Magnesium, Iron, Manganese and Zinc
b. Marginal chlorosis Potassium
2. Necrosis (Death of the tissue) Magnesium, Potassium, Calcium, Zinc,
Molybdenum and Copper.
3. Stunted growth Nitrogen, Phosphorus, Calcium,
Potassium and Sulphur.
4. Anthocyanin formation Nitrogen, Phosphorus, Magnesium and
Sulphur
5. Delayed flowering Nitrogen, Sulphur and Molybdenum
6. Die
back of shoot, Reclamation disease, Copper
Exanthema in citrus (gums on bark)
7. Hooked leaf tip Calcium
8. Little Leaf Zinc
9. B
rown heart of turnip and Boron
Internal cork of apple
10. Whiptail of cauliflower and cabbage Molybdenum
11. Curled leaf margin Potassium
10% Reduction 12.5.2 Mineral Toxicity

In Plant Growth
Adequate Zone a. Manganese toxicity

Increased Concentration of Manganese
Growth as a % of Maximum Rate
Toxic
Zone
Transition will prevent the uptake of Fe and Mg,
Zone prevent translocation of Ca to the shoot
Deficient Zone
apex and cause their deficiency. The

symptoms of manganese toxicity are
appearance of brown spots surrounded by
Critical Nutrient
chlorotic veins.
Toxic
Concentration Concentration b. Aluminium Toxicity
Aluminium toxicity causes precipitation
Concentration of Nutrient in Plant Tissue
of nucleic acid, inhibition of ATPase,
12.4 Critical Concentration and Mineral Toxicity
Figure 12.2: Critical Concentration
101

hydroponics. In hydroponics roots are
Iron and Manganese toxicity immersed in the solution containing
Iron and Manganese exhibit nutrients and air is supplied with help
competitive behaviour. Deficiency of of tube (Figure 12.3).
Fe and Mn shows similar symptoms. Aeroponics: This technique was
Iron toxicity will affect absorption of developed by Soifer Hillel and David
manganese. The possible reason for Durger. It is a system where roots
iron toxicity is excess usage of chelated are suspended in air and nutrients
iron in addition with increased acidity are sprayed over the roots by a motor
of soil (PH less than 5.8) Iron and driven rotor (Figure 12.4).
manganese toxicity will be solved by
using fertilizer with balanced ratio of
Fe and Mn.
Suspended
inhibition of cell division and binding of root
plasma membrane with Calmodulin. Mist
chamber
For theories regarding, translocation Nutrient
of minerals please refer Chapter- 11. Solution
Spray rotor
12.6 Hydroponics and Aeroponics Figure 12.4: Aeroponics
1.
Hydroponics or Soilless culture:
Von Sachs developed a method of 12.7 Nitrogen Fixation
growing plants in nutrient solution. Inspiring act of nature is self-regulation.
The commonly used nutrient solutions As all living organisms act as tools for
are Knop solution (1865) and Arnon bio geo chemical cycles, nitrogen cycle is
and Hoagland Solution (1940). highly regulated. Life on earth depends
Later the term Hydroponics was on nitrogen cycle. Nitrogen occurs in
coined by Goerick (1940) and he also atmosphere in the form of N2 (N≡N), two
introduced commercial techniques for nitrogen atoms joined together by strong
Diagram of Hydroponic System
Air pump
Buoyant pads to
support the plants Water
circulation
pump
Nutrient Air bubble

solution
Air pipe
Figure 12.3: Hydroponics
Figure 12.5 Hydroponics
102

12.7.2 Biological nitrogen fixation
Activity
Symbiotic bacterium like Rhizobium fixes
Preparation of Solution Culture to atmospheric nitrogen. Cyanobacteria
find out Mineral Deficiency found in Lichens, Anthoceros, Azolla and
coralloid roots of Cycas also fix nitrogen.
1. Take a glass jar or polythene bottle
non-symbiotic (free living bacteria) like
and cover with black paper (to
Clostridium also fix nitrogen.
prevent algal growth and roots
reacting with light). a. Symbiotic nitrogen fixation
2. Add nutrient solution. i. Nitrogen fixation with nodulation
3. Fix a plant with the help of split Rhizobium bacterium is found in leguminous
cork. plants and fix atmospheric nitrogen. This
4. Fix a tube for aeration. kind of symbiotic association is beneficial for
both the bacterium and plant. Root nodules
5.
Observe the growth by adding
are formed due to bacterial infection.
specific minerals.
Rhizobium enters into the host cell and
proliferates, it remains separated from the
triple covalent bonds. The process of host cytoplasm by a membrane (Figure 12.6).
converting atmospheric nitrogen (N2) into
Stages of Root nodule formation:
ammonia is termed as nitrogen fixation.
Nitrogen fixation can occur by two 1. Legume plants secretes phenolics
methods: 1. Biological; 2. Non-Biological which attracts Rhizobium.
(Figure 12.5). 2. Rhizobium reaches the rhizosphere
and enters into the root hair, infects
12.7.1 Non – Biological nitrogen fixation the root hair and leads to curling of
• Nitrogen fixation by chemical process root hairs.
in industry. 3. Infection thread grows inwards and
• Natural electrical discharge during separates the infected tissue from
lightening fixes atmospheric nitrogen. normal tissue.
NITROGEN FIXATION
Non-Biological Biological
Industrial Lightening Non symbiotic Symbiotic
Legume Non legume

Figure 12.5: Nitrogen fixation
103

b. Non-symbiotic Nitrogen fixation
Free living bacteria and fungi also fix
atmospheric nitrogen.
Aerobic Azotobacter, Beijer-

neckia and Derxia
Anaerobic Clostridium
Photosynthetic Chlorobium and
Rhodospirillum
Figure 12.6: Rhizobium (Bacteroid) in Chemosynthetic Disulfovibrio
root nodule Free living fungi Yeast and Pullularia
4. A membrane bound bacterium is Cyanobacteria Nostoc, Anabaena
formed inside the nodule and is called and Oscillatoria.
bacteroid.
5. Cytokinin from bacteria and auxin 12.8 Nitrogen cycle and nitrogen
from host plant promotes cell division metabolism
and leads to nodule formation
12.8.1 Nitrogen cycle
This cycle consists of
Activity following stages:
• C ollect roots of legumes with root 1. Fixation of

nodules. atmospheric nitrogen
• Take cross section of the root nodule. Di-nitrogen molecule
from the atmosphere
• Observe under microscope. Discuss
progressively gets reduced by addition of
your observations with your teacher.
a pair of hydrogen atoms. Triple bond
between two nitrogen atoms (N≡N) are
cleaved to produce ammonia (Figure 12.7).
Non-Legume nitrogen fixation process requires
Alnus and Casuarina contain the Nitrogenase enzyme complex, Minerals
bacterium Frankia. Psychotria contains (Mo, Fe and S), anaerobic condition, ATP,
the bacterium Klebsiella. electron and glucose 6 phosphate as H1
donor. Nitrogenase enzyme is active only
ii. Nitrogen fixation without nodulation
in anaerobic condition. To create this
The following plants and prokaryotes anaerobic condition a pigment known
are involved in nitrogen fixation. as leghaemoglobin is synthesized in the
Lichens - Anabaena and Nostoc nodules which acts as oxygen scavenger and
Anthoceros - Nostoc removes the oxygen. Nitrogen fixing bacteria
Azolla - Anabaena azollae in root nodules appears pinkish due to the
presence of this leghaemoglobin pigment.
Cycas - Anabaena and Nostoc
104

Enzyme Dinitrogen 3. Nitrate Assimilation
Nitrogenase N Molecule
The process by which nitrate is reduced
N to ammonia is called nitrate assimilation
and occurs during nitrogen cycle.
N
N Nitrate reductase
NO32 NO22
Mo
Nitrite reductase
N NO22 NH31
H Cu, Fe

N H
H 4. Ammonification
N H
Decomposition of organic nitrogen
N H (proteins and amino acids) from dead
H
plants and animals into ammonia is called
H
H ammonification. Organisim involved
N H
in this process are Bacillus ramosus and
N H
H Bacillus vulgaris.
H
5. Denitrification
H Nitrates in the soil are converted back into
N H
H atmospheric nitrogen by a process called
H denitrification. Bacteria involved in this
N H
H process are Pseudomonas, Thiobacillus
Nitrogenase Ammonia and Bacillus subtilis.
Figure 12.7: Nitrogenase enzyme function
Pseudomonas
Nitrate Molecular Nitrogen
Overall equation: (NO32) (N2)
N2 1 8e2 1 8H1 1 16ATP
The overall process of nitrogen cycle is
2NH31 1 H2 1 16ADP 1 16 Pi given in Figure 12.8.
2. Nitrification
12.8.2 Nitrogen Metabolism
Ammonia (NH31) is converted into Nitrite
(NO22) by Nitrosomonas bacterium. Ammonium Assimilation (Fate of
Nitrite is then converted into Nitrate Ammonia)
(NO32) by Nitrobacter bacterium. Ammonia is converted into amino acids
Plants are more adapted to absorb nitrate by the following processes:
(NO32) than ammonium ions from the soil.
1. Reductive amination
Nitrosomonas
2 NH31 1 3 O2 2 NO22 1 2 H1 1 2H2O Glutamic acid or glutamate is formed by
reaction of ammonia with α-ketoglutaric
Nitrobacter
2 NO22 1 O2 2 NO3- acid.
105

Atmospheric Nitrogen (N2)
Plants
Assimilation Denitrifying
bacteria
Nitrogen-fixing _
bacteria living in Nitrates (NO3)
legume root
nodules
Decomposers
(aerobic and anaerobic
bacteria and fungi) Nitrifying
bacteria
Ammonification Nitrification
Ammonification _
Nitrites (NO2)
(NH4+)
Nitrogen-fixing Nitrifying bacteria
soil bacteria
Figure 12.8: Nitrogen Cycle

Glutamate dehydrogenase (GDH)
α-Ketoglutaric acid 1 NH41 Glutamate 1 H2O
NADPH1H NADP1 1
2. Transamination
Transfer of amino group (NH31) from R1 R2 R1 R2
glutamic acid glutamate to keto group of keto
acid. Glutamic acid is the main amino acid C5O C5O CH–NH31
CH–NH31
from which other amino acids are synthesised 1 5 1
by transamination. Transamination requires COO2 COO 2
COO 2
COO2
the enzyme transaminase and co enzyme Amino Keto Keto Amino
acid 2 acid 1 acid 2
pyridoxal phosphate (derivative of vitamin acid 1
B6 -pyridoxine)
106

3. Catalytic Amination: (GS/GOGAT
Pathway)
Glutamate amino acid combines with
ammonia to form the amide glutamine.
Glutamine Synthetase (GS)
Glutamate 1 NH41 Glutamine.
ATP ADP 1 Pi
Glutamine reacts with α ketoglutaric acid

to form two molecules of glutamate.
GOGAT (enzyme) Neottia Monotropa

Glutamine 1 α Ketoglutaric acid 2 Glutamate (Bird's Nest Orchid) (Indian Pipe)
(2- oxoglutarate)
NADH1H1 NAD1 Figure 12.9: Saprophytic Mode of nutrition
(GOGAT- Glutamine-2-Oxoglutarate
aminotransferase) 12.9.2 Parasitic mode of nutrition in
angiosperms
12.9 Special modes of nutrition Organisms deriving their nutrient from
Nutrition is the process of uptake and another organism (host) and causing
utilization of nutrients by living organisms. disease to the host are called parasites.
There are two main types such as a. Obligate or Total parasite - Completely
autotrophic and heterotrophic nutrition. depends on host for their survival and
Autotrophic nutrition is further divided produces haustoria.
into photosynthetic and chemosynthetic i. Total stem parasite: The leafless stem
nutrition. Heterotrophic nutrition is twine around the host and produce
further divided into saprophytic, parasitic, haustoria. Example: Cuscuta (Dodder),
symbiotic and insectivorous type. In this a rootless plant growing on Zizyphus,
topic you are going to learn about special Citrus and so on.
mode of nutrition.
ii.
Total root parasite: They do not
12.9.1 Saprophytic mode of nutrition in have stem axis and grow in the roots
angiosperms of host plants produce haustoria.
Example: Rafflesia, Orobanche and
Saprophytes derive nutrients from dead
Balanophora.
and decaying matter. Bacteria and fungus
b. Partial parasite - Plants of this group
are main saprophytic organisms. Some
contain chlorophyll and synthesize
angiosperms also follow saprophytic mode
carbohydrates. Water and mineral
of nutrition. Example: Neottia. Roots of
requirements are dependent on host plant.
Neottia (Bird’s Nest Orchid) associate
with mycorrhizae and absorb nutrients i.
Partial Stem Parasite: Example:
as a saprophyte. Monotropa (Indian Loranthus and Viscum (Mistletoe)
Pipe) grow on humus rich soil found in Loranthus grows on fig and mango
thick forests. It absorbs nutrient through trees and absorb water and minerals
mycorrhizal association (Figure 12.9). from xylem.
107

Figure 12.11: Symbiotic mode of nutrition
Nostoc associates with its coralloid
roots. (Figure 12.11).
12.9.4 Insectivorous mode of nutrition

Plants which are growing in nitrogen
Figure 12.10: Parasitic Mode of Nutrition
deficient areas develop insectivorous habit
to resolve nitrogen deficiency.
ii.
Partial root parasite: Example: a. Nepenthes (Pitcher plant): Pitcher is
Santalum album (Sandal wood tree) a modified leaf and contains digestive
in its juvenile stage produces haustoria enzymes. Rim of the pitcher is provided
which grows on roots of many plants with nectar glands and acts as an attractive
(Figure 12.10). lid. When insect is trapped, proteolytic
enzymes will digest the insect.
12.9.3 Symbiotic mode of Nutrition
b. Drosera (Sundew): It consists of long
a. L ichens: It is a mutual association of club shaped tentacles which secrete
Algae and Fungi. Algae prepares food sticky digestive fluid which looks like a
and fungi absorbs water and provides sundew.
thallus structure.
c. Utricularia (Bladder wort): Submerged
b. Mycorrhizae: Fungi associated with plant in which leaf is modified into a
roots of higher plants including bladder to collect insect in water.
Gymnosperms. Example: Pinus.
d. Dionaea (Venus fly trap): Leaf of this
c. Rhizobium and Legumes: This symbiotic plant modified into a colourful trap. Two
association fixes atmospheric nitrogen folds of lamina consist of sensitive trigger
d. Cyanobacteria and Coralloid Roots: hairs and when insects touch the hairs it
This association is found in Cycas where will close (Figure 12.12).
108

higher concentration and micro nutrients
(Fe, Mn, Cu, Zn, B, Mo, Cl and Ni) are
required in lesser concentration. Minerals
like Sodium, Cobalt, Silicon and Selenium
are required by some plants for specific
functions and such minerals are grouped
as unclassified minerals. Actively mobile
elements are N, P, K, Mg, Cl, Na, Zn and
Mo. The deficiency symptoms for these
minerals first appear on old and senescent
leaves due to active movement of minerals
to younger leaves. Relatively immobile
elements are Ca, S, Fe, B and Cu. In such
minerals, deficiency symptoms first appear
on young leaves due to immobile nature.
Minerals and their deficiency symptoms
include chlorosis (loss of chlorophyll
Figure 12.12: Insectivorous mode of
pigments), necrosis (death of tissue),
nutrition anthocyanin formation, die back of shoot,
exanthema, hooked leaf tip, whiptail and
Lichens are indicators so on. A concentration at which 10% of
of SO2 pollution and dry weight is reduced is considered as
a pioneer species in critical concentration. Minerals used in
xeric succession. excess concentration become toxic.
Soil less cultivation alleviates problems
due to mineral deficiency. It includes
Check your grasp! hydroponics and aeroponics. Hydroponics
Mineral X required for the activation is a method of growing plants in a
of enzyme nitrogenase, Mineral Y nutrient solution. Aeroponics is the
involved in transport of sugar and technique in which roots are suspended
Mineral Z required for maintaining over the nutrient medium in air and
ribosome structure. Identify X, Y and Z. nutrient sprayed over the roots by motor
driven rotor. Nitrogen is an important
requirement for normal growth and
Summary functioning of a plant. Nitrogen fixing
Sources of minerals for plants are organisms fix nitrogen from atmosphere
atmosphere, water and soil. Minerals are naturally through symbiotic and non-
classified based on their quantity, mobility symbiotic modes. Special modes of
and functions. Macro nutrients (C, H, O, nutrition are seen in plant which grew in
N, P, K, Ca, Mg and S) are required in nutrient deficient soils and the character
becomes permanent.
109

Evaluation
1. Identify correct match.
1. Die back disease of citrus - (i) Mo
2. Whip tail disease - (ii) Zn
3. Brown heart of turnip - (iii) Cu
4. Little leaf - (iv) B
5. Identify the correct statement
a. 1 (iii) 2 (ii) 3 (iv) 4 (i) i. Sulphur is essential for amino acids
b. 1 (iii) 2 (i) 3 (iv) 4 (ii) Cystine and Methionine
ii. Low level of N, K, S and Mo affect
c. 1 (i) 2 (iii) 3 (ii) 4 (iv)
the cell division
d. 1 (iii) 2 (iv) 3 (ii) 4 (i)
iii. Non-leguminous plant Alnuswhich
2. If a plant is provided with all mineral contain bacterium Frankia
nutrients but, Mn concentration is iv. Denitrification carried out by
increased, what will be the deficiency? nitrosomonas and nitrobacter.
a. Mn prevent the uptake of Fe, Mg a. I, II are correct
but not Ca b. I, II, III are correct
b. Mn increase the uptake of Fe, Mg c. I only correct
and Ca d. all are correct
c. Only increase the uptake of Ca 6. The nitrogen is present in the
d. Prevent the uptake Fe, Mg, and Ca atmosphere in huge amount but
3. The element which is not remobilized? higher plants fail to utilize it. Why?
a. Phosphorous b. Potassium 7. Why is that in certain plants
c. Calcium d. Nitrogen deficiency symptoms appear first in
4. Match the correct combination. younger parts of the plants while in
others, they do so in mature organs?
Minerals Role
8. Plant A in a nutrient medium shows
A Molybdenum 1 Chlorophyll whiptail disease plant B in a nutrient
B Zinc 2 Methionine medium shows a little leaf disease.
Identify mineral deficiency of plant A
C Magnesium 3 Auxin and B?
D Sulphur 4 Nitrogenase 9. Write the role of nitrogenase enzyme
in nitrogen fixation?
a. A-1 B-3 C-4 D-2 10. Explain the insectivorous mode of
nutrition in angiosperms?
b. A-2 B-1 C-3 D-4
c. A-4 B-3 C-1 D-2
d. A-4 B-2 C-1 D-3
110

t ICT Corner
Role of minerals in plant growth
Let’s try to make the

plant blossom
Steps
• Scan the QR code
• Start a new game
• Add lime
• Test the Soil pH by test the sample press grows
• Do it for combination of minerals
Activity
• Change the combination of minerals and test the soil samples
• Find the correct proportion of chemical and specific pH for flowering
• Conclude your observations.
Step 1 Step 2
Step 3 Step 4
Web URL:
http://www.glencoe.com/sites/common_assets/science/virtual_labs/BL04/
BL04.html
111

Chapter
13 Photosynthesis
13.6 A
bsorption spectrum and Action
Learning Objectives spectrum
The learner will be able to, 13.7
Emerson’s experiments & Hill’s
reaction
• Learn the Ultra structure of
13.8 Modern concept of photosynthesis
Chloroplast .
13.9
Photo-oxidation phase of light
• Realise the importance of solar reaction
energy and properties of light. 13.10 Photochemical phase of light
• Acquire knowledge of Quantum, reaction
Quantum yield and Quantum 13.11 Photophosphorylation
requirement. 13.12 Chemiosmotic theory
• Develop curiosity for photosynthetic 13.13 Dark reaction or C3 cycle
experiments like Red drop, Emerson 13.14 Hatch & Slack Pathway or C4 Cycle
Enhancement effect and Hill’s 13.15 CAM cycle or Crassulacean Acid
Reaction. Metabolism
• Analyse the pathway of electron- 13.16 Photorespiration or C2 Cycle
PS I and PS II. 13.17 Factors affecting photosynthesis
13.18 Photosynthesis in bacteria
• Recognise the Photo-Oxidative and
Photo Chemical Pathway. Life on earth is made up of organic
• Develop skill in Photosynthetic compounds. How do we get these organic
pathways and ability to draw C3, compounds? Ultimately, plants are
C4, C2 and CAM cycle. the main source of all kinds of carbon
compounds in this planet. We directly or
indirectly depend on plants for this. Plants
Chapter Outline are the major machinery which produce
organic compounds like carbohydrates,
13.1 Historical events in photosynthesis lipids, proteins, nucleic acids and other
13.2 Definition, Significance and Site of biomolecules.
photosynthesis Though man has reached the glory of
13.3 Photosynthetic pigments achievements still he is not able to imitate
13.4 Spectrum of electromagnetic the metabolic activities of plants which
radiation produces energy resources and other
13.5 Photosynthetic unit
biomolecules.
112

atmospheric oxygen and carbon
A quest for future energy
dioxide level. Photosynthesis consumes
Hydrogen is considered as a promising
atmospheric carbon dioxide which is
energy vector for the next generation.
continuously added by the respiration of
It can be used for “green” electricity
organisms. Photosynthesis is the major
production or developing cogeneration
endergonic reaction. In this chapter,
systems such as fuel cells. The
we will study about the energy yielding
sustainability of its employment
process of photosynthesis and various
depends on the energy source used
types of energy utilization processes to
to synthesize it from hydrogen-rich
produce carbohydrates.
compounds such as water or biomass.
The splitting of water in hydrogen and
13.1 Historical Events in
oxygen by means of solar radiation in
Photolysis is common in plants. Water Photosynthesis
splitting is not an easy process to mimic • Van Helmont (1648) – Increase
artificially but preliminary success is in organic substances comes from
achieved so far. If young minds take water alone by growing a Willow tree
up this as their research ambition a that gains weight but soil loses only
revolution can be made in green energy. 2 ounces of the original weight.
• Stephen Hales (1727) – Father of
O2
In leaf cell Chloroplast Plant Physiology, Plants obtain
e-
PSII nourishment from air and light.
e-
e-
• Joseph Priestley (1772) – Performed
Hydrolysis
experiments with candle, mice
H2O H2 Fuel cell
e- and Mint plant and concluded that
H2 H2
4e- vegetation purifies the air.
2H2O 4H+ +O2
Hydrogen storage
• Jean-Ingen-Housz (1779) –
Confirmed Priestley’s experiment
The plants get energy from sun by that oxygen released by the plants is
converting solar or radiant energy possible only in light.
into chemical energy by the process
• Lavoisier (1783) – Purifying gas
of Photosynthesis, which acts as a
produced by plants in sunlight is
driving force for both biotic and abiotic
Oxygen (Phlogiston) and noxious gas
world. Photosynthesis produces 1700
produced by burning of candle (de
million tonnes of dry matter per year
Phlogiston) is Carbon di oxide.
by fixing 75 × 1012 Kg of carbon every
• Desaussure (1804)- Explained the
year. Photosynthetic organisms use
importance of water in the process of
only 0.2 % of incident solar light on
photosynthesis.
earth. Carbohydrates produced by
photosynthesis are the basic raw material • Dutrochet (1837) – Explained
for respiration and also to produce the importance of Chlorophyll in
many organic compounds. It maintains Photosynthesis.
113

• Von Mayer (1845) – Green plants • Melvin Calvin (1954) – Used
convert solar energy into chemical radioactive 14CO2 and traced path
energy of organic matter. of carbon in the dark phase of
CO2 1 H2O Organic matter 1 O2 photosynthesis or C3 Cycle.
• Emerson et al., (1957) – Reported
• Liebig (1845) – Organic matter of plants
existence of two photosystems
was derived from CO2.
• Hatch and Slack (1965) – Reported C4
• Julius Von Sachs (1854) – Discovered
pathway and CO2 fixation in C4 plants
that product of photosynthesis was starch.
Green substance (chlorophyll) is located • Huber, Michel and Dissenhofer
in special structures (Chloroplast). (1985) – Crystalized photosynthetic
reaction centre of Rhodobacter and
• T.W. Engelmann (1888)- Plotted a ction
received the Nobel Prize in 1988.
spectrum of photosynthesis
• Blackman (1905) – Proposed Law of
13.2 Definition, Significance and Site
Limiting factors.
of Photosynthesis
• Warburg (1920) – Used unicellular
green algae Chlorella for the study of 13.2.1 Definition of Photosynthesis
Photosynthesis. Photosynthesis is referred as photochemical
• Van Neil (1931) – Oxygen released oxidation and reduction reactions
during photolysis comes from water carried out with help of light, converting
and not from CO2. He also conducted solar energy into Chemical energy. It is
experiments in Purple green bacteria and the most important anabolic process.
demonstrated Photosynthesis. Plants and photosynthetic bacteria use
Light simple raw materials like carbon dioxide
2H2A 1 CO2
Chlorophyll water and with the help of light energy
(CH2O)n1H2O1 2A
synthesize carbohydrates and evolve
In Green Sulphur bacteria H2S is oxygen. The overall chemical equation for
the Hydrogen donor which releases photosynthesis is:
Sulphur instead of oxygen. Light
6CO2 1 6H2O C6H12O6 1 6O2↑
• Emerson and Arnold (1932) – Chlorophyll
Existence of light and dark reaction by Ruben and Kamen (1941) demonstrated
flashing light experiments. six molecules of water as insufficient for
• R. Hill (1937) – Explained photolysis the evolution of 6 molecules of O2 and
with the help of isolated chloroplasts and modified the equation as:
Light
electron acceptors in the presence of light. 6CO2 1 12H2O C6H12O6 1 6 H2O1 6O2↑
Chlorophyll
• Ruben and Kamen (1941) – Used
18
O radioactive Oxygen to prove that Photosynthesis is a collection of
oxygen evolves from water. oxidation and reduction reactions
• Arnon, Allen and Whatley (1954) – (Redox reaction).
Used radioactive 14CO2 to show fixation Oxidation- Water is oxidised into oxygen
of CO2 by isolated chloroplast. (loss of electrons).
114

Reduction – CO2 is reduced into
Bioluminescence is
Carbohydrates (gain of electrons).
the production and
In some bacteria, oxygen is not
emission of light by
evolved and is called as non-oxygenic and
a living organism.
anaerobic photosynthesis. Examples:
Bioluminescence is rare in true plants.
Green sulphur, Purple sulphur and green
A team of MIT engineers have created
filamentous bacteria.
living bioluminescent lamps out of
13.2.2 Significance of Photosynthesis watercress plants with the goal of one day
1. Photosynthetic organisms provide replacing conventional electrical lighting
food for all living organisms on earth with the glowing greenery.
either directly or indirectly.
2. It is the only natural process that A sac like membranous system called
liberates oxygen in the atmosphere thylakoid or lamellae is present in stroma
and balances the oxygen level. and they are arranged one above the other
forming a stack of coin like structure called
3. Photosynthesis balances the oxygen
granum (plural grana). Each chloroplast
and carbon cycle in nature.
contains 40 to 80 grana and each granum
4. Fuels such as coal, petroleum and consists of 5 to 30 thylakoids.
other fossil fuels are from preserved
Thylakoids found in granum are called
photosynthetic plants.
grana lamellae and in stroma are called
5. Photosynthetic organisms are the stroma lamellae. Thylakoid disc size is
primary producers on which all 0.25 to 0.8 micron in diameter. A thinner
consumers depend for energy. lamella called Fret membrane connects
6. Plants provide fodder, fibre, fire wood, grana. Pigment system I is located on
timber, useful medicinal products outer thylakoid membrane facing stroma
and these sources come by the act of and Pigment system II is located on inner
photosynthesis. membrane facing lumen of thylakoid.
13.2.3 Site of Photosynthesis Grana lamellae have both PS I and PS II
whereas stroma lamellae have only PS I.
Chloroplasts are the main site of
Chloroplast contains 30–35 Proteins,
photosynthesis and both energy yielding
20–30% phospholipids, 5–10% chlorophyll,
process (Light reaction) and fixation of
4–5% Carotenoids, 70S ribosomes, circular
carbon dioxide (Dark reaction)that takes
DNA and starch grains. Inner surface
place in chloroplast. It is a double wall
of lamellar membrane consists of small
membrane bounded organelle, discoid
spherical structure called as Quantasomes.
or lens shaped, 4–10 µm in diameter and
Presence of 70S ribosome and DNA
1–33 µm in thickness. The membrane
gives them status of semi-autonomy and
is a unit membrane and space between
proves endosymbiotic hypothesis which
them is 100 to 200 A °. A colloidal and
says chloroplast evolved from bacteria.
proteinaceous matrix called stroma is
Thylakoid contains pigment systems which
present inside.
produces ATP and NADPH 1 H1 using
115

(a) (b)
Figure 13.1: (a) 3D view of chloroplast (b) Sectional view of chloroplast
solar energy. Stroma contains enzyme or photosynthetic bacteria which

which reduces carbon dioxide into captures the light energy necessary for
carbohydrates. In Cyanobacteria thylakoid photosynthesis (Table 13.1).
lies freely in cytoplasm without envelope
(Figure 13.1). 13.3.1 Chlorophyll
Chlorophyll a is the primary pigment
13.3 Photosynthetic Pigments which acts as a reaction centre and all
other pigments act as accessory pigments
A photosynthetic pigment is a and trap solar energy and then transfer it
pigment that is present in chloroplasts
Table 13.1: Types of Photosynthetic pigments

Chlorophyll Carotenoids Phycobilins
1. Chlorophyll 'a' (C55H72O5N4Mg)
1. Carotene (C40H56) – 1. Phycocyanin –
– Green plants and
Lycopene (Red) Cyanobacteria
Cyanobacteria
2. Xanthophyll (C40H56O2)-
2. Chlorophyll 'b' (C55H70O6N4Mg)
Yellow colour – 2. Phycoerythrin –
– Green algae and all higher
Violaxanthin, Fucoxanthin Red Algae
plants
(Brown Algae) and Lutein
3. Chlorophyll 'c' (C55H32O5N4Mg) – Dinoflagellates, Diatoms and Brown Algae
4. Chlorophyll 'd' – Red Algae

5. Chlorophyll 'e' – Xathophycean Algae
6. Bacteriochlorophyll 'a'
7. Bacteriochlorophyll 'b'
8. Chlorobium Chlorophyll 650
9. Chlorobium Chlorophyll 666
116

to chlorophyll 'a'. Chlorophyll molecules photosynthesis. Succinic acid an
have a tadpole like structure. It consists of intermediate of Krebs cycle is activated by
Mg-Porphyrin head (Hydrophilic Head) the addition of coenzyme A and it reacts
and (Lipophilic tail) Phytol tail. The with a simple amino acid glycine and the
Porphyrin head consists of four pyrrol reaction goes on to produce chlorophyll 'a'.
rings linked together by C-H bridges. Each Bio synthesis of chlorophyll a requires
pyrrole ring comprises of four carbons Mg, Fe, Cu, Zn, Mn, K and nitrogen. The
and one nitrogen atom. Porphyrin ring absence of any one of these minerals leads
has several side groups which alter the to chlorosis (Recall what you have studied
properties of the pigment. Different side in ‘Mineral Nutrition’).
groups are indicative of various types of
chlorophyll. The Phytol tail made up of ii. Comparison of Chlorophyll – a with
20 carbon alcohol is attached to carbon other pigments
7 of the Pyrrole ring IV. It has a long 1. Chlorophyll 'b' differs from Chlorophyll 'a'
propionic acid ester bond. Long lipophilic in having CHO (aldehyde)group instead
tail helps in anchoring chlorophyll to the of CH3(Methyl) group at the 3rd C atom
lamellae (Figure 13.2). in II Pyrrol ring (Figure 13.2).
i. Biosynthesis of Chlorophyll 2. Chlorophyll 'c' differs from Chlorophyll
Chlorophyll is synthesized from 'a' by lacking phytol tail.
intermediates of respiration and
Figure 13.2: Structures of Chlorophyll 'a' and 'b'

117

3. Chlorophyll 'd' differs from
Chlorophyll 'a' in having O-CHO Separation of Chloroplast pigments
group instead of CH-CH2 group at 2nd by paper Chromatography method
Carbon in the 1st Pyrrol ring. Step 1. Extract chlorophyll pigment
4. Pheophytin resembles Chlorophyll 'a' from the leaves using 80% Acetone.
except that it lacks Mg atom. Instead it Step 2. Allow to concentrate by
has two H atoms. evaporation.
5. Phycobilins have open tetra pyrrols and Step 3. Apply few drops on one
they have neither Mg nor phytol chain. end above 2 cm from the edge of a
chromatographic paper.
13.3.2 Carotenoids
Step 4. A solvent with mixture of
Carotenoids are yellow to orange Petroleum ether and acetone in the
pigments, mostly tetraterpens and these ratio of 9:1 is prepared and poured
pigments absorb light strongly in the blue into development chamber.
to violet region of visible spectrum. These Step 5. Place the strip above the
pigments protect chlorophyll from photo- solvent by placing one end of the strip
oxidative damage. Hence, they are called as touching the solvent.
shield pigments. These pigments absorb
light and transfer these to chlorophyll. Observation
Almost all carotenoid pigments have After one hour observe the
40 carbon atoms. Ripening of fruits, floral chromatographic paper. You can find
colours and leaf colour change during the pigments being separated into
autumn is due to Carotenoids (Carotene four distinct spots (Figure 13. 4).
and Xanthophyll) (Figure 13.3). Chromatography
i. Carotenes: Paper
Orange, Red, Yellow and Brownish
Test tube
pigments, hydrocarbons (Lipids) and most
of them are tetraterpenes(C40H56). Carotene
Carotenes
Xanthophyll
Chlorophyll a
Chlorophyll b
Ether acetone
solvent
Figure 13.3: Changes in Fruit colour due to

Figure 13.4: Paper Chromatography
difference in pigmentation
118

is the most abundant Carotene in plants and 13.4 Spectrum of Electromagnetic
it is a precursor of Vitamin A. Lycopene Radiation
is the red pigment found in the fruits of
tomato, red peppers and roses. In the total electromagnetic
spectrum,visible light is the smallest
ii. Xanthophylls: part. The entire life on earth depends
Yellow (C40H56O2) pigments are like on light and is the driving force for all
carotenes but contain oxygen. Lutein is organisms. Plants have natural potential
responsible for yellow colour change of to utilize solar energy directly. In the
leaves during autumn season. Examples: given picture electromagnetic radiation
Lutein, Violaxanthin and Fucoxanthin. spectrum and components of visible
spectrum are mentioned. The wavelength
13.3.3 Phycobilins of solar radiation which reaches the earth
They are proteinaceous pigments, soluble is between 300 to 2600 nm. The visible
in water, and do not contain Mg and spectrum ranges between 390 to 763 nm
Phytol tail. They exist in two forms such (3900 å to 7630 å). The colour of the light
as 1. Phycocyanin found in cyanobacteria is determined by the wavelength. Energy
2. Phycoerythrin found in rhodophycean of the quantum is inversely proportional
algae (Red algae). to wavelength. Shorter wavelength has
Figure 13.5: Electromagnetic Spectrum

119

more energy than longer wavelength. 13.5 Photosynthetic Unit (Quantasome)
Electromagnetic spectrum consists of
Quantasomes are the morphological
8 types of radiations such as cosmic rays,
expression of physiological
gamma rays, X rays, U-V rays, Visible
photosynthetic units, located on the
light spectrum, infrared rays, electric rays
inner membrane of thylakoid lamellae.
and radio rays (Figure 13. 5).
Each quantasome measures about 180
å × 160 å and 100 åthickness. In 1952,
Light is extremely
Steinman observed granular structures
variable and if radiation
in chloroplast lamellae under electron
is evenly distributed
microscope. Later, Park and Biggins
over the globe it is
(1964) confirmed these granular structures
sufficient to melt 35 m thick ice layer.
as physiological units of photosynthesis
and coined the term Quantasome.
Properties of Light According to them one quantasome
1. Light is a transverse electromagnetic contains about 230 chlorophyll molecules.
wave. A minimum number of chlorophyll and
2. It consists of oscillating electric and other accessory pigments act together
magnetic fields that are perpendicular in a photochemical reaction to release
to each other and perpendicular to the one oxygen or to reduce one molecule of
direction of propagation of the light. CO2. It constitutes a photosynthetic unit.
(Figure 13.7) Emerson and Arnold (1932)
3. Light moves at a speed of 3 × 108 ms–1
based on flashing light experiment found
4.
Wavelength is the distance between 2500 chlorophyll molecules are required
successive crests of the wave. to fix one molecule of CO2. However, the
5. Light as a particle is called photon. reduction or fixation of one CO2 requires
Each photon contains an amount of 10 quanta of light and so each unit would
energy known as quantum. contain 1/10 of 2500 i.e. 250 molecules.
6. The energy of a photon depends on the Usually 200 to 300 chlorophyll molecules
frequency of the light (Figure 13. 6). are considered as a physiological unit of
photosynthesis. According to Emerson
8 quanta of light are required for the release
Electric-field of one oxygen molecule or reduction of one
Component
Carbon dioxide molecule. The quantum
yield is 1/8 or 12 %.
Direction of 13.6 Absorption Spectrum and Action

Propagation
Spectrum
Wavelenth
(λ) 13.6.1 Absorption Spectrum
Magnetic-field component
The term absorption refers to complete
Figure 13.6: Oscillation of electric and
retention of light, without reflection or
magnetic vectors in light
120

Chlorophyll a
Carotene
Thylakoid
Photosynthetic rate / Absorption rate

Chlorophyll b
100
80
Action
60
spectrum
Antenna 40
Molecule Absorption
20
spectrum
0
400 500 600 700
Chlorophyll b
Wavelength (nm)
Carotenoid
Figure 13. 8: Absorption and action
spectrum
Figure 13.7: Quantasome
spectrum is the absorption maxima for

transmission. Pigments absorb different Chlorophyll (a) and Chlorophyll (b).
wavelengths of light. A curve obtained The Action Spectrum is instrumental
by plotting the amount of absorption of in the discovery of the existence of
different wavelengths of light by a pigment two photosystems in O 2 evolving
is called its absorption spectrum. photosynthesis (Figure 13. 8).
• Chlorophyll 'a' and chlorophyll 'b'
absorb quanta from blue and red region 13.7 Emerson’s Experiments and Hill’s
• Maximum absorption peak for different Reaction
forms of chlorophyll 'a' is 670 to 673,
680 to 683 and 695 to 705nm. 13.7.1 Red Drop or Emerson’s First Effect
• Chlorophyll 'a' 680 (P680) and Emerson conducted experiment in
Chlorophyll 'a' 700 (P700) function as Chlorella using only one wavelength of
trap centre for PS II and PS I respectively. light (monochromatic light) at a time and
he measured quantum yield. He plotted
13.6.2 Action Spectrum a graph of the quantum yield in terms
The effectiveness of different wavelength of O2 evolution at various wavelengths of
of light on photosynthesis is measured light. His focus was to determine at which
by plotting against quantum yield. The wavelength the photochemical yield of
curve showing the rate of photosynthesis oxygen was maximum. He found that in
at different wavelengths of light is the wavelength of 600 to 680 the yield
called action spectrum. From the graph was constant but suddenly dropped in the
showing action spectrum, it can be region above 680 nm (red region). The fall
concluded that maximum photosynthesis in the photosynthetic yield beyond red
takes place in blue and red region of region of the spectrum is referred as Red
the spectrum. This wavelength of the drop or Emerson’s first effect.
121

13.7.2 Emerson’s Enhancement Effect Conclusions of Hill’s Reaction:
Emerson modified 1. During photosynthesis oxygen is
his first experiment evolved from water.
by supplying shorter 2. Electrons for the reduction of CO2
wavelength of light are obtained from water.
(red light) along with 3. Reduced substance produced, later
longer wavelength of helps to reduce CO2
light (far red light). He found that the
monochromatic light of longer wavelength 2H2O 1 2A 2 AH2 1 O2
(far red light) when supplemented with A is the Hydrogen acceptor, the common in
shorter wavelength of light (red light) vitro hydrogen acceptors are ferricyanide,
enhanced photosynthetic yield and benzoquinone and Di Chloro Phenol
recovered red drop. This enhancement Indole Phenol (DCPIP).
of photosynthetic yield is referred
to as Emerson’s Enhancement Effect 13.8 Modern Concept of Photosynthesis
(Figure 13.9).
Photosynthesis is an Oxidation and
Reduction process. Water
Rate of Photosynthesis
650 +710 (Red + Far red)

650nm (Red) is oxidised to release O2
710nm (Far red) and CO2 is reduced to
form sugars. The first
phase requires light and
is called light reaction or
λ of light exposed (nm)
Hill’s reaction.
Figure 13.9: Emerson’s Enhancement Effect
1. Light reaction: It is a photochemical
Photosynthetic rate at far red light reaction whereas dark reaction is a
(710 nm) 5 10 thermochemical reaction.
Photosynthetic rate at red light Solar energy is trapped by chlorophyll
(650 nm) 5 43.5 and stored in the form of chemical energy
Photosynthetic rate at red 1 far red (assimilatory power)as ATP and reducing
(650 1 710 nm) 5 72.5 (Enhancement power NADPH 1 H1. NADPH 1 H1
effect). alone are known as reducing powers. This
reaction takes place in thylakoid membrane
13.7.3 Hill’s Reaction of the chloroplast. Oxygen is evolved as a
R. Hill (1937) isolated chloroplasts result of splitting of water molecules by light.
and when they were illuminated in the Light reaction is discussed in two phases:
presence of suitable electron acceptors i. Photo-oxidation Phase:
such as ferricyanide, they were reduced • Absorption of light energy.
to ferrocyanide and oxygen is evolved.
• Transfer of energy from accessory
Hill’s Reaction is now considered to be
pigments to reaction centre.
equivalent to Light Reaction.
• Activation of Chlorophyll 'a' molecule.
122

ii. Photo Chemical Phase: molecule is in an excited state, this excitation
• Photolysis of water and oxygen evolution energy is utilised for the phosphorylation.
• Electron transport and synthesis of Phosphorylation takes place with the help
assimilatory power. of light generated electron and hence it is
known as photophosphorylation.
2. Dark reaction (Biosynthetic phase):
Fixation and reduction of CO2 into 13.9.1 Fluorescence and Phosphorescence
carbohydrates with the help of assimilatory
Normal state of an atom or molecule
power produced during light reaction. This
is called ground state. When a photon
reaction does not require light and is not
of light collides with the chlorophyll
directly light driven. Hence, it is called as
molecule, an electron from outer most
Dark reaction or Calvin-Benson cycle
orbit is moved to higher energy orbit
(Figure 13.10).
causing excitation of chlorophyll. This is
known as excited state. There are three
excited states such as:
1. First singlet state (S1)
2. Second singlet state (S2)
3. First Triplet Sate (T1)
When a red light strikes chlorophyll
molecule, one electron is released from
its ground level (S0) to first singlet state
(S1). It is in unstable state having half-life
period of 10-9 seconds. When a blue light
Figure 13.10: Light and Dark Reaction
strikes chlorophyll molecule, one electron
13.9 Photo-Oxidation Phase of Light is released from its ground level (S0) to
second singlet state (S2). It is because blue
Reaction
light has shorter wavelength and more
The action of photon plays a vital role in energy than red light. This state is also
excitation of pigment molecules to release unstable having half-life period of less
an electron. When the molecules absorb a than 10–12 seconds. Both S1 and S2 states
photon, it is in excited state. When the light being unstable move to ground state S0
source turned off, the high energy electrons by releasing energy through the several
return to their normal low energy orbitals as possible ways.
the excited molecule goes back to its original
stable condition known as ground state. i. Fluorescence
When molecules absorb or emit light they The electron from first singlet state (S1)
change their electronic state. Absorption of returns to ground state (S0) by releasing
blue light excites the chlorophyll to higher energy in the form of radiation energy
energy state than absorption of Red light, (light) in the red region and this is
because the energy of photon is higher when known as fluorescence. Fluorescence
their wavelength is shorter. When the pigment is the immediate emission of absorbed
123

S2 –12
(Half Life 10 Sec.) of absorbed radiations. Pathway of electron
during Phosphorescence: S2 → S1 → T1 → S0
S1 –9
(Half Life 10 Sec.)
13.9.2 Photosystem and Reaction Centre
Energy Level
T1(Half Life 10 –3 Sec.)

λ2 λ1 F P
• Thylakoid membrane contains
Photosystem I (PS I) and Photosystem
S0
II (PS II).
Ground State
• PS I is in unstacked region of granum
Figure 13.11: Fluorescence (F) and facing stroma of chloroplast.
Phosphorescence (P)
• PS II is found in
stacked region of thylakoid
radiations (Figure 13.11). Pathway of
membrane facing lumen of
electron during fluorescence: S1 → S0
thylakoid.
ii. Phosphorescence • Each Photosystem
Electron from Second Singlet State (S2) may consists of central core
return to next higher energy level (S1) by complex (CC) and light harvesting
losing some of its extra energy in the form Complex (LHC) or Antenna
of heat. From first singlet state (S1) electron molecules(Figure 13.12).
further drops to first triplet state (T1). Triplet • The core complex consists of respective
State is unstable having half life time of 10-3 reaction centre associated with proteins,
seconds and electrons returns to ground electron donors and acceptors.
state with emission of light in red region
• PS I – CC I consists of reaction centre
called as phosphorescence (Figure 13.11).
P700 and LHC I.
Phosphorescence is the delayed emission
Electron transfer Primary electron acceptor

Reaction
oton Reaction center center
Ph Chlorophyll
Transfer of energy
Antenna pigment molecules

Figure 13.12: Photosystem
124

Table 13.2: Differences between Photosystem I and Photosystem II
Photosystem I Photosystem II
1. The reaction centre is P700 1. Reaction centre is P680
2. PS I is involved in both cyclic and 2. PS II participates in Non-cyclic
non-cyclic. pathway
3. Not involved in photolysis of water and 3, Photolysis of water and evolution of
evolution of oxygen oxygen take place.
4. It receives electrons from PS II during 4. It receives electrons by photolysis of
non-cyclic photophosphorylation water
5. Located in unstacked region granum 5. Located in stacked region of thylakoid
facing chloroplast stroma membrane facing lumen of thylakoid.
6. Chlorophyll and Carotenoid ratio is 6. Chlorophyll and Carotenoid ratio is
20 to 30:1 3 to 7:1
• PS II – CC II consists of reaction centre of water is due to strong oxidant which is

P680 and LHC II (Table 13.2). yet unknown and designated as Z or Yz.
• Light Harvesting Complex consists of Widely accepted theory proposed by Kok
several chlorophylls, carotenoids and et al.,(1970) explaining photo-oxidation of
xanthophyll molecules. water is water oxidising clock (or) S’ State
• The main function of LHC is to harvest Mechanism. It consists of a series of 5 states
light energy and transfer it to their called as S0, S1, S2, S3 and S4. Each state
respective reaction centre. acquires positive charge by a photon (hv)
and after the S4 state it acquires 4 positive
charges, four electrons and evolution of
13.10 Photo chemical phase of light
oxygen. Two molecules of water go back to
reaction the S0. At the end of photolysis 4 H1,4 e- and
In this phase electrons pass through electron O2 are evolved from water (Figure 13.13).
carrier molecules and generate assimilatory
powers ATP and NADPH 1 H1. Splitting of 2H2O
water molecule generates electrons replacing s4 O2
electrons produced by the light.
13.10.1 Photolysis of Water H+

s3
The process of Photolysis is associated with s0
Oxygen Evolving Complex (OEC) or water H+
splitting complex in pigment system II and H+
is catalysed by the presence of Mn11 and H+
Cl–. When the pigment system II is active s2 s1
it receives light and the water molecule
splits into OH– ions and H1 ions. The OH–
ions unite to form water molecules again Figure 13.13: Oxygen Evolving
and release O2 and electrons. Photolysis Complex (OEC)
125

PQ acts as shuttle between PS II and
4 H 2O 4 H1 1 4 OH–
Cytochrome b6- f complex and PC
4 OH– 2 H 2O 1 O 2 1 4 e – connects
2H2O 4 H1 1 O2 1 4 e– • Cytochrome b6-f and PS I complex.
• ATPase complex or Coupling factor:
13.10.2 Electron Transport Chain of It is found in the surface of thylakoid
Chloroplast membrane. This complex is made up
Figure 13.14: Electron Transport Chain in Chloroplast
Electron transport chain in each of CF1 and CF0 factors. This complex
photosystem involves four complexes: utilizes energy from ETC and converts
• Core Complex (CC): CC I in PS I the ADP and inorganic phosphate (Pi) into
reaction centre is P700, CC II in PS II ATP (Figure 13.14).
the reaction centre is P680
• Light Harvesting Complex or Antenna 13.11 Photophosphorylation
complex (LHC): Phosphorylation taking place during
• Two types: LHC I in PS I and LHC II respiration is called as oxidative
in PS II. phosphorylation and ATP produced
• Cytochrome b6 f complex: It is the by the breakdown of substrate is known
non-pigmented protein complex as substrate level phosphorylation.
connecting PS I and PS II. In this topic, we are going to learn
Plastoquinone (PQ) and Plastocyanin about phosphorylation taking place in
(PC) are intermediate complexes chloroplast with the help of light. During
acting as mobile or shuttle electron the movement of electrons through carrier
carriers of Electron Transport Chain. molecules ATP and NADPH 1 H1 are
126

produced. Phosphorylation is the process ADP+ Pi 2e- FRS
of synthesis of ATP by the addition of ATP Ferredoxin
inorganic phosphate to ADP. The addition
2e- Light
of phosphate here takes place with the Cyt b6
2e
-
help of light generated electron and so 2e-
it is called as photophosphorylation. It ADP+ Pi

Cyt f
2e
-
takes place in both cyclic and non-cyclic P700

ATP PC 2e
-
electron transport.
PS I
LHC I
13.11.1 Cyclic Photophosphorylation
Figure 13.15: Cyclic Photophosphorylation
Cyclic photophosphorylation refers to
the electrons ejected from the pigment
13.11.2 Non-Cyclic Photophosphorylation
system I (Photosystem I) and again cycled
back to the PS I. When the photons When photons are activated reaction
activate P700 reaction centre photosystem centre of pigment system II(P680),
II is activated. Electrons are raised to electrons are moved to the high energy
the high energy level. The primary level. Electrons from high energy
electron acceptor is Ferredoxin Reducing state passes through series of electron
Substance (FRS) which transfers electrons carriers like pheophytin, plastoquinone,
to Ferredoxin (Fd), Plastoquinone (PQ), cytochrome complex, plastocyanin and
cytochrome b6-f complex, Plastocyanin finally accepted by PS I (P700). During
(PC) and finally back to chlorophyll this movement of electrons from PS II to
P700 (PS I). During this movement PS I ATP is generated (Figure 13. 16). PS I
of electrons Adenosine Di Phosphate (P700) is activated by light, electrons are
(ADP) is phosphorylated, by the addition moved to high energy state and accepted
of inorganic phosphate and generates by electron acceptor molecule ferredoxin
Adenosine Tri Phosphate (ATP). Cyclic reducing Substance (FRS). During the
electron transport produces only ATP and downhill movement through ferredoxin,
there is no NADPH 1 H1 formation. At electrons are transferred to NADP1 and
each step of electron transport, electron reduced into NADPH 1 H1 (H1 formed
loses potential energy and is used by from splitting of water by light).
the transport chain to pump H1 ions Electrons released from the
across the thylakoid membrane. The photosystem II are not cycled back. It
proton gradient triggers ATP formation is used for the reduction of NADP1 in
in ATP synthase enzyme situated on the to NADPH 1 H1. During the electron
thylakoid membrane. Photosystem I need transport it generates ATP and hence this
light of longer wave length (> P700 nm). type of photophosphorylation is called
It operates under low light intensity, less non-cyclic photophosphorylation. The
CO2 and under anaerobic conditions electron flow looks like the appearance
which makes it considered as earlier in of letter ‘Z’ and so known as Z scheme.
evolution (Figure 13.15). When there is availability of NADP1 for
reduction and when there is splitting of
127

-2.0 -
FRS 4e
-
Q 4e Ferredoxin
-1.0
Pheophytin
Light -
- -
4e PQ 4e 4e + +
2NADP 2NADPH+H
- Cyt b6,f
0 Light 4e complex 4e-
ADP+ Pi PC
P700
P680 4e
- PS I
+1.0 ATP LHC I
PS II -
2H2O LHC II 4e
Mn , Ca ,Cl
++ ++
-
O2 Evolving O2 +
4H
Complex
Figure 13.16: Non-Cyclic Photophosphorylation
water molecules both PS I and PS II are 13.11.3 Bio energetics of light reaction
activated (Table 13.3). Non-cyclic electron • To release one electron from pigment
transport PS I and PS II both are involved system it requires two quanta of light.
co-operatively to transport electrons from
• One quantum is used for transport of
water to NADP1 (Figure 13.6). In oxygenic
electron from water to PS I.
species non-cyclic electron transport takes
place in three stages. • Second quantum is used for transport
of electron from PS I to NADP1
i. Electron transport from water to P680:
• Two electrons are required to generate
Splitting of water molecule produce one NADPH 1 H1.
electrons, protons and oxygen. Electrons
• During Non-Cyclic electron transport
lost by the PS II (P680) are replaced by
two NADPH 1 H1 are produced and it
electrons from splitting of water molecule.
requires 4 electrons.
ii. Electron transport from P680 to P700:
• Transportation of 4 electrons requires
Electron flow starts from P680 through 8 quanta of light.
a series of electron carrier molecules
like pheophytin, plastoquinone (PQ), Check your grasp!
cytochrome b6-f complex, plastocyanin
Name the products produced from
(PC) and finally reaches P700 (PS I).
Non-Cyclic photophosphorylation?
iii. Electron transport from P700 to NADP1 Why does PS II require electrons from
PS I(P700) is excited now and the electrons water?
pass to high energy level. When electron Can you find the difference in the
travels downhill through ferredoxin, Pathway of electrons during PS I and
NADP1 is reduced to NADPH 1 H1. PS II?
128

Table 13.3 Differences between Cyclic Photophosphorylation and Non-Cyclic
Photophosphorylation
Cyclic Photophosphorylation Non-Cyclic Photophosphorylation
1. PS I only involved 1. PS I and PS II involved
2. Reaction centre is P700 2. Reaction centre is P680
3. Electrons released are cycled back 3. Electron released are not cycled back
4. Photolysis of water does not take place 4. Photolysis of water takes place
5. Only ATP synthesized 5. ATP and NADPH 1 H1are synthesized
6. Phosphorylation takes place at two 6. Phosphorylation takes place at only one
places place
7. It does not require an external electron 7. Requires external electron donor like
donor H2O or H2S
8. It is not sensitive to di chloro di methyl 8. It is sensitive to DCMI and inhibits
urea (DCMI) electron flow
13.12 Chemiosmotic Theory H+
NADP+
Chemiosmosis theory was proposed by +
Cytochromes NADPH+H
P. Mitchell (1966). According to this PS
PS I
b&f
theory electrons are transported along I I
MEN
LU
H+
the membrane through PS I and PS II and H
+
H+ Thylakoid
H membrane
+
connected by Cytochrome b6-f complex. The H

+
H
+
flow of electrical current is due to difference

STROMA
in electrochemical potential of protons across
the membrane. Splitting of water molecule ADP
ATP Synthase
ATP
takes place inside the membrane. Protons
Figure 13.17: Chemiosmotic Theory
or H1 ions accumulate within the lumen
of the thylakoid (H1 increase 1000 to 2000 membrane stimulates ATP generation
times). As a result, proton concentration (Figure 13.17).
is increased inside the thylakoid lumen. The evolution of one oxygen molecule
These protons move across the membrane (4 electrons required) requires 8 quanta
because the primary acceptor of electron is of light. C3 plants utilise 3 ATPs and
located outside the membrane. Protons in 2 NADPH 1 H1 to evolve one Oxygen
stroma less in number and creates a proton molecule. To evolve 6 molecules of
gradient. This gradient is broken down Oxygen 18 ATPs and 12 NADPH 1 H1
due to the movement of proton across the are utilised. C4 plants utilise 5 ATPs and
membrane to the stroma through CFO of the 2 NADPH 1 H1 to evolve one oxygen
ATP synthase enzyme. The proton motive molecule. To evolve 6 molecules of
force created inside the lumen of thylakoid Oxygen 30 ATPs and 12 NADPH 1 H1
or chemical gradient of H1 ion across the are utilised.
129

dioxide into carbohydrates. This reaction
Check your grasp! does not require light. Therefore, it is named
What will be the quanta requirement Dark reaction. Ribulose 1,5 bisphosphate
for complete light reaction which (RUBP) act as acceptor molecule of carbon
releases 6 oxygen molecules? dioxide and fix the CO2 by RUBISCO
Solution: Complete light reaction enzyme. The first product of the pathway is
releases 6 oxygen molecules. If a 3- carbon compound (Phospho Glyceric
one molecule of oxygen evolution Acid) and so it is also called as C3 Cycle. It
requires 8 quanta means, for 6 oxygen takes place in the stroma of the chloroplast.
molecules 6 × 8 5 48 quanta of light M. Melvin Calvin, A.A. Benson and their
required for complete light reaction. co-workers in the year 1957 found this path
way of carbon fixation. Melvin Calvin was
13.13 Dark Reaction or C3 Cycle or awarded Nobel Prize for this in 1961 and
Biosynthetic Phase or Photosynthetic this pathway named after the discoverers
as Calvin-Benson Cycle. Dark reaction
Carbon Reduction (PCR)Cycle
is temperature dependent and so it is also
Biosynthetic phase of photosynthesis called thermo-chemical reaction.
utilises assimilatory powers(ATP and Dark reaction consists of three phases:
NADPH 1 H1) produced during light (Figure 13.18).
reaction are used to fix and reduce carbon
Figure 13.18: Phases of Calvin Cycle
130

3C Dehydrogenase + 3C
(3)CO2 6 NADPH 6 NADP
Glyceraldehyde 3 -Phosphate (G3P) pool
(6) 3-Phospho
Glycerate G3P G3P G3P G3P G3P G3P
RUBISCO 6 ATP Kinase
TN_GOVT_BOTANY_XI_CH13_112-143.indd 131
5C 6 ADP+ 6 Pi
3C
(3) Ribulose DHAP DHAP
1,5-Bis Phosphate Dihydroxy
Acetone
3 ADP Phoshate
Kinase (DHAP)
5C 5C
3 ATP
(3) Ribulose Ribose 7C
5-Phosphate Isomerase 5-Phosphate Aldolase 6C Export
Sedoheptulose
131
7 Phosphate Fructose 1,6
Phosphatase Bis Phosphate
7C
Pi Phosphatase 6C
Pi
Sedoheptulose
5C
Epimerase
1,7 Bis Phosphate Aldolase Fructose 6 Phosphate
Xylulose 4C
5 Phosphate Erythrose
4 Phosphate Glucose 6 Phosphate
5C
phosphate pool
Stromal hexose
Glucose 1 Phosphate
Epimerase Xylulose
5 Phosphate
Starch
Figure 13.19: Calvin cycle
27-04-2018 11:56:34
1. Carboxylation (fixation) requires 3 ATPs and 2 NADPH 1 H1, and
2. Reduction (Glycolytic Reversal) for the fixation of 6 CO2 requires 18 ATPs
3. Regeneration and 12 NADPH 1 H1 during C3 cycle. One
6 carbon compound is the net gain to form
Phase 1- Carboxylation (Fixation) hexose sugar.
The acceptor molecule Ribulose 1,5 ATP ADP
Bisphosphate (RUBP) a 5 carbon compound
RU5P RUBP
with the help of RUBP carboxylase
oxygenase (RUBISCO) enzyme accepts Overall equation for dark reaction:
one molecule of carbon dioxide to form
an unstable 6 carbon compound. This 6CO2 1 18ATP 1 12NADPH 1 H1
6C compound is broken down into two C6H12O6 1 6H2O1 18ADP 1 18Pi 1
molecules of 3-carbon compound phospho 12NADP1
glyceric acid (PGA) (Figure 13.19).
RUBP 1 CO2
Rubisco
2 molecules PGA RUBISCO – RUBP
C a r b o x y l a s e
Oxygenase enzyme,
Phase 2 – Glycolytic Reversal /
is the most abundant
Reduction protein found on earth. It constitutes
Phospho glyceric acid is phosphorylated 16 % of the chloroplast protein. It acts
by ATP and produces 1,3 bis phospho as carboxylase in the presence of CO2
glyceric acid by PGA kinase. 1,3 bis phospho and oxygenase in the absence of CO2.
glyceric acid is reduced to glyceraldehyde
3 Phosphate (G-3-P) by using the reducing 13.14 Hatch & Slack Pathway or C4
power NADPH 1 H1. Glyceraldehyde Cycle or Dicarboxylic Acid
3 phosphate is converted into its isomeric Pathway or Dicarboxylation
form di hydroxy acetone phosphate (DHAP).
Pathway
Till 1965, Calvin cycle is the only pathway for
PGA PGA Kinase 1,3 bisphosphoglyceric acid
ATP ADP CO2 fixation. But in 1965, Kortschak, Hart
and Burr made observations in sugarcane
and found C4 or dicarboxylic acid pathway.
NADPH 1 H1 NADP1 Malate and aspartate are the major labelled
1,3 bisphosphoglceric acid products. This observation was confirmed
Glyceraldehyde-3-Phosphate by Hatch & Slack in 1967. This alternate
pathway for the fixation of CO2 was found in
Phase 3 – Regeneration several tropical and sub-tropical grasses and
Regeneration of RUBP involves the formation some dicots. C4 cycle is discovered in more
of several intermediate compounds of than 1000 species. Among them 300 species
6-carbon, 5-carbon,4-carbon and 7- carbon belong to dicots and rest of them are
skeleton. Fixation of one carbon dioxide monocots. C4 plants represent about 5% of
132

The C4 pathway
Photosynthetic
cells of C4 Mesophyll
cell PEP carboxylase CO2
plant leaf
C4 leaf anatomy
Oxaloacetate (4c) PEP (3c)
Mesophyll cell ADP
Malate (4c) ATP

Bundle-
sheath
cell Bundle Pyruvate (3c)
sheath CO2
vein cell
(vascular calvin
tissue) cycle
Sugar
stoma Vascular
tissue
Figure 13.20: C4 Cycle
Earth’s plant biomass and 1% of its known require 5 ATP and 2 NADPH 1 H1 to fix
plant species. Despite this scarcity, they one molecule of CO2.
account for about 30% of terrestrial carbon
fixation. Increasing the proportion of C4 13.14.1 Stage: I Mesophyll Cells
plants on earth could assist biosequestration
of CO2 and represent an important climate Phosphoenol Pyruvate 1 CO2
(PEP) (3C)
change avoidance strategy. PEP carboxylase
C4 pathway is completed in two
phases, first phase takes place in stroma of Oxaloacetic acid (OAA) (4C)
mesophyll cells, where the CO2 acceptor
molecule is 3-Carbon compound, phospho Oxaloacetic acid (OAA) is converted
enol pyruvate (PEP) to form 4-carbon Oxalo into malic acid or aspartic acid and is
acetic acid (OAA). The first product is a transported to the bundle sheath cells
4-carbon and so it is named as C4 cycle. oxalo through plasmodesmata.
acetic acid is a dicarboxylic acid and hence
13.14.2 Stage: II Bundle Sheath Cells
this cycle is also known as dicarboxylic
acid pathway (Figure 13.20). Carbon Malic acid undergoes decarboxylation
dioxide fixation takes place in two places and produces a 3 carbon compound
one in mesophyll and another in bundle Pyruvic acid and CO2. The released CO2
sheath cell (di carboxylation pathway). It is combines with RUBP and follows the
the adaptation of tropical and sub tropical calvin cycle and finally sugar is released
plants growing in warm and dry conditions. to the phloem. Pyruvic acid is transported
Fixation of CO2 with minimal loss is due to the mesophyll cells.
to absence of photorespiration. C4 plants
133

Rubisco Kranz Anatomy: It
RUBP 1 CO2 2 PGA
is the German term
(5C) (3C)
meaning a halo or
wreath. In C4 plants
Activity vascular bundles are surrounded by a
• Collect the leaves of Paddy (C3) layer of bundle sheath. Bundle sheath is
and Sugar cane (C4). surrounded by a ring of mesophyll cells.
The characteristic feature of C4 plants is
• Take the cross section.
the presence of dimorphic chloroplast:
• Observe the sections under the
Bundle sheath chloroplast: Larger
microscope.
chloroplast, thylakoids not arranged
• See the difference in their anatomy in granum and rich in starch.
(Dimorphic chloroplast and Kranz
Mesophyll Chloroplast: Smaller
anatomy).
chloroplast, thylakoids arranged in
granum and less starch.
Table 13.4: Differences between C3 and C4 plants

C3 Plants C4 Plants
1. CO2 fixation takes place in mesophyll 1. CO2 fixation takes place mesophyll and
cells only bundle sheath
2. CO2 acceptor is RUBP only 2. PEP in mesophyll and RUBP in bundle
sheath cells
3. First product is 3C- PGA 3. First product is 4C- OAA
4. Kranz anatomy is not present 4. Kranz anatomy is present
5. Granum is present in mesophyll cells 5. Granum present in mesophyll cells and
absent in bundle sheath
6. Normal Chloroplast 6. Dimorphic chloroplast
7. Optimum temperature 20o to 25oC 7. Optimum temperature 30o to 45oC
8. Fixation of CO2 at 50 ppm 8. Fixation of CO2 even less than 10 ppm
9. Less efficient due to higher 9. More efficient due to less
photorespiration photorespiration
10. RUBP carboxylase enzyme used for 10. PEP carboxylase and RUBP
fixation carboxylase used
11. 18 ATPs used to synthesize one 11. Consumes 30 ATPs to produce one
glucose glucose.
12. Efficient at low CO2 12. Efficient at higher CO2
13. Example: Paddy, Wheat, Potato and 13. Example: Sugar cane, Maize, Sorghum,
so on Amaranthus and so on
134

4.
Due to absence of photorespiration,
Check your grasp! CO2 Compensation Point for C4 is
C4 plants requires 30 ATPs and lower than that of C3 plants.
12 NADPH 1 H1 to synthesize one Differences between C3 Plants (C3 Cycle) and
glucose, but C3 plants requires only C4 Plants (C4 Cycle) are given in table 13.4.
18 ATPs and 12 NADPH 1 H1 to
synthesize one glucose molecule. If
13.15 Crassulacean Acid
then, how can you say C4 plants are
more advantageous? Metabolism or CAM cycle
Solution: C4 plants are more advantageous It is one of the carbon pathways identified
than C3 plants because most of the energy in succulent plants growing in semi-arid
lost during photo respiration in C3 plants. or xerophytic condition. This was first
observed in crassulaceae family plants like
13.14.3 Significance of C4 cycle Bryophyllum, Sedum, Kalanchoe and is the
1. Plants having C4 cycle are mainly of reason behind the name of this cycle. It is
tropical and sub-tropical regions and also noticed in plants from other families
are able to survive in environment Examples: Agave, Opuntia, Pineapple and
with low CO2 concentration. Orchids. The stomata are closed during day
2. C4 plants are partially adapted to and are open during night (Scotoactive).
drought conditions. This reverse stomatal rhythm helps to
conserve water loss through transpiration
3.
Oxygen has no inhibitory effect on
and will stop the fixation of CO2 during the
C4 cycle since PEP carboxylase is
day time. At night time CAM plants fix CO2
insensitive to O2.
Night: Open stomata Day: Closed stomata

Open stoma permits
CO2 uptake Decarboxylation of stored Closed stoma
Atmospheric entry of CO2 and malate and refixation of prevents H2O loss
and fixation loss of H2O internal CO2 deacidification and CO2 uptake
CO2
leaf acidification
NADP+ malk
PEP carboxylase enzyme
CO2 Malate
Phosphoenol- Oxaloacetate Malic acid
pyruvate NAD+ malic
NADH
dehydro Pyruvate
NAD+ Calvin
genase cycle
Triose Vacuole
phosphate Malate Starch
Starch Malic acid Chloroplast

Chloroplast Vacuole
Figure 13.21: CAM cycle
figure 13.26
135 CAM Cycle

with the help of Phospho Enol Pyruvic acid their photosynthesis.
(PEP) and produce oxalo acetic acid (OAA). 3. Stomata are closed during the day
Subsequently OAA is converted into malic time and help the plants to avoid
acid like C4 cycle and gets accumulated in transpiration and water loss.
vacuole increasing the acidity. During the
day time stomata are closed and malic acid 13.16 Photorespiration or C2 Cycle
is decarboxylated into pyruvic acid resulting
or Photosynthetic Carbon
in the decrease of acidity. CO2 thus formed
enters into Calvin Cycle and produces Oxidation (PCO) Cycle
carbohydrates (Figure13.21). Respiration is a continuous process for all
Significance of CAM Cycle living organisms including plants. Decker
(1959) observed that rate of respiration is
1. It is advantageous for succulent plants
more in light than in dark. Photorespiration
to obtain CO2 from malic acid when
is the excess respiration taking place in
stomata are closed.
photosynthetic cells due to absence of
2. During day time stomata are closed CO2 and increase of O2(Table 13.5). This
and CO2 is not taken but continue
(2) O2
5C
(2) Ribulose 1,5 (2) PGA

bis phosphate 2C
Calvin (2) Phospho Glycolate

Cycle
PGA 3C
ADP (2) Pi
3C 2C
ATP
Glycerate (2) Glycolate
3C
2C
PE
Glycerate
ROX I SOME
NAD+ + (2) Glycolate

3C + O2
NADH+H H O 2C
Hydroxy pyruvate 2 2 (2) Glyoxylate

3C H2O + ½ O2 2C
Serine (2) Glycine

3C 2C
M
ITO
Serine (2) Glycine

CHONDR
CO2
NH3
+
NADH+H NAD
+
O
I
N
Figure 13.22: Photorespiration
136

condition changes the carboxylase role conditions 50% of the photosynthetic
of RUBISCO into oxygenase. C2 Cycle potential is lost because of Photorespiration
takes place in chloroplast, peroxisome and (Figure 13.22).
mitochondria. RUBP is converted into PGA
13.16.1 Significance of photorespiration
and a 2C-compound phosphoglycolate by
Rubisco enzyme in chloroplast. Since the 1. Glycine and Serine synthesised during
first product is a 2C-compound, this cycle this process are precursors of many
is known as C2 Cycle. Phosphoglycolate biomolecules like chlorophyll, proteins,
by loss of phosphate becomes glycolate. nucleotides.
Glycolate formed in chloroplast enters into 2. It consumes excess NADH 1 H1 generated.
peroxisome to form glyoxylate and hydrogen 3. Glycolate protects cells from Photo
peroxide. Glyoxylate is converted into oxidation.
glycine and transferred into mitochondria.
In mitochondria, two molecules of glycine 13.16.2 Carbon Dioxide Compensation Point
combine to form serine. Serine enters into When the rate of photosynthesis equals
peroxisome to form hydroxy pyruvate. the rate of respiration, there is no exchange
Hydroxy pyruvate with help of NADH 1 H1 of oxygen and carbon dioxide and this is
becomes glyceric acid. Glyceric acid is cycled called as carbon dioxide compensation
back to chloroplast utilising ATP and point. This will happen at particular light
becomes Phosphoglyceric acid (PGA) and intensity when exchange of gases becomes
enters into the Calvin cycle (PCR cycle). zero. When light is not a limiting factor and
Photorespiration does not yield any free atmospheric CO2 concentration is between
energy in the form of ATP. Under certain 50 to 100 ppm the net exchange is zero.
Table 13.5: Differences between Photorespiration and Dark Respiration
Photorespiration Dark respiration
1. It takes place in photosynthetic green
1. It takes place in all living cells
cells
2. It takes place only in the presence of
2. It takes place all the time
light
3. It involves chloroplast, peroxisome
3. It involves only mitochondria
and mitochondria
4. It does not involve Glycolysis, Kreb’s
4. It involves glycolysis, Kreb’s Cycle and
Cycle, and ETS ETS
5. Substrate is carbohydrates, protein or
5. Substrate is glycolic acid
fats
6. It is not essential for survival 6. Essential for survival
7. Phosphorylation produces ATP
7. No phosphorylation and yield of ATP
energy
8. NADH2 is oxidised to NAD 1
8. NAD1 is reduced to NADH2
9. Hydrogen peroxide is produced 9. Hydrogen peroxide is not produced
10. End products are CO2 and PGA 10. End products are CO2 and water
137

13.17 Factors affecting Photosynthesis HIGH LIGHT
RATE OF PHOTOSYNTHESIS
D INTENSITY E
In 1860, Sachs gave three cardinal points
theory explaining minimum, optimum MEDIUM LIGHT
and maximum factors that control C INTENSITY F
photosynthesis. In 1905, Blackman put
LOW LIGHT
forth the importance of smallest factor. B INTENSITY
Blackman’s law of limiting factor is
actually a modified Law proposed by
Liebig’s Law of minimum. According to A
Blackman, “When a process is conditioned CO2 CONCENTRATION
as to its rapidity by a number of separate
Figure 13.23: Blackman’s Law of Limiting
factors, the rate of the process is limited by Factors
the pace of the lowest factor”. To conclude
in an easy way “at any given point of time
the lowest factor among essentials will limit directly controlled by light. Stomatal
the rate of photosynthesis”. For example, movement leading to diffusion of CO2 is
when even sufficient light intensity is indirectly controlled by light.
available, photosynthesis may be low
due to low CO2 in the atmosphere. Here, a. Intensity of Light:
CO2 acts as a limiting factor. If CO2 is Intensity of light plays a direct role in
increased in the atmosphere the rate of the rate of photosynthesis. Under low
photosynthesis also increases. Further intensity the photosynthetic rate is low
increase in photosynthesis is possible and at higher intensity photosynthetic rate
only if the available light intensity is also is higher. It also depends on the nature of
increased proportionately (Figure 13.23). plants. Heliophytes (Bean Plant) require
Factors affecting photosynthesis higher intensity than Sciophytes (Oxalis).
are further grouped into External or b. Quantity of Light:
Environmental factors and Internal factors. In plants which are exposed to light
I. External factors: Light, carbon for longer duration (Long day Plants)
dioxide, temperature, water, mineral photosynthetic rate is higher.
and pollutants.
c. Quality of light:
II. Internal factors: Pigments, protoplasmic
factor, accumulation of carbohydrates, Different wavelengths of light affect the rate of
anatomy of leaf and hormones. photosynthesis because pigment system does
not absorb all the rays equally. Photosynthetic
13.17.1. External factors rate is maximum in blue and red light.
Photosynthetically Active Radiation (PAR)
1. Light
is between 400 to 700 nm. Red light induces
Energy for photosynthesis comes only highest rate of photosynthesis and green light
from light. Photooxidation of water and induces lowest rate of photosynthesis.
excitation of pigment molecules are
138

2. Carbon dioxide 5. Water
CO2 is found only 0.3 % in the atmosphere Photolysis of water provides electrons and
but plays a vital role. Increase in protons for the reduction of NADP, directly.
concentration of CO2 increases the rate of Indirect roles are stomatal movement and
photosynthesis (CO2 concentration in the hydration of protoplasm. During water
atmosphere is 330 ppm). If concentration stress, supply of NADPH 1 H1 is affected.
is increased beyond 500ppm, rate of
photosynthesis will be affected showing the 6. Minerals
inhibitory effect. Deficiency of certain minerals affect
photosynthesis e.g. mineral involved in the
3. Oxygen synthesis of chlorophyll (Mg, Fe and N),
The rate of photosynthesis decreases Phosphorylation reactions (P), Photolysis
when there is an increase of oxygen of water (Mn and Cl), formation of
concentration. This Inhibitory effect of plastocyanin (Cu).
oxygen was first discovered by Warburg
(1920) using green algae Chlorella. 7. Air pollutants
Pollutants like SO2, NO2, O3 (Ozone) and
4. Temperature
Smog affects rate of photosynthesis.
The optimum temperature for photosynthesis
varies from plant to plant. Temperature is 13.17.2 Internal Factors
not uniform in all places. In general, the
1. Photosynthetic Pigments
optimum temperature for photosynthesis
is 25oC to 35oC. This is not applicable for It is an essential factor and even a
all plants. The ideal temperature for plants small quantity is enough to carry out
like Opuntia is 55oC, Lichens 20oC and photosynthesis.
Algae growing in hot spring photosynthesis 2. Protoplasmic factor
is 75oC. Whether high temperature or low
Hydrated protoplasm is essential for
temperature it will close the stomata as well
photosynthesis. It also includes enzymes
as inactivate the enzymes responsible for
responsible for Photosynthesis.
photosynthesis (Figure 13. 24).
Light intensity CO2 Concentration Temperature

Figure 13.24: Factors affecting Photosynthesis
139

3. Accumulation of Carbohydrates
Experiment to determine rate
Photosynthetic end products like
of photosynthesis by Wilmott’s
carbohydrates are accumulated in cells
bubbler
and if translocation of carbohydrates
Wilmott’s bubbler consists of a wide is slow then this will affect the rate of
mouth bottle fitted with single holed photosynthesis.
cork, a glass tube with lower end having 4. Anatomy of leaf
wider opening to insert Hydrilla plant,
Thickness of cuticle and epidermis,
the upper end fitted to a narrow bottle
distribution of stomata, presence or absence
with water (Figure 13.25).
of Kranz anatomy and relative proportion of
photosynthetic cells affect photosynthesis.
5. Hormones
Water Hormones like gibberellins and cytokinin
increase the rate of photosynthesis.
Test tube funnel experiment or

Experiment to prove oxygen evolved
during Photosynthesis
1. Place Hydrilla plant at the bottom

Specimen tube
of a beaker containing water.
2. Cover the plant with an inverted
funnel.
Hydrilla
3. Invert a test tube over the funnel.
4. Keep this setup in sunlight.
Figure 13.25: Wilmott’s Bubbler Note your observations (Figure 13. 26).
1. Fill the bottle with water and Gas collected by

y downward
displacement of water
w
insert Hydrilla twig into the wider
part of the tube Tesst tube
2. Hydrilla plant should be cut inside

the water to avoid entry of air
bubbles Po
ond water
3. Fix the tube with jar which acts as

water reservoir Invverted funnel
4. Keep the apparatus in sunlight Hyydrilla
5. Count the bubbles when they are

in same size.
Figure 13.26: Test tube funnel experiment
140

Table 13.6: Difference between photosynthesis in plants and photosynthesis in bacteria
Photosynthesis in Plants Photosynthesis in Bacteria
1. Cyclic and non-cyclic phosphorylation 1. Only cyclic phosphorylation takes
takes place place
2. Photosystem I and II involved 2. Photosystem I only involved
3. Electron donor is water 3. Electron donor is H2S
4. Oxygen is evolved 4. Oxygen is not evolved
5. Reaction centres are P700 and P680 5. Reaction centre is P870
6. Reducing agent is NADPH 1 H1 6. Reducing agent is NADH 1 H1
7. PAR is 400 to 700 nm 7. PAR is above 700 nm
8. Chlorophyll, carotenoid and 8. Bacterio chlorophyll and bacterio
xanthophyll viridin
9. Photosynthetic apparatus – chloroplast 9. It is chlorosomes and chromatophores
13.18 Photosynthesis in bacteria Summary

Though we study about bacterial Photosynthesis is an oxidation and reduction
photosynthesis as the last part, bacterial process. It has two phases: the light reaction and
photosynthesis formed first and foremost dark reaction. During light reaction water is
in evolution. Bacteria does not have oxidised to release O2 and during dark reaction
specialized structures like chloroplast. CO2is reduced to form sugars. Solar energy
It has a simple type of photosynthetic is trapped by pigment system I and pigment
apparatus called chlorosomes and system II. P700 and P680 act as reaction centres
chromatophores (Table 13.6). Van Neil for PS I and PS II respectively. Splitting of water
(1930) discovered a bacterium that molecule (Photolysis) produces electrons,
releases sulphur instead of oxygen during protons and oxygen. Photophosphorylation
photosynthesis. Here, electron donor is takes place through cyclic and non-cyclic
hydrogen sulphide (H2S) and only one mechanisms and generates energy and reducing
photosystem is involved (PS I) and the power. Dark reaction or biosynthetic phase of
reaction centre is P870. Pigments present photosynthesis use the products of light energy
in bacteria are bacteriochlorophyll a, b, c, (ATP and NADPH 1 H1) and carbon dioxide
d, e and g and carotenoids. Photosynthetic is reduced to Carbohydrates. Carbon pathway
bacteria are classified into three groups: in C3 cycle has RUBP as the acceptor molecule
and the first product is PGA (3C). Carbon
1. Green sulphur bacteria. Example: pathway in C4 plants involves mesophyll
Chlorobacterium and Chlorobium. and bundle sheath cells, Kranz anatomy.
2. Purple sulphur bacteria. Example: Dimorphic chloroplast, no photorespiration,
Thiospirillum and Chromatium. acceptor molecule as PEP and first product as
3. Purple non-sulphur bacteria. Example: OAA (4C) are some of the unique characters
Rhodopseudomonas and Rhodospirillum. of C4 cycle. C2 Cycle or photorespiration is
operated when less amount of CO2 is used for
reduction and O2 increases. Rubisco starts to
141

play oxygenase role. Succulent and xerophytic a. 2ATP 1 2NADPH
plants show reverse stomatal rhythm as they b. 2ATP 1 3NADPH
open during night time and close during c. 3ATP 1 2NADPH
day time and follow CAM cycle. Night time
d. 3ATP 1 3NADPH
produces malic acid and during day time
malate is converted into pyruvate and produces 5. Identify true statement regarding light
CO2 which is reduced to carbohydrates. reaction of photosynthesis?
Photosynthesis is affected by internal and a. Splitting of water molecule is associate
external factors. Bacterial photosynthesis is the with PS I.
primitive type of photosynthesis and it involves b. PS I and PS II involved in the
only photosystem I. formation of NDPH1H1.
Evaluation c. The reaction center of PS I is
1. Assertion (A): Chlorophyll a with absorption peak
Increase in Proton at 680 nm.
gradient inside lumen d. The reaction center of PS II is
responsible for ATP Chlorophyll a with absorption peak
synthesis at 700 nm.
Reason (R): Oxygen evolving complex 6. Two groups (A & B) of bean plants of
of PS I located on thylakoid membrane similar size and same leaf area were placed
facing Stroma, releases H1 ions in identical conditions. Group A was
a. Both Assertion and Reason are True. exposed to light of wavelength 400-450nm
& Group B to light of wavelength of 500-
b. Assertion is True and Reason is False. 550nm. Compare the photosynthetic rate
c. Reason is True and Assertion is False. of the 2 groups giving reasons.
d. Both Assertion and Reason are False. 7. A tree is believed to be releasing oxygen
2. Which chlorophyll molecule does not during night time. Do you believe the
have a phytol tail? truthfulness of this statement? Justify
a. Chl- a b. Chl-b c. Chl- c d. Chl -d your answer by giving reasons?
3. The correct sequence of flow of electrons 8. Grasses have an adaptive mechanism
in the light reaction is to compensate photorespiratory losses-
Name and describe the mechanism.
a. PS II, plastoquinone, cytochrome, PS
I, ferredoxin. 9. In Botany class, teacher explains, Synthesis
of one glucose requires 30 ATPs in C4
b. PS I, plastoquinone, cytochrome, PS
plants and only 18 ATPs in C3plants. The
II ferredoxin.
same teacher explains C4 plants are more
c. PS II, ferredoxin, plastoquinone, advantageous than C3 plants. Can you
cytochrome, PS I. identify the reason for this contradiction?
d. PS I, plastoquinone, cytochrome, PS 10. When there is plenty of light and higher
II, ferredoxin. concentration of O2, what kind of
4. For every CO2 molecule entering the C3 pathway does the plant undergo?Analyse
cycle, the number of ATP & NADPH the reasons.
required
142

t ICT Corner
Photosynthesis
Let’s play
photosynthesis
Steps
• Start a new game and tap
• Click light dependent reaction and follow the steps
• After completion – move back and Click Calvin cycle reaction and follow the steps
Activity
• Observe the cycle and record it
• Check your grasp by click the Quiz tap
Step 1 Step 2
Step 3 Step 4
Web URL:
https://biomanbio.com/HTML5GamesandLabs/PhotoRespgames/
photointeractivehtml5page.html
143

Chapter
14 Respiration
Plant and Animal Interdependence

Learning Objectives
In biosphere, plants and animals
are complementary systems which
• Recognize the stages of glucose are integrated to sustain life. In
breakdown and its redox system. plants, oxygen enters through the
• Differentiate aerobic respiration stomata and it is transported to
from anaerobic respiration. cells, where oxygen is utilized for
• Describe the conditions under energy production. Plants require
which respiration occurs. carbon dioxide to survive, to produce
• Realize the role of mitochondria as carbohydrates and to release oxygen
power house of the cell. through photosynthesis. These oxygen
• Understand, how ATP molecules molecules are inhaled by human
are generated during respiration. through the nose, which reaches the
lungs where oxygen is transported
through the blood and it reaches cells.
Chapter Outline Cellular respiration takes place inside
14.1 Gaseous exchange the cell. A specialized respiratory
system is present in animals but
14.2 Structure of ATP
is absent in plants for delivering
14.3 Redox reactions oxygen inside the cell. But the cellular
14.4 Types of Respiration respiration stages are similar in both
14.5 Stages of Respiration plants and animals which hint at
14.6 Respiratory Quotient evolutionary divergence.
14.7 Anaerobic Respiration
14.8 Factors Affecting Respiration O2
CO2
14.9 Pentose Phosphate Pathway
144

If you are sleeping under a tree organic substances which are oxidised
during night time you will feel difficulty during respiration are called respiratory
in breathing. During night, plants take substrates. Among these, glucose is
up oxygen and release carbon dioxide the commonest respiratory substrate.
and as a result carbon dioxide will be Breaking of C-C bonds of complex organic
abundant around the tree. This process compounds through oxidation within the
of CO2 evolution is called respiration. cells leads to energy release. The energy
This process takes place during day time released during respiration is stored in the
also (Figure 14.1). It is accompanied by form of ATP (Adenosine Tri Phosphate) as
breakdown of substrates and release of well as liberated heat. Respiration occurs
energy. In this chapter, respiration process in all the living cells of organisms. The
in plants at cellular level will be dealt with. overall process of respiration corresponds
to a reversal of photosynthesis.
C6H12O6 1 6O2 → 6CO2 1 6H2O 1 Energy

CO (686 K cal or 2868 KJ)
2
2
O
O
Depending upon the nature of
2
O
2
C
respiratory substrate, Blackman divided

respiration into,
1. Floating respiration
2. Protoplasmic respiration
When carbohydrate or fat or organic
acid serves as respiratory substrate and it is
called floating respiration. It is a common
2
CO
O
2
mode of respiration and does not produce

any toxic product. Whereas respiration
utilizing protein as a respiratory substrate,
it is called protoplasmic respiration.
Protoplasmic respiration is rare and
it depletes structural and functional
Figure 14.1: Gaseous exchange in plants proteins of protoplasm and liberates toxic
ammonia.
14.1 Gaseous Exchange
14.1.2 Compensation point
14.1.1 Respiration
At dawn and dusk the intensity of light
The term respiration was coined by is low. The point at which CO2 released
Pepys (1966). Respiration is a biological in respiration is exactly compensated by
process in which oxidation of various CO2 fixed in photosynthesis that means
food substances like carbohydrates, no net gaseous exchange takes place, it
proteins and fats take place and as a is called compensation point. At this
result of this, energy is produced where moment, the amount of oxygen released
O2 is taken in and CO2 is liberated. The from photosynthesis is equal to the
145

High energy bonds
Rate of
Photosynthesis NH2
Carbohydratre balance
Compensation N O O O
N
Point O P O P O P OH
N O
N
OH OH OH
Adenine Phosphate groups
Rate of OH OH
Respiration Ribose
Adenosine
Adenosine
Monophosphate (AMP)
Adenosine
Time in a day (hours) Diphosphate (ADP)
Adenosine
Figure 14.2: Compensation point Triphosphate (ATP)
amount of oxygen utilized in respiration. Figure 14.3: Molecular structure of ATP

The two common factors associated
with compensation point are CO2 and ATP is not only higher
light (Figure 14.2). Based on this there energy compound
are two types of compensation point. present in a cell. There
They are CO2 compensation point and are other higher energy
light compensation point. C3 plants have compounds also present. Example
compensation points ranging from 40-60 GTP (Guanosine Tri Phosphate) and
ppm (parts per million) CO2 while those UTP (Uridine Tri Phosphate).
of C4 plants ranges from 1-5 ppm CO2.
14.3 Redox Reactions

14.2 Structure of ATP
Respiration is responsible for generation NAD1 1 2e - 1 2H1 NADH 1 H1
of ATP. The discovery of ATP was made by FAD 1 2e- 1 2H1 FADH2
Karl Lohman (1929). ATP is a nucleotide When NAD1 (Nicotinamide Adenine
consisting of a base-adenine, a pentose Dinucleotide-oxidised form) and FAD
sugar-ribose and three phosphate groups. (Flavin Adenine Dinucleotide) pick up
Out of three phosphate groups the last two electrons and one or two hydrogen ions
are attached by high energy rich bonds (protons), they get reduced to NADH 1 H1
(Figure 14.3). On hydrolysis, it releases and FADH2 respectively. When they drop
energy (7.3 K cal or 30.6 KJ/ATP) and it is electrons and hydrogen off they go back
found in all living cells and hence it is called to their original form. The reaction in
universal energy currency of the cell. ATP which NAD1 and FAD gain (reduction) or
is an instant source of energy within the lose (oxidation) electrons are called redox
cell. The energy contained in ATP is used reaction (Oxidation reduction reaction).
in synthesis carbohydrates, proteins and These reactions are important in cellular
lipids. The energy transformation concept respiration.
was established by Lipman (1941).
146

energy is released. Aerobic respiration is a
Handy mnemonic very complex process and is completed in
four major steps:
1. Glycolysis
2. Pyruvate oxidation (Link reaction)
3. Krebs cycle (TCA cycle)
4. Electron Transport Chain
(Terminal oxidation).
14.4.2 Anaerobic respiration
LEO the lion says GER In the absence of molecular oxygen glucose
is incompletely degraded into either ethyl
LEO - Loss of Electrons is Oxidation
alcohol or lactic acid (Table 14.1). It
GER - Gain of Electrons is Reduction includes two steps:
1. Glycolysis
14.4 Types of Respiration 2. Fermentation
Respiration is classified into two types 14.5 Stages of Respiration
as aerobic and anaerobic respiration
(Figure 14.4) 1. Glycolysis-conversion of glucose into
pyruvic acid in cytoplasm of cell.
14.4.1 Aerobic respiration 2. Link reaction-conversion of pyruvic
acid into acetyl coenzyme-A in
Respiration occurring in the presence
mitochondrial matrix.
of oxygen is called aerobic respiration.
During aerobic respiration, food materials 3. Krebs cycle-conversion of acetyl
like carbohydrates, fats and proteins are coenzyme A into carbon dioxide and
completely oxidised into CO2, H2O and water in the mitochondrial matrix.
Respiration
Aerobic Respiration Anaerobic Respiration
Alcoholic Lactic acid Mixed acid

fermentation fermentation fermentation
Figure 14.4: Types of Respiration
147

Table 14.1: Differences between aerobic and anaerobic respiration
Aerobic respiration Anaerobic Respiration
1. It occurs in all living cells of higher
It occurs yeast and some bacteria.
organisms.
2. It requires oxygen for breaking the Oxygen is not required for breaking the
respiratory substrate. respiratory substrate.
The end products are alcohol, and CO2
3. The end products are CO2 and H2O.
(or) lactic acid. .
4. Oxidation of one molecule of glucose
Only 2 ATP molecules are produced.
produces 36 ATP molecules.
5. It consists of four stages-glycolysis,
It consists of two stages-glycolysis and
link reaction, TCA cycle and electron
fermentation.
transport chain.
6. It occurs in cytoplasm and mitochondria. It occurs only in cytoplasm.
4. Electron transport chain and oxidative molecule with energy in the form of
phosphorylation remove hydrogen atoms ATP in mitochondrial inner membrane
from the products of glycolysis, link (Figure 14.5).
reaction and Krebs cycle release water
Glucose
Glycolysis
ADP+Pi
Ethyl alcohol + CO2 ATP
Anaerobic 2 molecules
of Pyruvic acid
Lactic acid
Aerobic
Link react
2NA
ion
DH+
2NA H+
o-A DH+
lC H+ P
ty 6NA AT
Ace
DH+
H+
Pi
2x
P+
2FA
O
AD
2
ETC
DH2
2C Krebs
Cycle
2 ADP+2
Pi O2
2 ATP
4CO2 H 2O
Figure 14.5: Overall stages of Respiration

148

PREPARATORY PHASE
Glucose c c c c c c
1. Phosphorylaon 1 ATP
Hexokinase
ADP
P
Glucose-6-Phosphate c c c c c c
2. Isomerisaon 2 Phosphohexose isomerase

Mg++ P
Fructose-6-Phosphate c c c c c c
ATP Phosphofructo kinase
3. Phosphorylaon 3 ADP Mg++ P P
Fructose-1,6-Bisphosphate c c c c c c
4. Spling into Aldolase
two molecules P
4 P
c c c c c c
Triose phosphate
Glyceraldehyde- isomerase Dihydroxy Acetone
5.Isomerisaon
3-Phosphate 5 Phosphate
2NAD
+ 2Pi Glyceraldehyde-3-Phosphate
6.Oxidaon and + dehydrogenase
Phosphorylaon
2NADH+H 6
P P
2x 1,3 Bisphospho Glycerate c c c
2ADP
7. Dephosphorylaon 2ATP
7 Phosphoglycerate kinase
Mg++
PAY OFF PHASE

P
2x 3-Phospho Glycerate c c c
8. Shi
ing P from 8 Phosphoglyceromutase
3rd C to 2nd C Mg++ P
2x 2-Phospho Glycerate c c c
2H2O
9 Enolase
9. Dehydraon
Mg
++
P
2x Phospho Enol Pyruvate c c c
2ADP Pyruvate kinase
10 Mg ++
2ATP
10. Dephosphorylaon K++
2x Pyruvate c c c
Figure 14.6: Glycolysis or EMP pathway
149

14.5.1 Glycolysis
Check your grasp!
(Gr: Glykos 5 Glucose, Lysis 5 Splitting)
Glycolysis is a linear series of reactions in How many ATP molecules are
which 6-carbon glucose is split into two produced from one sucrose molecule?
molecules of 3-carbon pyruvic acid. The
enzymes which are required for glycolysis 2. Pay off phase
are present in the cytoplasm (Figure 14.6).
Two molecules of glyceraldehyde-3-
The reactions of glycolysis were worked
phosphate oxidatively phosphorylated into
out in yeast cells by three scientists Gustav
two molecules of 1,3 - bisphospho glycerate.
Embden (German), Otto Meyerhoff
During this reaction 2NAD1 is reduced
(German) and J Parnas (Polish) and so
to 2NADH 1 H1 by glyceraldehyde-
it is also called as EMP pathway. It is the
3- phosphate dehydrogenase at step 6.
first and common stage for both aerobic
Further reactions are carried out by
and anaerobic respiration. It is divided
different enzymes and at the end two
into two phases.
molecules of pyruvate are produced. In
1. Preparatory phase or endergonic this phase, 2ATPs are produced at step 7
phase or hexose phase (steps 1-5). and 2 ATPs at step10 (Figure 14.6). Direct
2. Pay off phase or oxidative phase or transfer of phosphate moiety from substrate
exergonic phase or triose phase (steps molecule to ADP and is converted into
6-10). ATP is called substrate phosphorylation
or direct phosphorylation or trans
1. Preparatory phase phosphorylation. During the reaction at
Glucose enters the glycolysis from sucrose step 9, 2phospho glycerate dehydrated into
which is the end product of photosynthesis. Phospho enol pyruvate a water molecule is
Glucose is phosphorylated into glucose-6- removed by the enzyme enolase. As a result,
phosphate by the enzyme hexokinase, and enol group is formed within the molecule.
subsequent reactions are carried out by This process is called Enolation.
different enzymes (Figure 14.6). At the end
of this phase fructose-1, 6 - bisphosphate is 3. Energy Budget
cleaved into glyceraldehyde-3- phosphate In the pay off phase totally 4ATP and
and dihydroxy acetone phosphate by the 2NADH 1 H1 molecules are produced.
enzyme aldolase. These two are isomers. Since 2ATP molecules are already
Dihydroxy acetone phosphate is isomerised consumed in the preparatory phase, the
into glyceraldehyde-3- phosphate by the net products in glycolysis are 2ATPs and
enzyme triose phosphate isomerase, now 2NADH 1 H1.
two molecules of glyceraldehyde 3 phosphate The overall net reaction of glycolysis
enter into pay off phase. During preparatory
C6 H12O6 1 2ADP 1 2Pi 1 2NAD1
phase two ATP molecules are consumed in
step-1 and step-3 (Figure 14.6). 2x CH3COCOOH 1 2ATP 12NADH12H1
150

14.5.2 Pyruvate Oxidation (Link reaction)
Sir Hans Adolf
Two molecules of pyruvate formed by Krebs was born in
glycolysis in the cytosol enters into Germany on 25th
August 1900. He was
the mitochondrial matrix. In aerobic
awarded Nobel Prize
respiration this pyruvate with coenzyme for his discovery of
A is oxidatively decarboxylated into acetyl Citric acid cycle in
CoA by pyruvate dehydrogenase complex. Physiology in 1953.
This reaction is irreversible and produces
two molecules of NADH 1 H1 and 2CO2.
It is also called transition reaction or
Link reaction. The reaction of pyruvate
oxidation is
2x CH3COCOOH 1 2NAD1 1 2CoA

Pyruvate dehydrogenase
complex/ Mg11
2xCH3CO.CoA1 2NADH12H11 F1
2CO2↑ Stalk
F0
Pyruvate dehydrogenase complex
consist of three distinct enzymes,
Figure 14.7: Structure of Mitochondrion
such as
1. Pyruvate dehydrogenase TCA cycle starts with condensation
2. Dihydrolipoyil transacetylase of acetyl CoA with oxaloacetate in the
3. Dihydrolipoyil dehydrogenase presence of water to yield citrate or citric
and five different coenzymes, TPP acid. Therefore, it is also known as Citric
(Thymine Pyro Phosphate), NAD1, Acid Cycle (CAC) or Tri Carboxylic Acid
FAD, CoA and lipoate. (TCA) cycle. It is followed by the action of
different enzymes in cyclic manner. During
14.5.3 Krebs cycle or Citric acid cycle or the conversion of succinyl CoA to succinate
TCA cycle: by the enzyme succinyl CoA synthetase
or succinate thiokinase, a molecule of
Two molecules of acetyl CoA formed from
ATP synthesis from substrate without
link reaction now enter into Krebs cycle.
entering the electron transport chain is
It is named after its discoverer, German
called substrate level phosphorylation. In
Biochemist Sir Hans Adolf Krebs (1937). The
animals a molecule of GTP is synthesized
enzymes necessary for TCA cycle are found
from GDP1Pi. In a coupled reaction GTP
in mitochondrial matrix except succinate
is converted to GDP with simultaneous
dehydrogenase enzyme which is found in
synthesis of ATP from ADP1Pi. In three
mitochondrial inner membrane (Figure 14.7).
151

Pyruvate c c c
Oxidation and
Link Reaction NAD+ Co A Pyruvate
decarboxylation
+
NADH+H CO2 dehydrogenase
CoA
Acetyl CoA c c
H2O
Krebs cycle Co A
C 1. Condensation
1
c c c c
c c Citrate synthase c c
+ Oxaloacetate Citrate c c 2. Dehydration
NADH+H H2O
10. Oxidation Malate Aconitase 2 c c
+
10 c c
NAD dehydrogenase Cis aconitate c c
152
c c c c Malate H2O
++ 3 3. Rehydration
H2O 9 Aconitase Fe
Fumarase c c
9. Hydration Isocitrate + c c
Isocitrate ++
NAD c c
c c c c Fumarate dehydrogenase Mn 4 NADH+H+ 4. Oxidation
Succinate c c
FADH2 8 dehydrogenase
Oxalosuccinate c c
Oxalosuccinate 5 c c
8.Oxidation FAD decarboxylase CO2
Succinate 5. Decarboxylation
c c c c Succinyl α-ketoglutarate c c
Co-A synthetase α−ketoglutarate Co A c
ATP dehydrogenase 6 c c
+ CO2
7. Hydration and ADP+Pi 7 Succinyl
c CoA NAD 6. Oxidation and
Phosphorylation c c c c +
NADH+H decarboxylation
CoA
Co A H2O
Figure 14.8: Krebs cycle or Citric acid cycle
27-04-2018 11:58:42
steps (4, 5, 9) in this cycle NAD1 is reduced Two molecules of pyruvic acid formed
to NADH1 H1 and at step 7 (Figure14.8) at the end of glycolysis enter into the
where FAD is reduced to FADH2. mitochondrial matrix. Therefore, Krebs
The summary of link reaction and cycle is repeated twice for every glucose
Krebs cycle in Mitochondria is molecule where two molecules of pyruvic
acid produces six molecules of CO2, eight
Pyruvic acid 1 4NAD1 1 FAD 1 4H2O 1 ADP1Pi
molecules of NADH 1 H1, two molecules
Mitochondrial matrix.
of FADH2 and two molecules of ATP.
3CO21 4NADH14H1 1FADH2 1H2O1ATP.
Fats Carbohydrates Proteins

Proteases
Fatty acids Glycerol Glucose Amino acids
Deamination
Fructose-1,6-Bisphosphate
DHAP Glyceraldehyde
-3-Phospate
Pyruvic acid
CO2
Acetyl CoA
Krebs NH2
cycle
H2O CO2
Figure 14.9: Alternative substrates for respiration
153

1. Significance of Krebs cycle: through pyruvic acid or acetyl CoA and it
1. TCA cycle is to provide energy in the depends upon the structure. So respiratory
form of ATP for metabolism in plants. intermediates form the link between
2.
It provides carbon skeleton or raw synthesis as well as breakdown. The citric
material for various anabolic processes. acid cycle is the final common pathway
for oxidation of fuel molecules like amino
3.
Many intermediates of TCA cycle are
acids, fatty acids and carbohydrates.
further metabolised to produce amino
Therefore, respiratory pathway is an
acids, proteins and nucleic acids.
amphibolic pathway (Figure 14.9).
4.
Succinyl CoA is raw material for
formation of chlorophylls, cytochrome, 14.5.4 Electron Transport Chain (ETC)
phytochrome and other pyrrole (Terminal oxidation)
substances. During glycolysis, link
5.
α-ketoglutarate and oxaloacetate reaction and Krebs
undergo reductive amination and cycle the respiratory
produce amino acids. substrates are oxidised
6. It acts as metabolic sink which plays a at several steps and as a
central role in intermediary metabolism. result many reduced coenzymes NADH
1 H1 and FADH2 are produced. These
2. Amphibolic nature
reduced coenzymes are transported to
Krebs cycle is primarily a catabolic
inner membrane of mitochondria and
pathway, but it provides precursors for
are converted back to their oxidised
various biosynthetic pathways there by
forms produce electrons and protons. In
an anabolic pathway too. Hence, it is
mitochondria, the inner membrane is
called amphibolic pathway. It serves as
folded in the form of finger projections
a pathway for oxidation of carbohydrates,
towards the matrix called cristae. In cristae
fats and proteins. When fats are respiratory
many oxysomes (F1 particles) are present
substrate they are first broken down
which have electron transport carriers are
into glycerol and fatty acid. Glycerol is
present. According to Peter Mitchell’s
converted into DHAP and acetyl CoA.
Chemiosmotic theory this electron
This acetyl CoA enter into the Krebs
transport is coupled to ATP synthesis.
cycle. When proteins are the respiratory
Electron and hydrogen(proton) transport
substrate they are degraded into amino
takes place across four multiprotein
acids by proteases. The amino acids after
complexes(I-IV). They are
deamination enter into the Krebs cycle
1. Complex-I (NADH dehydrogenase).
The synthesis of It contains a flavoprotein(FMN) and
glucose from certain associated with non-heme iron Sulphur
non-carbohydrate protein (Fe-S). This complex is responsible
carbon substrates for passing electrons and protons from
such as proteins and lipids are called mitochondrial NADH (Internal) to
gluconeogenesis. Ubiquinone(UQ).
154

NADH 1 H1 1 UQ NAD1 1 UQH2 (A and B) and cytochromes a and a3.
In plants, an additional NADH Complex IV is the terminal oxidase and
dehydrogenase (External) complex is brings about the reduction of 1/2 O2 to
present on the outer surface of inner H2O.Two protons are needed to form a
membrane of mitochondria which can molecule of H2O (terminal oxidation).
oxidise cytosolic NADH 1 H1. 2Cyt coxidised 1 2H1 1 1/2 O2 2Cyt creduced 1H2O
Ubiquinone (UQ) or Coenzyme
Quinone(Co Q) is a small, lipid soluble The transfer of electrons from
electron, proton carrier located within the reduced coenzyme NADH to oxygen
inner membrane of mitochondria. via complexes I to IV is coupled to the
synthesis of ATP from ADP and inorganic
2. Complex-II (Succinic dehydrogenase)
phosphate (Pi) which is called Oxidative
It contains FAD flavoprotein is associated
phosphorylation. The F0F1-ATP synthase
with non-heme iron Sulphur (Fe-S)
(also called complex V) consists of F0
protein. This complex receives electrons
and F1. F1 converts ADP and Pi to ATP
and protons from succinate in Krebs cycle
and is attached to the matrix side of the
and is converted into fumarate and passes
inner membrane. F0 is present in inner
to ubiquinone.
membrane and acts as a channel through
Succinate 1 UQ → Fumarate 1 UQH2 which protons come into matrix.
3. Complex-III (Cytochrome bc1 com- Oxidation of one molecule of
plex) This complex oxidises reduced ubi- NADH 1 H1 gives rise to 3 molecules
quinone (ubiquinol) and transfers the elec- of ATP and oxidation of one molecule
trons through Cytochrome bc1 Complex FADH2 produces 2 molecules of ATP
(Iron Sulphur center bc1 complex) to cy- within a mitochondrion. But cytoplasmic
tochrome c. Cytochrome c is a small pro- NADH 1 H1 yields only two ATPs
tein attached to the outer surface of inner through external NADH dehydrogenase.
membrane and act as a mobile carrier to Therefore, two reduced coenzyme
transfer electrons between complex III to (NADH 1 H1) molecules from glycolysis
complex IV. being extra mitochondrial will yield
UQH2 12Cyt coxidised UQ12Cyt creduced 12H1 2 3 2 5 4 ATP molecules instead of
6 ATPs (Figure 14.10). The Mechanism
of mitochondrial ATP synthesis is based
Ubiquinone and
on Chemiosmotic hypothesis. According
cytochrome bc1 complex
to this theory electron carriers present
are structurally and
in the inner mitochondrial membrane
functionally similar
allow for the transfer of protons (H1).
to plastoquinone and cytochrome b6,f
For the production of single ATP,
complex respectively in the photosynthetic
3 protons (H1) are needed. The terminal
electron transport chain.
oxidation of external NADH bypasses
the first phosphorylation site and hence
4. Complex IV (Cytochome c oxidase) only two ATP molecules are produced
This complex contains two copper centers per external NADH oxidised through
155

288.247 pt
Figure 14.10: Electron Transport Chain and Terminal Oxidation
mitochondrial electron transport chain. Recent view

However, in those animal tissues in which When the cost of transport of ATPs from
malate shuttle mechanism is present, the matrix into the cytosol is considered,
oxidation of external NADH will yield the number will be 2.5 ATPs for each
almost 3 ATP molecules. NADH 1 H1 and 1.5 ATPs for each FADH2
oxidised during electron transport system.
Abnormal rise in Therefore, in plant cells net yield of 30 ATP
respiratory rate of molecules for complete aerobic oxidation of
ripening in fruits is one molecule of glucose. But in those animal
called Climacteric. cells (showing malate shuttle mechanism)
Examples are apple, banana, mango, net yield will be 32 ATP molecules.
papaya, pear.
Electron transport chain inhibitors
1. 2,4 DNP (Dinitrophenol) - It prevents
Complete oxidation of a glucose
synthesis of ATP from ADP, as it directs
molecule in aerobic respiration results in
electrons from Co Q to O2
the net gain of 36 ATP molecules in plants
2. Cyanide - It prevents flow of electrons
as shown in table 14.2. Since huge amount
from Cytochrome a3 to O2
of energy is generated in mitochondria
in the form of ATP molecules they are 3. Rotenone - It prevents flow of electrons
called ‘power house of the cell’. In the from NADH 1 H1/FADH2 to Co Q
case of aerobic prokaryotes due to lack of 4. Oligomycin – It inhibits oxidative
mitochondria each molecule of glucose phosphorylation
produces 38 ATP molecules.
156

Volume of CO2 liberated
Peter Mitchel, a British RQ 5
Volume of O2 consumed
Biochemist received Nobel
1. The respiratory substrate is a
prize for Chemistry in 1978
carbohydrate, it will be completely
for his work on the coupling
oxidised in aerobic respiration and the
of oxidation and
value of the RQ will be equal to unity.
hosphorylation in mitochondria.
p
C6H12O6 1 6O2 6CO2 ↑ 1 6H2O 1 Energy
Glucose
6 molecules of CO2
Cyanide resistant RQ of glucose 5
6 molecules of O2
respiration is believed
to be responsible for 5 1 (unity)
the climacteric in fruits 2. If the respiratory substrate is a
Cyanide resistant respiration is carbohydrate it will be incompletely
known to generate heat in thermogenic oxidised when it goes through anaerobic
tissues. respiration and the RQ value will be
The amount of heat produced in infinity.
thermogenic tissues may be as high as C6H12O6 2CO2↑1 2C2H5OH 1 Energy
51°C. Glucose Ethyl alcohol
14.6 Respiratory Quotient (RQ)

RQ of glucose
Anaerobically }
5
2 molecules of CO2
zero molecule of O2
5 ∞ (infinity)
The ratio of volume of carbon dioxide
3. In some succulent plants like Opuntia,
given out and volume of oxygen taken in
Bryophyllum carbohydrates are partially
during respiration is called Respiratory
oxidised to organic acid, particularly
Quotient or Respiratory ratio. RQ value
malic acid without corresponding release
depends upon respiratory substrates and
of CO2 but O2 is consumed hence the RQ
their oxidation.
value will be zero.
Table 14.2: Net Products gained during aerobic respiration per glucose molecule.
Reduced Total ATP

Stages CO2 ATP Reduced NAD1
FAD Production
2
Glycolysis 0 2 0 6
(2 3 2 5 4)
2
Link reaction 2 0 0 6
(2 3 3 5 6)
6 2
Krebs cycle 4 2 24
(6 3 3 5 18) (2 3 2 5 4)
Total 6 4 ATPs 28 ATPs 4 ATPs 36 ATPs
157

2C6H12O6 1 3O2 3C4H6O5 1 3H2O 1 Energy
Glucose Malic acid
RQ of glucose zero molecule of CO2

5
in succulents 3 molecules of O2
5 0 (zero)
4. When respiratory substrate is protein
or fat, then RQ will be less than unity.
2(C51H98O6) 1 145O2 102CO2↑1 98H2O 1 Energy
Tripalmitin(Fat)
RQ of 102 molecules of CO2
5
Tripalmitin 145 molecules of O2
5 0.7 (less than unity)
5. When respiratory substrate is an

organic acid the value of RQ will be more
than unity.
C4H6O5 1 3O2 4CO2 ↑1 3H2O 1 Energy
Malic acid The apparatus used for determining
RQ of 4 molecules of CO2 respiration and RQ is called Ganong’s
5
malic acid 3 molecules of O2 Respirometer.
5 1.33 (more than unity)
Respiratory quotients of some other
Significance of RQ
substances
1. RQ value indicates which type of
Proteins : 0.8–0.9
respiration occurs in living cells, either
Oleic acid (Fat) : 0.71
aerobic or anaerobic.
Palmitic acid (Fat) : 0.36
2. It also helps to know which type of Tartaric acid : 1.6
respiratory substrate is involved. Oxalic acid : 4.0
Red colour in various

Experiment to demonstrate the
parts of plants is
production of CO2 in aerobic
due to the presence
respiration
of anthocyanin,
synthesis of which require more O2 Take small quantity of any seed
than CO2 evolved. RQ will be less (groundnut or bean seeds) and allow
than one. them to germinate by imbibing them.
While they are germinating place them
in a conical flask. A small glass tube
containing 4 ml of freshly prepared
158

Potassium hydroxide (KOH) solution Activity
is hung into the conical flask with the
help of a thread and tightly close the Take a test tube with some germinated
one holed cork (Figure 14.11). Take seeds and fill with water. Keep this test
a bent glass tube, the shorter end of tube after some time until liberation of
which is inserted into the conical CO2. When the carbon dioxide from
flask through the hole in the cork, respiration is mixed to water, carbonic
while the longer end is dipped in a acid (H2CO3) is produced. Therefore,
beaker containing water. Observe the as more carbon dioxide is released,
position of initial water level in bent the solution becomes more acidic. You
glass tube. This experimental setup will see changes in pH as an indicator
is kept for two hours and the seeds using blue litmus paper changed into
were allowed to germinate. After red that respiration has occurred
two hours, the level of water rises in CO21H2O H2CO3
the glass tube. It is because, the CO2
evolved during aerobic respiration by 14.7 Anaerobic Respiration
germinating seeds will be absorbed by
14.7.1 Fermentation
KOH solution and the level of water
will rise in the glass tube. Some organisms can
CO2 1 2KOH —> K2CO3 1H2O respire in the absence of
oxygen. This process is
called fermentation or
anaerobic respiration
(Figure 14.12). There are
three types of fermentation:
1. Alcoholic fermentation
2. Lactic acid fermentation
Figure 14.11: Demonstration of
production of CO2 during respiration 3. Mixed acid fermentation
In the case of groundnut or bean 1. Alcoholic fermentation

seeds, the rise of water is relatively The cells of roots in water logged soil respire
lesser because these seeds use fat and by alcoholic fermentation because of lack
proteins as respiratory substrate and of oxygen by converting pyruvic acid into
release a very small amount of CO2. ethyl alcohol and CO2. Many species of yeast
But in the case of wheat grains, the (Saccharomyces) also respire anaerobically.
rise in water level is greater because This process takes place in two steps:
Pyruvate
they use carbohydrate as respiratory (i) 2CH3COCOOH
decarboxylase
2CH3CHO 12CO2↑

substrate. When carbohydrates are Pyruvic acid TPP
Alcohol Acetaldehyde
dehydrogenase
used as substrate, equal amounts of (ii) 2CH3CHO 1 2NADH12H1 2CH3CH2OH 1 2NAD1
Acetaldehyde Ethyl alcohol
CO2 and O2 are evolved and consumed.
159

Table 14.3: Comparison of alcoholic fermentation and lactic acid fermentation
Alcoholic fermentation Lactic acid fermentation
1.
It produces alcohol and releases CO2 It produces lactic acid and does not release
from pyruvic acid. CO2 from pyruvic acid.
2.
It takes place in two steps. It takes place in single step.
3.
It involves two enzymes, pyruvate
It uses one enzyme, lactate dehydrogenase
decarboxylase with Mg11 and alcohol
with Zn11.
dehydrogenase.
4.
It forms acetaldehyde as intermediate Does not form any intermediate
compound. compound.
Occurs in bacteria, some fungi and

5.
It commonly occurs in yeast.
vertebrate muscles.
Industrial uses of alcoholic 3. In producing vinegar and in tanning,

fermentation: curing of leather.
1. In bakeries, it is used for preparing 4. Ethanol is used to make gasohol (a fuel
bread, cakes, biscuits. that is used for cars in Brazil).
2. In beverage industries for preparing
wine and alcoholic drinks.
Glucose
Net gain of 2 ATP
+
2NAD
+
2NADH+H
2 x Pyruvic Acid
+ +
2 x NADH+H 2 x NADH+H
+ +
2 x NAD 2 x NAD
Alcohol dehydrogenase Lactate dehydrogenase
2 x Ethyl alcohol + CO2 2 x Lactic Acid

Alcoholic fermentation Lactic acid fermentation
Figure 14.12: Anaerobic Respiration
160

2. Lactic acid fermentation Table 14.5: Net products from one
Some bacteria (Bacillus), fungi and molecule of Glucose under Glycolysis and
muscles of vertebrates produce lactic acid Anaerobic respiration.
from pyruvic acid (Table 14.3). Substrate
Reduced Total
2CH3COCOOH 1 2NADH12H1 Stage level ATP
NAD1 ATP
Pyruvic acid Lactate dehydrogenase production
2CH3CHOHCOOH 1 2NAD1 Glycolysis 2 2* 8

Lactic acid 2 reduced
Anaerobic
3. Mixed acid fermentation 2 NAD1 re- 2
respiration
oxidised
This type of fermentation is a characteristic
feature of Enterobacteriaceae and results *One reduced NAD1 equivalent to 3 ATPs
in the formation of lactic acid, ethanol,
formic acid and gases like CO2 and H2. Check your grasp!
Characteristics of Anaerobic Respiration • Why Microorganisms respire
1. Anaerobic respiration is less efficient anaerobically?
than the aerobic respiration (Figure 14. 12) • Does anaerobic respiration take
(Table 14.4). place in higher plants?
2. Limited number of ATP molecules
is generated per glucose molecule
(Table 14.5). Demonstration of alcoholic
3. It is characterized by the production of fermentation
CO2 and it is used for Carbon fixation in Take a Kuhne’s fermentation tube
photosynthesis. which consists of an upright glass
tube with side bulb. Pour 10% sugar
Table 14.4: Comparison between glycolysis solution mixed with baker’s yeast
and fermentation into the fermentation tube the side
Glycolysis Fermentation tube is filled plug the mouth with lid.
After some time, the glucose solution
1. Glucose is Starts from pyruvic
acid and is converted will be fermented. The solution will
converted into
into alcohol or lactic give out an alcoholic smell and level
pyruvic acid.
acid. of solution in glass column will fall
2. It takes place in It takes place in the due to the accumulation of CO 2 gas.
the presence or absence of oxygen. It is due to the presence of zymase
absence of oxygen. enzyme in yeast which converts the
3. Net gain is 2ATP. No net gain of ATP glucose solution into alcohol and
molecules. CO 2. Now introduce a pellet of KOH
4. 2NADH 1 H 1
2NADH 1 H1 into the tube, the KOH will absorb
molecules are molecules are CO 2 and the level of solution will
produced. utilised. rise in upright tube (Figure 14.13).
161

Activity
CO2 Take a bottle filled with warm water

mixed with baker’s yeast and sugar. After
some time, you will notice water bubbling
as yeast produces carbon dioxide. Attach
a balloon to the mouth of the bottle.
Sugar solution and Yeast
After 30 minutes you’ll notice balloon
standing upright (Figure 14.14).
Why the balloon has inflated?
Yeast & sugar in warm
water were poured
into a bottle After 15 minutes. After 30 minutes.
Sugar Sugar Sugar
Figurefermentation
Figure 14.17 Kuhne’s 14.13: Kuhne’s
experiment Figure: 14.14: Air balloon activity
fermentation experiment
14.8 Factors Affecting Respiration
Factors affecting Respiration

The amount of protoplasm
and its state of activity Optimum temperature for
influence the rate of Internal External respiration is 30º C. At low
respiration Factors Factors temperatures and very high
temperatures rate of respiration
Concentration of respiratory decreases
substrate is proportional to
the rate of respiration W h e n s u ffic i e n t a m o u n t o f
O 2 is available the rate of
Wounding of plant
aerobic respiration will be
organs stimulates
optimum and anaerobic
the rate of respiration
respiration is completely stopped.
in that region.
This is called Extinction point.
Rate of respiration
decreases with
decreasing amount
of water. Proper hydration High concentration of CO2
is essential for respiration reduces the rate of respiration
Some chemical
s u b s t a n c e A plant or tissue transferred
acts as inhibitors. from water to salt solution
Example: Cyanides will increase the rate of
Light is an indirect factor
respiration. It is called
affecting the rate of respiration
salt respiration
162

or Direct Oxidative Pathway. It consists
How alcoholic beverages
of two phases, oxidative phase and non-
like beer and wine is
oxidative phase. The oxidative events
made?
convert six molecules of six carbon
The conversion of Glucose-6-phosphate to 6 molecules
pyruvate to ethanol takes place in malted of five carbon sugar Ribulose-5
barley and grapes through fermentation. phosphate with loss of 6CO2 molecules
Yeasts carryout this process under and generation of 12 NADPH 1 H1
anaerobic conditions and this conversion (not NADH). The remaining reactions
increases ethanol concentration. If the known as non-oxidative pathway, convert
concentration increases, it’s toxic effect Ribulose-5-phosphate molecules to
kills yeast cells and the left out is called various intermediates such as Ribose-5-
beer and wine respectively. phosphate(5C), Xylulose-5-phosphate(5C),
Glyceraldehyde-3-phosphate(3C),
14.9 Pentose Phosphate Pathway Sedoheptulose-7-Phosphate(7C), and
(Phospho Gluconate Pathway) Erythrose-4-phosphate(4C). Finally, five
molecules of glucose-6-phosphate is
During respiration breakdown of glucose regenerated (Figure 14.16). The overall
in cytosol occurs both by glycolysis reaction is:
(about 2/3) as well as by oxidative pentose
phosphate pathway (about 1/3). Pentose 6 x Glucose-6-Phosphate 1 12NADP1 1 6H2O
phosphate pathway was described by
Warburg, Dickens and Lipmann (1938). 5 x Glucose-6-Phosphate 1 6CO2 1 Pi 1
Hence, it is also called Warburg-Dickens- 12NADPH 112H1
Lipmann pathway. It takes place in The net result of complete oxidation
cytoplasm of mature plant cells. It is an of one glucose-6-phosphate yield 6CO2
alternate way for breakdown of glucose and 12NADPH 1 H1. The oxidative
(Figure 14.15). pentose phosphate pathway is controlled
It is also known as Hexose by glucose-6-phosphate dehydrogenase
monophosphate shunt (HMP Shunt) enzyme which is inhibited by high ratio of
NADPH to NADP1.
Starch
Oxidation via Pentose

phosphate Pathway Glucose Oxidation via glycolysis
Ribulose- 5-phosphate Pyruvic acid
Figure 14.15: Fate of Glucose in HMP shunt and Glycolysis

163

6C
Glucose
ATP
Hexokinase
Phosphorylation ADP
6C
36 C
Glucose-6-Phosphate
6 X Glucose-6-Phosphate +
6 x NADP
Glucose-6-Phosphate 1 1. Oxidation
Phospho hexose isomerase +
dehydrogenase 6 x NADPH+H
6. Isomerisation 36 C
30 C 6
6 X 6-Phospho Gluconolactone
5 X Fructose-6-Phosphate
2 6H2O 2. Hydration
5. Conversion 5 OXIDATIVE
Lactonase
164
PHASE 36 C
NON OXIDATIVE
30 C PHASE 6 X 6-Phospho Gluconate
+
Various intermediate compounds 6 x NADP
6-Phospho gluconate 3 +
such as 3C, 4C, 5C and 7C dehydrogenase 6 x NADPH+H
phosphorylated sugars
6CO2
30 C
6 X Ribulose-5-Phosphate
3. Oxidation and
Decarboxylation
4
4. Formation of
phosphorylated
compounds
Figure 14.16: Pentose phosphate pathway or HMP shunt
27-04-2018 11:58:45
Significance of pentose phosphate pathway Summary
1 HMP shunt is associated with the Respiration is a biological process in which
generation of two important products, energy is released by breaking down of
NADPH and pentose sugars, which play a vital complex organic substances into simple
role in anabolic reactions. compounds. The respiratory substrates may
2 Coenzyme NADPH generated is used for be carbohydrate, protein or fats. Respiration is
reductive biosynthesis and counter damaging of two types, aerobic (with O2) and anaerobic
the effects of oxygen free radicals (without O2). All plants, animals and most
3 Ribose-5-phosphate and its derivatives of the microbes derive energy from aerobic
are used in the synthesis of DNA, RNA, ATP, respiration. Some bacteria and fungi like yeast
NAD1, FAD and Coenzyme A. show anaerobic respiration. Aerobic respiration
4 Erythrose is used for synthesis of consists of four stages and they are glycolysis,
anthocyanin, lignin and other aromatic link reaction, TCA cycle and ETS. Glycolysis
compounds. is the first stage which occurs in cytosol and
common for both aerobic and anaerobic
respiration and it involves breaking down of
165

glucose into two molecules of pyruvic acid. molecules produced in plants are
Acetyl CoA formed from pyruvic acid, acts as a a. 3 b. 4 c. 6 d. 8
link between glycolysis and Krebs cycle. Krebs 3. The compound which links glycolysis and
cycle takes place in matrix of mitochondria Krebs cycle is
and also called as citric acid cycle in which CO2
a. succinic acid b. pyruvic acid
and H2O were produced. Hydrogen removed
c. acetyl CoA d. citric acid
from the substrates is received by coenzymes
which get reduced. They are again oxidised by 4. Assertion (A): Oxidative phosphorylation
removal of hydrogen. This hydrogen splits into takes place during the electron transport
protons and electrons. The electrons transferred chain in mitochondria.
through various electron transport carriers Reason (R): Succinyl CoA is
present in inner membrane of mitochondria is phosphorylated into succinic acid by
used for the synthesis of ATP with the help of substrate phosphorylation.
ATP synthase. This process is called oxidative a. A and R is correct. R is correct
phosphorylation. explanation of A
Anaerobic respiration involves incomplete b. A and R is correct but R is not the
breaking down of the substrate glucose correct explanation of A
into ethyl alcohol or lactic acid. In aerobic c. A is correct but R is wrong
respiration 36 ATP molecules are produced d. A and R is wrong.
in plant mitochondria but in animals 38 ATP 5. Which of the following reaction is not
molecules are produced per glucose molecule. involved in Krebs cycle.
During anaerobic respiration only 2 ATP a. Shifting of phosphate from 3C to 2C
molecules are produced, therefore anaerobic b. Splitting of Fructose 1,6 bisphosphate
respiration is less efficient than aerobic of into two molecules 3C compounds.
respiration. The respiratory quotient (RQ)
c. Dephosphorylation from the
is the ratio of carbon dioxide production to
substrates
oxygen consumption and reflects the relative
d. All of these
contributions of fat, carbohydrate, and protein
to the oxidation. Pentose phosphate pathway is 6. What are enzymes involved in
an alternative pathway to glycolysis and TCA phosphorylation and dephosphorylation
cycle for oxidation of glucose. It occurs in reactions in EMP pathway?
cytoplasm of both prokaryotes and eukaryotes. 7. Respiratory quotient is zero in succulent
plants. Why?
Evaluation
8. Explain the reactions taking place in
1. The number of ATP mitochondrial inner membrane.
molecules formed by 9. What is the name of alternate way of
complete oxidation of glucose breakdown? Explain the process
one molecule of pyruvic involved in it?
acid is
10. How will you calculate net products of
a. 12 b. 13 c. 14 d. 15 one sucrose molecule upon complete
2. During oxidation of two molecules of oxidation during aerobic respiration as
cytosolic NADH 1 H1, number of ATP per recent view?
166

t ICT Corner
Rate of respiration
Let’s estimate rate of

respiration
Steps
• Scan the QR code or go to google play store
• Select biology and select rate of respiration
• Click theory to know the basic about respiration
• Register yourself with mail-id and create password to access online lab simulations
Activity
• Press simulation to do the rate of respiration.
Step 2
Step 4
Step 1 Step 3
URL:
https://play.google.com/store/apps/details?id=in.edu.olabs.olabs&hl=en
Alternate web:
http://www.sumanasinc.com/webcontent/animations/content/
cellularrespiration.html
167

Chapter
15 Plant Growth and Development
seedling? How does a new plant structure

Learning Objectives arise from the pre-existing structure? Growth
The learner will be able to, is defined as an irreversible permanent
increase in size, shape, number,volume
• Define growth.
and dry weight. Plant growth occurs by cell
• List out and differentiate the phases division, cell enlargement, differentiation
of growth. and maturation.
• Understand the ways of measuring Bamboos are evergreen
growth. grasses and certain
• Explain the structure, precursor, species of it can grow
bioassay and physiological effects at the rate of growth 91
of plant growth regulators. cm per day. The Saguaro
Cactus is a tree like cactus and is a slow
Chapter Outline growing plant. The rate of growth is one
inch in the first ten years and it does not
15.1 Characteristics of growth begin to flower until it is about 60 years
15.2 Plant growth regulators old. It’s lifespan exceeds 150 years and
takes 75–100 years to grow a side arm.
15.3 Plant movements
15.4 Photoperiodism
15.5 Vernalization
15.6 Seed germination and dormancy
15.7 Senescence
15.8 Stress physiology

15.1 Characteristics of Growth

The Banyan tree continues to grow for
thousands of years and some others • Growth increases in protoplasm at
particularly annual plants cease growth cellular level.
within a season or within a year. Can you • Stem and roots are indeterminate in
understand the reasons? How does a zygote growth due to continuous cell division
give rise to an embryo and an embryo to a and is called open form of growth.
168

15.1.2 Phases of growth
Growth is
measurable, it is There are three phases of growth,
amazing to know that 1. Formative phase
one single maize root 2. Elongation phase
apical meristem can give rise to more 3. Maturation phase
than 17,500 new cells per hour and 1. Formative phase: Growth in this phase
cells in a watermelon may increase in occurs in meristematic cells of shoot and
size upto 3,50,000 times. root tips. These cells are small in size,
have dense protoplasm, large nucleus and
• The primary growth of the plant is due small vacuoles. Cells divide continuously
to the activity of apical meristem where, by mitotic cell division. Some cells retain
new cells are added to root and shoot capability of cell division while other cells
apex causing linear growth of plant body. enter the next phase of growth (Figure 15.1).
• The secondary vascular cambium and 2. Elongation Phase: Newly formed
cork cambium add new cells to cause daughter cells are pushed out of the
increase in girth. meristematic zone and increases the volume.
• Leaves, flowers and fruits are limited It requires auxin and food supply, deposition
in growth or of determinate or closed of new cell wall materials (intussusception),
form growth. addition of protoplasm and development of
• Monocarpic annual plants produce central vacuole take place.
flowers only once during lifetime and 3. Maturation Phase: During this
dies. Example: Paddy and Bean stage cells attain mature form and size.
• Monocarpic perennials produce Thickening and differentiation takes
flowers only once during life time place. After differentiation, the cells do
but the plants survive for many years. not grow further.
Example: Bamboo.
Activity
• Polycarpic perennials produce flowers
every year during life time. Example: Demonstration of phases of growth
Coconut. To demonstrate and study the phases
15.1.1 Indication of growth of growth, germinate a few seeds of
bean on a circular filter paper soaked
Growth in plants can be measured in with water in a petridish. After two
terms of, days of growth, select a few seedlings
i. Increase in length or girth (roots and with straight radical of 2 to 3 cm
stems) length. Dry the surface of radical with
ii. Increase in fresh or dry weight a blotting paper and mark the radical
iii. Increase in area or volume (fruits and from tip to base with at least 2 mm
leaves) gap using water proof ink. Replace the
seedlings in filter paper and observe
iv. Increase in number of cells produced.
further growth.
169

tip of the stem, root and branches. It is
the initial stage of growth. In other words,
growth starts from this period (Figure 15.2).
Maturation Phase ii. Log phase or exponential growth
Here, the newly formed cell increases
Elongation Phase
in size rapidly by deposition of cell wall
? Formative Phase
material. Growth rate is maximum and
Figure 15.1: Phases of growth in root reaches top because of cell division and
physiological processes are quite fast.
15.1.3 Kinetics of growth
The volume of protoplasm also increases.
It is an analysis of the motion of cells or It results in rapid growth and causes
expansion. elongation of internode in the stem.
1. Stages in Growth rate iii. Decelerating phase or Decline phase
The total period from initial to the final or slow growth phase
stage of growth is called the grand period The rate of growth decreases and becomes
of growth. The total growth is plotted limited owing to internal and external or
against time and ‘S’ shaped sigmoid both the factors because the metabolic
curve (Grand period curve) is obtained. process becomes slow.
It consists of four phases (Figure 15.2).
They are: iv. Steady state period or maturation
i. Lag phase phase
ii. Log phase In this phase cell wall thickening due
iii. Decelerating phase to new particle deposition on the inner
surface of the cell wall takes place. The
iv. Maturation phase
overall growth ceases and becomes
i. Lag phase constant. The growth rate becomes zero.
In this phase new cells are formed from
2. Types of growth rate
pre-existing cells slowly. It is found in the
The increased growth per unit time is
termed as growth rate. An organism or
Maturation Phase
Size / Weight of the organ
part of an organism can produce more cells

Decelerating Phase through arithmetic growth or geometric
growth or both.
se
i. Arithmetic Growth Rate

ha
gP
If the length of a plant organ is plotted

Lo
against time, it shows a linear curve and

Lag Phase this growth is called arithmetic growth.
Time • The rate of growth is constant and it
increases in an arithmetic manner.
Figure 15.2: Stages in growth rate
170

• Only one cell is allowed to divide
between the two-resulting progeny cell.
• One continues to divide but the other
undergoes cell cycle arrest and begins
Height of the plant

to develop, differentiate and mature.
C
• After each round of cell division, only
a single cell remains capable of division D
and one new body cell forms.
For example, starting with a single cell
after round 1 of cell division there is one
dividing cell and one body cell. After round
Time
2 there are two body cells, after round 3
there are three and so on (Figure 15.3). Figure 15.4: Constant Linear Growth
Dividing cell hair next to other epidermal cells. Hair may

Body Cell contain 5 to 10 cells by the division of the
basal cell. So, all its cells could be produced
in just five to ten days. In the figure 15.4, on
plotting the hight of the plant against time a
linear curve is obtained. Mathematically it is
expressed as:
Lt 5 Lo + rt
Lt 5 length at time ‘t’
Lo 5 length at time zero
r 5 growth rate of elongation per unit
ii. Geometric growth rate:
This growth occurs in many higher plants
Figure 15.3: Arithmetic Growth Rate
and plant organs and is measured in size
The plants single dividing cell would or weight. In plant growth, geometric cell
undergo one million rounds of nuclear and division results if all cells of an organism
cellular division. If each round requires one or tissue are active mitotically. Example:
day, this type of arithmetic increase would Round three in the given figure 15.5,
require one million days or 2739.7 years. produces 8 cells as 23 5 8 and after round
This arithmetic rate is capable of producing 20 there are 220 5 1,048,576 cells.
small number of cells present in very small The large plant or animal parts are
parts of plants. For example the hair on produced this way. In fact, it is common
many leaves and stems consists of just a in animals but rare in plants except when
single row of cells produced by the division they are young and small. Exponential
of the basal cell, the cell at the bottom of the growth curve can be expressed as,
171

Mother cell
2 Progeny cells
4 Progeny
cells
8 Progeny cells
Figure 15.5: Geometric growth
W1 5 W0ert
W1 5
Final size (weight, height and
number)
W0 5 Initial size at the beginning of the
period
r 5 Growth rate
t 5 Time of growth
e 5 Base of the natural logarithms Figure 15.6: Arithmetic and geometric
Here ‘r’ is the relative growth rate and growth of embryo
also a measure of the ability of the plant to Quantitative comparisons between the
produce new plant material, referred to as growth of living system can also be made
efficiency index. Hence, the final size of in two ways and is explained in the table 1.
W1 depends on the initial size W0. In figure 15.7, two leaves A and B are
iii. Arithmetic and Geometric Growth drawn at a particular time. Then A1and
B1 are drawn after a given time. A and
of Embryo
B 5 Area of leaves at a particular time. A1
Plants often grow by a combination and B1 5 Area of leaves after a given time.
of arithmetic and geometric growth (A1-A) and (B1-B) represents an absolute
patterns. A young embryonic plant grows increase in area in the given time. Leaf A
geometrically and cell division becomes
restricted to certain cells at the tips of roots Table 1: Comparison between absolute and
relative growth rates
and shoots. After this point, growth is of
Absolute growth rate Relative growth rate
the slower arithmetic type, but some of the
Increase in total The growth of the
new cells that are produced can develop into growth of two organs given system per unit
their mature condition and begin carrying measured and time expressed per
out specialized types of metabolism compared per unit unit initial parameter
(Figure 15. 6). Plants are thus a mixture of time is called absolute is called relative
older, mature cells and young, dividing cells. growth rate. growth rate.
172

increases from 5 cm2 to 10 cm2; 5 cm2 in a in carbon-di-oxide and hydrogen in water
given time. Leaf B increases from 50 cm2 are assimilated in photosynthesis.
to 55 cm2 ; 5 cm2 in a given time. Hence, c. Temperature
both leaves A and B increase their area
Temperature plays a significant role in
by 5 cm2 in a given time. This is absolute
the growth of the plant. Proper growth
growth. Relative growth is faster in leaf A
of a plant occurs at a about 28o C to 30o C
because of initial small size. It decreases
temperature and above 45o C will damage
with time (Figure 15.7).
the protoplasm and hinders the growth.
d. Oxygen
Oxygen has a vital role in the growth of
the plant. It helps in releasing metabolic
energy essential for growth activities. It is
necessary for respiration.
e. Light
Light has its own contribution in the
growth of the plant. Light is important
for growth and photosynthesis. Light
stimulates healthy growth. Absence of
Figure 15.7: Diagrammatic comparision of light may lead to yellowish in colour. This
absolute and relative growth rates is called etiolation.
II. Internal Factors
3.Conditions of growth a.
Genes are intracellular factors for
Plant growth is influenced by a variety growth.
of external and internal factors. A brief b. Phytohormones are intracellular factors
account of these factors is given below: for growth. Example: auxin, gibberellin,
I. External Factors cytokinin.
a. Water c. C/N ratio.
Water is essential for cell enlargement The ratio of carbohydrates and nitrogenous
as well as growth in the size of the compounds regulate the specific pattern
cell. Turgidity of cells helps in growth of growth in plants. For example, if a plant
extension. Water provides the medium for contains more nitrogenous compounds as
enzymatic activities needed for growth. compared to carbohydrates it produces
more protoplasm less mechanical tissues
b. Nutrition and vigorous vegetative growth. On the
Nutrition plays an important role in the other hand, less nitrogenous compounds
formation of protoplasm. Macro and micro and more carbohydrates favour the
elements are very important as sources of synthesis of more wall material, less
energy. For example, carbon and oxygen protoplasm, and more mechanical tissues.
173

4. Measurement of growth 5. Sequence of developmental process
in a plant cell
Activity
Development is a term that includes
Measurement of growth by direct
all the changes that an organism goes
method.
through during life cyle from germination
Step 1: Take ordinary scale. of a seed to senescence. Diagrammatic
Step 2: Measure ground stem up to representation of the sequence of processes
the growing point of the plant. which constitute the development of a cell
Step 3: Use Indian ink and mark at of a higher plant is given in the figure. It is
regular intervals to measure the length also applicable to tissues/organ.
of root, stem, and girth of the trunk.
Experiment: 1. Arc auxanometer:

The increase in the length of the stem tip can easily be measured by an arc auxanometer
which consists of a small pulley to the axis of which is attached a long pointer sliding
over a graduated arc. A thread one end of which is tied to the stem tip and another end
to a weight passes over the pulley tightly. As soon as the stem tip increases in length, the
pulley moves and the pointer slide over the graduated arc (Figure 15.8). The reading is
taken. The actual increase in the length of the stem is then calculated by knowing the
length of the pointer and the radius of the pulley. If the radius of the pulley is 4 inches
and the length of pointer 20 inches the actual growth is measured as follows:
Arc
Pulley
Pointer
Weight
Potted plant
Stand
Figure 15.8: Arc auxanometer

Actual growth in length 5 Distance travelled by the pointer × radius of the pulley
Length of the pointer
For example, actual growth in length 5 10 × 4 inches

20 inches
5 2 inches
174

1. Differentiation This ability is called plasticity. Example:
The process of maturation of meristematic Heterophylly in cotton and coriander.
cells to specific types of cells performing In such plants, the leaves of the juvenile
specific functions is called differentiation. plant are different in shape from those
in mature plants. On the other hand,
2. Dedifferentiation the difference in shapes of leaves
The living differentiated cells which had produced in air and those produced in
lost capacity to divide, regain the capacity water in buttercup also represent the
to divide under certain conditions. Hence, heterophyllous development due to
dedifferentiation is the regaining of the the environment. This phenomenon of
ability of cell division by the differentiated heterophylly is an example of plasticity.
cells. Example: Interfascicular cambium
and Vascular cambium. 15.2 Plant Growth Regulators
3. Redifferentiation Plant Growth Regulators
Differentiated cells, after multiplication (chemical messenger)
again lose the ability to divide and mature are defined as organic
to perform specific functions. This is called substances which are
redifferentiation (Figure 15.9). Example: synthesized in minute
Secondary xylem and Secondary phloem. quantities in one part
of the plant body and transported to
4. Plasticity another part where they influence specific
Plants follow different pathways in physiological processes. Five major groups
response to environment or phases of of hormones viz., auxins, gibberellins,
life to form different kinds of structures. cytokinins, ethylene and abscisic acid
are presently known to coordinate and
regulate growth and development in
plants. The term phytohormones is
implied to those chemical substances
which are synthesized by plants and thus,
naturally occurring. On the other hand,
there are several manufactured chemicals
which often resemble the hormones in
physiological action and even in molecular
structure. Recently, another two groups,
the brassinosteroids and polyamines were
also known to behave like hormones.
1. Plant growth regulators –

classification
Plant Growth Regulators are classified
Figure 15.9: Sequences of developmental
process in a plant cell as natural and synthetic based on their
175

Plant Growth Regulators (PGRs)
Natural (Phytohormones) Synthetic
Plant Growth Promoters Growth inhibitors
Auxin Ethylene NAA

Gibberellin Abscisic acid 2,4 -D
Cytokinin 2,4,5 - T
Figure 15.10: Classification of Plant Growth Regulators
source and a detailed flow diagram is ii. Antagonistic effects: The effect of two
given in Figure 15.10. substances in such a way that they have
opposite effects on the same process.
2. Characteristics of phytohormones
One accelerates and other inhibits.
i. Usually produced in tips of roots, stems
Example: ABA and gibberellins during
and leaves.
seed or bud dormancy. ABA induces
ii. Transfer of hormones from one place to dormancy and gibberellins break it.
another takes part through conductive
systems. 15.2.1 Auxins
iii. They are required in trace quantities.
1. Discovery
iv. All hormones are organic in nature.
During 1880, Charles Darwin noted the
v. There are no specialized cells or organs unilateral growth and curvature of Canary
for their secretion. grass (Phalaris canariensis) coleoptile to light.
vi. They are capable of influencing The term auxin (Greek: Auxin – to Grow)
physiological activities leading to was first used by F. W. Went in 1926 using
promotion, inhibition and modification Oats (Avena) coleoptile and isolated the
of growth. auxin. F. W. Went in 1928 collected auxin in
agar jelly. Kogl and Haugen Smith (1931)
3. Synergistic and Antagonistic effects isolated Auxin from human urine, and called
i. Synergistic effects: The effect of one or it as Auxin A. Later on in 1934, similar active
more substance in such a way that both substances was isolated from corn grain oil
promote each others activity. Example: and was named as Auxin B. Kogl et al., (1934)
Activity of auxin and gibberellins or found heteroauxin in the plant and chemically
cytokinins. called it as Indole Acetic Acid (IAA)
176

Types of Auxin
Natural Synthetic
Auxin occuring in plants are called These are synthesized artificially and have
“Natural auxin” properties like Auxin.
1. Indole Acetic Acid (IAA) 1. 2,4-Dichloro Phenoxy Acetic Acid (2,4-D)
2. Indole Propionic Acid (IPA) 2. 2,4,5-Trichloro Phenoxy Acetic Acid (2,4,5-T)
3. Indole Butyric Acid (IBA) 3. Napthalene Acetic Acid (NAA)
4. Phenyl Acetic Acid (PAA)
Figure 15.11: Classification of Auxins
2. Occurrence 7. Chemical structure

Auxin is generally produced by the growing Auxin has similar chemical structure of
tips of the stem and root, from where they IAA.
migrate to the region of the action.
8. Transport in Plants
3. Types of Auxin
Auxin is polar in transport. It includes
Auxins are divided into two categories basipetal and acropetal transport.
Natural auxins and Synthetic auxins Basipetal means transport through
(Figure 15.11). phloem from shoot to root and acropetal
means transport through xylem from root
Anti-auxins to shoot.
Anti-auxin compounds when applied
to the plant inhibit the effect of auxin. 9. Bioassay (Avena Curvature Test /
Example: 2, 4, 5-Tri Iodine Benzoic Went Experiment)
Acid (TIBA) and Napthylpthalamine. Bioassay means testing of substances for
their activity in causing a growth response
4. Free auxin in a living plant or its part.
They move out of tissues as they are easily The procedure involves the following
diffusible. Example: IAA. steps:
5. Bound Auxin When the Avena seedlings have attained

They are not diffusible. Example: IAA- a height of 15 to 30 mm, about 1mm of
Aspartic acid the coleoptile tip is removed. This apical
part is the source of natural auxin. The
6. Precursor tip is now placed on agar blocks for few
The amino acid Tryptophan is the hours. During this period, the auxin
precursor of IAA and zinc is required for diffuses out of these tips into the agar.
its synthesis. The auxin containing agar block is now
177

Auxin in the Auxin containing agar block Diffusion of Auxin
Avena coleoptile in one side of stump from agar block
Decapited stump
Coleoptile placed on
Agar Block Auxin diffuses
in to agar block
Figure 15.12: Avena Curvature Test
placed on one side of the decapitated and for the formation of callus.
stump of Avena coleoptile. The auxin • Auxin stimulates respiration.
from the agar blocks diffuses down • Auxin induces vascular differentiation.
through coleoptile along the side to
which the auxin agar block is placed. An
Agent Orange
agar block without auxin is placed on
another decapitated coleoptile. Within Mixture of two phenoxy herbicides
an hour, the coleoptiles with auxin agar 2,4-D and 2,4,5-T is given the name
block bends on the opposite side where ‘Agent orange’ which was used by
the agar block is placed. This curvature USA in Vietnam war for defoliation
can be measured (Figure 15.12). of forest (chemical warfare).
10. Physiological Effects

• They promote cell elongation in stem
and coleoptile.
• At higher concentrations auxins inhibit
the elongation of roots but induce more
lateral roots. Promotes growth of root
only at extremely low concentrations.
• Suppression of growth in lateral
bud by apical bud due to auxin
produced by apical bud is termed as
In botanical gardens and tea gardens,
apical dominance.
gardeners trim the plants regularly
• Auxin prevents abscission. so that they remain bushy. Does this
• It is responsible for initiation and practice have any scientific explanation?
promotion of cell division in cambium, Yes, trimming of plants removes
which is responsible for the secondary apical buds and hence apical
growth and tumor. This property of dominance. The lateral buds sprout
induction of cell division has been and make the plants bushy.
exploited for tissue culture techniques
178

11. Agricultural role and steroids) formed by 5-C precursor,
• It is used to eradicate weeds. Example: an Isoprenoid unit called Iso Pentenyl
2,4-D and 2,4,5-T. Pyrophosphate (IPP) through a number
• Synthetic auxins are used in the formation of intermediates. The primary precursor
of seedless fruits (Parthenocarpic fruit). is acetate.
• It is used to break the dormancy in seeds. 4. Chemical structure

• Induce flowering in Pineapple by NAA All gibberellins have gibbane ring
& 2,4-D. structure.
• Increase the number of female flowers
and fruits in cucurbits. 5. Transport in plants
The transport of gibberellins in plants is
15.2.2 Gibberellins non-polar. Gibberellins are translocated
through phloem and also occur in xylem
1. Discovery
due to lateral movement between vascular
The effect of gibberellins had been known bundles.
in Japan since early 1800 where certain
rice plants were found to suffer from 6. Bioassay (Dwarf Pea assay)
‘Bakanae’ or foolish seedling disease. Seeds of dwarf pea are allowed to germinate
This disease was found by Kurosawa till the formation of the coleoptile. GA
(1926) to be caused by a fungus Gibberella solution is applied to some seedlings. Others
fujikuroi. The active substance was are kept under control. Epicotyle length
separated from fungus and named as is measured and as such, GA stimulating
gibberellin by Yabuta (1935). These are epicotyle growth can be seen.
more than 100 gibberellins reported from
7. Physiological Effects
both fungi and higher plants. They are
noted as GA1, GA2, GA3 and so on. GA3 • It produces extraordinary elongation
is the first discovered gibberellin. In 1938, of stem caused by cell division and cell
Yabuta and Sumiki isolated gibberellin in elongation.
crystalline form. In1955, Brain et al., gave • Rosette plants (genetic dwarfism)
the name gibberellic acid. In 1961, Cross plants exhibit excessive internodal
et al., established its structure. growth when they are treated with
gibberellins. This sudden elongation
2. Occurrence of stem followed by flowering is called
The major site of gibberellin production bolting (Figure 15.13).
in plants is parts like embryo, roots and • Gibberellin breaks dormancy in potato
young leaves near the tip. Immature seeds tubers.
are rich in gibberellins. • Many biennials usually flower during
second year of their growth. For
3. Precursors flowering to take place, these plants
The gibberellins are chemically related to should be exposed to cold season. Such
terpenoids (natural rubber, carotenoids plants could be made to flower without
179

(liquid endosperm of coconut) which
contains cell division inducing substances.
In 1954, Skoog and Miller discovered
that autoclaved DNA from herring
Rosette leaves sperm stimulated cell division in tobacco
pith cells. They called this cell division
inducing principle as kinetin (chemical
structure: 6-Furfuryl Amino Acid).
(b) Treated plant This does not occur in plants. In 1963,
(a) Untreated plant
showing bolting. Lethan introduced the term cytokinin.
Figure 15.13: Bolting In 1964, Lethan and Miller isolated and
identified a new cytokinin called Zeatin
exposure to cold season in the first
from unripe grains of maize. The most
year itself, when they are treated with
widely occurring cytokinin in plants is
gibberellins.
Iso Pentenyl adenine (IPA).
8. Agricultural role
2. Occurrence
• Formation of seedless fruits without
Cytokinin is formed in root apex, shoot
fertilization is induced by gibberellins
apex, buds and young fruits.
Example: Seedless tomato, apple and
cucumber. 3. Precursor
• It promotes the formation of male Cytokinins are derivatives of the purine
flowers in cuccurbitaceae. It helps in adenine.
crop improvement.
4. Bioassay (Neem Cotyledon Assay)
• Uniform bolting and increased uniform
seed production. Neem cotyledons are measured and placed
in cytokinin solution as well as in ordinary
• Improves number and size of fruits in
water. Enlargement of cotyledons is an
grapes. It increase yield.
indication of cytokinin activity.
• Promotes elongation of inter-node in
sugarcane without decreasing sugar 5. Transport in plants
content. The distribution of cytokinin in plants
• Promotion of flowering in long day is not as wide as those of auxin and
plants even under short day conditions. gibberellins but found mostly in roots.
• It stimulates the seed germination. Cytokinins appear to be translocated
through xylem.
15.2.3 Cytokinins (Cytos – cell, 6. Physiological effect
Kinesis – division)
• Cytokinin promotes cell division in the
1. Discovery presence of auxin (IAA).
The presence of cell division inducing • Induces cell enlargement associated
substances in plants was first demonstrated with IAA and gibberellins
by Haberlandt in 1913 in Coconut milk
180

• Cytokinin can break the dormancy 4. Precursor
of certain light-sensitive seeds like It is a derivative of amino acid methionine,
tobacco and induces seed germination. linolenic acid and fumaric acid.
• Cytokinin promotes the growth of lateral
bud in the presence of apical bud. 5. Bioassay (Gas Chromatography)
Ethylene can be measured by gas
• Application of cytokinin delays the
chromatography. This technique helps in
process of aging by nutrient mobilization.
the detection of exact amount of ethylene
It is known as Richmond Lang effect.
from different plant tissues like lemon
• Cytokinin (i) increases rate protein and orange.
synthesis (ii) induces the formation
of inter-fascicular cambium 6. Physiological Effects
(iii) overcomes apical dominance • Ethylene stimulates respiration and
(iv) induces formation of new leaves, ripening in fruits.
chloroplast and lateral shoots. • It stimulates radial growth in stem and
• Plants accumulate solutes very actively root and inhibits linear growth.
with the help of cytokinins. • It breaks the dormancy of buds, seeds
and storage organs.
15.2.4 Ethylene • It stimulates formation of abscission
(Gaseous Phytohormone) zone in leaves, flowers and fruits. This
makes the leaves to shed prematurely.
Almost all plant tissues produce ethylene
• Inhibition of stem elongation
gas in minute quantities.
(shortening the internode).
1. Discovery • In low concentration, ethylene helps in
In 1924, Denny found that ethylene stimulates root initiation.
the ripening of lemons. In 1934, R. Gane • Growth of lateral roots and root hairs.
found that ripe bananas contain abundant This increases the absorption surface of
ethylene. In 1935, Cocken et al., identified the plant roots.
ethylene as a natural plant hormone. • The growth of fruits is stimulated by
ethylene in some plants. It is more
2. Occurrence marked in climacteric fruits.
Maximum synthesis occurs during • Ethylene causes epinasty.
climacteric ripening of fruits (see Box
info) and tissues undergoing senescence. Agricultural role
It is formed in almost all plant parts like • Ethylene normally reduces flowering in
roots, leaves, flowers, fruits and seeds. plants except in Pine apple and Mango.
• It increases the number of female
3. Transport in plants flowers and decreases the number of
Ethylene can easily diffuse inside the plant male flowers.
through intercellular spaces. • Ethylene spray in cucumber crop
produces female flowers and increases
the yield.
181

Ethylene Synergistic effects Auxin
Auxin, GA3 and
X
Cytokinin induces
X
x
Plant growth
Fruit
InduceEthylene Apical Prevents
Abscission ripening dominance abscission Weedicide
Plant Growth Cytokinins

Regulators
182
12
9 3
Radial growth 6 x
ABA Root /Shoot Delaying Promote

initiation from ageing lateral bud
ABA GA3 Callus process growth
ABA
Induces Breaks Gibberellins
seed seed
ABA
dormancy dormancy
Yellowing Induce Closure of

GROWTH PROMOTERS
GROWTH INHIBITORS
of leaf Abscission stomata
Antagonistic effects Bolting
27-04-2018 12:12:11
3. Precursors
Climacteric fruits: In most of the The hormone is formed from mevalonic
plants, there is sharp rise in respiration acid pathway or xanthophylls.
rate near the end of the development
of fruit, called climacteric rise. Such 4. Transport in plants
fruits are called climacteric fruits. The Abscisic acid is transported to all parts
ripening on demand can be induced of the plant through diffusion as well as
in these fruits by exposing them to through phloem and xylem.
normal air containing about 1 ppm
5. Chemical structure
of ethylene. A liquid called ethephon
is being used in fruit ripening as it It has carotenoid structure.
continuously releases ethylene. 6. Bioassay (Rice Coleoptile)
Example: Tomato, Apples, Banana, The inhibition of IAA induces straight
Mango. growth of rice seedling coleoptiles.
Non climacteric fruits: All fruits
7. Physiological effects
cannot be ripened by exposure to
ethylene. Such fruits are called non- • It helps in reducing transpiration rate
climacteric fruits and are insensitive by closing stomata. It inhibits K1 uptake
to ethylene. by guard cells and promotes the leakage
of malic acid. It results in closure of
Example: Grapes, Watermelon,
stomata.
Orange.
• It spoils chlorophylls, proteins and
nucleic acids of leaves making them
15.2.5 Abscisic Acid (ABA) yellow.
(Stress Phyto Hormone)
• Inhibition of cell division and cell
1. Discovery elongation.
In 1963, the hormone was first isolated • ABA is a powerful growth inhibitor. It
by Addicott et al., from young cotton causes 50% inhibition of growth in Oat
bolls and named as Abscission II. Eagles coleoptile.
and Wareing during 1963–64 isolated a • It induces bud and seed dormancy.
dormancy inducing substance from leaves • It promotes the abscission of leaves,
of Betula and called it as dormin. In 1965, flowers and fruits by forming abscission
it was found by Cornsforth et al., that both layers.
dormin and abscission are chemically • ABA plays an important role in plants
same compounds and called Abscisic during water stress and during drought
Acid (ABA). conditions. It results in loss of turgor
2. Occurrence and closure of stomata.
This hormone is found abundantly inside • It has anti-auxin and anti-gibberellin
the chloroplast of green cells. property.
• Abscisic acid promotes senescence in
183

leaves by causing loss of chlorophyll 15.3 Plant Movements
pigment decreasing the rate of
Plants have the capacity for
photosynthesis and changing the rate
changing their positions
of proteins and nucleic acid synthesis.
in response to external
8. Agricultural Role or internal stimuli, which
• In Cannabis sativa, induces male flower are known as plant
formation on female plants. movements. Movements
are basically of two types: I. Vital movements
• Induction of flowers in short day plants.
and II. Physical movements (hygroscopic)
• It promotes sprouting in storage organs (Figure 15.14).
like Potato.
• ABA plays an important role in plants I. Vital movements
during water stress drought conditions. Vital movements are those which are
• It inhibits the shoot growth and exhibited by the living cells or plants or
promotes growth of root system. This organs and they are always related to
character protect the plants from water the irritability of the protoplasm. These
stress. Hence, ABA is called as stress movements are of two types:
hormone. A. Movements of locomotion
B. Movements of curvature
Plant Movement
Vital Movement Physical Movement
Movement of locomotion Movement of curvature
Autonomic Paratonic Autonomic Paratonic

(Spontaneous) (Tactic/ Induced) (Spontaneous) (Tactic/ Induced)
Ciliary Phototactic
Amoeboid Chemotactic
Cyclosis Thermotactic Tropic Nastic
(Growth Movement) (Variation Movement)
Geotropic Nyctinastic
Phototropic Seismonastic
Thigmotropic Thigmonastic
Growth Movement Variation Movement Hydrotropic
Hyponastic Chemotropic
Epinastic Thermotropic
Nutational Aerotropic
Figure 15.14: Types of Plant Movements

184

Paratonic or Tactic (induced) movement of locomotion
Phototactic Chemotactic Thermotactic

These movements It occurs in response to It occurs in response
occur in response to chemical stimulus. to heat stimulus.
light. Example: Antherozoids in Example:
Example: Zoospores Bryophytes and Chlamydomonas
of Chlamydomonas Pteridophytes are moves from cold to
warm water
attracted to chemical
substances of Archegonia
Figure 15.15: Types of Paratonic movements

A. Movements of locomotion i. Autonomic movements of curvature
These movements include the movement The movement arising from internal
of protoplasm inside the cell or movement changes or internal stimuli of plant
of whole unicellular or multicellular plant body is called autonomic movement
body as in Chlamydomonas, gametes and of curvature. This does not require any
zoospores. external stimulus. They are two types:
i. Autonomic movements of locomotion a. A
utonomic movement of growth: It is of
the following types:
The movements arising from internal
changes or internal stimuli of plant 1. Hyponasty: When growth is more
body is called autonomic movements of on lower surface, petals show curvature
locomotion. This movement takes place on upper side and ultimately the flower
due to the presence of cilia or flagella and becomes closed. Such type of movement
movement of cytoplasm (Cyclosis). is called hyponasty.
2. Epinasty: When the growth is more
ii. Paratonic or Tactic (induced) on upper surface, petals show curvature
movements of locomotion on the lower side and ultimately the
The movements due to external factors flower opens. Such movement is called
or stimuli like light, temperature and epinasty. The flower usually opens at
chemicals are called paratonic movement high temperature and remains closed at
of locomotion (Figure 15.15). low temperature (Figure 15.16).
B. Movement of curvature 3. Nutation: The growth of the stem
apices occurs in a zig-zag manner. It is
In higher plants they are restricted only
because the two sides of the stem apex
to bending or curvature of some of their
alternatively grow more. Such growth
parts. There are mainly two types:
movements are called as nutational
They are i) Autonomic movement of
movements. In some plants nutational
curvature and ii) Paratonic movement of
movements allow the shoots apex
curvature.
to grow in helical path in upward
185

Hyponasty Epinasty
Figure 15.16: Hyponasty and Epinasty
direction. This movement is called

circumnutation. It is commonly found
in the stems of climbers of Cucurbitaceae
(Figure 15. 17).
b.
Autonomic Movement of variation:
It happens in Indian telegraph plant.
(Desmodium gyrans). The compound
leaf consists of a larger terminal and
two smaller lateral leaflets. During
day time, the two lateral leaflets move
upward at an angle of 90° and come to
lie parallel to the rachis. Again, they may
move downward at 180° so that they are
Figure 15.17: Circumnutation
parallel to the rachis. They may again
move upward at 90° to come in their
original position. All these movements
occur with jerks after intervals, each
movement being completed in about
2 minutes (Figure 15.18).
ii. Paratonic (induces) movements of
curvature
The movement arising from external
stimulus is called Paratonic (Induced)
movements of curvature. They are of
two types. 1) Tropic movements 2) Nastic
movements (Table 2)
a. Tropic movements
A movement that occurs in response to
Figure 15.18: Autonomic Movement of
an unidirectional stimulus is called tropic Variation
186

Table 2: Differences between Tropic Movements and Nastic movements
Tropic Movements Nastic movements
1. Movement occurs due to unidirectional These movements occur due to a
stimulus. diffused stimulus.
2. The stimulus acts on protoplasm from one The stimulus acts on the protoplasm
direction only. from all sides.
3. The response is directly related to the The response has no relation to the
direction of the stimulus. direction of the stimulus but with
organ.
4. These are movements of curvature caused These are also the movement of
by unilateral growth. curvature but they are caused by
reversible turgor changes.
5. Tropic movements may be phototropic, Nastic movements may be
geotropic, hydrotropic, thigmotropic, seismonastic, photonastic or
chemotropic, thermotropic or aerotropic. thermonastic
movement or tropism. There are seven flowers move towards the stimulus of light
types in tropic movements (Geotropic, and are said to be positively phototropic
Phototropic, Thigmotropic, Chemotropic, while others such as roots and rhizoids
Hydrotropic, Thermotropic and Aerotropic) which move away from the stimulus of
1. Geotropism light are called negatively phototropic.
The movements which take place in b. Nastic Movements
response to gravity stimulus are called
When growth movements occur in
geotropic movements. The primary
response to an external stimulus which is
roots growing down into soil are
not unidirectional but diffused, they are
positive geotropic. Primary stems that
called nastic or paratonic movements of
grow away from soil (against gravity)
variation. Paratonic variation movements
are negative geotropic. Secondary roots
are determined by some external stimuli,
growing at right angles to the force of
light, temperature, chemicals and touch.
gravity are Diageotropic. Secondary
They are:
lateral roots which grow obliquely
downwards are Plagiogeotropic. 1. Nyctinastic movement (or) sleep
Lateral roots and branches which are movement
not sensitive to gravitational stimulus
The diurnal (change in day-night)
are Apogeotropic.
movements of leaves and flowers of some
2. Phototropism species which take up sleep position at
The tropic movement taking place as night are called nyctinastic movements.
a response to light stimulus is called They are caused by relative changes in
phototropism. Some of the plant parts such cell size on the opposite sides of the leaf
as stems, branches, leaves and pedicels of base called pulvinus. The movements
187

are attributed to the amount of auxin and
Experiment to demonstrate K1 ions. The entry of water to the lower
negative geotropism in aerial stem
side of the pulvinus causes the leaves to
The Clinostat has a rotating pot like
container mounted on an axis rod. stand erect and exit of water causes them
A potted plant is fitted horizontally to drop. They are of two types:
on the Clinostat and rotated slowly
which completely eliminates gravity i. Photonasty
as all the sides of the plant are equally The nastic movement caused in response to
stimulated. If the rotation of the light is called photonasty or photonastic
Clinostat is stopped for a considerable
period of time then the tip of the movement. The opening of leaves and
stem is observed to curve and grow flowers during daytime and their close at
upwards this proves that the stem tip night is an example.
is negatively geotropic (Figure 15.19).
ii. Thermonasty
Clinostat
The nastic movement taking place
in response to temperature is called
thermonasty or thermonastic movement.
In Crocus the flowers open at high
temperature and close at low temperature.
Figure 15.19: Clinostat 2. Seismonastic movement
This means a response to shaking. The
Experiment to demonstrate positive best example is Mimosa pudica (Touch-
phototropism in shoot tips
A darkened black box is taken having
a small window on one side. A well-
watered potted plant is placed inside the
box. This is referred to as a phototropic
chamber or heliotropic chamber. If the
window is kept closed for about 24 hours
the plant shows normal growth. If the
window is kept opened, it is found after
two days that the shoot tip bends and
grows towards light proving that it is
positively phototropic (Figure 15.20).
Figure 15.21: Mimosa pudica showing
Closed Window Open Window response to touch
Direction of
Darkness Light me-not plant) which is a sensitive plant
(Figure 15.21). Such plants respond to
stimuli such as touch, blow or metallic
shock by folding their leaflets and
lowering their leaves .This effect is caused
Figure15.20: Experiment to demonstrate by the movement of water in and out of
Photropism the parenchymatous cells of the pulvinus
(Figure 15.22).
188

b c
(d) (e)
plasma
H 2O membrane
cell wall
flexor cells
turgid extensor Ca2+
n us cells TnV
lvi O
pu N H2
H+
K+
Cl - turgid state
shrinking swelling
+
Cl-
stretched K flaccid state
flexor cells H+
flaccid extensor H 2O H 2 O H O
2
O
cells
2
H
H 2O H 2O
TnV
T nV
N
Figure 15.22: Mechanism of Seismonastic

Figure 15.22
movement in Mimosa pudica
3. Thigmonastic movement hairs are activated. Similarly, in dionaea,

The movements found in the leaves of the two halves of the leaf curve upwards
Drosera and Dionaea (Venus fly trap) result along the midrib. These parts of the leaves
in response to the touch stimulus of insects. come to their normal position after the
As soon as an insect sits on the leaf the cilia insect has been digested (Figure 15.23).
curve inward to trap the insect and trigger
II Physical Movement (Hygroscopic
Movements)
Physical movements are those which are
found in dead parts of the plants and
they are not related to any irritability
of the protoplasm. They are also called
hygroscopic movements or mechanical
movements. Dispersal of spores and
seeds, dehiscence of sporangia, bursting
of seeds and movement of elaters are
the examples of physical or hygroscopic
Figure 15.23: Thigmonasty in Dionaea movement.
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15.4 Photoperiodism
Intermediate plants Day neutral plants
Trees take several years for initiation of
flowering whereas an annual herb flowers
within few months. Each plant requires Photoperiodism in plants
a specific time period to complete their
vegetative phase which will be followed
by reproductive phase as per their Long day plants Short day plants
internal control points through Biological
Clock. The physiological mechanisms in
relation to flowering are controlled by Short long day Long short day
(i) light period (Photoperiodism) and plants plants
(ii) temperature (Vernalization). The
Figure: 15.24 Classification of Plants based
physiological change on flowering due on Photoperiodism
to relative length of light and darkness
iii. Short day plants: The plants that require
(photoperiod) is called Photoperiodism.
a short critical day length for flowering
The term photoperiodism was coined
are called short day plants or long night
by Garner and Allard (1920) when they
plants. Example: Tobacco, Cocklebur,
observed this in ‘Biloxi’ variety of soybean
Soybean, Rice and Chrysanthemum.
(Glycine max) and ‘Maryland mammoth’
variety of tobacco (Nicotiana tabacum). iv. Long short day plants: These are
The photoperiod required to induce actually short-day plants but they have
flowering is called critical day length. to be exposed to long days during their
Maryland mammoth (tobacco variety) early periods of growth for flowering.
requires 12 hours of light and cocklebur Example: Some species of Bryophyllum
(Xanthium pensylvanicum) requires and Night jasmine.
15.05 hours of light for flowering. v. Intermediate day plants: These require
a photoperiod between long day and
1. Classification of plants based on short day for flowering. Example:
Photoperiodism Sugarcane and Coleus.
Depending upon the photoperiodic vi. Day neutral plants: There are a
responses plants are classified as given in number of plants which can flower in
Figure 15.24. all possible photoperiods. They are also
i. Long day plants: The plants that require called photo neutrals or indeterminate
long critical day length for flowering plants. Example: Potato, Rhododendron,
are called long day plants or short night Tomato and Cotton.
plants. Example: Pea, Barley and Oats.
2. Photoperiodic induction
ii. Short long day plants: These are long
day plants but should be exposed to An appropriate photoperiod in 24 hours’
short day lengths during early period of cycle constitutes one inductive cycle.
growth for flowering. Example: Wheat Plants may require one or more inductive
and Rye. cycles for flowering. The phenomenon of
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conversion of leaf primordia into flower 5. Phytochrome
primordia under the influence of suitable
X
inductive cycles is called photoperiodic Day
induction. Example: Xanthium (SDP) – 1 Pr

660nm
Pfr Pfr X Physiological response
730nm
inductive cycle and Plantago (LDP) – 25 Night
inductive cycles.
Phytochrome is a bluish biliprotein pigment
3. Site of Photoinductive perception responsible for the perception of light in
Photoperiodic stimulus is perceived by the photo physiological process. Butler et al.,
leaves. Floral hormone is synthesised in (1959) named this pigment and it exists
leaves and translocated to the apical tip to in two interconvertible forms: (i) red light
promote flowering. This can be explained absorbing pigment which is designated as
by a simple experiment on Cocklebur Pr and (ii) far red light absorbing pigment
(Xanthium pensylvanicum), a short day which is designated as Pfr. The Pr form
plant. Usually Xanthium will flower under absorbs red light in 660nm and changes to Pfr.
short day conditions. If the plant is defoliated The Pfr form absorbs far red light in 730nm
and kept under short day conditions it will and changes to Pr. The Pr form is biologically
not flower. Flowering will occur even when inactive and it is stable whereas Pfr form is
all the leaves are removed except one leaf. biologically active and it is very unstable. In
If a cocklebur plant is defoliated and kept short day plants, Pr promotes flowering and
under long day conditions, it will not flower. Pfr inhibits the flowering whereas in long
If one of its leaves is exposed to short day day plants flowering is promoted by Pfr and
condition and rest are in long day condition, inhibited by Pr form. Pfr is always associated
flowering will occur (Figure 15.25). with hydrophobic area of membrane systems
while Pr is found in diffused state in the
The nature of flower producing cytoplasm. The interconversion of the two
stimulus has been elusive so far. It is forms of phytochrome is mainly involved in
believed by many physiologists that it flower induction and also additionally plays
is a hormone called florigen. The term a role in seed germination and changes in
florigen was coined by Chailakyan membrane conformation.
(1936) but it is not possible to isolate.
Short Day Long Day
4. Importance of photoperiodism
Short
1. The knowledge of photoperiodism Day
plays an important role in
hybridisation experiments.
2. Photoperiodism is an excellent
example of physiological
pre-conditioning that is using
an external factor to induce A B C D E F
physiological changes in the plant. Figure 15.25: Experiment on Cocklebur

plant showing photoperiodic stimulus
191

192

15.5 Vernalization (Vernal – Spring
Like) C Devernalization
Besides photoperiod certain plants require High

temperature
a low temperature exposure in their B

Vernalin D Chilling
earlier stages for flowering. Many species Translocation of flower
inducing substance
A
of biennials and perennials are induced to Florigen F
flower by low temperature exposure (0oC to Precursor
5oC). This process is called Vernalization.

The term Vernalization was first used by
T. D. Lysenko (1938).
1. Mechanism of Vernalization: Figure 15.26: Vernalization and Flowering

Two main theories to explain the
seeds are transferred to low temperature
mechanism of vernalization are:
(3oC to 5oC) from few days to 30 days.
i. Hypothesis of phasic development Germinated seeds after this treatment are
ii. Hypothesis of hormonal involvement allowed to dry and then sown. The plants
will show quick flowering when compared
i. Hypothesis of phasic development
to untreated control plants.
According to Lysenko, development of an
annual seed plant consists of two phases. 3. Devernalization
First phase is thermostage, which is Reversal of the effect of vernalization is
vegetative phase requiring low temperature called devernalization.
and suitable moisture. Next phase is photo
4. Practical applications
stage which requires high temperature for
1. Vernalization shortens the vegetative
synthesis of florigen (flowering hormone).
period and induces the plant to
ii. Hypothesis of hormonal involvement flower earlier.
According to Purvis (1961), formation 2. It increases the cold resistance of the
of a substance A from its precursor, plants.
is converted into B after chilling. The 3. It increases the resistance of plants to
substance B is unstable. At suitable fungal disease.
temperature B is converted into stable 4. Plant breeding can be accelerated.
compound D called Vernalin. Vernalin is
converted to F (Florigen). Florigen induces 15.6 Seed Germination and Dormancy
flower formation. At high temperature B is
converted to C and devernalization occurs I. Seed Germination
(Figure 15.26). The activation and growth of embryo
from seed into seedling during favourable
2. Technique of Vernalization: conditions is called seed germination.
The seeds are first soaked in water and
allowed to germinate at 10o C to 12o C. Then
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1. Types of germination b. Temperature: Seeds fails to germinate
There are two methods of seed at very low and high temperature.
germination. Epigeal and hypogeal. The optimum temperature is 25oC to
35oC for most tropic species.
i. Epigeal germination
c. Oxygen: It is necessary for
During epigeal germination cotyledons germination. Since aerobic
are pushed out of the soil. This happens respiration is a physiological
due to the elongation of the hypocotyl. requirement for germination most
Example: Castor and Bean. will germinate well in air containing
ii. Hypogeal germination 20% oxygen.
During hypogeal germination cotyledons d. Light: There are many seeds which
remain below the soil due to rapid elongation respond to light for germination and
of epicotyls (Figure 15.27). Example: Maize these seeds said to be photoblastic.
e. Soil conditions: Germination of seed
Foliage leaves
in its natural habit is influenced by
Epicotyl
soil conditions such as water holding
Hypocotyl Hypocotyl
capacity, mineral composition and
aeration of the soil.
Cotyledon
Radicle
(a) Bean ii. Internal factors
a. Maturity of embryo: The seeds of
some plants, when shed will contain
Foliage leaves
immature embryo. Such seeds
Hypocotyl
germinate only after maturation of
Epicotyl
Cotyledon
embryo.
Hypocotyl b. Viability: Usually seeds remain viable
Radicle or living only for a particular period.
(b) Pea Viability of seeds range from a few
Figure 15.27: (a) Epigeal Germination days (Example: Oxalis) to more than
(b) Hypogeal Germination hundred years. Maximum viability
(1000 years) has been recorded in
2. Factors affecting germination
lotus seeds. Seeds germinate only
Seed germination is directly affected by within the period of viability.
external and internal factors:
c. Dormancy: Seeds of many plants
i. External factors are dormant at the time of shedding.
a. Water: It activates the enzymes which A detailed treatment is given below.
digest the complex reserve foods of
the seed. If the water content of the II. Seed Dormancy
seed goes below a critical level, seeds The seeds of most plants germinate under
fail to germinate. favourable environmental conditions
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but some seeds do not germinate when due to blockage by cork cells. These
suitable conditions like water, oxygen and seeds are shaken vigorously to remove
favourable temperature are not available. the plug which is called Impaction.
Germination of such seeds may be delayed iii. Stratification: Seeds of rosaceous
for days, months or years. The condition plants (Apple, Plum, Peach and Cherry)
of a seed when it fails to germinate even in will not germinate until they have been
suitable environmental condition is called exposed to well aerated, moist condition
seed dormancy. There are two main under low temperature (0oC to 10oC)
reasons for the development of dormancy: for weeks to months. Such treatment is
Imposed dormancy and innate dormancy. called Stratification.
Imposed dormancy is due to low moisture iv. Alternating temperatures: Germination
and low temperature. Innate dormancy is of some seeds is strongly promoted
related to the properties of seed itself. by alternating daily temperatures. An
alternation of low and high temperature
1. Factors causing dormancy of seeds:
improves the germination of seeds.
i. Hard, tough seed coat causes barrier
v. Light: The dormancy of photoblastic
effect as impermeability of water, gas
seeds can be broken by exposing them
and restriction of the expansion of
to red light.
embryo prevents seed germination.
ii. Many species of seeds produce 15.7 Senescence
imperfectly developed embryos called
Plant life comprises some sequential events,
rudimentary embryos which promotes
viz: germination, juvenile stage, maturation,
dormancy.
old age and death. Old age is called
iii. Lack of specific light requirement leads
senescence in plants. Senescence refers to
to seed dormancy.
all collective, progressive and deteriorative
iv. A range of temperatures either higher processes which ultimately lead to complete
or lower cause dormancy. loss of organization and function. Unlike
v. The presence of inhibitors like phenolic animals, plants continuously form new
compounds which inhibits seed organs and older organs undergo a highly
germination cause dormancy. regulated senescence program to maximize
nutrient export.
2. Methods of breaking dormancy:
The dormancy of seeds can be broken by 1. Types of Senescence
different methods. These are: Leopold (1961) has recognised four types
i. Scarification: Mechanical and chemical of senescence:
treatments like cutting or chipping of i. Overall senescence
hard tough seed coat and use of organic ii. Top senescence
solvents to remove waxy or fatty iii. Deciduous senescence
compounds are called as Scarification. iv. Progressive senescence
ii. Impaction: In some seeds water and
oxygen are unable to penetrate micropyle
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secretes hydrolytic enzymes.
The branch of botany which deals with
• The starch content is decreased in the
ageing, abscission and senescence is
cells.
called Phytogerontology
• Photosynthesis is reduced due to loss of
i. Overall senescence: This kind of chlorophyll accompanied by synthesis
senescence occurs in annual plants and accumulation of anthocyanin
when entire plant gets affected and dies. pigments, therefore the leaf becomes red.
Example: Wheat and Soybean. It also • There is a marked decrease in protein
occurs in few perennials also. Example: content in the senescing organ.
Agave and Bamboo. • RNA content of the leaf particularly
ii. Top senescence: It occurs in aerial parts rRNA level is decreased in the cells
of plants. It is common in perennials, due to increased activity of the enzyme
underground and root system remains RNAase.
viable. Example: Banana and Gladiolus. • DNA molecules in senescencing leaves
iii. Deciduous senescence: It is common degenerate by the increased activity of
in deciduous plants and occurs only in enzyme DNAase.
leaves of plants, bulk of the stem and 3. Factors affecting Senescence:
root system remains alive. Example:
• ABA and ethylene accelerate senescence
Elm and Maple.
while auxin and cytokinin retard
iv. Progressive senescence: This kind of senescence.
senescence is gradual. First it occurs
• Nitrogen deficiency increases
in old leaves followed by new leaves
senescence whereas nitrogen supply
then stem and finally root system. It is
retards senescence.
common in annuals (Figure 15.28).
• High temperature accelerates senescence
2. Physiology of Senescence but low temperature retards senescence.
• Cells undergo changes in structure. • Senescence is rapid in dark than in
• Vacuole of the cell acts as lysosome and light.
Overall senescence Top senescence Deciduous senescence Progressive senescence
Figure 15.28: Different types of senescence in plants

196

• Water stress leads to accumulation of its vascular system to prevent loss of water
ABA leading to senescence. and nutrients. Final stage of senescence
is abscission. In temperate regions all the
4. Programmed cell death (PCD) leaves of deciduous plants fall in autumn
Senescence is controlled by plants own and give rise to naked appearance, then the
genetic programme and death of the plant new leaves are developed in the subsequent
or plant part consequent to senescence is spring season. But in evergreen plants there
called Programmed Cell Death. In short is gradual abscission of leaves, the older
senescence of an individual cell is called PCD. leaves fall while new leaves are developed
The proteolytic enzymes involving PCD continuously throughout the year.
in plants are phytaspases and in animals
are caspases. The nutrients and other 6. Morphological and Anatomical
substrates from senescing cells and tissues changes during abscission
are remobilized and reallocated to other parts Leaf abscission takes place at the base of
of the plant that survives. The protoplasts petiole which is marked internally by a
of developing xylem vessels and tracheids distinct zone of few layers of thin walled
die and disappear at maturity to make them cells arranged transversely. This zone is
functionally efficient to conduct water for
called abscission zone or abscission layer.
transport. In aquatic plants, aerenchyma
An abscission layer is greenish-grey in
is normally formed in different parts of the
colour and is formed by rows of cells of 2 to
plant such as roots and stems which encloses
large air spaces that are created through PCD. 15 cells thick. The cells of abscission layer
In the development of unisexual flowers, separate due to dissolution of middle lamella
male and female flowers are present in earlier and primary wall of cells by the activity of
stages, but only one of these two completes enzymes pectinase and cellulase resulting
its development while other aborts through in loosening of cells. Tyloses are also formed
PCD (Figure 15.29). blocking the conducting vessels. Degrading
of chlorophyll occur leading to the change
5. Abscission in the colour of leaves, leaf detachment
Abscission is a physiological process of from the plant and leaf fall. After abscission,
shedding of organs like leaves, flowers, fruits outer layer of cells becomes suberized by the
and seeds from the parent plant body. When development of periderm (Figure 15.30).
these parts are removed the plant seals off
Mitochondria
Vacuole
Nucleus
Plastid
Figure 15.27
Figure 15.29: Programmed
Programmed cell
Celldeath
Death
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functioning of plants under adverse
Pholoem environmental conditions is called stress
Xylem physiology. Jacob Levitt (1972) first used
the term biological stress in relation to
plants and according to him stress is “any
change in environmental condition that
might adversely change the growth and
Cortex
development of a plant”.
Abscission layer
The reaction of plants facing stress is called
strain. For example, if a normal plant growing
under favourable light conditions is subjected
to low light intensity, its photosynthesis is
reduced. Thus, low light intensity is referred
as stress and reduction of photosynthesis is
referred as strain. Biological strains are of
Figure 15.30: L.S of petiolar base showing two types; Elastic biological strain and Plastic
abscission layer
biological strain. If the reaction of plant
function is temporary and when it returns to
7. Hormones influencing abscission its original state it is called elastic biological
All naturally occurring hormones strain. Example: Temporary wilting. If
influence the process of abscission. Auxins the reaction is permanent and the plant
and cytokinins retard abscission, while function does not return to the normal state
abscisic acid (ABA) and ethylene induce it. it is called plastic biological strain. Example:
Permanent wilting. Some plants get adapted
8. Significance of abscission
to stress condition and are not adversely
1. Abscission separates dead parts of the affected by stress. Such plants are called
plant, like old leaves and ripe fruits. stress resistant or stress tolerant plants.
2. It helps in dispersal of fruits and Example: Mangroves. Some plants cannot
continuing the life cycle of the plant. face stress and they pass their adverse period
3. Abscission of leaves in deciduous in dormant state and so they are called stress
plants helps in water conservation enduring plants. Ephemeral plants are short
during summer. lived desert plants, which complete their life
4. In lower plants, shedding of vegetative cycle during the seasonal rains before the
parts like gemmae or plantlets help in onset of dry season. These ephemeral plants
vegetative reproduction. are called stress escapers. Stress in plants can
be classified as given in figure 15.31.
15.8 Stress Physiology
1. Biotic Stresses
Like all other organisms, plants are also
These are adverse effects on plants caused
subjected to various environmental stresses
by other living organisms such as viruses,
such as water deficit, drought, cold, heat,
bacteria, fungi, parasites, insects, weeds and
salinity and air pollution. The study of
198

Environmental Stress
Positive Allelopathy Allelopathic plant

Biotic Abiotic
Negative Allelopathy
Allelopathy Pathogenecity Atmospheric Edaphic
Water Salt
Promote Retard
Light Temperature Air pollution growth growth
Allelochemicals
Figure 15.31: Classification of Stress types SOIL
in plants
Figure 15.32: Allelopathy in plants
competing plants. Biotic environmental stress
is also caused due to the activity of man by allelochemicals exhibit symptoms such as
cutting herbs and trees, twigs for fodders, fuels wilting, chlorosis and death.
and agricultural purposes. The biotic stresses
caused by bacteria, fungi and nematodes that Check your grasp! Are all plants
are ever present in the environment are called allelopathic? Can allelopathic
potential biotic stresses. These are divided into chemicals affect animals and humans?
two types. They are:
i) Allelopathy Tree of heaven (Ailanthus altissima) is
An organism producing one or more a recent addition to the list of allelopathic
biochemical substances that greatly trees. Ailanthone an allelochemical
influence the germination, growth and extracted from the root of Ailanthus acts
reproduction of other organisms is as potent herbicide. In Sorghum plant
called Allelopathy. These biochemicals the allelochemical sorgolone possess
are known as allelochemicals. They allelopathic activity. It is found in root
are beneficial (positive allelopathic) exudates of most Sorghum species. Root
or detrimental (negative allelopathic). exudation of maize inhibits the growth of
These allelochemicals are obtained from some weeds such as Chenopodium album
leaf after leaching on the ground and and Amaranthus retroflexus. The seed
also from roots. The term allelopathy is exudates of oat (Avena fatua) affect the
from Greek words allelon-each other and germination of wheat seedling.
pathos-to suffer and first used in 1937 by ii. Pathogenecity
Hans Molisch. Allelopathic effect may
The effect of microbes that cause diseases
occur with weeds on crops and vice versa
in plants. Example: Xanthomonas citri
(Figure 15.32).
One of the most famous allelopathic 2. Abiotic Stresses
plants is Black walnut (Juglans nigrum). The Abiotic stress may occur due to an
chemical which is present in Black walnut atmospheric condition (atmospheric
is Juglone and it is a respiratory inhibitor. stress) or soil condition (edaphic stress).
Solanaceous plants such as tomato, Atmospheric stresses may occur due
capsicum and eggplant are susceptible to to excess and deficient levels of light
juglone. These plants when exposed to these temperature and air pollutants.
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i. Light Stress b. Low Temperature
Light limits the distribution of species. Low temperature stress is quite harmful to
In low light intensity Sciophytes (shade plants and the temperature near freezing
loving plants) develop, while in high light point causes irreversible damage so that
intensity Heliophytes (high light loving the plants fail to survive under extreme
plants) develop. In low light intensity, cold conditions. However, some plants
stomata do not fully open hence there is growing in alpine and arctic regions can
survive under low temperature and such
less diffusion of gases. As a result, there
plants are said to be cold resistant. Stress
is less photosynthesis and the chlorophyll
due to freezing temperature is called frost
synthesis is also affected. High light
stress. Temperature below 10oC, decreases
intensity also inhibits photosynthesis. root growth, increases leakage of ions and
Change in photoperiod inhibits flowering. ethylene production.
ii. Temperature
Plants are adapted to a particular region and Some plant parts like
they face temperature stress in another region. Seeds, pollen grains
and embryos can be
a. High temperature
stored at very low
High temperature causes soil and temperature (–196oC).
atmospheric drought. Plants are subjected
to permanent wilting in soil drought and
temporary wilting in atmospheric drought. iii. Air pollutants
Plants generally die above temperature Important atmospheric pollutants
of 44oC. However, some organisms like prevalent in the Indian sub-continent
Mastigocladus (a cyanobacterium) grow are CO2, CO, SO2, NO2, O3, fluoride
well at 85oC to 90oC in hot springs. At 42oC and H2S. These pollutants do not cause
synthesis of normal protein declines and new visible injury but cause hidden injury.
protein called Heat Shock Proteins (HSPs) If the concentration of these pollutants
appears. These proteins were discovered increases visible injury like chlorotic
in fruit fly (Drosophila melanogaster) and and necrotic spots appear on leaves as
since then they have also been discovered well as inhibit photosynthetic carbon
in animals, plants and microorganisms. metabolism and biomass formation. Some
At high temperature all physiological pollutants at low concentration stimulate
processes decline. Photosynthesis decreases plant growth. Example: SO2, NO2 and
and respiration increases. So, plants face a NO. Respiration and photorespiration
are sensitive to air pollutants. If the
shortage of organic substances.
concentration of air pollutants is high
it inhibits respiration whereas at lower
Apple, a temperate concentration stimulates respiration.
plant, when planted in Nitrogenous air pollutants under chronic
tropical condition fails exposure increases chlorophyll content
to produce fruits and while NO2 reduces pigment content at
growth is also affected. acute exposure.
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iv. Edaphic Stress the activity of certain enzymes; Increase in
They are divided into two types. They are abscisic acid level ultimately closes down the
water stress and salt stress: stomatal apparatus to the minimum, hence,
a. Water stress transpiration declines; Protochlorophyll
formation is inhibited and photosynthetic
A common stress condition arising from
process declines; Levels of proline increases;
lack of water or excess of water is called
water stress. The abundance of water Respiration and translocation of assimilates
leads to a stress called flood stress and decreases; Loss of water leads to increase in
scarcity of water leads to a stress called the activity of hydrolytic enzymes, followed by
drought stress. destruction of RNA and disruption of protein;
Wilting in mature leaves is associated with
I. Flood Stress carbohydrate depletion due to mobilization
The temporary inundation of plants export, followed by leaf senescence.
and its parts by flooding causes oxygen Mechanism of drought resistance
deficiency to the roots and soil borne Xerophytes are well adapted for drought
microorganisms. Effects of flooding are either because,
as follows: Nitrogen turnover in the soil i. the protoplasm of such plants does not
is reduced; Abscisic acid, ethylene and die when it faces extreme or prolonged
ethylene precursors are formed in larger desiccation (dehydration) hence, it
amount; Stimulation of partial stomatal tolerates or endures such conditions.
closure, epinasty and abscission in Example: Creosote bush (Larrea
leaves; Cellular membrane systems break tridentata) can survive water content
down, mitochondria and microbodies drops upto 30% whereas, in most
disintegrate and enzymes are partially plants the lethal level is below 50–70%
inhibited. Flood tolerant plants include or these plants are able to avoid or
those found on permanently wet soils. postpone the lethal level of desiccation
Examples: Marsh plants, shore plants and because they have developed structural
hydrophytes. Tree species found dominant or physiological adaptations. Plants
in flooded sites are also tolerant. Examples: that avoid or postpone desiccation have
Taxodium disticum, Mangroves and palms evolved an alternative path by developing
are tolerant to flood stress. following mechanisms: Improved water
II. Drought Stress uptake by roots which penetrate deep
down up to the water source; Efficient
The term ‘drought’ denotes a period without water conduction by increasing and
appreciable precipitation, during which enlarging the conductive tissues in
the water content of the soil is reduced to terms of producing more number of
such an extent that plants suffer from water xylem elements, dense leaf venation and
deficiency. Effects of drought are as follows: reducing the transport distance (short
Decrease in cellular growth and synthesis internodes); Restriction of transpiration
of cell wall components cause the cells to brought about by stomata present only
become smaller in size; Nitrogen fixation on the lower epidermis and covered
and its reduction are decreased by decreasing
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by dense trichomes; Rolling of leaves of toxic sodium carbonate and
also help to reduce water uptake by chloride ions.
minimizing the transpiring surface; On the basis of salt tolerance, they are
Water storage in succulent tissue of Agave grouped into two categories:
americana and other CAM plants have 1. Halophytes
been found to use water conservatively. 2. Non-halophytes or glycophytes
Halophytes are native to saline soils. The
Resurrection plants,
halophytes which can resist a range of salt
those plants having
concentration are called as euryhaline and
ability to survive near
those with narrow range of resistance are
total drying which
called stenohaline. Non-halophytes cannot
causes them to appear dead. They
resist salts as the halophytes. Helianthus
recover when water is available.
annus tolerates high Mn21 ions. Those which
Example: Selaginella lepidophylla
are present in salt regions face two problems:
During drought stress an essential • One is high concentration of salts in
protection mechanism that stabilizes the soil water leads to decrease in water
cell structure is induced gene expression potential so they grow in opposite
of stress protein (dehydrin and osmotin). direction. Example: Salicornia.
These proteins protect the macromolecules • Injuries in salt affected plants caused by
in the cytoplasm and in the nucleus, the both osmotic effects and specification
cytoskeleton (biomembranes) against effects. Accumulation of chloride
denaturation. High desiccation tolerance ions reduces water absorption and
transpiration.
implies that the protoplasm rehydrates when
water becomes available. Plants growing in Salt stress due to deficiency of mineral
deserts and arid regions are usually drought elements (K, P, S, Fe, Mo, Zn, Mg, Mn)
resistant. causes physiological disorders which lead
to reduced growth and yields.
b. Salt Stress
Presence of high salt concentration in the 1. Salt accumulators absorb and store
soil restricts the growth and development salts so that the osmotic potential
of their cells continues to remain
of plants. Most commonly the plants which
negative throughout the growing
are present near the seashore and estuaries
region.
are subjected to salt stress. According to
2. In some salt hardy plants, the excess
an estimate about one third of irrigated
salt is excluded on the surface of leaves.
land on earth is affected by salt stress.
Some plants have salt glands which
Na1, Cl2, K1, Ca11 and Mg11 ions usually
secrete salt (mostly NaCl). The exuded
contribute to soil salinity. Plants growing
salt absorbs water hygroscopically
in such areas face two problems: from the atmosphere.
1. Absorption of water from the soil
3. Some plants lose their excess salt by
with negative water potential
leaching into the soil or by dropping
2. Interaction with high concentration their salt filled leaves.
202

4. Salt tolerant plants (true halophytes) otherwise known as exponential phase. The
synthesize large amounts of the amino three phases are collectively called Grand
acid proline, galactosyl glycerol and period of growth. Plant exhibits plasticity in
some organic acids which function development. Plant growth and development
in osmotic adjustments. are controlled by both internal and external
Mechanism of salt tolerance factors. The internal factors are chemical
substances called Plant Growth Regulators
The plants growing in salty habitats like
(PGRs). The hormones are classified
halophytes face the problem of excessive
into five groups: Auxins, gibberellins,
dissolved salts in the solution. Excess of
cytokinins, abscisic acid and ethylene.
salt creates comparatively more negative
These PGRs are synthesized in various parts
osmotic potential so that the plants tend to
of the plant. PGRs may act synergistically
lose water into surrounding medium. Under
or antagonistically. The external factors
such conditions the plants tend to lose water
affecting growth includes water, nutrition,
only when their water potential becomes
temperature, oxygen and light. Mechanism
more negative. It is possible only if they
of flowering is controlled by light period
absorb excess of salt and accumulate it in
(photoperiodism) and temperature
their cell saps to maintain the same or higher
(vernalization). The physiological changes
concentrations as those of outside plants.
on flowering with effect from relative length
The drawbacks: of light and darkness (photoperiodism) are
1. Salt accumulates in the vacuoles called photoperiodism. A bluish biliprotein
2. The plants become succulents responsible for the perception of light in
3. Accumulated salt dehydrates the photophysiological process (induction
cytoplasm and inhibition of flowering) is called
4. Sodium chloride cannot be Phytochrome. Besides photoperiod certain
tolerated in the cytoplasm and it plants require a low temperature in the
denatures several enzymes earlier stages for flowering. Many biennial
Thus, absorption and accumulation of and perennial plants are induced to flower by
inorganic salts fail to solve the problem. The low temperature (0oC to 5oC). This process is
plants however tolerate the salt stress by called vernalization and the reversal effect
synthesizing organic compounds that can exist of vernalization is called devernalization.
at high salt concentrations without denaturing The condition of a seed when it fails to
the enzymes. These organic compounds are germinate even in suitable environmental
called nontoxic organic osmotica. Examples: condition is called seed dormancy. Thus,
Proline and Betalin (osmoregulators). dormancy can be overcome by following
methods such as scarification, impaction,
Summary stratification, alternating temperatures and
Growth occurs by cell division, cell elongation light. Senescence refers to all collective,
and cell maturation. The first phase is lag progressive and deteriorative processes
phase, the second is log phase and the final which ultimately lead to complete loss of
phase is steady state phase. The log phase is organization and function. Senescence is of
four types and they are overall, top, deciduous
203

and progressive. Senescence is controlled by c. ABA d. Auxin
plant’s own genetic programme. Death of the 4. Select the correctly matched one
plant or its parts consequent to senescence A) Human urine i) Auxin –B
is called Programmed Cell Death (PCD). B) Corn gram oil ii) GA3
The final stage of senescence is abscission. C) Fungus iii) Abscisic acid II
Abscission is a physiological process of D) Herring fish iv) Kinitin
shedding of organs from the parent plant sperm
body. The study of functioning of plants E) Unripe maize v) Auxin A
under adverse environmental conditions is grains
called stress physiology. The environmental F) Young cotton vi) Zeatin
stress may broadly be divided into biotic and bolls
abiotic stress. The adverse effect on plants a) A-iii, B-iv, C-v, D-vi, E-i, F-ii,
caused by other living organisms such as
b) A-v, B-i, C-ii, D-iv, E-vi, F-iii,
viruses, bacteria, fungi, parasites, insects,
c) A-iii, B-v, C-vi, D-i, E-ii, F-iv,
weeds is competing plants are called biotic
d) A-ii, B-iii, C-v, D-vi, E-iv, F-i
stress. Abiotic stress may occur due to an
atmospheric condition or soil condition. 5. Seed dormancy allows the plants to
a. overcome unfavourable climatic
conditions
Evaluation
b. develop healthy seeds
1. Select the wrong
c. reduce viability
statement from the
following: d. prevent deterioration of seeds
a. Formative phase of 6. Which one of the following method are
the cells retain the used to break the seed dormancy?
capability of cell division. a) Scarification b) Impaction
b. In elongation phase development of c) Stratification d) All the above.
central vacuole takes place. 7. What are the parameters used to
c. In maturation phase thickening and measure growth of plants?
differentiation takes place. 8. What is plasticity?
d. In maturation phase, the cells grow 9. Write the physiological effects of
further. Cytokinins.
2. If the diameter of the pulley is 6 inches, 10. Describe the mechanism of
length of pointer is 10 inches and photoperiodic induction of
distance travelled by pointer is flowering.
5 inches. Calculate the actual growth 11. Give a brief account on Programmed
in length of plant. Cell Death (PCD)
a. 3inches b. 6 inches 12. What are the physiological effects of
c. 12 inches d. 30 inches plants facing drought condition?
3. In unisexual plants, sex can be 13. Explain the mechanism of biotic
changed by the application of stress.
a. Ethanol b. Cytokinins
204

t ICT Corner
How do Plants respond to different stimuli?
Let’s Stimulate the Plants.
Steps
• Click Exploring plant responses
• Select items and complete the check list
• Follow the procedure – 1 to 10 steps
• Record your prediction and not your observation in lab note – Right top
Activity
• Observe the movements of plant seedlings and plant parts.
Step 1 Step 2
Step 3 Step 4
Web URL:
https://www.classzone.com/books/hs/ca/sc/bio_07/virtual_labs/virtualLabs.html
205

References
Unit – 4 Plant Anatomy
1. Fahn.A, (1990), Plant Anatomy, 3rd edition, Oxford; New York; Pergamon Press
2. Gangulee,Das& Data, (2011) College Botany,Vol-II, New Central Bool Agency
3. Katherine Esau, (2006), Anatomy of Seed Plants, 2nd Edition, John Wiley & Sons, Inc.
4. PandeyB.P, (2015), A Textbook of Botany: Angiosperms, New Delhi, S. Chand & Company Ltd.
5. Pijush Roy, (2012), Plant Anatomy, New Central Book Agency (P) Ltd.
6. Ray.F.Evert, (2007), Esau’s Plant Anatomy, 3rd Edition. Wiley-Liss
Unit – 5 Plant Physiology

1. Campbell and Reece (2005) Biology Vol I, 7th Edition, Boston, Pearson,.
2. Clegg C J (2014) Biology, London, Hooder Education,.
3. Data.S.C (1990) Plant Physiology, New Delhi,Willey Eastern.
4. Devlin, R. M. (2017). Outline of Plant Physiology. Medtech Pubs.
5. Dey P.M & Harborne J.B (1997) Plant Bio chemistry, London, Academic press
6. Dey, P. M. and Harborne, J. B. (2013). Plant Bio chemistry. Elsevier.
7. Helgiopik and Stephan Rolfe (2005) The Physiology of Flowering Plants, 4th Edition, London,
Cambridge University Press.
8. Jain V.K. (2017) Fundamentals of Plant Physiology, 19th Edition, New Delhi, S.Chand & Co.
9. Jain. J L., Sunjay Jain and Nitin Jain. (2005). Fundamentals of Biochemistry, 6th Edition. New
Delhi S. Chand and Co.,.
10. Jane B Reece etal. (2011) Campbell Biology, 10th Edition, Pearson.
11. K.N.Rao, G. Sudhakara Rao, S. Bharatan (1987) The functioning plant, S. Viswanathan Pvt.Ltd.
12. Kumar.A & Purohit S.S (2002) Plant Physiology: Fundamentals and Applications, 2nd Edition,
Agro-Bios.
13. Leninger, Nelson and Cox. (2017). Principles of Biochemistry, 7th Edition. NewDelhi, Macmillan
Learning.
14. Maria Duca (2015) Plant Physiology, Switzerland, Springer international publishing house.
15. Mukherji, S. and Ghosh, A. K. (2015). Plant Physiology. London, New Central Book Agency Pvt.
Ltd.,
16. Noggle, G. R. and Fritz, G. J. (1983). Introductory Plant Physiology, Second edition. Prentice Hall
India.
17. R.K.Sinha (2004) Modern plant Physiology, Alpha Publishing
18. Salisbury, F. and Ross, C. (1991). Plant Physiology, 4th Edition. India, Thomson Publications.
19. Sinha, R. K. (2003). Modern Plant Physiology, 2nd Ed. Kolkata,Narosa Publishing House.
20. Srivastava H.N (2004) Plant Physiology, Pradeep publication, Jalandhar.
21. Stern, Jansky, Bidlack (2003) Introductory Plant Biology, 9th Edition, New York, McGraw Hill,.
22. SundaraRajan.S (2000) Plant Physiology, New Delhi, Anmol Publication,.
23. Taiz.L and Zeigar.E (2010) Plant Physiology, 3rd Edition, Sunderland, Sinauer Associates,
24. Taiz, L., Zeiger, E., Moller, I. M. and Murphy, A. (2014). Plant Physiology and Development,
Sixth Edition. Ingram International Inc.
25. Verma S.K and MohitVerma, (2016) A Text Book of Plant Physiology, Biochemistry and
Biotechnology, New Delhi, S.Chand& Co,.
26 Walter Larcher, (2003). Physiological Plant Ecology, 4th Edition. New York, Springer International
Edition,.
206

Glossary
Abscission zone A region near the base of petiole of leaf which contains
abscission layer.
Absorption Spectrum A curve obtained by plotting the amount of absorption
of different wavelengths of light by a pigment is called its
absorption spectrum.
Action Spectrum A graphic representation showing the rate of photosynthesis
at different wavelengths of light is called action spectrum
Aeroponics A technique of growing plants suspended over the nutrient
solution in a mist chamber. Nutrient sprayed by motor
driven rotor on the roots.
Agar Jelly-like substance, derived from red algae
Allelopathy The chemical substances released by one plant species which
affect or benefit another plant
Amphicribal/ Xylem in the centre with phloem surrounding it. Example:
Hadrocentric Ferns ( Polypodium)
Amphivasal /Leptocentric Phloem in the centre with xylem surrounding it. Example:
Dragon plant – Dracena and Yucca
Anabolic It is an enzyme catalyzed reaction in a cell that involves
synthesis of complex molecules from simple molecules
which uses energy.
Apical cell theory Single apical cell growing into whole plant
Axil Parenchyma Parenchyma arranged longitudinally along the axis
Callose Sieve pores are blocked by substances called callose
Carbonic acid A weak acidic solution of carbon-di-oxide dissolved in
water
Catabolic It is an enzyme catalyzed reaction in a cell that involves
degradation of molecules into simple subunits which
release energy.
Chelating agents A chelate is the soluble product formed when certain atoms
in an organic ligand donate electrons to the cation.
Chlorosis Breakdown of chlorophylls leads to yellowing of leaves
Closed vascular bundle Cambium absent between xylem and phloem Example:
Monocot stem
Coenzyme A non-protein molecule involved in enzyme catalyzed
reactions serves as transfer of protons or electrons between
various molecules
Colloidal An evenly distributed mixture of two different particles in
a system without losing its own properties.
Deamination The enzymatic removal of an amino group from an amino
acid to form its corresponding keto acid.
Desiccation tolerance Ability of plants which can tolerate extreme water stress
without being killed.
Drought resistance Capacity of a plant to limit and control consequences of
water deficit.
207

EDTA Ethylene Diamine Tetra Acetic acid, chelating agent makes
iron uptake possible by forming soluble complex in an
alkaline soil.
Endergonic A chemical reaction with a positive free energy charge or
ATP utilizing reactions.
Exergonic A chemical reaction with a negative free energy charge or
ATP producing reactions.
Extra stellar ground tissue Tissues outside the stele
Fibre-Tracheids Transitional form between fibre and tracheids
Fluorescence Emission of light by a substance that has absorbed light in
the form luminescence.
Gelatin An animal-based product used as a gelling agent.
Granum A stack of thylakoid in a stroma of chloroplast
Hadrome Xylem-by Haberlandt
Halophytes Plants native to saline soils and complete their life cycle
Heliophytes Plants which are adapted to light
Histogenesis Differentiate tissues from undifferentiated cells of meristem
Indeterminate growth Plants grow throughout their life
Intrastelar ground tissue Tissues within the stele
Isomerisation Rearrangement of atomic groups within the same molecule
without any loss or gain of atoms.
Leptome Phloem – by Haberlandt
Lumen Space inside the tracheid/vessel/fibres
Malate Shuttle mechanism It is a biochemical system for translocating electrons
produced from glycolysis across inner membrane of
mitochondrion for oxidative phosphorylation.
Mass meristem Meristem which divides in all planes
Necrosis Death of tissue
Non heme iron An iron porphyrin prosthetic group of heme proteins from
plant origin
Nutation The growing stems of twiner and tendrils show automatic
movement
Open vascular bundle Cambium present between xylem and phloem Example:
Dicot stem
Oxidation Water is oxidised into Oxygen (loss of electrons)
PAR The wavelength at which the rate of photosynthesis is more
is called ‘Photosynthetically Active Radiations’ which falls
between 400 to 700 nm.
Phosphorescence Phosphorescence is the delayed emission of absorbed
radiations.
Photolysis Splitting of water molecules by light which generate
protons, electrons and oxygen.
Photon Light is electromagnetic radiant energy and travels as tiny
particles called photons. A discrete Physical unit of light energy.
208

Photoperiodism The response of plants to the photoperiod expressed in the
form of flowering.
Phytochrome A photo reversible proteinaceous plant pigment in very
low concentration that absorbs red and far red light which
controls flowering.
Pitted thickening Uniformly thick except at their pits
Preparatory phase First half of glycolysis comprising five enzymatic reactions
in which one molecule of glucose splitting into two
molecules of glyceraldehyde 3 phosphate with consumption
of two ATP molecules.
Prickles Stiff and sharp outgrowth
Quantasome Morphological expression of physiological photosynthetic
units, located on the inner membrane of thylakoid lamellae.
Act as photosynthetic unit contains 200 to 300 chlorophyll
molecules.
Quantum The energy contained in a photon is represented as quantum
Quantum requirement The number of photons or quanta required to release one
molecule of oxygen during photosynthesis
Quantum yield The number of oxygen molecules produced per quantum
of light absorbed.
Quiescent centre concept Inactive region of root meristem
Radial vascular bundles Xylem and phloem present on different radii
Ray Parenchyma Parenchyma cells arranged in radial rows
Redox reactions Oxidation (loss of electrons) and Reduction (gain of
electrons) reactions are called redox reactions.
Reduction CO2 is reduced into Carbohydrates (gain of electrons)
Rib-meristem Meristem which divides anticlinally in two planes
RUBISCO Enzyme responsible for fixation of Carbon dioxide, the most
abundant protein (Ribulose 1,5 bisphosphate Carboxylase
Oxygenase)
Salt stress Adverse effects of excess mineral salts on plants
Sap It is a fluid consist of water and dissolved minerals
Slime body A special protein (Phloem Protein) in sieve tubes
Stellate hairs Star shaped hairs
Stratification A process of breaking the dormancy of some plants
resulting from chilling requirements
Subsidiary cells Surrounding guard cells in the leaf epidermis
Sucrose Non-reducing disaccharide composed of glucose and
fructose
Trichoblasts One type of epidermal cells that is also called short cell
Trichomes Unicellular or multicellular appendages
Tunica-carpus theory Two zones of apical meristem Tunica and Carpus
Xylos Wood
209

English – Tamil Terminology
Abscission உதிர்தல்
Abscission zone உதிரும் அடுக்கு
Absorption spectrum ஒளி ஈர்ப்பு நிறமாலை
Action spectrum ஒளி செயல்திறன் நிறமாலை
Activated diffusion மேம்படுத்தப்பட்ட பரவல்
Active transport ஆற்றல்சார் கடத்தல்
Adhesion ஒட்டிணைவு
Aeroponics காற்றூடக வளர்ப்பு
Anabolic சேர்க்கைச் செயல்
Annual rings ஆண்டு வளையங்கள்
Antenna molecules ஏற்பி மூலக்கூறுகள்
Apical cell theory நுனி செல் க�ொள்கை
Arithmetic growth எண் கணித வளர்ச்சி
Ascent of sap சாறேற்றம்
Assimilatory power தன்மயமாக்கும் ஆற்றல்
Autonomous movement தன்னிச்சையான அசைவுகள்
Autumn wood or late wood குளிர்க்காலக் கட்டை அல்லது பின்பருவக் கட்டை
Axial parenchyma அச்சு பாரங்கைமா
Bicollateral vascular bundle இருபக்க ஒருங்கமைந்த வாஸ்குலக் கற்றை
Biosynthetic phase உயிர்மத�ோற்ற நிலை
Biosequestration உயிர்வளி தனிமைப்படுத்துதல்
Brown heart disease மைய கருக்கல் ந�ோய்
Callus திசுத்திரள்
Carbon fixation கார்பன் நிலைநிறுத்தம்
Carbon di oxide compensation point கார்பன்-டை-ஆக்ஸைட் ஈடு செய்யும் புள்ளி
Carrier protein தாங்கிப் புரதம்/கொண்டு செல்லும் புரதம்
Catabolic சிதைக்கும் செயல்
Catalytic amination வினையூக்க அமைன�ோவாக்கம்
Cavitation குமிழாதல்
Channel protein கால்வாய் புரதம்
Chelating agents பிணைக்கும் காரணி
Chemiosmotic theory வேதி சவ்வூடு பரவல் க�ோட்பாடு
Chlorophyll பச்சையம்
Chloroplast பசுங்கணிகம்
Chlorosis பச்சைய ச�ோகை
Closed collateral vascular bundles மூடிய ஒருங்கமைந்த வாஸ்குலக் கற்றைகள்
Cohesion கூட்டிணைவு
Collateral vascular bundles ஒருங்கமைந்த வாஸ்குலக் கற்றைகள்
Companion cells துணைச் செல்கள்
Compensation point ஈடுசெய்யும் புள்ளி
210

Concentration gradient செறிவு சரிவு வாட்டம்
Concentric vascular bundles சூழமைந்த வாஸ்குலக் கற்றைகள்
Core complex மைய ஆதார கூட்டமைப்பு
Critical concentration தீர்வுக் கட்ட செறிவு
Day neutral plants நாள் நடுநிலை தாவரங்கள்
Deamination அமின�ோ நீக்கம்
Dendrochronology மர வயதியல்
Deplasmolysis பிளாஸ்மா சிதைவு மீட்சி
Dicarboxylic acid pathway டைகார்பாக்சிலிக் அமில சுழற்சி
Die back of shoot தண்டின் நுனி அடி இறப்பு
Diffusion பரவல்
Dimorphic chloroplast இருவடிவ பசுங்கணிகம்
Drought resistance வறட்சியை எதிர்ப்பவை
Efflux அயனி வெளிப்புகல்
Electro magnetic spectrum மின்காந்த நிறமாலை
Electron transport chain எலக்ட்ரான் கடத்து சங்கிலி
Emerson’s enhancement effect எமர்சனுடைய மேம்படுத்தப்பட்ட விளைவு
Endergonic ஆற்றல் ஏற்கும் வினை
Endosymbiotic hypothesis அக கூட்டுயிர் க�ோட்பாடு
Eutrophication மிகை ஊட்ட நிலை
Exarch Xylem வெளிந�ோக்கு சைலம்
Exergonic ஆற்றல் வெளியிடும் வினை
Extinction point அழிவுப் புள்ளி
Fermentation ந�ொதித்தல்
Fibre Tracheids நார் டிரக்கீடுகள்
Flourescence உடன் ஒளிர்தல்
Flux அயனிபுகல்
Geometric growth ஜிய�ோமித வளர்ச்சி
Grand period of growth ம�ொத்த வளர்ச்சிக் காலம்
Growth rate பெரும வளர்ச்சி வீதம்
Halophiles உவர்நாட்டவுயிரிகள்
Halophytes உவர்நிலை தாவரங்கள்
Heart wood வைரக்கட்டை
Heliophytes ஒளியை விரும்பும் தாவரங்கள்
Histogen theory ஹிஸ்டோஜன் க�ொள்கை
Histogenesis ஹிஸ்டோஜெனிசிஸ்
HMP shunt HMP மாற்றுவழிப் பாதை
Hydathode நீர்கசிவுத் துளை
Hydroponics நீர் ஊடக வளர்ப்பு
Imbibition உள்ளீர்த்தல்
Influx அயனி உட்புகல்
211

Interveinal chlorosis நரம்பிடை பச்சைய ச�ோகை
Isomerisation மாற்றியமாதல்
Lag phase உருவாக்க நிலை
Lenticel பட்டைத்துளை
Light harvesting complex ஒளி அறுவடை கூட்டமைப்பு
Link reaction இணைப்பு வினை
Log phase நீட்சியுறு நிலை
Macro nutrients பெரும ஊட்ட மூலங்கள்
Malate Shuttle mechanism மாலேட் திருப்பு செயல்
Mass meristem ப�ொருண்மை ஆக்குதிசு
Matric potential ஊடக உட்திறன்
Micro nutrients நுண் ஊட்ட மூலங்கள்
Mineral Nutrition கனிம ஊட்டம்
Mitochondrial matrix மைட்டோகாண்ட்ரிய உட்கூழ்மம்
Necrosis நைவுப் புண்கள்
Nitrate Assimilation நைட்ரேட் தன்மயமாதல்
Nitrogen metabolism நைட்ரஜன் வளர்சிதை மாற்றம்
Non-porous wood துளைகளற்ற கட்டை
Nutation சுழலசைவு
Obligate parasite கட்டாய ஒட்டுண்ணி
Open vascular bundle திறந்த வாஸ்குலக் கற்றை
Oxygen evolving complex (OEC) ஆக்ஸிஜன் உருவாக்கும் கூட்டமைப்பு
Paper chromatography வண்ண பிரிகைதாள் வரைப்படம்
Paratonic movement தூண்டப்படும் அசைவுகள்
Parthenocarpy விதையிலாக் கனி
Passive transport ஆற்றல்சாரா கடத்தல்
Pay off phase விளை நிலை
Phosphorescence நின்றொளிர்தல் / தாமத மறு ஒளிர்தல்
Photo chemical phase ஒளி வேதிநிலை
Photo oxidation phase ஒளி ஆக்ஸிஜனேற்ற நிலை
Photo respiration ஒளி சுவாசம்
Photolysis ஒளியின் நீராற் பகுப்பு
Photon ஒளித்துகள்
Photoperiodic induction ஒளிக் காலத்துவ தூண்டுதல்
Photoperiodism ஒளிக்காலத்துவம்
Photophosphorylation ஒளி பாஸ்பரிகரணம் / ஒளி பாஸ்பரஸ் சேர்க்கை
Photosynthetic carbon reduction cycle ஒளிச்சேர்கையின் கார்பன் ஒடுக்க சுழற்சி
Photosynthetic unit (Quantasome) ஒளிச்சேர்க்கை அலகு (குவாண்டோச�ோம்)
Photosystem நிறமி அமைப்பு / ஒளி அமைப்பு
Plant antitranspirants நீராவிப்போக்குத் தடுப்பான்கள்
Plasmolysis பிளாஸ்மா சிதைவு
212

Plasticity உருமாறும் தன்மை
Porous woods துளைக்கட்டை
Preparatory phase ஆயத்த நிலை
Pressure potential அழுத்தயியல் திறன்
Primary growth முதல்நிலை வளர்ச்சி
Programmed cell death திட்டமிடப்பட்ட செல் இறப்பு
Proton gradient புர�ோட்டான் சரிவு
Pumps உந்திகள்
Quiescent centre concept உறக்க மையக் க�ொள்கை
Radial vascular bundles ஆரப்போக்கமைந்த வாஸ்குலக் கற்றைகள்
Ray parenchyma கதிர் பாரங்கைமா
Reaction Centre வினை மையம்
Red drop சிவப்பு வீழ்ச்சி
Redox reaction ஆக்ஸிஜனேற்ற ஒடுக்கவினை
Reducing power ஒடுக்கும் ஆற்றல்
Respiratory quotient சுவாச ஈவு
Reverse osmosis பின்னோக்கிய சவ்வூடு பரவல்
Rib meristem வரிசை ஆக்குத்திசு
Ring Bark வளைய பட்டை
Sap wood சாற்றுக்கட்டை
Scale Bark செதில் பட்டை
Seed dormancy விதை உறக்கம்
Semi autonomy பாதி சுயசார்புதன்மை
Senescence மூப்படைதல்
Sink தேங்கிடம்
Slime bodies ஸ்லைம் உடலங்கள்
Solute potential கரைப�ொருள் திறன்
Source த�ோற்றுவாய்
Spring wood or early wood வசந்தக்காலக் கட்டை அல்லது முன்பருவக் கட்டை
Stress escapers நெருக்கடியை தப்பித்துக் க�ொள்ளும் தாவரங்கள்
Stress physiology நெருக்கடி சார் வாழ்வியல்
Substrate phosphorylation தளப்பொருள் பாஸ்பரிகணம்
Sunken stomata உட்குழிந்த இலைத்துளை
Terminal oxidation இறுதி ஆக்ஸிஜனேற்றம்
Thermonastic வெப்ப தூண்டல்
Thigmotactic த�ொடு உணர்வு அசைவு
Transamination அமைன�ோ மாற்றம்
Tunica corpus theory டூனிகா கார்பஸ் க�ொள்கை
Vernalization தட்பப்பதனம்
Water potential நீரியல் திறன்
Xeric Succession வறள் தாவர படிநிலை வளர்ச்சி
213

Competitive Exam Questions
Unit -4 – Plant Anatomy 6. The annular and spirally thickened

conducting elements generally develop in
1. The balloon – shaped structures called the protoxylem when the root or stem is
tyloses (NEET II – 2016 ) (CBSE -AIPMT 2009)
a. originate in the lumen of vessels a. maturing b. elongating
b. characterise the sap wood c. widening d. differentiating
c. are extensions of xylem parenchyma
cells into vessels 7. Anatomically fairly old dicotyledonous root
d. are linked to the ascent of sap through is distinguished from the dicotyledonous
xylem vessels stem by the (CBSE- AIPMT 2009)
a. absence of secondary xylem
2. Cortex is the region found between (NEET b. absence of secondary phloem
II – 2016) c. presence of cortex
a. epidermis and stele d. position of protoxylem
b. pericycle and endodermis
c. endodermis and pith 8. In barley stem, vascular bundles are (CBSE
d. endodermis and vascular bundle -AIPMT 2009)
a. open and scattered
3. Read I – IV and find the correct order of b. closed and scattered
components from outer side to inner side c. open and in a ring
in a woody dicot stem (CBSE -AIPMT – d. closed and radial
2015)
(I) secondary Cortex (II) wood 9. Palisade parenchyma is absent in the leaves
(III) secondary phloem (IV) phellem of (CBSE- AIPMT 2009)
a. III, IV, II and I b. I, II, IV and III a. sorghum b. mustard
c. IV, I, III and II d. IV, III, I and II c. soyabean d. gram
4. You are given a fairly old piece of a dicot 10. Sugarcane plant has (AIIMS 2009)
stem and a dicot root. Which of the a. reticulate venation
following anatomical structures will you b. capsular fruits
use to distinguish between the two? (CBSE c. pentamerous flowers
-AIPMT 2014) d. dump-bell shaped guard cells
a. secondary xylem
11. Vascular tissues in flowering plants develop
b. secondary phloem
from (CBSE- AIPMT 2008 & JIPMER
c. protoxylem
2012)
d. cortical cells
a. phellogen b. plerome
5. Heart wood differs from sapwood in (CBSE
c. periblem d. dermatogen
-AIPMT 2010)
a. the presence of rays and fibres 12. The length of different internodes in a culm
b. the absence of vessels and parenchyma of sugarcane is variable because of (CBSE
c. having dead and non-conducting -AIPMT 2008)
elements a. short apical meristem
d. being susceptible to hosts and pathogens b. position of axillary buds
214

c. size of leaf lamina at the node below each characterized by (CBSE -AIPMT 2003)
internode a. having dense cytoplasm and
d. intercalary meristems prominent nucleus
b. having light cytoplasm and small
13. Passage cells are thin-walled cells found in nucleus
(CBSE -AIPMT 2007) c. dividing regularly to add to the corpus
a. endodermis of roots facilitating rapid d. dividing regularly to add to tunica
transport of water from cortex to
pericycle 18. P. Protein is found in (CBSE- AIPMT 2000)
b. phloem elements that serve as entry a. parenchyma b. collenchyma
points for substances for transport to c. sieve tube d. xylem
other plant parts
19. Specialized epidermal cells surrounding the
c. testa of seeds to enable emergence of
guard cells are called (NEET (I) 2016)
growing embryonic axis during seed
a. bulliform cells
germination
b. lenticels
d. central region of style through which
c. complementary cells
the pollen tube grows towards the ovary
d. subsidiary cells
14. Which one of the following is not a lateral
Directions:
meristem (CBSE -AIPMT 2010)
The following questions 20 & 21 consist of two
a. interfascicular cambium
statements, one labelled Assertion and the
b. phellogen
another labelled Reason. Select the correct
c. intercalary meristem
answer from the codes given below:
d. intrafascicular cambium
a) Both assertion and reason are true and
15. A common feature of vessel elements and reason is the correct explanation of assertion
sieve tube elements is (CBSE- AIPMT 2007) b) Both assertion and reason are true, but
a. enucleate condition reason is not the correct explanation of
b. presence of P. Protein assertion
c. thick secondary wall c) Assertion is true but reason is false
d. pores on lateral walls d) Assertion and reason are false
16. In a longitudinal section of a root, starting 20. Assertion: Conducting tissues, especially
from the tip upward, the four zones occur xylem show greatest reduction in submerged
in the following order (CBSE -AIPMT hydrophytes.
2004)
Reason: Hydrophytes live in water. So no
a. root cap, cell division, cell enlargement,
need of tissues. (AIIMS – 2010)
cell maturation
Ans: c.
b. root cap, cell division, cell maturation,
21. Assertion: Long distance flow of photo
cell enlargement
assimilates in plants occurs through sieve
c. cell division, cell enlargement, cell
tubes.
maturation, root cap
Reason: Mature sieve tubes have partial
d. cell division, cell maturation, cell
cytoplasm and perforated sieve plates
enlargement, root cap
(AIIMS – 2012)
17. The cells of the quiescent centre are
Ans: a.
215

22. Duramen is present in (JIPMER 2016) a. phelloderm b.primary phloem
a. the inner region of secondary wood c. secondary xylem d. periderm
b. a part of sap wood
30. Which of the following plants shows multiple
c. the outer region of secondary wood
epidermis? (Manipal 2012)
d. region of pericycle
a. Croton b. Allium
23. The interxylary phloem is found in the stem c. Nerium d. Cucurbita
of (JIPMER 2013)
a. Cucurbita b. Salvia UNIT -5 PLANT PHYSIOLOGY
c. Calotropis d. none of these
1. The water potential of pure water is (NEET
24. Wound healing is due to (JIPMER 2013) 2017)
a. ventral meristem a. Less than zero
b. secondary meristem b. More than zero but less than one
c. primary meristem c. More than one
d. all of these d. Zero
25. Which of the following tissues consists of 2. Transpiration and root pressure cause
living cells (JIPMER 2012) water to rise in plants by (NEET 2015)
a. vessels b. tracheids a. pulling it upward
c. companion cell d. sclerenchyma b. pulling and pushing it, respectively
c. pushing it upward
26. The Quiescent centre in root meristem
d. pushing and pulling it, respectively
serves as a (JIPMER 2011)
a. site for storage of food, which is utilized 3. Movement of ions or molecules in a
during maturation direction opposite to that of prevailing
b. reservoir of growth hormones electro-chemical gradient is known as
c. reserve for replenishment of damaged (C.B.S.E. 2000)
cells of the meristem a. Active transport
d. region for absorption of water b. Pinocytosis
27. In the sieve elements, which one of the c. Brownian movement
following is the most likely function of d. Diffusion
P.Proteins? (JIPMER 2011)
4. Correct sequence of events in wilting?
a) Deposition of callose on sieve plates
(P.M.T. Kerala 2001)
b. Providing energy for active translocation
a. Exosmosis-deplasmolysis-temporary
c. Autolytic enzymes
and permanent wilting
d. Sealing-off mechanism on wounding
b. Exosmosis-plasmolysis-temporary
28 .Which of the following is made up of dead and permanent wilting
cells? (NEET 2017) c. Endosmosis-plasmolysis-temporary
a. Xylem parenchyma b. Collenchyma and permanent wilting
c. Phellem d. Phloem d. Endosmosis-deplasmolysis - temporary
29. The vascular cambium normally gives rise to and permanent wilting
(NEET 2017) e. Exosmosis-deplasmolysis-plasmolysis -
temporary and permanent wilting
216

5. What will be the direction of net osmotic b. Water plus minerals
movement of water if a solution 'A', c. Water plus enzymes
enclosed in a semi permeable membrane, d. All of these
having an osmotic potential of '- 30' bars
and turgor pressure of '5' bars is submerged 12. Stomata of a plant open due to (CBSE 2003)
in a solution 'B' with an osmotic potential of a. Influx of potassium ions
'- 10' bars and '0' turgor pressure ? (C.E.T. b. Efflux of potassium ions
Karnataka 2002) c. Influx of hydrogen ions
a. Equal movement in both directions d. Influx of calcium ions
b. 'B' to 'A'
13. Potometer works on the principle of
c. No movement
(CBSE 2000)
d. 'A' to 'B'
a. Osmotic pressure
6. The pressure exerted by a swollen vacuole b. Amount of water absorbed equals the
on the cell wall is (C.M.C. Vellore 2002) amount transpired
a. OP b. WP c. Potential difference between the tip of
c. TP d. DPD the tube and then of the plant
d. Root pressure
7. Who said that ‘transpiration is a necessary
evil’? (JIPMER-2006) 14. Most suitable theory for ascent of sap is
a. Curtis b. Steward (CBSE 1991, CPMT-UP 1995)
c. Anderson d. J.C.Bose a. Transpirational pull and cohesion
theory of Dixon and Jolly
8. Which one gives the most valid and recent b. Pulsation theory of J.C. Bose
explanation for stomatal movements? c. Relay pump theory of Godlewski
(NEET 2015) d. None of these
a. Transpiration
15. If a cell kept in a solution of unknown
b. Potassium influx and efflux
concentration gets deplasmolysed, the
c. Starch hydrolysis
solution is, (CPMT-UP 1996)
d. Guard cell photosynthesis a. Detonic b. Hypertonic
9. Carrier proteins are involved in ( P M T - c. Isotonic d. Hypotonic
UP-1998) 16. Which is essential for the growth of root tip
a. Active transport of ions ? (NEET PHASE II 2016)
b. Passive transport of ions a. Zn b. Fe
c. Water transport c. Ca d. Mn
d. Water evaporation 17. On the basis of symptoms of chlorosis in
leaves, a student inferred that this was due to
10. Active transport of ions in the cell requires
deficiency of nitrogen. The inference could
(PMT MP 2002)
be correct only if we assume that yellowing
a. High temperature b. ATP
of leaves appeared first in (AIIMS 2007)
c. Alkaline pH d. Salts
a. old leaves b. young leaves
11. Guttated liquid is (AFMC 2002) c. young leaves followed by mature leaves
a. Pure water d. mature leaves followed by young leaves.
217

18. Cytochrome oxidase contains (UP CPMT atmospheric nitrogen in leguminous plants
2006) is _____ (AIPMT 2013)
a. Iron b. Magnesium a. NO-3 b. glutamate
c. Zinc d. Copper c. NO-2 d. ammonia
19.
Which is correct to saprophytic 26. C4 plants are more efficient in photosynthesis
angiosperms? (UP CPMT 2006) than C3 plants due to (AIPMT 2010)
a. They secrete enzyme outside the body a. presence of thin cuticle
and absorb b. lower rate of photorespiration
b. They have mycorrhizae fungi c. higher leaf area
c. They take food and then digest it d. presence of larger number of chloroplast
d. They are photosynthetic in the leaf cells.
20. The ability of the venus fly trap to capture 27. Chlorophyll b is (JIPMER 1980)
insects is due to (JIPMER 2008) a. C54H70 O6 N4 Mg
a. chemical stimulation by the prey b. C55H70 O6 N4 Mg
b. a passive process requiring no special c. C55H72 O5 N4 Mg
ability on the part of the plant. d. C45H72 O5 N4 Mg
c. Specialized muscle like cells
d. rapid turgor pressure changes 28. Synthesis of ADP + Pi → ATP in grana is
(AIIMS 1993)
21. Boron in green plants assists in (RPMT a. phosphorylation
2007) b. photophosphorylation
a. photosynthesis c. oxidative phosphorylation
b. Sugar transport d. photolysis
c. activation of enzyme
d. acting as enzyme cofactor 29. In chloroplast, chlorophyll is present in the
(AIPMT 2004)
22. Which of the following elements is very a. stroma
essential for the uptake of Ca2+ and b. outer membrane
membrane function? (Kerala CEE 2007) c. inner membrane
a. phosphorus b. molybdenum d. thylakoids
c. manganese d. boron
30. Electrons from the excited chlorophyll
23. Sulphur is not a constituent of (AMU 2011) molecule of photosystem II are accepted
a. cysteine b. methionine first by (AIPMT 2008)
c. ferredoxin d. pyridoxine a. quinone b. ferredoxin
c. cytochrome-b d. cytochrome-f
24.
Deficiency symptoms of nitrogen and
potassium are visible first in _____ (AIPMT 31. Read the following four statements A,B,C
2014) and D. Select the right option (AIPMT 2010)
a. senescent leaves b. young leaves A. Z scheme of light reaction takes place in
c. roots d. buds the presence of PS I only
B. only PS I is functional in cyclic
25. The first stable product of fixation of
photophosphorylation
218

C. cyclic photophosphorylation results into improved efficiency of nitrogen utilization.
synthesis of ATP and NADPH2 In which of the following physiological
D. stroma lamellae lack PS II as well as groups would you assign this plant? (NEET
NADP PHASE I 2016)
a. A and B b. B and C a. C4 b. CAM
c. C and D d. B and D c. Nitrogen fixer d. C3
32. Photolysis of each water molecule in light 37. Emerson's enhancement effect and Red
reaction will yield ___ (Kerala CEE 2007) drop have been instrumental in the
a. 2 electrons and 4 protons discovery of (NEET PHASE I 2016)
b. 4 electrons and 4 protons a.
two photosystems operating
simultaneously
c. 4 electrons and 3 protons
b. photophosphorylation and cyclic
d. 2 electrons and 2 protons
electron transport
33. Photosynthetic active radiation (PAR) has c. oxidative phosphorylation
the following range of wavelength (AIPMT d. photophosphorylation and non-cyclic
2005) electron transport
a. 400-700 nm b. 450-920 nm
c. 340-450 nm d. 500-600 nm 38. The process which makes major difference
between C3 and C4 plants is (NEET PHASE
34. Phosphoenol pyruvate (PEP) is the primary II 2016)
CO2 acceptor in __ (NEET 2017) a. glycolysis b. calvin cycle
a. C3 plants b. C4 plants c. photorespiration d. respiration
c. C2 plants d. C3 and C4 plants
39. In a chloroplast the highest number of
35. With reference to factors affecting the rate protons are found in (NEET PHASE I 2016)
of photosynthesis, which of the following a. lumen of thylakoids
statements is not correct? (NEET 2017) b. inter membrane space
a light saturation for CO2 fixation occurs at c. antennae complex
10 % of full sunlight d. stroma
b. increasing atmospheric CO2
concentration up to 0.05% can enhance 40. Oxidative phosphorylation is ( N E E T
CO2 fixation rate 2016)
c. C3 plants respond to higher temperature a. formation of ATP by transfer of phosphate
with enhanced photosynthesis while C4 group from a substrate to ADP
plants have much lower temperature b. oxidation of phosphate group in ATP
optimum. c. Aaddition of phosphate group to ATP
d. tomato is a greenhouse crop which can d. formation of ATP by energy released
be grown in CO2 enriched atmosphere from electrons during substrate
for higher yield oxidation.
36.
A plant in your garden avoids 41. Which of the biomolecules is common to
photorespiratory losses, has improved respiration-mediated breakdown of fats,
water use efficiency, shows high rates of carbohydrates and proteins? (NEET
photosynthesis at high temperatures and has 2013, 2016)
219

a. glucose-6-phosphate a. malic acid and acetyl coenzyme
b. fructose1,6-bisphosphate b. oxaloacetic acid and acetyl coenzyme
c. pyruvic acid c. succinic acid and pyruvic acid
d. acetyl CoA d. fumaric acid and pyruvic acid
42 Which statement is wrong for Krebs cycle? 46. Respiration is a process in which (CPMT
(NEET 2017) 1980)
a. there is one point in the cycle where FAD a. energy is used up
is reduced to FADH2 b. energy is stored in the form of ADP
b. during conversion of succinyl CoA c. energy is released and stored in the
to succinic acid, a molecule of GTP is form of ATP
synthesised. d. energy is not released at all
c. the cycle starts with condensation of
acetyl group a.cetyl CoA. with pyruvic 47. The common phase between aerobic and
acid to yield citric acid anaerobic respiration is called (CPMT
d. there are three points in the cycle where 1984)
NAD+ is reduced to NADH+H+ a. glycolysis
b. krebs cycle
43. The three boxes in this diagram represents c. tricarboxylic acid cycle
the three major biosynthetic pathways in d. oxidative phosphorylation
aerobic respiration and arrows represent
net reacts or products. (NEET 2013) 48. ATP synthesis occurs on/in the ( A I I M S
1984)
a. matrix
b. outer membrane of mitochondrion
Arrows numbered 4, 8 and 12 can be c. innermembrane of mitochondrion
a. ATP d. none of the above
b. H2O
49. Which 5-carbon organic acid of the
c. FAD or FADH2 Krebs cycle is a key compound in the N2
d. NADH metabolism of a cell (AIIMS 1989)
a. citric acid
44.
The energy released metabolic process
in which substrate is oxidised without an b. fumaric acid
external electron acceptor is called c. oxalosuccinic acid
(AIPMT 2010) d. α-Ketoglutaric acid
a. glycolysis
50. Which one of the following acts as a
b. fermentation
hormone involved in ripening of fruits
c. aerobic respiration (CBSE PMT 2000)
d. photorespiration a. naphthalene acetic acid
45. Krebs cycle starts with the formation of six b. ethylene
carbon compound by a reaction between c. indole acetic acid
(CPMT 1980) d. zeatin
220

51. Coconut milk factor is (PMT 2003) 59. Root development is promoted by
a. auxin b. gibberellin (AIPMT 2010)
c. abscisic acid d. cytokinin a. Auxin b. Gibberellin
c. Ethylene d. Abscisic acid
52. Banana is seedless because (JIPMER
2004) 60. Senscence as an active developmental cellular
a. it produces asexually process in the growth and functioning of
b. auxin is sprayed a flowering plant is indicated in (AIPMT
c. both A and B 2008)
d. none of the above a. Annual plants
b. Floral plants
53. Pruning of plants promotes branching due c. Vessels and Tracheid differentiation
to sensation of axillary buds by (AIIMS d. Leaf abscission
2004)
a. Ethylene b. Gibberellin 61. You are given a tissue with its potential
c. IAA d. Cytokinin for differentiation in an artificial culture.
Which of the following pairs of hormones
54 Avena curvature test is bioassay for activity would you add to the medium to secure
of (AIIMS 2006) (NEET 2016) shoots as well as roots?
a. Auxin b. Ethylene (NEET 2016)
c.Cytokinin d. Gibberellin a. Gibberellin and abscissic acid
b. IAA and gibberellins
55. One of the synthetic auxin is (AIPMT
c. Auxin and cytokinin
2009)
a. IBA b. NAA d. Auxin and abscisic acid
c. IAA d. GA 62. Phytochrome is a (NEET 2016)
a. Chromo protein
56
Which one of the following acids is
derivative of carotenoids (AIPMT 2009) b. Flavo protein
a. Abscisic acid c. Glyco protein
b. Indole butyric acid d. Lipo protein
c. Indole – 3 acetic 63. Typical growth curve in plants is
d. Gibberellic acid (NEET 2016)
a. Linear
57. Photoperiodism was first characterized in
(AIPMT 2010) b. Stair – steps shaped
a. Cotton b. Tobacco c. Parabolic
c. Potato d. Tomato d. Sigmoid
58. One of the commonly used plant growth

hormone in tea plantations is (AIPMT
2010)
a. Abscisic acid b. Zeatin
c. Indole – 3 – acetic acid
d. Ethylene
221

Botany - Class XI
List of Authors and Reviewers
Reviewers Authors
Dr. K.V. Krishnamurthy, P. Senthil, P.G. Assistant in Botany,
Professor and Head (Rtd), GBHSS, Uthangarai, Krishnagiri.
Bharathidasan University, Trichy
P. Saravanakumaran,
Dr. P. Ravichandran, P.G. Assistant in Botany, GHSS, Koduvilarpatti, Theni.
Associate Professor and Head,
Department of Botany, MS University, Tirunelveli Dr. N. Maheshkumar, Dist. Environmental Coordinator,
Chief Educational Office, Namakkal.
Dr. R. Ravindhran,
Associate Professor and Head, P. Anandhimala, P.G. Assistant in Botany,
Department of Plant Biology and Biotechnology, GGHSS,Pochampalli, Krishnagiri.
Loyola College, Chennai. Dr. P. Sivashankar,
Dr. M.P. Ramanujam, P.G. Assistant in Botany, GGHSS, Nachiyar Koil. Thanjavur.
Associate Professor of Botany G. Muthu, P.G. Assistant in Botany,
Kanchi Mamunivar Center for Post Graduate Studies GHSS (ADW) Achampatti, Madurai.
Pondichery J. Mani, P.G. Assistant in Botany,
Academic Coordinators GHSS, R Gobinathampatti, Dharmapuri.
U. Kalirajan,P.G. Assistant in Botany,
K. Manjula, ADWHSS, Meenambakkam, Kancheepuram.
Lecturer in Botany, DIET, Triplicane, Chennai.
G. Sathiyamoorthy,
J.Radhamani, PGTGHSS, Jayapuram, Vellore.
Lecturer in Botany, DIET, Kaliyampoondi.Kancheepuram S.B. Amuthavalli, P.G. Assistant in Botany,
Domain Experts GHSS, Ottery (Extension), Vandalur, Kancheepuram.
S. Malar Vizhi,P.G. Assistant in Botany,
Dr. S.S. Rathinakumar,Principal (Rtd.), GHSS, Chenbagaramanputhoor, Kannyakumari.
Sri Subramania Swamy Government Arts College, Thiruthani.
G. Bagyalakshmi, P.G. Assistant in Botany,
Dr. D. Narashiman, Professor and Head (Rtd.), GGHSS, Jalagandapuram, Salem.
Plant Biologly & BioTechnology, MCC College, Tambaram,
Kancheepuram. M. Chelladurai,
P.G. Assistant in Botany, GGHSS, Samuthiram, Salem.
Dr. Mujeera Fathima, Associate Professor of Botany,
Govt. Arts & Science College, Nandanam, Chennai. C. Kishore Kumar,
P.G. Assistant in Botany, GHSS, Thattaparai,Vellore.
Dr. K.P. Girivasan, Associate Professor of Botany,
Govt. Arts & Science College, Nandanam, Chennai. M. Vijayalakshmi , P.G. Assistant in Botany,
Dr. C.V. Chitti Babu, Associate Professor of Botany, Model School, Asthinapuram, Ariyalur.
Presidency College, Chennai. M. Lakshmi, P.G. Assistant in Botany,
Dr. Renu Edwin, Associate Professor of Botany, Sri Sankara Senior Secondary School, Adyar, Chennai.
Presidency College, Chennai. M. Chamundeswari, P.G. Assistant in Botany,
Dr. D. Kandavel, Associate Professor of Botany, Prince MHSS, Nanganallur, Kancheepuram.
Periyar EVR College, Trichy.
Dr. T. Sekar, Associate Professor of Botany, Content Readers
Pachaiyappa's College, Chennai.
Dr. T. S. Subha, Associate Professor in Botany,
Dr. D. Kathiresan, Assosiate Professor of Botany, Bharathi Women’s College, Chennai.
Saraswathi Narayana College, Madurai.
Dr. M. Pazhanisami,
Dr. S. Nagaraj, Assistant Professor of Botany, Associate Professor in Botany,
University of Madras, Guindy Campus, Chennai. Govt. Arts College, Nandanam, Chennai
Dr. M. Kumar, Assistant Professor of Botany, Dr. G. Rajalakshmi, Assistant Professor in Botany,
MCC College, Tambaram, Kancheepuram. Bharathi Women’s College, Chennai.
Dr. R. Kavitha, Assistant Professor in Botany,
Bharathi Women’s college, Chennai.
Art and Design Team
Chief Co-ordinator and Creative Head
Srinivasan Natarajan ICT Coordinator
N. Rajesh Kumar, B.T. Assistant,
CCMAGGHSS, Coimbatore
Graphics
Gopu Rasuvel,
Karthik kalaiarasu
Illustration
A. Jeyaseelan, Art Teacher
GBHSS, Uthangarai, Krishnagiri.
S.Gopu, Dr. N. Maheshkumar, Sathish, This book has been printed on 80 G.S.M.
Srinivasan
Elegant Maplitho paper.
Layout Printed by offset at:
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Ramesh Munisamy
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Pavithran, SCERT, Chennai
222

NOTES
223

NOTES
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GOVERNMENT OF TAMIL NADU

HIGHER SECONDARY FIRST YEAR

Untouchability is Inhuman and a Crime

Department of School Education

TN_GOVT_BOTANY_LV_XI_Page I-IV Front.indd 1 27-04-2018 15:21:40

First Edition - 2018

NOT FOR SALE

State Council of Educational

Printing & Publishing

Tamil NaduTextbook and Educational

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Chapter Outline Illustrate the complete overview of chapter

Amazing facts, Rhetorical questions to lead students

Activity Directions are provided to students to conduct activities

Infographics Visual representation of the lesson to enrich learning .

Evaluation Assess students to pause, think and check their understanding

ICT To enhance digital Science skills among students

Conceptual diagram that depicts relationships between

References List of related books for further details of the topic

Web links Digital resources

Glossary Explanation of scientific terms

Tamil terminology for Botanical terms given for easy

Competitive Model questions to face various competitive exams

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TN_GOVT_BOTANY_LV_XI_Page I-IV Front.indd 5

TN_GOVT_BOTANY_LV_XI_Page I-IV Front.indd 6

National Centre for Integrated Pest Pest Management www.ncipm.org.in

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UNIT IV: Plant Anatomy (Structural Organisation)

UNIT V: Plant Physiology (Functional Organisation)

E-book Assessment DIGI links

Lets use the QR code in the text books ! How ?

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9 Tissue and Tissue System

This chapter introduces the internal

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Position Origin Function Plane of division

Apical meristem Protoderm

Theories of Meristem Organization and Shoot Apical Meristem

Leaf primodium Leaf primordia

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Epidermis Stele Cortex Cortex Stele Cortex

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9.2 Permanent Tissues Figure 9.4: Parenchyma

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5. Prosenchyma: 2. Storage Parenchyma:

4. Chlorenchyma 3. How?.... Stellate

Intercellular Intercellular small chloroplast or none. Tannin maybe

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Figure 9.6: Types of Collenchyma a) Angular collenchyma, b) Lacunar collenchyma,

1. Branchysclereids or Stone cells: 2. Macrosclereids:

3. Osteosclereids (Bone cells):

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Lumen cell Thick

Figure 9.7: Types of Sclereids a) Brachysclereids, b) MacroSclereids, c) Osteosclereids,

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Student Activity Annular Spiral Reticulate Scalariform Pitted thickening

Xylem Consists of Four Types of Cells

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Vessels are found in 4. Phloem fibres

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9.4 Epidermal Tissue System Stem Epidermis