Chapter 35

Plant Structure, Growth, and

Development
Overview: Plastic Plants?

• To some people, the fanwort is an intrusive weed,

but to others it is an attractive aquarium plant
• This plant exhibits developmental plasticity, the
ability to alter itself in response to its environment
Fig. 35-1
• Developmental plasticity is more marked in plants
than in animals
• In addition to plasticity, plant species have by
natural selection accumulated characteristics of
morphology that vary little within the species
Concept 35.1: The plant body has a
hierarchy of organs, tissues, and cells
• Plants, like multicellular animals, have organs
composed of different tissues, which in turn are
composed of cells
The Three Basic Plant Organs: Roots,
Stems, and Leaves
• Basic morphology of vascular plants reflects their
evolution as organisms that draw nutrients from
below ground and above ground
• Three basic organs evolved: roots, stems, and leaves
• They are organized into a root system and a shoot
system
• Roots rely on sugar produced by photosynthesis in
the shoot system, and shoots rely on water and
minerals absorbed by the root system
Fig. 35-2
Reproductive shoot (flower)
Apical bud

Node
Internode

Apical
bud
Shoot
system
Vegetative
shoot

Blade
Leaf
Petiole

Axillar
y
bud
Stem
Taproot

Lateral Root
branch syste
roots m
Roots

•Roots are multicellular organs with important

functions:
– Anchoring the plant
– Absorbing minerals and water
– Storing organic nutrients
•A taproot system consists of one main vertical root
that gives rise to lateral roots, or branch roots
•Adventitious roots arise from stems or leaves
•Seedless vascular plants and monocots have a
fibrous root system characterized by thin lateral
roots with no main root
•In most plants, absorption of water and minerals
occurs near the root hairs, where vast numbers of
tiny root hairs increase the surface area
Fig. 35-3
• Many plants have modified roots
• Fibrous roots
• Taproots
• Storage roots
• Prop roots
• Adventitious roots
Fig. 35-4
Prop roots

“Strangling
”
aerial roots

Storage roots

Buttress roots

Pneumatophores
Fig. 35-4a

Prop roots
Fig. 35-4b

Storage roots
Fig. 35-4c

“Strangling” aerial roots

Fig. 35-4d

Pneumatophores
Fig. 35-4e

Buttress roots
Stems

• A stem is an organ consisting of

– An alternating system of nodes, the points at which leaves are
attached
– Internodes, the stem segments between nodes
• An axillary bud is a structure that has the potential
to form a lateral shoot, or branch
• An apical bud, or terminal bud, is located near the
shoot tip and causes elongation of a young shoot
• Apical dominance helps to maintain dormancy in
most nonapical buds
• Many plants have modified stems
Fig. 35-5
Rhizomes

Bulbs

Storage leaves
Stem
Stolons

Stolon

Tubers
Fig. 35-5a

Rhizomes
Fig. 35-5b

Storage leaves

Stem

Bulb
Fig. 35-5c

Stolon

Stolons
Fig. 35-5d

Tubers
Leaves

• The leaf is the main photosynthetic organ of most

vascular plants
• Leaves generally consist of a flattened blade and a
stalk called the petiole, which joins the leaf to a
node of the stem
• Monocots and eudicots differ in the arrangement of
veins, the vascular tissue of leaves
– Most monocots have parallel veins
– Most eudicots have branching veins
• In classifying angiosperms, taxonomists may use leaf
morphology as a criterion
Fig. 35-6

(a) Simple leaf

Petiole
Axillary bud

Leaflet
(b) Compound
leaf

Petiole
Axillary bud

(c) Doubly
compound Leaflet
leaf
Petiole
Axillary bud
Fig. 35-6a

(a) Simple leaf

Petiole
Axillary bud
Fig. 35-6b

Leaflet
(b
) Compo
und
leaf Petiole
Axillary bud
Fig. 35-6c

(c Doubly
)
compound
Leaflet
leaf
Petiole
Axillary bud
• Some plant species have evolved modified leaves
that serve various functions
Fig. 35-7
Tendrils

Spines

Storage
leaves

Reproductive leaves

Bracts
Fig. 35-7a

Tendrils
Fig. 35-7b

Spines
Fig. 35-7c

Storage leaves
Fig. 35-7d

Reproductive leaves
Fig. 35-7e

Bracts
Dermal, Vascular, and Ground Tissues

• Each plant organ has dermal, vascular, and ground

tissues
• Each of these three categories forms a tissue system
Fig. 35-8

Dermal
tissue
Ground
tissue Vascular
tissue
• In non-woody plants, the dermal tissue system
consists of the epidermis
• A waxy coating called the cuticle helps prevent
water loss from the epidermis
• In woody plants, protective tissues called periderm
replace the epidermis in older regions of stems and
roots
• Trichomes are outgrowths of the shoot epidermis
and can help with insect defense
Fig. 35-9
EXPERIMENT

Very hairy pod Slightly hairy pod Bald pod

(10 trichomes/ (2 trichomes/ (no trichomes)
mm2) mm2)

RESULTS

Very hairy pod: Slightly hairy Bald pod:

10% damage pod: 40% damage
25% damage
• The vascular tissue system carries out long-distance
transport of materials between roots and shoots
• The two vascular tissues are xylem and phloem
• Xylem conveys water and dissolved minerals
upward from roots into the shoots
• Phloem transports organic nutrients from where
they are made to where they are needed
• The vascular tissue of a stem or root is collectively
called the stele
• In angiosperms the stele of the root is a solid central
vascular cylinder
• The stele of stems and leaves is divided into vascular
bundles, strands of xylem and phloem
• Tissues that are neither dermal nor vascular are the
ground tissue system
• Ground tissue internal to the vascular tissue is pith;
ground tissue external to the vascular tissue is
cortex
• Ground tissue includes cells specialized for storage,
photosynthesis, and support
Common Types of Plant Cells

• Like any multicellular organism, a plant is

characterized by cellular differentiation, the
specialization of cells in structure and function
• Some major types of plant cells:
– Parenchyma
– Collenchyma
– Sclerenchyma
– Water-conducting cells of the xylem
– Sugar-conducting cells of the phloem
Parenchyma Cells
• Mature parenchyma cells
– Have thin and flexible primary walls
– Lack secondary walls
– Are the least specialized
– Perform the most metabolic functions
– Retain the ability to divide and differentiate

BioFlix: Tour of a Plant Cell

Fig. 35-10a

Parenchyma cells in Elodea leaf,

with chloroplasts (LM) 60 µm
Collenchyma Cells
•Collenchyma cells are grouped in strands and help
support young parts of the plant shoot
•They have thicker and uneven cell walls
•They lack secondary walls
•These cells provide flexible support without
restraining growth
Fig. 35-10b

5 µm

Collenchyma cells (in Helianthus stem) (LM)

Sclerenchyma Cells
•Sclerenchyma cells are rigid because of thick
secondary walls strengthened with lignin
•They are dead at functional maturity
•There are two types:
– Sclereids are short and irregular in shape and have thick
lignified secondary walls
– Fibers are long and slender and arranged in threads
Fig. 35-10c

5 µm

Sclereid cells in pear

(LM)

25 µm

Cell wall

Fiber cells (cross section from ash tree) (LM)

Water-Conducting Cells of the Xylem
•The two types of water-conducting cells, tracheids
and vessel elements, are dead at maturity
•Tracheids are found in the xylem of all vascular
plants
• Vessel elements are common to most angiosperms
and a few gymnosperms
• Vessel elements align end to end to form long
micropipes called vessels
Fig. 35-10d

100 µm
Vessel Tracheids

Pits

Tracheids and vessels

(colorized SEM)
Perforation
plate
Vessel
element

Vessel elements, with

perforated end walls Tracheid
s
Fig. 35-10d1
100 µm
Vessel Tracheids

Tracheids and vessels

(colorized SEM)
Fig. 35-10d2

Pits

Perforation
plate

Vessel
element

Vessel elements, with

perforated end walls
Tracheid
s
Sugar-Conducting Cells of the
Phloem
•Sieve-tube elements are alive at functional
maturity, though they lack organelles
•Sieve plates are the porous end walls that allow
fluid to flow between cells along the sieve tube
•Each sieve-tube element has a companion cell
whose nucleus and ribosomes serve both cells
Fig. 35-10e
Sieve-tube elements:
3 µm longitudinal view (LM)

Sieve plate
Sieve-tube element (left)
Companio
and companion cell: n
cross section (TEM) cells

Sieve-tube
elements

Plasmodesma

Sieve
plate 30 µm

Nucleus of
companio
n
cells

15 µm
Sieve-tube elements:
longitudinal view Sieve plate with pores (LM)
Fig. 35-10e1

3 µm

Sieve-tube element (left)

and companion cell:
cross section (TEM)
Fig. 35-10e2
Sieve-tube elements:
longitudinal view (LM)

Sieve plate

Companion
cells

Sieve-tub
e
elements

30 µm
Fig. 35-10e3

Sieve-tube
element

Plasmodesma

Sieve
plate

Nucleus of
companion
cells

15
µ
Sieve-tube elements: m
longitudinal view Sieve plate with pores (LM)
Concept 35.2: Meristems generate cells
for new organs
• A plant can grow throughout its life; this is called
indeterminate growth
• Some plant organs cease to grow at a certain size;
this is called determinate growth
• Annuals complete their life cycle in a year or less
• Biennials require two growing seasons
• Perennials live for many years
• Meristems are perpetually embryonic tissue and
allow for indeterminate growth
• Apical meristems are located at the tips of roots and
shoots and at the axillary buds of shoots
• Apical meristems elongate shoots and roots, a
process called primary growth
• Lateral meristems add thickness to woody plants, a
process called secondary growth
• There are two lateral meristems: the vascular
cambium and the cork cambium
• The vascular cambium adds layers of vascular tissue
called secondary xylem (wood) and secondary
phloem
• The cork cambium replaces the epidermis with
periderm, which is thicker and tougher
Fig. 35-11

Primary growth in stems

Epidermis
Cortex
Shoot tip (shoot Primary phloem
apical meristem
and young leaves) Primary xylem
Pith
Lateral meristems:
Vascular cambium Secondary growth in stems
Cork cambium
Axillary bud Periderm
meristem Cork
cambium

Cortex

Pith Primary
phloem
Primary
Root apical xylem Secondary
meristems Secondary phloem
xylem
Vascular cambium
• Meristems give rise to initials, which remain in the
meristem, and derivatives, which become
specialized in developing tissues
• In woody plants, primary and secondary growth
occur simultaneously but in different locations
Fig. 35-12
Apical bud
Bud scale

Axillary buds

This year’s growth

(one year old) Leaf
scar

Bud Node
One-year-old side
scar branch formed
Internode from axillary bud
near shoot tip

Last year’s growth

(two years old) Leaf scar

Stem

Bud scar left by apical

bud scales of previous
winters
Growth of two
years ago
(three years old) Leaf scar
Concept 35.3: Primary growth lengthens
roots and shoots
• Primary growth produces the primary plant body,
the parts of the root and shoot systems produced by
apical meristems
Primary Growth of Roots

• The root tip is covered by a root cap, which protects

the apical meristem as the root pushes through soil
• Growth occurs just behind the root tip, in three
zones of cells:
– Zone of cell division
– Zone of elongation
– Zone of maturation

Video: Root Growth in a Radish Seed (Time

Lapse)
Fig. 35-13
Cortex Vascular cylinder
Key
to labels
Epidermis
Dermal
Zone of Ground
Root hair differentiation Vascular

Zone of
elongation

Zone of cell Mitotic

division cells
(including
apical
meristem)

100 µm
Root cap
• The primary growth of roots produces the
epidermis, ground tissue, and vascular tissue
• In most roots, the stele is a vascular cylinder
• The ground tissue fills the cortex, the region
between the vascular cylinder and epidermis
• The innermost layer of the cortex is called the
endodermis
Fig. 35-14
Epidermis

Cortex

Endodermi
s
Vascular
cylinder

Pericycle

Core of
parenchyma
cells

Xylem
100 µm
Phloem
(a) Root with xylem and phloem in the center 100 µm
(typical of eudicots)
(b) Root with parenchyma in the center (typical of
monocots)

Endodermi Key
s to labels
Pericycle
Dermal
Ground
Vascular
Xylem

Phloem

50 µm
Fig. 35-14a1

Epidermis Key
to labels
Cortex
Dermal
Endodermis Ground
Vascular
Vascular
cylinder

Pericycle

Xylem
100 µm
Phloem
(a) Root with xylem and phloem in the center
(typical of eudicots)
Fig. 35-14a2

(a) Root with xylem and phloem in the center

(typical of eudicots)

Endodermis Key
to labels
Pericycl
e Dermal
Ground
Vascular
Xylem

Phloem

50 µm
Fig. 35-14b

Epidermis

Cortex

Endodermis

Vascular
Key cylinder
to labels
Pericycle
Dermal
Groun Core of
d parenchyma
Vascular cells

Xylem

Phloem
100 µm

(b) Root with parenchyma in the center

(typical of
monocots)
• Lateral roots arise from within the pericycle, the
outermost cell layer in the vascular cylinder
Fig. 35-15-1

100 µm
Emerging
lateral
root

Cortex
1 Vascular
cylinder
Fig. 35-15-2

100 µm Epidermis
Emerging
lateral Lateral root
root

Cortex
1 Vascular 2
cylinder
Fig. 35-15-3

100 µm Epidermis
Emerging
lateral Lateral root
root

Cortex
1 Vascular 2 3
cylinder
Primary Growth of Shoots

• A shoot apical meristem is a dome-shaped mass of

dividing cells at the shoot tip
• Leaves develop from leaf primordia along the sides
of the apical meristem
• Axillary buds develop from meristematic cells left at
the bases of leaf primordia
Fig. 35-16
Shoot apical meristem Leaf primordia

Young
leaf

Developin
g
vascular
strand

Axillary bud
meristems

0.25 mm
Tissue Organization of Stems

• Lateral shoots develop from axillary buds on the

stem’s surface
• In most eudicots, the vascular tissue consists of
vascular bundles that are arranged in a ring
Fig. 35-17

Phloem Xylem

Sclerenchyma Ground
Ground tissue
(fiber cells) tissue
connecting
pith to cortex

Pith Epidermis

Key
to labels

Epidermis Cortex Vascular

Dermal bundles
Vascular
bundle Ground
1 mm Vascular 1 mm
(a) Cross section of stem with vascular bundles forming (b) Cross section of stem with scattered vascular bundles
a ring (typical of eudicots) (typical of monocots)
Fig. 35-17a
Phloem Xylem

Sclerenchyma Ground
(fiber cells) tissue
connecting
pith to cortex

Pith

Key
to labels

Epidermis Cortex
Dermal
Vascular
bundle Groun
d
1 mm Vascular
(a) Cross section of stem with vascular bundles forming
a ring (typical of eudicots)
Fig. 35-17b

Ground
tissue

Epidermis

Key
to
lab
Vascular
els
Dermal bundles
Ground
Vascular 1 mm
(b) Cross section of stem with scattered vascular bundles
(typical of monocots)
• In most monocot stems, the vascular bundles are
scattered throughout the ground tissue, rather than
forming a ring
Tissue Organization of Leaves

•The epidermis in leaves is interrupted by stomata,

which allow CO2 exchange between the air and the
photosynthetic cells in a leaf
•Each stomatal pore is flanked by two guard cells,
which regulate its opening and closing
•The ground tissue in a leaf, called mesophyll, is
sandwiched between the upper and lower
epidermis
• Below the palisade mesophyll in the upper part of
the leaf is loosely arranged spongy mesophyll,
where gas exchange occurs
• The vascular tissue of each leaf is continuous with
the vascular tissue of the stem
• Veins are the leaf’s vascular bundles and function as
the leaf’s skeleton
• Each vein in a leaf is enclosed by a protective bundle
sheath
Fig. 35-18

Guard
cells
Key
to labels
Stomatal

50 µm
pore
Dermal
Epidermal
Ground
Cuticle Sclerenchyma cell
Vascular fibers
Stoma (b) Surface view of a spiderwort
(Tradescantia) leaf (LM)

Upper
epidermis

Palisade
mesophyll

Bundle- Spongy
sheath mesophyll
cell

100 µm
Lower
epidermis
Cuticle
Xylem
Phloem Vein
Guard Vein Air spaces Guard cells
(a) Cutaway drawing of leaf tissues cells (c) Cross section of a lilac
(Syringa)) leaf (LM)
Fig. 35-18a
Key
to labels

Dermal
Ground
Cuticle Sclerenchyma
Vascular fibers
Stoma

Upper
epidermis

Palisade
mesophyll

Bundle- Spongy
sheath mesophyll
cell
Lower
epidermis
Cuticle
Xylem
Phloem Vein
Guard
(a) Cutaway drawing of leaf tissues cells
Fig. 35-18b

Guard
cells

Stomatal

50 µm
pore
Epidermal
cell

(b) Surface
view of a
spiderwo
rt
(Tradescant
ia) leaf
Fig. 35-18c

Upper
epidermis
Key
to labels
Palisade
Dermal mesophyl
l
Ground
Vascular
Spongy
mesophyl
l

100 µm
Lower
epidermis

Vein Air spaces Guard

cells
(c) Cross section of a lilac
(Syringa) leaf (LM)
Concept 35.4: Secondary growth adds
girth to stems and roots in woody plants
• Secondary growth occurs in stems and roots of
woody plants but rarely in leaves
• The secondary plant body consists of the tissues
produced by the vascular cambium and cork
cambium
• Secondary growth is characteristic of gymnosperms
and many eudicots, but not monocots
Fig. 35-19

(a) Primary and secondary

growth
in a two-year-old stem

Epidermis
Pith
Cortex
Primary
Primary xyle
Vascular cambium Epidermis
phloem m
Primary phloem Cortex
Vascular
cambiu
m
Primary wth
xylem Gro
Vascular
Pith ray

Primary
xylem
Secondary
xylem
Vascular
Secondary cambiu
m
phloem
Primary phloem
First cork cambium Cork

Periderm
(mainly cork wth
cambia Gro
and cork)
Secondary Bark
Vascular phloem
Primary Late woodcambiu Cork
Secondary m cambiu
phloem xylem Early wood Periderm
m
Secondary Cork
phloem Secondary
Xylem (two
Vascular years of

0.5 mm
cambium
production)
Secondary Vascular
xylem cambiu Bark
Secondary m
Primary phloem Layers of Vascular ray Growth ring
Most recent
xylem cork cambium Cork periderm (b) Cross section of a three-year-
old Tilia (linden) stem (LM)
Pith 0.5 mm
Fig. 35-19a1
(a) Primary and secondary growth Pith
in a two-year-old stem Primary xylem
Vascular cambium
Epidermis Primary phloem
Cortex Cortex
Primary phloem Epidermis
Vascular cambium
Primary xylem
Pith

Periderm (mainly
cork cambia
and cork)

Secondary phloem

Secondary
xylem
Fig. 35-19a2
(a) Primary and secondary growth Pith
in a two-year-old stem Primary xylem
Vascular cambium
Epidermis Primary phloem
Cortex Cortex
Primary phloem Epidermis
Vascular cambium
wth Vascular ray
Primary xylem Gro
Secondary xylem
Pith
Secondary
phloem
First cork cambium
Cork

Periderm (mainly
cork cambia
and cork)

Secondary phloem

Secondary
xylem
Fig. 35-19a3
(a) Primary and secondary growth Pith
in a two-year-old stem Primary xylem
Vascular cambium
Epidermis Primary phloem
Cortex Cortex
Primary phloem Epidermis
Vascular cambium
wth Vascular ray
Primary xylem Gro
Secondary xylem
Pith
Secondary
phloem
First cork cambium
Cork

Periderm (mainly Most recent cork

cork cambia cambium
and cork)
Cork

Secondary phloem Bark

Layers of
periderm
Secondary
xylem
Fig. 35-19b

Secondary phloem Bark

Vascular cambium
Late wood Cork
Secondary xylem cambium Periderm
Early wood
Cork

0.5 mm
Vascular ray Growth ring
(b) Cross section of a three-year-
old Tilia (linden) stem (LM)
0.5 mm
The Vascular Cambium and Secondary
Vascular Tissue
• The vascular cambium is a cylinder of meristematic
cells one cell layer thick
• It develops from undifferentiated parenchyma cells
• In cross section, the vascular cambium appears as a
ring of initials
• The initials increase the vascular cambium’s
circumference and add secondary xylem to the
inside and secondary phloem to the outside
Fig. 35-20

Vascular cambium Growth

Vascular
X X C P P cambium
Secondary
Secondary
X X C P phloem
xylem

X C P
C
C

C C C X C

C
After one year After two years
C C C
of growth of growth
• Secondary xylem accumulates as wood, and consists
of tracheids, vessel elements (only in angiosperms),
and fibers
• Early wood, formed in the spring, has thin cell walls
to maximize water delivery
• Late wood, formed in late summer, has thick-walled
cells and contributes more to stem support
• In temperate regions, the vascular cambium of
perennials is dormant through the winter
• Tree rings are visible where late and early wood
meet, and can be used to estimate a tree’s age
• Dendrochronology is the analysis of tree ring growth
patterns, and can be used to study past climate
change
Fig. 35-21

RESULTS

Ring-width 2
1.5
indexes

1
0.5
0
1600 1700 1800 1900 2000
Year
• As a tree or woody shrub ages, the older layers of
secondary xylem, the heartwood, no longer
transport water and minerals
• The outer layers, known as sapwood, still transport
materials through the xylem
• Older secondary phloem sloughs off and does not
accumulate
Fig. 35-22

Growth
ring
Vascular
ray

Heartwood
Secondary
xylem Sapwood

Vascular cambium

Secondary phloem
Bark
Layers of
periderm
Fig. 35-23
The Cork Cambium and the Production
of Periderm
• The cork cambium gives rise to the secondary plant
body’s protective covering, or periderm
• Periderm consists of the cork cambium plus the
layers of cork cells it produces
• Bark consists of all the tissues external to the
vascular cambium, including secondary phloem and
periderm
• Lenticels in the periderm allow for gas exchange
between living stem or root cells and the outside air
Concept 35.5: Growth, morphogenesis,
and differentiation produce the plant
body
• Morphogenesis is the development of body form
and organization
• The three developmental processes of growth,
morphogenesis, and cellular differentiation act in
concert to transform the fertilized egg into a plant
Molecular Biology: Revolutionizing the Study of
Plants
• New techniques and model systems are catalyzing
explosive progress in our understanding of plants
• Arabidopsis is a model organism, and the first plant
to have its entire genome sequenced
• Studying the genes and biochemical pathways of
Arabidopsis will provide insights into plant
development, a major goal of systems biology
Fig. 35-24
DNA or RNA metabolism (1%)
Signal transduction (2%)
Development (2%)
Energy pathways (3%)
Unknow
n Cell division and
Other organization (3%)
(24%)
metabolism
(18%) Transport (4%)
Transcription
(4%)
Response to
environment
(4%)
Protein
metabolism
(7%)

Other biological
processes (11%)
Other cellular
processes (17%)
Growth: Cell Division and Cell Expansion

• By increasing cell number, cell division in meristems

increases the potential for growth
• Cell expansion accounts for the actual increase in
plant size
The Plane and Symmetry of Cell Division

• The plane (direction) and symmetry of cell division

are immensely important in determining plant form
• If the planes of division are parallel to the plane of
the first division, a single file of cells is produced
Fig. 35-25

Plane of
cell division

(a) Planes of cell division

Developing
guard cells

Unspecialized Guard cell

epidermal cell “mother cell”

(b) Asymmetrical cell division

Fig. 35-25a

Plane of
cell division

(a) Planes of cell division

• If the planes of division vary randomly, asymmetrical
cell division occurs
Fig. 35-25b

Developing
guard cells

Unspecialized Guard cell

epidermal cell “mother cell”

(b) Asymmetrical cell division

• The plane in which a cell divides is determined
during late interphase
• Microtubules become concentrated into a ring
called the preprophase band that predicts the
future plane of cell division
Fig. 35-26

Preprophase 10 µm
bands
of microtubules

Nuclei

Cell plates
Orientation of Cell Expansion

• Plant cells grow rapidly and “cheaply” by intake and

storage of water in vacuoles
• Plant cells expand primarily along the plant’s main
axis
• Cellulose microfibrils in the cell wall restrict the
direction of cell elongation
Fig. 35-27

Cellulose
microfibrils

Nucleus Vacuoles 5 µm
Microtubules and Plant Growth

• Studies of fass mutants of Arabidopsis have

confirmed the importance of cytoplasmic
microtubules in cell division and expansion
Fig. 35-28

0.3 mm
(b) fass seedling

m
m

m
m
2

(a) Wild-type seedling (c) Mature fass mutant

Morphogenesis and Pattern Formation

• Pattern formation is the development of specific

structures in specific locations
• It is determined by positional information in the
form of signals indicating to each cell its location
• Positional information may be provided by gradients
of molecules
• Polarity, having structural or chemical differences at
opposite ends of an organism, provides one type of
positional information
• Polarization is initiated by an asymmetrical first
division of the plant zygote
• In the gnom mutant of Arabidopsis, the
establishment of polarity is defective
Fig. 35-29
• Morphogenesis in plants, as in other multicellular
organisms, is often controlled by homeotic genes
Fig. 35-30
Gene Expression and Control of Cellular
Differentiation
• In cellular differentiation, cells of a developing
organism synthesize different proteins and diverge
in structure and function even though they have a
common genome
• Cellular differentiation to a large extent depends on
positional information and is affected by homeotic
genes
Fig. 35-31

Cortical
cells

20 µm
Location and a Cell’s Developmental
Fate
• Positional information underlies all the processes of
development: growth, morphogenesis, and
differentiation
• Cells are not dedicated early to forming specific
tissues and organs
• The cell’s final position determines what kind of cell
it will become
Shifts in Development: Phase Changes

• Plants pass through developmental phases, called

phase changes, developing from a juvenile phase to
an adult phase
• Phase changes occur within the shoot apical
meristem
• The most obvious morphological changes typically
occur in leaf size and shape
Fig. 35-32

Leaves produced
by adult phase
of apical meristem

Leaves produced
by juvenile phase
of apical meristem
Genetic Control of Flowering

• Flower formation involves a phase change from

vegetative growth to reproductive growth
• It is triggered by a combination of environmental
cues and internal signals
• Transition from vegetative growth to flowering is
associated with the switching on of floral meristem
identity genes
• Plant biologists have identified several organ
identity genes (plant homeotic genes) that regulate
the development of floral pattern
• A mutation in a plant organ identity gene can cause
abnormal floral development
Fig. 35-33

Pe
Ca
St
Se

Pe

Se

(a) Normal Arabidopsis flower Pe

Pe

Se

(b) Abnormal Arabidopsis flower

• Researchers have identified three classes of floral
organ identity genes
• The ABC model of flower formation identifies how
floral organ identity genes direct the formation of
the four types of floral organs
• An understanding of mutants of the organ identity
genes depicts how this model accounts for floral
phenotypes
Fig. 35-34
Sepals
Petals

Stamens

A Carpels (a) A schematic diagram of the ABC hypothesis

B
C

C gene
activity
B+C Carpel
A+B gene
gene activity
activity
Petal
A gene
activity

Stamen

Sepal

Active B B B B B B B B A A A A
genes: A A C C CC AA C C C C C C C C A A C CCC A A A B B A A B B A
Whorls:

Carpel
Stamen Petal

Sepal
Wild type Mutant lacking A Mutant lacking B Mutant lacking C

(b) Side view of flowers with organ identity mutations

Fig. 35-34a
Sepals
Petals
Stamens
A (a) A schematic diagram of the ABC hypothesis
B Carpels
C

C gene
activit
B+C y Carpel
A+B gene
gene activit
activit y Petal
y
A gene
activit
y
Stamen

Sepal
Fig. 35-34b

Active BB B B BB BB AA AA
genes: AACCCC AA CCCCCCCC A ACCCC AA ABBAABBA
Whorls:

Carpel
Stamen Petal

Sepa
l
Wild type Mutant lacking A Mutant lacking B Mutant lacking C

(b) Side view of flowers with organ identity mutations

Fig. 35-UN1

Shoot tip
(shoot apical
meristem and
young leaves)

Vascular
cambium Lateral
Cork meristems
Axillary bud cambium
meristem

Root apical
meristems
Fig. 35-UN2
Fig. 35-UN3
You should now be able to:

1. Compare the following structures or cells:

– Fibrous roots, taproots, root hairs, adventitious roots
– Dermal, vascular, and ground tissues
– Monocot leaves and eudicot leaves
– Parenchyma, collenchyma, sclerenchyma,
water-conducting cells of the xylem, and
sugar-conducting cells of the phloem
– Sieve-tube element and companion cell
2. Explain the phenomenon of apical dominance
3. Distinguish between determinate and
indeterminate growth
4. Describe in detail the primary and secondary
growth of the tissues of roots and shoots
5. Describe the composition of wood and bark
6. Distinguish between morphogenesis,
differentiation, and growth
7. Explain how a vegetative shoot tip changes into a
floral meristem