THE PLANT SHOOT

SYSTEM
FORM 3

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 ► 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

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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

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• 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

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• In most plants, absorption of water and
minerals occurs near the root hairs,
where vast numbers of tiny root hairs
increase the surface area

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► Many plants have modified roots

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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

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► 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

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► Many plants have modified stems

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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

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► 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

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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

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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

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Fig. 35-8

Dermal
tissue
Ground
tissue Vascular
tissue
► In nonwoody 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
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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
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► 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

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► 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

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Common Types of Plant Cells

► Like any multicellular organism, a plant is

characterized by cellular differentiation, the
specialization of cells in structure and function

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► Some major types of plant cells:
► Parenchyma
► Collenchyma
► Sclerenchyma
► Water-conducting cells of the xylem
► Sugar-conducting cells of the phloem

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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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

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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

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Fig. 35-10c

5 µm

Sclereid cells in pear

(LM)

25 µm

Cell wall

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

Fig. 35-10d

Vessel Tracheids 100 µm

Pits

Tracheids and vessels

(colorized SEM)
Perforation
plate
Vessel
element

Vessel elements, with

perforated end walls Tracheid
s
Fig. 35-10d1

Vessel Tracheids 100 µm

Tracheids and vessels

(colorized SEM)
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

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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

10 µm
Nucleus of
companio
n
cells

Sieve-tube elements:
longitudinal view Sieve plate with pores (SEM)
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
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Fig. 35-10e3

Sieve-tube
element

Plasmodesma

Sieve
plate
10
Nucleus of µ
companion m
cells

Sieve-tube elements:
longitudinal view Sieve plate with pores (SEM)
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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

► 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

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► 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
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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

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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

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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)
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Fig. 35-13
Cortex Vascular cylinder

Epidermis
Key
to labels
Zone of
Root hair differentiation
Dermal
Ground
Vascular

Zone of
elongation

Apical
meristem Zone of cell
division
Root cap

100 µm
► 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

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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)

Endodermi Key
s
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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

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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

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► 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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Fig. 35-18

Guard
cells
Key
to labels Stomatal
pore

50 µm
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
pore

50 µm
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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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 Gr o
Vascular
Pith ray

Primary
xylem
Secondary
xylem
Vascular
Secondary cambiu
m
Primary phloem
First corkphloe
cambium Cork
m
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
w th Vascular ray
Primary xylem Gr o
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
w th Vascular ray
Primary xylem Gr o
Secondary xylem
Pith
Secondary
phloem
First cork cambium
Cork

Periderm (mainly Most recent

cork cambia cork
and cork) cambium
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

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► 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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
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► 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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 35-21

RESULTS

2
Ring-width

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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

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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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 35-24
DNA or RNA metabolism (1%)
Signal transduction (2%)
Development (2%)
Energy pathways (3%)
Unknow
n Cell division and
Other (24%) organization (3%)
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

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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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
► Polarization is initiated by an
asymmetrical first division of the plant
zygote
► In the gnom mutant of Arabidopsis, the
establishment of polarity is defective

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 35-29
► Morphogenesis in plants, as in other
multicellular organisms, is often
controlled by homeotic genes

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

► 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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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