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Plant Structure
Plants, like multicellular animals
– Have organs composed of different tissues,
which are in turn composed of cells
• Three basic organs
evolved: roots, stems,
and leaves
• They are organized into
a root
system and a shoot
system
Roots
• A root
– Is an organ that anchors the vascular plant
– Absorbs minerals and water
– Often stores organic nutrients
In most plants
– The absorption of water and minerals occurs
near the root tips, where vast numbers of tiny
root hairs increase the surface area of the
root
Many plants have modified roots
(a) Prop roots (b) Storage roots (c) “Strangling” aerial
roots
(d) Buttress roots (e) Pneumatophores
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
Stems
• An axillary bud
– Is a structure that has the potential to form a
lateral shoot, or branch
• A terminal bud
– Is located near the shoot tip and causes
elongation of a young shoot
Stems
Many plants have modified stems
Rhizomes. The edible base
of this ginger plant is an example
of a rhizome, a horizontal stem
that grows just below the surface
or emerges and grows along the
surface.
(d)
Tubers. Tubers, such as these
red potatoes, are enlarged
ends of rhizomes specialized
for storing food. The “eyes”
arranged in a spiral pattern
around a potato are clusters
of axillary buds that mark
the nodes.
(c)
Bulbs. Bulbs are vertical,
underground shoots consisting
mostly of the enlarged bases
of leaves that store food. You
can see the many layers of
modified leaves attached
to the short stem by slicing an
onion bulb lengthwise.
(b)
Stolons. Shown here on a
strawberry plant, stolons
are horizontal stems that grow
along the surface. These “runners”
enable a plant to reproduce
asexually, as plantlets form at
nodes along each runner.
(a)
Storage leaves
Stem
Root Node
Rhizome
Root
Leaves
• The leaf
– Is the main photosynthetic organ of most
vascular plants
• Leaves generally consist of
– A flattened blade and a stalk
– The petiole, which joins the leaf to a node of
the stem
In classifying angiosperms
– Taxonomists may use leaf morphology as a
criterion
Petiole
(a) Simple leaf. A simple leaf
is a single, undivided blade.
Some simple leaves are
deeply lobed, as in an
oak leaf.
(b) Compound leaf. In a
compound leaf, the
blade consists of
multiple leaflets.
Notice that a leaflet
has no axillary bud
at its base.
(c) Doubly compound leaf.
In a doubly compound
leaf, each leaflet is
divided into smaller
leaflets.
Axillary bud
Leaflet
Petiole
Axillary bud
Axillary bud
Leaflet
Petiole
Some plant species
• Have evolved
modified leaves
that serve various
functions
(a) Tendrils. The tendrils by which this
pea plant clings to a support are
modified leaves. After it has “lassoed”
a support, a tendril forms a coil that
brings the plant closer to the support.
Tendrils are typically modified leaves,
but some tendrils are modified stems,
as in grapevines.
(b) Spines. The spines of cacti, such
as this prickly pear, are actually
leaves, and photosynthesis is
carried out mainly by the fleshy
green stems.
(c) Storage leaves. Most succulents,
such as this ice plant, have leaves
modified for storing water.
(d) Bracts. Red parts of the poinsettia
are often mistaken for petals but are
actually modified leaves called bracts
that surround a group of flowers.
Such brightly colored leaves attract
pollinators.
(e) Reproductive leaves. The leaves
of some succulents, such as Kalanchoe
daigremontiana, produce adventitious
plantlets, which fall off the leaf and
take root in the soil.
The Three Tissue Systems: Dermal,
Vascular, and Ground
Dermal
tissue
Ground
tissue Vascular
tissue
• The dermal tissue system
– Consists of the epidermis and periderm
• The vascular tissue system
– Carries out long-distance transport of
materials between roots and shoots
– Consists of two tissues, 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
• Ground tissue
– Includes various cells specialized for
functions such as storage, photosynthesis,
and support
Some of the major types of plant cells include
– Parenchyma
– Collenchyma
– Sclerenchyma
– Water-conducting cells of the xylem
– Sugar-conducting cells of the phloem
Parenchyma, collenchyma, and sclerenchyma cells
Parenchyma cells 60 µm
PARENCHYMA CELLS
80 µm Cortical parenchyma cells
COLLENCHYMA CELLS
Collenchyma cells
SCLERENCHYMA CELLS
Cell wall
Sclereid cells
in pear
25 µm
Fiber cells
5 µm
• Water-conducting cells of the xylem and
sugar-conducting cells of the phloem
WATER-CONDUCTING CELLS OF THE XYLEM
Vessel Tracheids 100 µm
Tracheids and vessels
Vessel
element
Vessel elements with
partially perforated
end walls
Pits
Tracheids
SUGAR-CONDUCTING CELLS OF THE PHLOEM
Companion cell
Sieve-tube
member
Sieve-tube members:
longitudinal view
Sieve
plate
Nucleus
Cytoplasm
Companion
cell
30 µm
15 µm
Meristems generate cells for new organs
• Apical meristems
– Are located at the tips of roots and in the buds
of shoots
– Elongate shoots and roots through primary
growth
• Lateral meristems
– Add thickness to woody plants through
secondary growth
• An overview of primary and secondary
growth
In woody plants,
there are lateral
meristems that
add secondary
growth, increasing
the girth of
roots and stems.
Apical meristems
add primary growth,
or growth in length.
Vascular
cambium
Cork
cambium
Lateral
meristems
Root apical
meristems
Primary growth in stems
Epidermis
Cortex
Primary phloem
Primary xylem
Pith
Secondary growth in stems
Periderm
Cork
cambium
Cortex
Primary
phloem
Secondary
phloem
Vascular cambium
Secondary
xylem
Primary
xylem
Pith
Shoot apical
meristems
(in buds)
The cork
cambium adds
secondary
dermal tissue.
The vascular
cambium adds
secondary
xylem and
phloem.
• In woody plants
– Primary and secondary growth occur
simultaneously but in different locations
This year’s growth
(one year old)
Last year’s growth
(two years old)
Growth of two
years ago (three
years old)
One-year-old side
branch formed
from axillary bud
near shoot apex
Scars left by terminal
bud scales of previous
winters
Leaf scar
Leaf scar
Stem
Leaf scar
Bud scale
Axillary buds
Internode
Node
Terminal bud
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
delicate apical meristem as the root pushes through soil
during primary growth
Dermal
Ground
Vascular
Key
Cortex Vascular cylinder
Epidermis
Root hair
Zone of
maturation
Zone of
elongation
Zone of cell
division
Apical
meristem
Root cap
100 µm
• The primary growth of roots
– Produces the epidermis, ground tissue, and
vascular tissue
Organization of primary tissues in young roots
Cortex
Vascular
cylinder
Endodermis
Pericycle
Core of
parenchyma
cells
Xylem
50 µm
Endodermis
Pericycle
Xylem
Phloem
Key
100 µm
Vascular
Ground
Dermal
Phloem
Transverse section of a root with parenchyma
in the center. The stele of many monocot roots
is a vascular cylinder with a core of parenchyma
surrounded by a ring of alternating xylem and phloem.
(b)
Transverse section of a typical root. In the
roots of typical gymnosperms and eudicots, as
well as some monocots, the stele is a vascular
cylinder consisting of a lobed core of xylem
with phloem between the lobes.
(a)
100 µm
Epidermis
• Lateral roots
– Arise from within the pericycle, the outermost
cell layer in the vascular cylinder
Cortex
Vascular
cylinder
Epidermis
Lateral root
100 µm
1 2
3 4
Emerging
lateral
root
Primary Growth of Shoots
• A shoot apical meristem
– Is a dome-shaped mass of dividing cells at the tip of
the terminal bud
– Gives rise to a repetition of internodes and leaf-
bearing nodes Apical meristem Leaf primordia
Developing
vascular
strand
Axillary bud
meristems
0.25 mm
Tissue Organization of Stems
• In gymnosperms and most eudicots
– The vascular tissue consists of vascular
bundles arranged in a ring
XylemPhloem
Sclerenchyma
(fiber cells)
Ground tissue
connecting
pith to cortex
Pith
Epidermis
Vascular
bundle
Cortex
Key
Dermal
Ground
Vascular
1 mm
(a) A eudicot stem. A eudicot stem (sunflower), with
vascular bundles forming a ring. Ground tissue toward
the inside is called pith, and ground tissue toward the
outside is called cortex. (LM of transverse section)
Ground
tissue
Epidermis
Vascular
bundles
1 mm
(b) A monocot stem. A monocot stem (maize) with vascular
bundles scattered throughout the ground tissue. In such an
arrangement, ground tissue is not partitioned into pith and
cortex. (LM of transverse section)
• In most monocot stems
– The vascular bundles are scattered
throughout the ground tissue, rather than
forming a ring
Tissue Organization of Leaves
• The epidermal barrier in leaves
– Is interrupted by stomata, which allow CO2
exchange between the surrounding air and the
photosynthetic cells within a leaf
• The ground tissue in a leaf
– Is sandwiched between the upper and lower
epidermis
• The vascular tissue of each leaf
– Is continuous with the vascular tissue of the
stem
Key
to labels
Dermal
Ground
Vascular
Guard
cells
Stomatal pore
Epidermal
cell
50 µm
Surface view of a spiderwort
(Tradescantia) leaf (LM)
(b)
Cuticle
Sclerenchyma
fibers
Stoma
Upper
epidermis
Palisade
mesophyll
Spongy
mesophyll
Lower
epidermis
Cuticle
Vein
Guard
cells
Xylem
Phloem
Guard
cells
Bundle-
sheath
cell
Cutaway drawing of leaf tissues(a)
Vein Air spaces Guard cells
100 µmTransverse section of a lilac
(Syringa) leaf (LM)
(c)
Figure 35.17a–c
• Leaf anatomy
• 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
The Vascular Cambium and Secondary
Vascular Tissue
• The vascular cambium
– Is a cylinder of meristematic cells one cell
thick
– Develops from parenchyma cells
Vascular cambium
Pith
Primary xylem
Secondary
xylem
Vascular
cambium
Secondary
phloem
Primary
phloem
Periderm
(mainly cork
cambia
and cork)
Pith
Primary
xylem
Vascular
cambium
Primary
phloem
Cortex
Epidermis
Vascular cambium
4 First cork cambium
Secondary
xylem (two
years of
production)
Pith
Primary xylem
Vascular cambium
Primary phloem
2
1
6
Growth
Primary
xylem
Secondary xylem
Secondary phloem
Primary phloem Cork
Phloem ray
3
Xylem
ray
Growth
Bark
8 Layers of
periderm
7 Cork5 Most recent
cork cambium
Cortex
Epidermis
9
In the youngest part of the stem, you can see the primary
plant body, as formed by the apical meristem during primary
growth. The vascular cambium is beginning to develop.
As primary growth continues to elongate the stem, the portion
of the stem formed earlier the same year has already started
its secondary growth. This portion increases in girth as fusiform
initials of the vascular cambium form secondary xylem to the
inside and secondary phloem to the outside.
The ray initials of the vascular cambium give rise to the xylem
and phloem rays.
As the diameter of the vascular cambium increases, the
secondary phloem and other tissues external to the cambium
cannot keep pace with the expansion because the cells no
longer divide. As a result, these tissues, including the
epidermis, rupture. A second lateral meristem, the cork
cambium, develops from parenchyma cells in the cortex. The
cork cambium produces cork cells, which replace the epidermis.
In year 2 of secondary growth, the vascular cambium adds to
the secondary xylem and phloem, and the cork cambium
produces cork.
As the diameter of the stem continues to increase, the
outermost tissues exterior to the cork cambium rupture and
slough off from the stem.
Cork cambium re-forms in progressively deeper layers of the
cortex. When none of the original cortex is left, the cork
cambium develops from parenchyma cells in the
secondary phloem.
Each cork cambium and the tissues it produces form a
layer of periderm.
Bark consists of all tissues exterior to the vascular
cambium.
1
2
3
4
5
6
7
8
9
Secondary phloem
(a) Primary and secondary growth
in a two-year-old stem
• Primary and secondary growth of a stem
Figure 35.18a
Secondary phloem
Vascular cambium
Late wood
Early wood
Secondary
xylem
Cork
cambium
Cork
Periderm
(b) Transverse section
of a three-year-
old stem (LM)
Xylem ray
Bark
0.5 mm0.5 mm
Figure 35.18b
• Viewed in transverse section, the vascular
cambium
– Appears as a ring, with interspersed regions
of dividing cells called fusiform initials and ray
initials
Figure 35.19a, b
Vascular
cambium
C X C
P
C
X
C
X
C
P
P
PC
X
X P
C
XX
C C
Types of cell division. An initial can divide
transversely to form two cambial initials (C)
or radially to form an initial and either a
xylem (X) or phloem (P) cell.
(a)
Accumulation of secondary growth. Although shown here
as alternately adding xylem and phloem, a cambial initial usually
produces much more xylem.
(b)
• 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
Growth ring
Vascular
ray
Heartwood
Sapwood
Vascular cambium
Secondary phloem
Layers of periderm
Secondary
xylem
Bark
Figure 35.20
Cork Cambia and the Production of
Periderm
• The cork cambium
– Gives rise to the secondary plant body’s
protective covering, or periderm
• Bark
– Consists of all the tissues external to the
vascular cambium, including secondary
phloem and periderm

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Plant structure

  • 2. Plants, like multicellular animals – Have organs composed of different tissues, which are in turn composed of cells
  • 3. • Three basic organs evolved: roots, stems, and leaves • They are organized into a root system and a shoot system
  • 4. Roots • A root – Is an organ that anchors the vascular plant – Absorbs minerals and water – Often stores organic nutrients
  • 5. In most plants – The absorption of water and minerals occurs near the root tips, where vast numbers of tiny root hairs increase the surface area of the root
  • 6. Many plants have modified roots (a) Prop roots (b) Storage roots (c) “Strangling” aerial roots (d) Buttress roots (e) Pneumatophores
  • 7. 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
  • 8. Stems • An axillary bud – Is a structure that has the potential to form a lateral shoot, or branch • A terminal bud – Is located near the shoot tip and causes elongation of a young shoot
  • 10. Many plants have modified stems Rhizomes. The edible base of this ginger plant is an example of a rhizome, a horizontal stem that grows just below the surface or emerges and grows along the surface. (d) Tubers. Tubers, such as these red potatoes, are enlarged ends of rhizomes specialized for storing food. The “eyes” arranged in a spiral pattern around a potato are clusters of axillary buds that mark the nodes. (c) Bulbs. Bulbs are vertical, underground shoots consisting mostly of the enlarged bases of leaves that store food. You can see the many layers of modified leaves attached to the short stem by slicing an onion bulb lengthwise. (b) Stolons. Shown here on a strawberry plant, stolons are horizontal stems that grow along the surface. These “runners” enable a plant to reproduce asexually, as plantlets form at nodes along each runner. (a) Storage leaves Stem Root Node Rhizome Root
  • 11. Leaves • The leaf – Is the main photosynthetic organ of most vascular plants
  • 12. • Leaves generally consist of – A flattened blade and a stalk – The petiole, which joins the leaf to a node of the stem
  • 13. In classifying angiosperms – Taxonomists may use leaf morphology as a criterion Petiole (a) Simple leaf. A simple leaf is a single, undivided blade. Some simple leaves are deeply lobed, as in an oak leaf. (b) Compound leaf. In a compound leaf, the blade consists of multiple leaflets. Notice that a leaflet has no axillary bud at its base. (c) Doubly compound leaf. In a doubly compound leaf, each leaflet is divided into smaller leaflets. Axillary bud Leaflet Petiole Axillary bud Axillary bud Leaflet Petiole
  • 14. Some plant species • Have evolved modified leaves that serve various functions (a) Tendrils. The tendrils by which this pea plant clings to a support are modified leaves. After it has “lassoed” a support, a tendril forms a coil that brings the plant closer to the support. Tendrils are typically modified leaves, but some tendrils are modified stems, as in grapevines. (b) Spines. The spines of cacti, such as this prickly pear, are actually leaves, and photosynthesis is carried out mainly by the fleshy green stems. (c) Storage leaves. Most succulents, such as this ice plant, have leaves modified for storing water. (d) Bracts. Red parts of the poinsettia are often mistaken for petals but are actually modified leaves called bracts that surround a group of flowers. Such brightly colored leaves attract pollinators. (e) Reproductive leaves. The leaves of some succulents, such as Kalanchoe daigremontiana, produce adventitious plantlets, which fall off the leaf and take root in the soil.
  • 15. The Three Tissue Systems: Dermal, Vascular, and Ground Dermal tissue Ground tissue Vascular tissue
  • 16. • The dermal tissue system – Consists of the epidermis and periderm
  • 17. • The vascular tissue system – Carries out long-distance transport of materials between roots and shoots – Consists of two tissues, xylem and phloem
  • 18. • 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
  • 19. • Ground tissue – Includes various cells specialized for functions such as storage, photosynthesis, and support
  • 20. Some of the major types of plant cells include – Parenchyma – Collenchyma – Sclerenchyma – Water-conducting cells of the xylem – Sugar-conducting cells of the phloem
  • 21. Parenchyma, collenchyma, and sclerenchyma cells Parenchyma cells 60 µm PARENCHYMA CELLS 80 µm Cortical parenchyma cells COLLENCHYMA CELLS Collenchyma cells SCLERENCHYMA CELLS Cell wall Sclereid cells in pear 25 µm Fiber cells 5 µm
  • 22. • Water-conducting cells of the xylem and sugar-conducting cells of the phloem WATER-CONDUCTING CELLS OF THE XYLEM Vessel Tracheids 100 µm Tracheids and vessels Vessel element Vessel elements with partially perforated end walls Pits Tracheids SUGAR-CONDUCTING CELLS OF THE PHLOEM Companion cell Sieve-tube member Sieve-tube members: longitudinal view Sieve plate Nucleus Cytoplasm Companion cell 30 µm 15 µm
  • 23. Meristems generate cells for new organs • Apical meristems – Are located at the tips of roots and in the buds of shoots – Elongate shoots and roots through primary growth
  • 24. • Lateral meristems – Add thickness to woody plants through secondary growth
  • 25. • An overview of primary and secondary growth In woody plants, there are lateral meristems that add secondary growth, increasing the girth of roots and stems. Apical meristems add primary growth, or growth in length. Vascular cambium Cork cambium Lateral meristems Root apical meristems Primary growth in stems Epidermis Cortex Primary phloem Primary xylem Pith Secondary growth in stems Periderm Cork cambium Cortex Primary phloem Secondary phloem Vascular cambium Secondary xylem Primary xylem Pith Shoot apical meristems (in buds) The cork cambium adds secondary dermal tissue. The vascular cambium adds secondary xylem and phloem.
  • 26. • In woody plants – Primary and secondary growth occur simultaneously but in different locations This year’s growth (one year old) Last year’s growth (two years old) Growth of two years ago (three years old) One-year-old side branch formed from axillary bud near shoot apex Scars left by terminal bud scales of previous winters Leaf scar Leaf scar Stem Leaf scar Bud scale Axillary buds Internode Node Terminal bud
  • 27. 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
  • 28. Primary Growth of Roots • The root tip is covered by a root cap, which protects the delicate apical meristem as the root pushes through soil during primary growth Dermal Ground Vascular Key Cortex Vascular cylinder Epidermis Root hair Zone of maturation Zone of elongation Zone of cell division Apical meristem Root cap 100 µm
  • 29. • The primary growth of roots – Produces the epidermis, ground tissue, and vascular tissue
  • 30. Organization of primary tissues in young roots Cortex Vascular cylinder Endodermis Pericycle Core of parenchyma cells Xylem 50 µm Endodermis Pericycle Xylem Phloem Key 100 µm Vascular Ground Dermal Phloem Transverse section of a root with parenchyma in the center. The stele of many monocot roots is a vascular cylinder with a core of parenchyma surrounded by a ring of alternating xylem and phloem. (b) Transverse section of a typical root. In the roots of typical gymnosperms and eudicots, as well as some monocots, the stele is a vascular cylinder consisting of a lobed core of xylem with phloem between the lobes. (a) 100 µm Epidermis
  • 31. • Lateral roots – Arise from within the pericycle, the outermost cell layer in the vascular cylinder Cortex Vascular cylinder Epidermis Lateral root 100 µm 1 2 3 4 Emerging lateral root
  • 32. Primary Growth of Shoots • A shoot apical meristem – Is a dome-shaped mass of dividing cells at the tip of the terminal bud – Gives rise to a repetition of internodes and leaf- bearing nodes Apical meristem Leaf primordia Developing vascular strand Axillary bud meristems 0.25 mm
  • 33. Tissue Organization of Stems • In gymnosperms and most eudicots – The vascular tissue consists of vascular bundles arranged in a ring XylemPhloem Sclerenchyma (fiber cells) Ground tissue connecting pith to cortex Pith Epidermis Vascular bundle Cortex Key Dermal Ground Vascular 1 mm (a) A eudicot stem. A eudicot stem (sunflower), with vascular bundles forming a ring. Ground tissue toward the inside is called pith, and ground tissue toward the outside is called cortex. (LM of transverse section)
  • 34. Ground tissue Epidermis Vascular bundles 1 mm (b) A monocot stem. A monocot stem (maize) with vascular bundles scattered throughout the ground tissue. In such an arrangement, ground tissue is not partitioned into pith and cortex. (LM of transverse section) • In most monocot stems – The vascular bundles are scattered throughout the ground tissue, rather than forming a ring
  • 35. Tissue Organization of Leaves • The epidermal barrier in leaves – Is interrupted by stomata, which allow CO2 exchange between the surrounding air and the photosynthetic cells within a leaf • The ground tissue in a leaf – Is sandwiched between the upper and lower epidermis • The vascular tissue of each leaf – Is continuous with the vascular tissue of the stem
  • 36. Key to labels Dermal Ground Vascular Guard cells Stomatal pore Epidermal cell 50 µm Surface view of a spiderwort (Tradescantia) leaf (LM) (b) Cuticle Sclerenchyma fibers Stoma Upper epidermis Palisade mesophyll Spongy mesophyll Lower epidermis Cuticle Vein Guard cells Xylem Phloem Guard cells Bundle- sheath cell Cutaway drawing of leaf tissues(a) Vein Air spaces Guard cells 100 µmTransverse section of a lilac (Syringa) leaf (LM) (c) Figure 35.17a–c • Leaf anatomy
  • 37. • 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
  • 38. The Vascular Cambium and Secondary Vascular Tissue • The vascular cambium – Is a cylinder of meristematic cells one cell thick – Develops from parenchyma cells
  • 39. Vascular cambium Pith Primary xylem Secondary xylem Vascular cambium Secondary phloem Primary phloem Periderm (mainly cork cambia and cork) Pith Primary xylem Vascular cambium Primary phloem Cortex Epidermis Vascular cambium 4 First cork cambium Secondary xylem (two years of production) Pith Primary xylem Vascular cambium Primary phloem 2 1 6 Growth Primary xylem Secondary xylem Secondary phloem Primary phloem Cork Phloem ray 3 Xylem ray Growth Bark 8 Layers of periderm 7 Cork5 Most recent cork cambium Cortex Epidermis 9 In the youngest part of the stem, you can see the primary plant body, as formed by the apical meristem during primary growth. The vascular cambium is beginning to develop. As primary growth continues to elongate the stem, the portion of the stem formed earlier the same year has already started its secondary growth. This portion increases in girth as fusiform initials of the vascular cambium form secondary xylem to the inside and secondary phloem to the outside. The ray initials of the vascular cambium give rise to the xylem and phloem rays. As the diameter of the vascular cambium increases, the secondary phloem and other tissues external to the cambium cannot keep pace with the expansion because the cells no longer divide. As a result, these tissues, including the epidermis, rupture. A second lateral meristem, the cork cambium, develops from parenchyma cells in the cortex. The cork cambium produces cork cells, which replace the epidermis. In year 2 of secondary growth, the vascular cambium adds to the secondary xylem and phloem, and the cork cambium produces cork. As the diameter of the stem continues to increase, the outermost tissues exterior to the cork cambium rupture and slough off from the stem. Cork cambium re-forms in progressively deeper layers of the cortex. When none of the original cortex is left, the cork cambium develops from parenchyma cells in the secondary phloem. Each cork cambium and the tissues it produces form a layer of periderm. Bark consists of all tissues exterior to the vascular cambium. 1 2 3 4 5 6 7 8 9 Secondary phloem (a) Primary and secondary growth in a two-year-old stem • Primary and secondary growth of a stem Figure 35.18a
  • 40. Secondary phloem Vascular cambium Late wood Early wood Secondary xylem Cork cambium Cork Periderm (b) Transverse section of a three-year- old stem (LM) Xylem ray Bark 0.5 mm0.5 mm Figure 35.18b
  • 41. • Viewed in transverse section, the vascular cambium – Appears as a ring, with interspersed regions of dividing cells called fusiform initials and ray initials Figure 35.19a, b Vascular cambium C X C P C X C X C P P PC X X P C XX C C Types of cell division. An initial can divide transversely to form two cambial initials (C) or radially to form an initial and either a xylem (X) or phloem (P) cell. (a) Accumulation of secondary growth. Although shown here as alternately adding xylem and phloem, a cambial initial usually produces much more xylem. (b)
  • 42. • 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
  • 43. Growth ring Vascular ray Heartwood Sapwood Vascular cambium Secondary phloem Layers of periderm Secondary xylem Bark Figure 35.20
  • 44. Cork Cambia and the Production of Periderm • The cork cambium – Gives rise to the secondary plant body’s protective covering, or periderm • Bark – Consists of all the tissues external to the vascular cambium, including secondary phloem and periderm