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Stem Structure

and Function
Topic Outline

• External Organization of the stem


(Node, Internode, Leaf Axil and Bud)
• Internal Organization of the stem
• Longitudinal Section (Primary Growth,
Differentiation, and Secondary Growth)
• Cross section (Arrangement of Tissues in the
Dicot and Monocot Young and Old stems)
FUNCTIONS

• Stems support leaves and


branches.
• Stems transport water and
solutes between roots and
leaves.
• Stems in some plants are
photosynthetic.
• Stems may store materials
necessary for life (e.g., water,
starch, sugar).
• In some plants, stems have
become adapted for specialized
functions.
Used with permission from http://education-portal.com
– May be vegetative (leaf bearing)
or reproductive (flower
bearing).

– Node- area of stem where leaf


is born

– Internodes- stem area between


nodes

– Buds: Stem elongation.


Embryonic tissue of leaves and
stem (not flower bud)
– Terminal bud- Located at tip
of stems or branches.
– Axillary bud- Gives rise to
branches

– Apical Dominance: Prevention


of branch formation by terminal
bud
Plant Tissues

1) Dermal Tissue System


• Outer covering
• Protection
2) Vascular Tissue System
• “Vessels” throughout plant
• Transport materials

3) Ground Tissue System


• “Body” of plant
• Photosynthesis; storage; support

Used with permission from http://education-portal.com


Stems – Structure and Development
• Stems have all three types of
plant tissue
• Grow by division at meristems
– Develop into leaves, other
shoots, and even flowers

• Leaves may be arranged in


one of three ways
Modified Stems
• Modified shoots with diverse functions have
evolved in many plants.
– These shoots, which include stolons, rhizomes,
tubers and bulbs, are often mistaken for roots.

– Stolons, such as the “runners” of strawberry


plants, grow on the
surface and enable
a plant to colonize large
areas asexually when a
parent plant fragments
into many smaller
offspring. Fig. 35.4a
Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings
– Rhizomes, like those of ginger, are horizontal stems
that grow underground.

– Tubers, including potatoes, are the swollen ends of


rhizomes specialized for food storage.

– Bulbs, such as onions, are vertical, underground


shoots consisting mostly of the swollen bases of
leaves that store food.

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


Plant Tissues – Ground Tissue
• Some major types of plant cells:
– Parenchyma
– Collenchyma
– Sclerenchyma
• Tissues that are neither dermal nor vascular are ground
tissue

• 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
Vascular Tissue
Vascular tissue:
Runs continuous throughout the plant
• transports materials between roots and
shoots.
– Xylem transports water and dissolved
minerals upward from roots into the shoots.
(water the xylem)

– Phloem transports food from the leaves to


the roots and to non-photosynthetic parts
of the shoot system.
(feed the phloem)
Xylem
– Main water-conducting
tissue of vascular plants.
– arise from individual
cylindrical cells oriented
end to end.
– At maturity the end walls
of these cells dissolve away
and the cytoplasmic
contents die.
– The result is the xylem
vessel, a continuous
nonliving duct.
– carry water and some
dissolved solutes, such as
inorganic ions, up the plant
Phloem

– The main components of phloem


are
• sieve elements
• companion cells.
– Sieve elements have no nucleus and
only a sparse collection of other
organelles . Companion cell
provides energy
– so-named because end walls are
perforated - allows cytoplasmic
connections between vertically-
stacked cells .
– conducts sugars and amino acids -
from the leaves, to the rest of the
plant
Phloem transport requires
specialized, living cells
• Sieve tubes elements join
to form continuous tube
• Pores in sieve plate
between sieve tube
elements are open
channels for transport
• Each sieve tube element
is associated with one or
more companion cells.
– Many plasmodesmata
penetrate walls between sieve
tube elements and companion
cells
– Close relationship, have a
ready exchange of solutes
between the two cells
Phloem transport requires
specialized, living cells
• Companion cells:
– Main role is the transport of
photosynthesis products from
producing cells in mature leaves
to sieve plates of the small vein
of the leaf
– Synthesis of the various
proteins used in the phloem
– Contain many mitochondria for
cellular respiration to provide
the cellular energy required for
active transport
– There ate three types
• Ordinary companion cells
• Transfer cells
• Intermediary cells
Vasculature - Comparisons
• In most monocot stems, the vascular bundles are
scattered throughout the ground tissue, rather than
forming a ring as with Dicots
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 dicots) (typical of monocots)

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


epidermis
Dicot Stem
Anatomy phloem

cortex
vascular
bundle
pith vascular
cambium

xylem
Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings
Monocot Stem
phloem
Anatomy

epidermis
vascular
bundles

ground
tissue
xylem

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


Plant stem growth
 Vegetative development is based on
meristems, in which cell division occurs
throughout life, producing cells that go on to
differentiate.

 When a meristem is converted from


vegetative to reproductive development,
regulatory transcription factors are
activated that control the identity and
position of floral organs.
Plant Growth

1) Primary Growth:
• Apical Meristems:
Mitotic cells at “tips” of roots / stems length

1) Increased length
2) Specialized structures (e.g.
fruits)
2) Secondary Growth: girth

• Lateral Meristems:
Mitotic cells “hips” of plant

Responsible for increases in stem/root diameter


Plant Growth
1) Indeterminate: Grow throughout life
2) Growth at “tips” (length) and at “hips” (girth)

Growth patterns in plant:


1) Meristem Cells: Dividing Cells
2) Differentiated Cells: Cells specialized in structure & role
• Form stable, permanent part of plant
Shoot apical meristem Leaf primordia

Young
leaf

Developing
vascular
strand

Axillary bud
meristems

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Meristems
• The tissue in most plants consisting of
undifferentiated cells (meristematic
cells), found in zones of the plant
where growth can take place.

• Meristematic cells are analogous in


function to stem cells in animals, are
incompletely or not differentiated, and
are capable of continued cellular
division.

• Furthermore, the cells are small


and protoplasm fills the cell completely. Tunica-Corpus model of the apical
meristem (growing tip). The epidermal

(L1) and subepidermal (L2) layers
• The vacuoles are extremely small. form
The cytoplasm does not contain the outer layers called the tunica.
chloroplasts although they are present The inner L3 layer is called the
in rudimentary form (proplastids). corpus.
Cells in the L1 and L2 layers divide in
• Meristematic cells are packed closely a sideways fashion which keeps these
together without intercellular cavities. layers distinct, while the L3 layer
divides in a more random fashion.
Plant Growth
Two lateral meristems: vascular cambium and cork cambium
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

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


Plant Growth
primary phloem
vascular cambium
Stem – Secondary Growth:
• thicker, stronger stems primary xylem

Vascular Cambium: between epidermis


primary xylem and phloem
pith
cortex
Produces inside stem:
primary xylem
A) Secondary xylem
- moves H2O, inward dividing
B) Secondary phloem vascular
- moves sugars, outward cambium

primary phloem
Vascular cambium
• Is a lateral meristem in the vascular
tissue of plants.

• It is a cylinder of unspecialized
meristematic cells that divide to give
rise to cells that further divide,
differentiate and specialize to form
the secondary vascular tissues.

• The vascular cambium is the source


of both the secondary xylem
(inwards, towards the pith)

• And the secondary phloem


– (outwards),

• And is located between these tissues


in the stem and root.
Vascular cambium
• Made from, procambium that remains
undifferentiated between the primary
xylem and primary phloem.

• Upon maturity, this region is known as


the fascicular cambium, and the area
of cells between the vascular bundles
(fascicles) called pith rays becomes
what is called the interfascicular
cambium.

• The fascicular and inter-fascicular


cambiums, therefore, represent a
continuous ring which bisects the
primary xylem and primary phloem.

• The vascular cambium then produces


secondary xylem on the inside of the
ring, and secondary phloem on the
outside, pushing the primary xylem and
phloem apart.
Vascular cambium
• The vascular cambium usually consists
of two types of cells:
– Fusiform initials (tall cells, axially orientated.
– Ray initials (almost isodiametric cells - smaller
and round to angular in shape).

Note:
• The vascular cambium is a type
of meristem - tissue consisting of
embryonic (incompletely
differentiated) cells from which
other (more differentiated) plant
tissues originate.

• Primary meristems are the apical


meristems on root tips and shoot
tips.
Vascular Cambium: primary
xylem
Secondary growth
new
secondary
secondary phloem xylem dividing
primary phloem vascular
cambium
primary xylem
secondary xylem new
secondary primary
vascular cambium phloem phloem

pith
cortex

Vascular cambium Growth


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

C X C

C
After one year After two years
C of growth of growth
Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings
Production of Secondary Xylem and Phloem
– The accumulation of this tissue over the years
accounts for most of the increase in diameter of a
woody plant.
– Secondary xylem forms to the interior and secondary
phloem to the exterior of the vascular cambium.
C=cambium cell
X=2o xylem
P=2o phloem
D=derivative
Cork cambium
• Another lateral meristem is the cork
cambium, which produces cork, part
of the bark. Growth
• Together, the secondary vascular
ring
tissues (produced by the vascular
cambium) and periderm (formed by Vascular
the cork cambium) makes up
the secondary plant body. ray

Heartwood
• Vascular cambia are found
in dicots and gymnosperms but Secondary
not monocots, which usually lack
secondary growth. xylem Sapwood

• In wood, the vascular cambium is the


obvious line separating the bark and
wood. Vascular cambium

Secondary phloem
• Capon, Brian (2005). Botany for
Gardeners (2nd ed.). Portland, OR: Bark
Timber Publishing
Layers of periderm
Cork Cambium
• The cork cambium is a lateral
meristem and is responsible for
secondary growth that replaces
the epidermis in roots and stems.

• It is found in woody and many


herbaceous dicots, gymnosperms an
d some monocots, which usually lack
secondary growth.

• Growth and development of cork


cambium is very variable between
different species, and is also highly
dependent on age, growth
conditions, etc. as can be observed
from the different surfaces of
bark:
– smooth, fissured, tesselated,
scaly, flaking off, etc.
Stem – Secondary Growth:

heartwood
(xylem)
sapwood
(xylem)
vascular
cambium
phloem
annual ring
Sapwood = Young xylem, water
late Heartwood = Old xylem, support
xylem
Seasonal Growth = annual rings
early Secondary phloem = grows outward
xylem older phloem crushed

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


Secondary Growth of a Stem

Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings


Recall
Primary growth is responsible for:
1. Growth in length of the stem
2. Produces the basic tissue pattern in the
stem

Secondary growth is responsible for:


1. Growth in girth of the stem
2. Produces secondary (new and more)
vascular tissues
3. Provides a continuous connection of
meristematic cells between the primary tissues
of the roots and the primary tissues of the
shoots.
Development of the Vascular Cambium

A. The function of the vascular cambium is to produce


secondary growth, thus the vascular
cambium must be formed before secondary growth can
occur

B. Two regions of the primary stem contribute to the


vascular cambium
1. Fasciscular cambium - the meristematic cells within the
vascular bundle
2. Interfascicular cambium - the meristematic cells
between the vascular bundles
The differentiation of the fascicular cambium

1. Not all of the procambium in the vascular bundle


differentiates into xylem or phloem
2. This undifferentiated procambium is called the
residual procambium
3. The residual procambium is 2-4 cells wide
and remains meristematic
4. The residual procambium will be called vascular
cambium when the vascular cambium begins to
divide to form secondary tissue
5. Thus, fascicular cambium is the regions of the
vascular cambium that originated within the
vascular bundle
Legend:

1 epidermis
3 primary phloem
4 fascicular cambium
5 interfascicular cambium
7 pith
2 cortex
6 primary xylem
Legend:

1 fascicular cambium
3 phloem
4 xylem http://www.vcbio.science.ru.nl/en/virtuallessons/fascicularcambium/
2 interfascicular cambium
The differentiation of the interfascicular cambium

1. This portion of the vascular


cambium originates in the pith
rays between the vascular
bundles
2. A band of parenchyma cells,
about 2-4 cells wide de-
differentiate and become
meristematic
Two kinds of initials in the vascular cambium

• Ray initials
*In xylem and phloem these are the
parenchyma cells
*small cells perpendicular to the
axis of stem
*from sclerenchyma cells or ray
parenchyma that radially divide the
stem (like slices of pie)
*rays elongate perpendicular to axis
to stem
*cells transport water and dissolved
solutes radially

http://www.cas.muohio.edu/~meicenrd/mudescd/dendrology/SCDRYGTH/VC.gif
• Fusiform initials
a. tapered, prism-shaped cell, periclinal division
b. oriented parallel to stem axis
c. produce secondary xylem and secondary phloem
between rays
d. conducting cells of this tissue transport water and
dissolved solute longitudinally
e. Produces the axial (vertical) transport system
• Generally the xylem-producing cells are
more active than the phloem producing
cells
• In temperate areas, the cambium is active
from the spring to the fall and is inactive in
the winter
• The yearly activity of the cambium
produces the annual rings in the xylem
Periderm (cork)
A. Function of the periderm:
– Increase in diameter of the stem occurs with the activity of the
vascular cambium
– This causes the protective epidermis to crack and split open
– Thus, a need for a meristematic layer at the outer edge of the
phloem for the internal protection of the stem
– Thus a layer of cork cambium forms outside of the phloem.
– The cork cambium forms a layer of waxy cork cells
– The cylinder of cork cambium increase in diameter as the stem
increases in diameter
Formation of the cork
In the young stem (1 year old or less)
a. Cortical cells just under the epidermis become
meristematic
b. Produces a layer 1-2 cells thick of cork cambium
(phellogen)
c. Phellogen produces a layer of cork cells 4-6 cells
thick external (toward the epidermis) to the phellogen
d. Phellogen produces a single layer of cells,
phelloderm, internal (toward the xylem and phloem) to
the phellogen
Structure of Periderm

a. Cells are flattened


b. Cell walls contain suberin,
a waxy substance
c. phellogen (cork cambium) +
phellem + phelloderm ⇒
periderm
d. periderm + primary phloem
⇒ outer bark
e. outer bark + secondary
phloem ⇒ inner bark
In old stems (more than 1 year old, generally 3-4 years)
a. A new phellogen forms because the former phellogen dies
b. The new phellogen forms in the outer region of the still-living
phloem
c. New phellogens will form about every one to four years depending
upon the species of tree
Bark and Wood
a. bark: aggregation of tissues outside vascular cambium
– As the layers of cells outside the vascular cambium die, they are
sloughed off as bark
– In the young stem the bark contains: epidermis, cork, cork
cambium, phelloderm, cortex, and phloem
– In the old stem the bark contains: cork, cork cambium,
phelloderm, and phloem

b. wood: aggregation of tissues inner to the vascular


cambium
WOOD
• xylem tissue
• Softwood - wood with only tracheids in it
(gymnosperm wood)
• Hardwood - wood with both tracheids and
vessels in it (angiosperm wood)
• Heartwood - wood in the center of the tree, no
longer conducting; darker
• - usually with tylose (balloon-like outgrowths from ray or axial parenchyma cells
through pit cavities in vessel wall): may serve as defense mechanism by inhibiting
spread of pathogen through the plant via xylem

• Sapwood - wood at the periphery of the stem,


actively conducting; lighter
External Features of
Woody Stems
Heartwood vs. Sapwood
• Heartwood- the part of
the wood in a living tree
that contains dead cells;
nonconducting wood.
• Sapwood- the part of the
wood in a living tree that
contains living cells and
reserve materials;
conducting wood.
Seasonal Growth Cycles
• Annual- a plant whose life cycle is
completed in a single growing season.
• Biennial- a plant whose life cycle is
completed in two growing seasons;
flowering and fruiting occurs in the second
year.
• Perennial- a plant whose vegetative portion
of the life cycle lives year after year.
Secondary Growth
• At the beginning of each growing season
primary growth is resumed and secondary
tissues are added.
• Secondary Growth- an increase in thickness
(girth) to the plant body as a result of the
activity of 2 lateral meristems:
– Vascular Cambium
– Cork Cambium
WOOD CUTS

Transverse section (c.s.)

section cut at right angles to the


main axis of stem or root;

cells of the axial system are cut


transversely and reveal their
smallest dimensions
• radial
– when stem or root are cut
lenghtwise, parallel to a
radius;
– Expose rays as horizontal
bands lying across the axial
system
– At its median plane, it
reveals the height of the ray
• Tangential
– when stem or root are cut
lenghtwise,perpendicular to
the radius;
– Cuts a ray perpendicular to
its horizontal extent and
reveals its height and width
– Easy to measure the height
of a ray, (usually done in
terms of number of cells)
and to see if ray is one or
more cells wide
Summary
Vascular Cambium
• Vascular cambium- a Tangential Section
cylindrical sheath of
meristematic cells that
produces secondary
xylem & phloem.
• Consists of two forms of
highly vacuolated cells:
– Fusiform initials
– Ray initials
• Vascular rays
– Pathways for
the movement
of food
substances and
water.
– Storage of
starch, protein,
& lipids.
Wood- Secondary Xylem
• Wood uses- shelter, fire,
weapons, furniture, tools,
paper, boats, wheels.
• Wood is classified as:
– Hardwood- magnoliids
and eudicots.
– Softwood- conifers.
Conifers- softwoods
• Tracheary
elements-
tracheids only.
Pinus- radial and tangential sections
Magnoliids and Eudicots- hardwoods
• Tracheary elements-
tracheids and vessel
elements.
Periderm
• Periderm- outer tissue that replaces the epidermis
as the protective covering of the plant.
– Cork cambium- meristem that produces the periderm.
– Cork (phellem)- secondary tissue that cuts toward the
outside of the cork cambium; dead at maturity;
suberin; impermeable to water and gases.
– Phelloderm- secondary tissue that is cut towards the
inside of the cork cambium; living at maturity; no
suberin; permeable.
• Lenticel- spongy regions on
the cork surfaces of stems,
roots, and other plant parts
that allow for gas exchange.
• Bark- collective term
for all tissues outside
the vascular cambium.
– Secondary phloem
– Periderm
Growth Rings Result from the Periodic
Activity of the Vascular Cambium
• Growth rings- a layer of growth in
secondary xylem or phloem.
• Annual rings- a growth layer that
represents one season’s growth.
• Early wood
– Less dense than late wood.
– Produced during period of rapid growth.
– Wide cells with thin walls.
• Late wood
– Dense.
– Produced during periods of slow growth.
– Narrow cells with thick walls.
Quiz
In diagrammatic form, summarize the
development of root and stem from
primary meristem to primary tissues to
secodary meristem to secondary tissue
Next meeting
• Long distance Transport in Phloem
(Translocation) and Xylem (Conduction)
• Practical Application of the Knowledge
on stem Structure and function
(Propagation: Grafting, Marcoting, Stem
Cuttings and Tissue Culture)