Secondary Growth

Roots and leaves together are sufficient to take up all

essential resources, so why make stems?

Stem functions
1. Support leaves
2. Conductance (connect root and leaf vasculature)
3. Storage (some species)
4. Photosynthesis (some species)

But lots of plants don’t have any stems at all – if not

required, why “waste” the resources?

While all the above functions are

important, the most general benefit
is improved light environment for
the leaves – especially due to
competition between plants.
 Shoot growth – primary plant body

Shoot – above ground plant structure

stems, branches, leaves

Apical Meristems make primary shoot

increase in length

Lateral Meristems make secondary growth

increase in girth

35.11/35.10
Apical Meristem makes
primary shoot
1. stem growth and tissues
2. leaf primordia – become
leaves
3. bud primordia – become
lateral branches

35.16/35.15
This process
creates the primary
plant body with
it’s node –
internode structure

Node – leaf/branch
attachment

Internode –
between nodes

35.2
Apical dominance
Apical meristem at the leading shoot tip
inhibits bud primordia (lateral buds) nearer to the tip,
releasing them later (farther) resulting in “Christmas
tree” like growth form.

39.9
 35.17/35.16
Stem sections show all three main tissue types
vasculature - often in bundles or sometimes rings
epidermis
ground tissue – often called cortex, or “pith” if inside
 Modified Stems
Tendrils and twining stems
Thorns (vs. spines)
Stolons – above ground runners
Rhizomes – below surface runners
Food storage
Tubers – swellings of stolons & rhizomes
Corms – swellings at base of stem
Water storage (succulence)
Cacti – stem modified for water storage and
photosynthesis (leaves are spines). Two other families,
Spurge (Euphorbiaceae) and milkweeds (Asclepiadaceae)
have also evolved this. Classic example of convergent
evolution.
 SECONDARY GROWTH

Secondary growth – increase in girth

(width) of stems and roots resulting
from lateral meristems

[aside]
Simple non-rigorous categories, widely used
1.Woody plants – with 2o growth – trees and shrubs
2. Herbaceous plants – little or no 2o growth, “herbs”
a. Grasses
b. Forbs (herbaceous dicots)
Two main Lateral Meristems control secondary growth
1. Vascular cambium – makes 2o xylem and phloem
2. Cork cambium – makes periderm

In the secondary plant body, these form concentric

cylinders or sheaths (rings in cross section) of meristem

35.11/35.10
Vascular Cambium
Fusiform initials – key meristematic cells, vertically
elongated. They produce xylem cells to the inside and phloem
outside, causing increase in girth.
Vascular cambium
(cross-section) phloem
xylem

35.20/35.19
 Wood – secondary xylem

Xylem – remains in place and

continues to function for many
years. Rigid cells build on one
another expanding the stem.

35.22/35.20

Eventually the tree may stop using the inner xylem.

1. Sapwood – outer conducting xylem
2. Heartwood – inner older xylem, no longer
conducting. Often darker due to deposits into
vessels to block conductance and pathogen transmission
Growth rings

In seasonal climates (cold/warm,

wet/dry), early season growth, “early
wood”, is less dense (larger cells,
thinner walls) than “late wood”,
producing annual “rings” visible to
the eye.
Used to create detailed
climate records.

Bristlecone pine can live

4900 years. Used to
reconstruct climate to 8000
years ago.
 Phloem – usually only the current year phloem
functions in sugar transport. It is eventually pushed
outward and collapses due to the expanding stem.
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

35.19/35.18
 Cork Cambium – the Periderm
A second sheath of meristem develops in the secondary
phloem, outside the vascular cambium, called the cork
cambium. It produces the periderm, which replaces the
epidermis in secondary growth.
Cork cells – main component of the
periderm, produced by the cork
cambium to the outside. They are
lined with suberin and dead at
maturity. Impermeable to water (and
gases).

Lenticels – openings in the cork layer to

allow the living cells of the stem to respire
 Bark – all tissue outside the vascular cambium
-what can be removed without killing the tree

As the tree expands, this tissue is stretched and

eventually split and sloughed off. Structure of
periderm determines the pattern.
Cork oaks in Portugal – a renewable resource
 Cork cambium

This figure nicely summarizes the structure of a woody stem

 xylem phloem

tylose

Spring wood Summer wood bark cork

 tylose
Pith rays
How many years’ growth?
cork

phloem

xylem

heartwoo

sapwood
 • Most monocots do not have secondary growth.
• However, some, like palms, produce new vascular bundles
and ground tissue, but not wood.

• Palms have diffuse secondary growth.

• This does not involve a vascular cambium, and no new vascular tissue is formed.
• Instead, parenchyma cells in the ground tissue divide and enlarge, increasing the girth of the
stem.
END