Leaves

Origin
Differentiation
Leaf structure of lower vascular plants,
gymnosperms, angiosperms
C3, C4 plants
 Enations/microphylls – evolved as simple
outgrowths from stem; vascularized by a
single vein; in stems, with no leaf gaps

Macrophyll - larger leaves; more

complex of veins; in stems, with
leaf gaps
Gaps do not
constitute breaks in
the continuity of
axial vascular system.
Lateral connections
occur between the
tissues above and
below the gap.
 Lateral leaf
unilacunar trilacunar trace

Unilacunar, trilacunar, multilacunar - depending on whether

a given leaf is associated with one, three or several gaps at a node.

If leaf has more than one leaf trace, and is confronted by 3 or more
gaps, the gap (and associated trace) in the median position with
reference to the leaf is called median, the others are called laterals.
 Recall
• As shoot apex grows, older leaf
primordia are left behind.
• New primordia are continuously borne
above existing ones.
• In angiosperms specially, as young
primordium develops from SAM,
(grows into radially symmetrical lateral
outgrowth), preparations are under
way for emergence of the next
primordium
• The Initiation of a leaf primordium is associated
with an increase in the frequency of periclinal
divisions at the initiation site
1. In the eudicots and monocots with a two-layered tunica, the first
periclinal divisions occur most frequently in the L2 layer followed by
similar divisions in the L3 layer and by anticlinal divisions in the L1
layer
2. In some monocots, leaf primordia are initiated by periclinal divisions
in the L1 layer. Periclinal divisions then spread laterally in both
layers, forming a ring that encircles the meristem.

Thus leaf primordia are initiated by groups of cells that span

two or more cell layers in the meristem.
 The total number of cells involved has been estimated
– about 100 in cotton, Impatiens, to 250 in maize 30 in
Arabidopsis.
• Immediate precursors of the leaf primordia—are referred to
by some workers as founder cells
 procambium
Either concomitant with or preceding the periclinal divisions
associated with initiation of a leaf primordium, one or more
procambial strands (leaf traces), which will differentiate upward
into the developing leaf, may already be present at the base of
the leaf site
 SAM - a small mound
before initiation of
leaf primordium

Sam widens

Leaf buttress
initiated
on the sides
Early stage of
leaf pair
development
 Differentiation of leaf cells : Mesophyll
•Different cells and tissues contain different proteins

Example: difference between an epidermal cell and mesophyll

cell = Leaf mesophyll have high levels of proteins essential parts
of photosynthetic apparatus such as:
–Rubisco
–Chlorophyll a/b-binding proteins
 Differentiated plastid
• As the chloroplastid differentiates, internal
membranes (thylakoids) are not connected with the
plastid.

• The internal structure of the chloroplast is

complex.The stroma is traversed by an elaborate
system of thylakoids.

• Thylakoids consist of grana—stacks of disk-like

thylakoids that resemble a stack of coins—and
stroma thylakoids (or intergrana thylakoids) that
traverse the stroma between grana and interconnect
them.

• Grana and stroma thylakoids and their internal

compartments are believed to constitute a single,
interconnected system. The thylakoids are not
physically connected with the plastid envelope but
are completely embedded in the stroma.

• Chlorophylls and carotenoid pigments—both of which

are involved in light harvesting—are embedded,
along with proteins, in the thylakoid membranes in
discrete units of organization called photosystems.
 enation microphyll
enation

Enation (prophyll) - leaflike with no vascular tissue

Microphyll - small leaf, simple vascular system

*At the node of a microphyllous shoot a single strand of vascular tissue
called a leaf trace diverges from the edge of stele or vascular cylinder
Microphylls Megaphylls
- leaves of small size - leaves of generally larger size
- with simple venation (one vein) - with complex venation
- associated with steles that lack leaf gaps - associated with leaf gaps in the stele
(protosteles)

17
Microphyll

18
Evolution of macrophyllous leaves
According to telome theory:
1.Change from dichotomous to sympodial branching in an erect leafless shoot
resulting in an axis bearing a succession of telome trusses.
Telome truss - consists of: telomes (ultimate dichotomies) and mesomes
(internode below telomes)
2. Orientation of members of a truss was that telome trusses were in one plane
3. Fusion of members lateral to one another resulted in a leaf composed of a
petiole and a blade with a dichotomously branched, freely ending vein
system.
20
21
Pendant aerial branches of
Tmesipteris

Pendulous epiphyte
Larger leaves supplied
with single, unbranched
vein or unbranched
vascular bundle
 Selaginella
• Foliage leaves small, simple and symmetrical or asymmetrical
in outline

Leaves are
microphylls
 • Mesophyll between veins and epidermis contain chloroplastids
and have prominent intercellular spaces. According to the
species, each mesophyll cell may contain just a single
chloroplast.
• Stomates may be found on both surfaces.
Intercellular spaces

Spongy cells

Annular and
spiral
vessels

Epidermis
Selaginella kraussiana
Lycopodium

• ~ 2-3 cm, lanceolate,with broad base

• has single unbranched vein running from
base to apex
• single vein connects to a leaf trace
• xylem consists of few thick-walled
tracheids surrounded by parenchyma
• no leaf gaps
Lycopodium with unbranched vein Spirally
Spongy cells thickened
stomate
Mesarch bundle tracheids

• Well-developed epidermal layer

• Stomates in both surfaces or in some species in only one
surface
• Isodiametric cells with conspicuous intercellular spaces
28
29
Equisetum

At each node is a whorl of small slender, scalelike leaves, laterally united
with one another to form a brownish sheath around the node
Each leaflet has one trace which
connects with the stele of the
stem
FERNS
Leaf venation
in ferns

Reticulate venation
33
Form and organization of fern fronds

A. Compound pinnate
B. Tripinnate
C. Bipinnate
D. Once/simple pinnate
E. Simple leaf with
open
dichotomous
venation
F. Simple leaf with
D pinnate
reticulate venation
Fern fronds

Pinnules
(subdivisions
of a pinna)
fiddlehead
 Fern leaf with sorus
Each sorus has a
central axis to which the
sporangia are attached;
there is a common
covering underneath the
sporangia, and it is
called an indusium.
Each sporangium is
encircled by a thick
walled ring of cells
called the annulus. The
annulus is hygroscopic
(readily absorbs and
annulus indusium retains water);
Fern pinna with sorus

photosynthetic cells

epidermis

annulus

spores
•

upper
epidermis
palisade layer
Intercellular space
spongy cells
phloem
xylem
stomate
Lower
epidermis
X-section of fern pinna
 Pine seeds

Female cone

Mature female cone

Male cone
 Pine leaf
Epidermis with thick periclinal walls and cuticle
Hypodermis-thick-walled fibers
Resin duct
Lobed mesophyll with
chloroplast
endodermis
Xylem with tracheids
Phloem with albuminous
cells
Transfusion tissue
Guard cell of sunken
stomate

Substomatal cavity
Amphistomatic- stomates occur on all surfaces of the leaf
Details of pine mesophyll
guard cells of
sunken stomate

hypodermis (thick-
walled)
lobed mesophyll
cell

Infolded cell walls

44
45
46
 Transfusion tissue
▪Closely associated with the
vascular bundles
▪With 2 principal types of cells

Cells of transfusion tissues:

1.Transfusion tracheids - thick, lignified secondary walls with circular bordered
pits
2.Transfusion parenchyma - produce tannin-like substances
Both types in direct contact with the endodermis.
Function: conduction of materials between the vascular bundle and the mesophyll.
X-section of Cycas leaflet
Cycas leaf: x-section of a part of leaflet

epidermis

hypodermis
chlorophyll-bearing
palisade layer

Middle elongated
Colorless cells

Few layers of
mesophyll
50
 Leathery simple leaves, well-developed
Gnetum opposite leaves, pinnately net-veined
52
53
X-section of midrib in Gnetum gnemon

Separate collateral
bundles; tracheids and
vessels present in
xylem

xylem
phloem
55
gymnosperms

Vascular bundles in the midrib of Gnetum

vessels
auricle
Simple
leaves
Three axes
Adaxial-abaxial axis
Parallel venation
Cross-section of a dicot leaf
Reticulate/netted venation
Thin part
of the
blade

midrib
62
Dorsiventral Dicot leaves- palisade layer occurs
adaxially, spongy layer occurs abaxially
epidermis collenchyma
parenchyma
Palisade layers

xylem
phloem

Spongy layer
DORSIVENTRAL

64
ISOBILATERAL

65
Isobilateral leaf: A leaf in which the palisade
parenchyma occurs adaxially and abaxially
beneath the epidermis of the leaf

Palisade layer

Spongy layer

Palisade layer
Bundle sheath

Mesophyll ce
68
Kranz type of leaf anatomy “wreathlike

Single layer of
chloroplast –
bearing bundle
sheath

radially-oriented
mesophyll cells
69
Fig. 10-19b
The C4 Mesophyll
pathway cell CO2
PEP carboxylase

Oxaloacetate (4C) PEP (3C)

ADP
Malate (4C) ATP

Pyruvate (3C)
Bundle-
sheath CO2
cell

Calvin
Cycle

Sugar

Vascular
tissue
71
Development of a dicot leaf

Leaf arises as a
dorsiventral
Protuberance that
grows in height and
width.
Procambium present
before leaf primordium
has emerged as
protuberance.
Origin of leaf from SAM
 Ontogeny of a monocot leaf.

Collar or sheath
•

A. emergence of protuberance on the side of apical meristem

B-D. extension of protuberance laterally around apical meristem that forms
a collar or sheath
E. Enclosure of apical meristem and elongation of blade or sheath by
intercalary meristem
Monocot leaf- corn leaf
78
Monocot leaf Bundle sheath or
border parenchyma

xylem

phloem

Photosynthetic
cells
In times of water stress bulliform cells lose water allowing the
leaf to roll up. This results in a small evaporative surface area
being exposed to the atmosphere. As a result, a slower rate
of transpirational or evaporative water loss.