Lecture 9

Pollen Grains
Seed bearing
In the foregoing kinds of life cycle, fertilization still
requires
q water to bear the sperm p on its uncertain travels.
Pressure to occupy ground during the late Paleozoic,
however, encouraged some ferns to short-circuit the life
cyclel slightly.
li htl Thi
This was achievedhi dbby iimprisoning
i i th
the
megaspore within the megasporangium and letting it
undergo its development there in the confinement of the
ovule. Development of the spore was also telescoped, so
that it divided into a tiny, y few-celled p prothallus before
liberation. This multicellular spore is the pollen grain.
The function of a pollen grain, like a spore, is one
off dispersal
di lb
butt itits aim
i iis tto reach
h th
the ffemale
l cone di
directly
tl
and to proceed with fertilization varies between the two
main groups of seed bearers bearers, the gymnosperms and the
angiosperms.
 Life cycle of seed-bearing plants

Function of pollen grains: (male sex organs)

The function of pollen grains is to accomplish the transport of the male
gametophyte to the female flower so that fertilization can take place.
Gymnosperms
The gygymnosperms
p ((i.e. conifers and their allies)) bear more
or less exposed megasporangia in cones or in flower-like
structures. The pollen grains alights on the ovule and
grows a long
l pollen
ll tubet b tot connectt withith the
th egg cells
ll via
i
a slit called micropyle. Sperm cells then travel down this
tube and fertilization takes place leading to the growth of a
young sporophyte (i.e. seed) within the cone. This seed is
provided with a p
p protective and nutritious covering g and is
now the main mode of propagation and dispersal. The first
seed-bearing gymnosperms plants (seed ferns) appear to
h
have evolved
l dbby mid-carboniferous
id b if titimes, although
lth h seed-
d
like structures are known in upper Devonian rocks.
Angiosperms

The angiosperms (i.e. flowering plants) have gone a stage

f th to
further t protect
t t the
th female
f l ovule,
l by
b enclosing
l i it within
ithi an
outer jacket called a carpel. This passes upwards into a
prominent projection called the style,
style with a terminal
knoblike stigma to catch the passing pollen. The
microsporangia (anthers) are also borne on stalks, called
stamens, and liberate their pollen to the wind, to water, or
to visiting insects. Pollination involves the growth of a
pollen
ll ttube
b d down ththe style,
t l th
throughh th
the micropyle
i l tto th
the
ovule. After fertilization, the seed is able to develop,
protected from the climate and from ravaging insects
insects. Such
flowering plants are known first in lower Cretaceous rocks
and by late Cretaceous times were a principal part of the
land flora.
Pollen Morphology and Terminology
Pollen grains are distinguished primarily by their form,
aperture and their surface sculpture as follows:

Aperture: Aperture is an opening or thin part of the spore or

pollen coat which permits germination. Apertures are
described with regards to position, number and shape or
character.
1 Pollen in groups of four grains..TETRADS
1. grains TETRADS

2 Pollen in groups of more than four grains..POLYADS

2. grains POLYADS

3. Pollen grains free:

3.1. With bladders.............................VESICULATE

3.2. With deep meridional ridges..............POLYPLICATE

3.3. With No distinct apertures.....................INAPERTURATE

4. One aperture:
4.1. Aperture elongate.....................MONOCOLPATE

4.2. Aperture +/- circular................MONOPORATE

5. Two apertures:
5 1 Two furrows…………….
5.1. furrows DICOLPATE

5.2. Two pores………………..DIPORATE

6. Three apertures:
6.1. Three furrows..................TRICOLPATE

6.2. Three pores.......................TRIPORATE

6.3. Three colpi + pore............TRICOLPORATE

6.4. Apertures fuses to rings etc. ...........SYNCOLPATE

7 More than three furrows:

7.
7.1. All furrows meridional....STEPHANOCOLPATE

7.2. Furrows not meridional....PERICOLPATE

8. More than three pores:
8.1. Pores in an equatorial zone....STEPHANOPORATE

8.2. Pores evenly distributed.........PERIPORATE

9. More than three colpi + pores:

9.1. Colpi
p + pores
p in an equatorial
q zone…….STEPHANOCOLPORATE
9.2. Colpi + pores evenly distributed………. PERICOLPORATE
 POLLEN CLASSES
GYMNOSPERM POLLEN CLASSES
VESICULATE: with hollow vesicles (alveoli) (often two) (Pinus, Abies, Picea)

POLYPLICATE: football shaped with longitudinal ridges (Ephedra)

•INAPTERTURATE: without an opening

p g (Juniperus
p )

MONOCOLPATE: (Cycas):
ANGIOSPERM POLLEN CLASSES

•DYAD: pollen grain comprised of 2 cells

•TETRAD: pollen grain comprised of 4 cells (Ericaceae)

•POLYAD: pollen grain comprised of 8 or more cells (Acacia)

ANGIOSPERM APERTURE TYPES

INAPERTURATE without an opening (Populus)

(FORATE = with apertures)

PORATE: p
polar distance = equatorial
q distance (Betula))
COLPATE: polar distance > equatorial distance Acer

COLPORATE: pore in (equatorial plain of) furrow Salix

•HETEROCOLPORATE
HETEROCOLPORATE (3) furrows
f with
ith pores alternating
lt ti with
ith (3)
•furrows without pores. Examples: heliotrope (Phacelia),
•dwarf mistletoe (Arceuthobium)

FENESTRATE: large openings (fenestra, lacunae) in wall

(Li lifl
(Liguliflorae, Pl t
Platyopuntia
ti )
CLASSIFICATION OF FORATE POLLEN:
POLLEN Aperture
A t Number
N b and
d Arrangement
A t

1. Mono-: one aperture (furrow or pore)

2. (Di:) two apertures

3. Tri-: three apertures

4. Stephano-: more than three apertures on equatorial plane

5. Peri-: more than three apertures over the grains surface

6. Syn-: a single aperture (fused apertures) encircling the grain

 Pollen Shape & Contour
P

3
8 E 8 8 E
4 6
3
Peroblate 6
8 Oblate Suboblate

Spheroidal
p

6 8 4 8 3

88 8 8

6 4 3

Subprolate Perprolate
Prolate
Shape of pollen grains depend on the polar/equatorial ratio (P/E) as follows:

Shape classes P/E

Perprolate < 4/8
Oblate 4/8 – 6/8
Subspheroidal 6/8 – 8/6
Suboblate 6/8 – 7/8
Oblate spheroidal 7/8 – 8/8
Prolate spheroidal
p 8/8 – 8/7
Subprolate 8/7 – 8/6
Prolate 8/6 – 8/4
P
Perprolate
l t > 8/4
Sporoderm Stratification

The pollen wall is designed to protect the sperm nucleus from

desiccation and irradiation during transport from the anther to
the stigma. The tiny (20 - 100 µm) pollen grain is coated with
waxes and p proteins held in place
p y sculpture
by p elements. The
wall of pollen grain is constructed of two layers: an outer
resistant exine, which is made of the biopolymer sporopollenin
and an inner cellulosic intine immediately surrounding the
living protoplasm. If fertilization does not occur, the protoplasm
and intine quickly decay, leaving only the outer resistant layer,
the exine,
exine which can be fossilized
fossilized. Escape of the pollen tube
through the wall takes places through apertures, which further
facilitate shrinking and swelling.
1. Exine:
Outer granular (atectate-primitive) or baculate (tectate-advanced)
layer. It may be differentiated into an outer sculptured
(ornamented) layer, the Sexine and a structureless inner layer, the
Nexine. The sexine being g usually y sporopollenin
p p in nature,, it resists
acetolysis, stains readily with basic Fuchsin and does not take
ruthenium red. It dissolves in 2-aminoethanol. It protects the pollen
grain against desiccation and injury
injury.

2. Intine:
Innermost transparent thin layer
layer. Being cellulosic it does not resist
acetolysis, does not take Fuchsin and takes ruthenium red. Usually
even and closely invests the protoplasm, protecting it during
germination.
i ti
 SCULPTURING TYPES

WITHOUT SCULPTURING ELEMENTS

Psilate without sculpturing ELEMENTS Pseudotsuga
ELEMENTS ISODIAMETRIC
ELEMENTS <1µ
ƒScabrate with small (<1µ) isodiametric ELEMENTS
ƒPopulus
P l aspen - cottonwood
tt d

ELEMENTS >1µ
ƒGemmate
G t ELEMENTS >1µ, h i l Juniperus
>1 + spherical J i j i
juniper

ƒClavate ELEMENTS club-shaped, taller than wide

ƒGeranium geranium

ƒVerrucate ELEMENTS >1u, + isodiametric, not

ƒconstricted at base Quercus oak
 ƒBaculate ELEMENTS taller than wide, end rounded
ƒCucurbita squash

ƒEchinate
E hi ELEMENTS taller
ll than
h wide,
id end
d pointed
i d
ƒCompositae sunflowers

ELEMENTS ELONGATED PARALLEL TO SURFACE

Rugulate ELEMENTS irregularly distributed Erodium filaree

ƒStriate ELEMENTS regular

regular, parallel Mentzelia blazing star

ƒReticulate ELEMENTS forming a net-like pattern

Cercidium paloverde
Salix Cercidium Xanthoxylum
y Erodium