COURSE: BOCOR T205

Group B: PALYNOLOGY

UNIT 1: Spore-pollen morphology

units, polarity, symmetry, shape, size, aperture

NPC system for numerical expression of apertural details
Evolution of aperture types
 H. A. Hyde & D. A. Williams (1944) coined the term Palynology
but
the study of pollen analysis was started in 1916 by E. J. Lennart Von Post
The branch of science concerned with the study of pollen grains,
spores, and similar palynomorphs, both living and fossil
E. J. Lennart Von Post Harold Augustus Hyde
Angiosperm pollen

Gymnosperm pollen Pollen grains- male reproductive unit of the

flowering plants

Spores- various types of reproductive bodies

in non flowering plants formed by sexual
and asexual methods
Pteridophytic spore
SPORE-POLLEN MORPHOLOGY

•Pollen units
•Polarity
•Symmetry
•Shape
•Size
•Aperture
MICROSPOROGENESIS

A. Cross Section of anther showing

the pollen sacs with the
sporogenous tissue

B. Pollen mother cell

C. Meiosis stage

D.-H. Tetrad type

I. Polyad

J. Individual pollen grains

K.-P. Post meiotic stage

 POLLEN UNIT

Monad

Dyad Tetrahedral tetrad Tetragonal tetrad

(arranged in two diff. planes) (arranged in one plane)

Rhomboidal tetrad Decussate tetrad T-shaped tetrad Linear tetrad

(in 1 plane forming a rhomboidal shape) (pair-wise at right angel to other) (1st div. transverse, upper (Transverse div.)
and lower cells undergo vertical)

Polyad
Crypto tetrad (tetrad cells divide once or more) Pollinia
(Cell div. without wall formation; 3 obliterated) (whole contents of anther shed in a mass)
MONAD
DYAD
TETRAD
 TETRAHEDRAL DECUSSATE
TETRAD TETRAD

UNIT OF FOUR GRAINS IN WHICH THE

CENTRA OF THE GRAINS DEFINE A
TETRAHEDRON UNIT OF FOUR GRAINS
Drosera, Rhododendron etc. ARRANGED IN TWO PAIRS IN
TWO DIFFERENT PLAINS
(Magnolia grandiflora etc.)
TETRAGONAL
TETRAD

T-SHAPED
TETRAD
Typha latifolia etc.

LINEAR
TETRAD
Aristilochia, Polyanthes etc.

Mimosa pudica etc.

 POLYAD

Acacia auriculiformis,
Samania saman etc.

INTERCONNECTED
POLLINIUM LOCULIFORM POLLEN
Calotropis, Daemia etc. MASS

POLLINARIUM: Dispersal unit of pollinium plus secretions and/or tissues

that aid in the removal of the structure from the sower
 POLARITY
Polarity of individual pollen grains is expressed in terms of their
arrangement in the tetrad; end of the grains directed towards the centre of
the tetrad is proximal pole & towards the outside is the distal pole

Fig. A= Polarity in tetrad stage; B= The length of polar axis (PA): hypothetical line connecting two
poles; perpendicular to it is the equatorial axis; ED: Breadth of equatorial diameter in a monad
 POLARITY
Apolar spore: In individual spore, poles or polar regions can
not distinguished after separation from tetrad stage

Isopolar grain: If equatorial axis demarcates two equal polar

faces

Paraisopolar or subisopolar: slight differences exist between

the distal and proximal faces (equatorial plane is usually curved)

Heteropolar grain: If equatorial axis demarcates two unequal

polar faces so that two faces are distinctly different either in
shape, ornamentation of apertural system

Cryptopolar: Distal and proximal faces have dissimilar

sculpturing and having no tetrad mark (in some bryophyte spores)
Apolar Isopolar Heteropolar Heteropolar

Paraisopolar Paraisopolar Cryptopolar

 SYMMETRY
Asymmetric grain: It has no plane of symmetry
Nonfixiform: not having fixed shape
Fixiform: having fixed shape
Symmetric grain:
Radiosymmetric- having a radial symmetry; any plane including the
polar axis if passes through will produce identical halves-more than two vertical plane of
symmetry; isopolar grain: 1 horizontal, 2 or more vertical planes; heteropolar grain: no
horizontal plane of symmetry)
Bilateral symmetry- kidney/boat shaped: 1 vertical plane of symmetry;
isopolar: 1 horizontal & 2 vertical planes; heteropolar: 2 vertical planes of symmetry

Radial symmetry Bilateral symmetry

SHAPE
Outline of
view
Amb
it may be

NON
ANGULAR
circular, elliptic

ANGULAR
triangular or in
other geometric
forms
 SHAPE
Determination of Shape: the ratio between the polar
and equatorial diameters multiplied by 100
Shape classes (PA/ED) x 100

Per-oblate < 50
Oblate 50-75
Sub-oblate 75-88
Oblate-spheroidal 88-99
Spheroidal 100
Prolate- spheroidal 101-114
Sub-prolate 114-133
Prolate 133-200
Per-prolate > 200
 SHAPE CLASS

Peroblate Oblate Sub-oblate Oblate- Spheroidal

spheroidal

Prolate-spheroidal Sub-prolate Prolate Perprolate

 SIZE CLASS
Pollen Size Class (after Erdman 1945): expressed as length of the longest axis

SIZE CLASS LENGTH OF LONGEST AXIS

Very Small <10 µm

Small 10-25 µm

Medium 25-50µm

Large 50-100 µm

Very large 100-200 µm

Gigantic >200 µm
Measurement criteria:
For Size: PA, ED, EB for bilateral grains; PA & Greatest ED for radially
symmetrical grains in equatorial view; EB for radially grains in polar view

For Exine: thickness of exine; sexine/nexine ratio; thickness of the exine

projection if >0.5 µm
 APERTURE
A specialized region of the sporoderm, that is thinner than the remainder of the
sporoderm and generally differs in ornamentation and/or in structure.

In living pollen grains or spores the apertures usually function as

•sites of germination;
•they may also provide routes for transfer of water and other substances;
•play a part in harmomegathy

Simple Aperture: Apertures are described as

simple if they are present in only one wall layer,

Compound Aperture: if they affect more than

one layer of the wall. In compound apertures the
shape of apertures may differ between layers.

Ectoapertures occur in the sexine/ectexine,

endoapertures in the nexine/endexine, and
mesoapertures are sometimes found in an
intermediate position between an ecto- and
endoaperture.
STRUCTURE OF POLLEN WALL
 TYPES OF APERTURE
The term is often used in conjunction with a prefix or suffix, as
for example, ectoaperture, endoaperture, inaperturate,
omniaperturate, pseudoaperture, triaperturate.

Various types of apertures are recognised on the basis of their

shape

Simple
Lete: slit like; present on proximal end; 1: monolete; 3: trilete

Monolete
Trilete
Porus: circular or elliptic
aperture with a
length/breadth ratio less than
2

Ulcus: A rounded Porus

ectoaperture situated at the
distal or proximal pole of a
pollen grain. Examples:
Sparganium (Sparganiaceae),
Typha (Typhaceae). Pollen
grains are called ulcerate Ulcus

Ulculus: A rounded
ectoaperture not situated at a
pole. Example: Poaceae
Ulculus
Colpus: An elongated, aperture with a
length/breadth ratio greater than 2
Colpus
Margo (margin): An area of exine
around an ectocolpus that is
differentiated from the remainder of the
sexine, either in ornamentation or by
difference in thickness Margo

Sulcus: An elongated latitudinal

ectoaperture situated at the distal or
proximal pole of a pollen grain
Sulcus

A sulcus has the same shape as a colpus, but differs in

orientation. Sulci are essentially latitudinal apertures whereas
colpi are essentially longitudinal apertures
Trichotomosulcate: pollen grain with a
three-armed sulcus. Example: some
Palmae
Trichotomosulcate

Syncolpate: pollen grain with two or

more simple colpi; the ends of which
anastomose at the pole. Example:
Primula farinosa

Syncolpate
Parasyncolpate: syncolpate pollen
grains in which the apices of the colpi
divide into two branches and
anastomose towards the poles,
delimiting an isolated area known as
the apocolpial field. Examples:
Nymphoides peltata Parasyncolpate
Brevicolpate: brevi is a prefix for short; it is pollen grain with more
or less shorter length of colpi

Brevicolpate

Pseudoaperture: A thinning of the exine which, although

superficially resembling an aperture, is not associated with a
thickening of the intine and is presumed not to function as an exitus
(the site of exit of the pollen tube from the sporoderm)
pseudocolpus- colpus like pseudoaperture;
pseudopore- pore like pseudoaperture
Compound aperture: at ektexine- (ektoaperture-Colpi or pore
at endexine (endoaperture-ora)

Colporus: A compound aperture consisting of an ectocolpus with one or more

endoapertures; aperture type colporate; Asteraceae, Fabaceae etc.

Colporus

Pororus: pollen grain with compound apertures in which both the ectoaperture and the
endoaperture are pores and the two are not congruent.
aperture type pororate; Example: Myrica gale (Myricaceae)

Pororus
 EDGES OF APERTURES

Annulus
Distinct ring like
thickening/thinning of ektexine
(sexine) encircling a pore; Poaceae
Margo
Costa Distinct ring like
rib-like thickening/thinning that thickening/thinning of
occurs on endexine (nexine) ektexine (sexine) encircling
bordering aperture; Melia colpus; Cassia sophera

Arcus Laesura
Vestibulum a kind of
Oncus Laesura is the
Operculum a structure that
Aperture or at least thickening of scar mark of a
Circular, elliptical, occurs beneath the
their outer parts sexine in the spore. The mark
annular or bridge like
borne on small form of a band aperture of certain represents the
thick membrane either of unacetolysed pollen original contact
circular shield that occurs
ektexine (sexine), grains. lens- shaped of spores at their
shaped areas between the
endexine (nexine) or both structure, not
protruding as apertures of a tetrad stage;
covering the aperture; resistant to acetolysis;
round domes; pollen grain; trilete, monolete
Calectasia Corylus
Betula Alnus spore
 VIEWS OF GRAINS
A: Equatorial View

B: Polar View

Equatorial view : apertures are viewed meridionally i.e. pole to pole at right angles
to the equator. In this view, both PA & ED will be viewed & measured

Polar View : pollen or spores are viewed from one of the poles. In this view,
lete, sulcus, ulcus will be observed centrally. PA will not be viewed. Zonal apertures
– colpi, pore etc. will be viewed in the circumference.
 DIFFERENT TYPES OF AMB

Peritreme Goniotreme Pleurotreme Ptychotreme

(radiosymmetric, circular amb) (anguloaperture) (radiosymmetric, angular amb, (radiosymmetric, deep
mostly triangular) meridional fold, usually 3)

Angulaperturate Planaperturate Sinuaperturate Fossaaperturate

(at angles of amb; side may be(at midpoint of the sides)(equally half between the
(at ditch like indentation
Straight, concave, convex) angles; sides concave) between lobes)
 NPC classification
A classification based on the number (N-single, two or many), position (P: polar: distal or
proximal; global; meridional) and character (C: circular or elongated) of the apertures of pollen
grains and spores for identification proposed by G. Erdtman (1969)
treme (aperture) is used for preparing keys; cata: aperture in proximal face, ana: in distal face; anacata: in both faces; zone:
equatorial zone; dizono: equatorial zones but in two rows; panto: all over the surface; c0 character is not known; c1: aperture like
thin area or leptoma; pollen with one leptoma-monolept; c2: three slit like colpus-trichotomocolpate; c3: colpa; c4: pore; c5:
colpa+ora; c6: pore+ora
 Numerical expression of apertural details

Numerical value: Pollen type Example

244 Dizonoporate Tofieldia

344 Trizonoporate Betula
443 Tetrazonocolpate Hippuris
445 Tetrazonocolporate Rumex
544 Pentazonoporate Alnus
764 Polypantoporate Chenopodium
 Evolution of aperture types
POSITIONAL CHANGE OF APERTURE

Bryophytes, Pteridophytes: Aperture (either trilete or monolete) is proximal

in position

Primitive extinct gymnosperm like pteridosperm (Lyginopterids-trilete,

Medullosans-monolete), Cordaitales (trilete), voltziales (trilete): Pre-pollen
which is characterized by proximal aperture presuming proximal
germination rather than distal as in case of other seed plants.

Extant gymnosperm: either inaperturate or if aperture present, it is in

distal position

Angiosperm: along with distal and zonal position; global distribution of

apertures are also found which increase the probability of pollen
germination in any part of the pollen surface and its adherence to
stigmatic surface may accelerate pollen germination

Positional change of aperture from proximal to distal, zonal, global:

required to meet up the physiological and ecological needs that occurred
due to diversification of new plant taxa.
APERTURE TYPES IN EXTINCT PRE-ANGIOEPERMIC PLANTS
Mid. Silurian to Early Devonian vascular Most primitive
plants- Rhyniales, Zosterophyllales, trilete spores
Trimerophytales, Lepidodendrales,
Sphenophyllales, Calamitales, Primofilicales,
some Cycadeoids
Pteridosperms, Cordaitales, Voltziales Pre-pollen
Bennettitales Monosulcate pollen
APERTURE TYPES IN EXTANT PRE-ANGIOEPERMIC PLANTS
Lycopsids & some members of Filicales Trilete spore
Psilotales & some members of Filicales Monolete spore
Equisetales Inaperturate spore
Cycadales, Coniferales, Ginkgoales Monosulcate pollen
As such, pre-angiospermic plants have following morpho-
forms:
a. Basic form-trilete, non-perinous, non saccate
b. Derived form-i. Trilete, perinous non-saccate
ii. Trilete, saccate
iii. Monolete, non perinous, non saccate
iv. Monolete, perinous
v. Monosulcate, saccate
vi. Inaperturate

Perine (occurs especially in certain pteridophytes. It seems to develop after the exine proper and forms usually
and saccus (occurs in most of the gymnospermous
wrinkled, loosely attached covering of the mature spore)
pollen grains; winged like expansion made as a windborne mechanism) developed later during the
morphological evolution of spore and pollen which indicates-

Origin of monosulcate and inaperturate forms – through reduction of

the basic structural organization i.e. trilete

Origin of perine and sacci - through modification in form of expansion

of the basic structural organization
Angiospermic Plants Clavatipollenites
Earliest record: Lower Cretaceous
Most primitive angiosperms: monosulcate aperture (pollen grains were
named as Clavatipollenites, Retimonocolpites, Liliacidites – these may
have originated from a common Mesozoic ancestor Benettitales)

This monosulcate aperturate condition is distributed among

followings:
• Most gymnosperms: Cycas (Cycadales); Pinus, Abies, Cedrus
(Coniferlales); Ginkgo (Gikgoales)
and
• Primitive dicotyledonous angiosperms: Magnolia, Degeneria etc.
and
• Many monocotyledonous angiosperms: members of Arecaceae

Distal sulcate aperture then modified to a few other types –

• distal porate (some monocots-poaceae);
• distal trichotomosulcate (both in dicots & monocots);
• equatorial ring like aperture (Nymphaeaceae);
• Inaperturate (by having reduced thin exine and thick intine)
As such, the main trend of evolution of pollen aperture-

• Transformation of the distal monosulcate (monocolpate)

into tricolpate form. Takhtajan (1980) hypothesized that
tricolpate condition developed as a result of evolutionary
deviation of the primitive stage of sporoderm development
and this is derived form of monocolpate condition

• Tricolpate type independently given rise to other types

namely polycolpate, triporate, pantoporate

• Highest stage of evolution of pollen aperture: compound

apertures like tricolporate, or tripororate etc. are also
derived from simple tricolpate type
EVOLUTION OF APERTURES IN POLLEN (Takhtajan 1980)