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A natural system of classification should be based on the analysis and harmonization of evidence from
all organs, tissues and parts.
Traditionally, plant taxonomy has been mainly dependent upon comparative external morphological
characters.
Morphological characters of plants have been used extensively both for classification and for diagnostic
purposes and they are still indispensable to the taxonomists.
However, external morphology study alone is not adequate and other branches of study are of
considerable value in proper assessment of the systematic status of a taxon and its phylogeny.
Thecontributions to systematics can come from any branch of botany viz.,Anatomy, Palynology,
Cytology, Embryology, Phytochemistry, Chemotaxonomy, Molecular Systematics and they have
played significant role in plant taxonomy.
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PALYNOLOGY(Palynology in relation to Taxonomy)
PALYNOLOGY IS THE STUDY OR SCIENCE OF POLLEN GRAINS AND SPORES.
The term Pollen is derived from Greek verb PALYNEIN means “to scatter”.
Description of the fundamental features of pollen morphology by early botanists has demonstrated the
potential value of palynology in Systematics and Phylogeny. The contribution of Woodhouse (1935)
through thepublication of the book “Pollen Grains”, marked the beginning of purposeful studies on
angiosperm pollen. Eardtman’s (1952) book “Pollen Morphology and Plant Taxonomy in
Angiosperms” laid a solid foundation of Palynology and demonstrated the importance of pollen
morphology in plant taxonomy. Erdtman, G (1969) proposed NPC system of pollen classification
based on Number, Position and Character of the pollen apertures.
In India pioneer workers in the field of Palynology are Birbal Sahni, P. K. K. Nair, Vishnu Mittre and
S. K. Srivastava. P.K.K. Nair considered as Father of Indian Palynology.
The main centres of research in Palynology in India are – National Botanical Garden (NBG), Lucknow;
Birbal Sahni Institute of Paleobotany (BSIP), Lucknow and Bose Institute, Kolkata.
Pollen grains are found in every nook and corner, e.g., in glacier ice, in the air over the poles and over
the Oceans.
Fossil spores are found in peat and other sediments, in lignite, coal and shales. They are evident since
Pre-Cambrian Times, hundreds of millions of years ago.
The Exine of Pollen grains are extremely resistant andremain as such for a very long period.
Therefore, the data provided by them as fossil has been used successfully in the interpretation of the
past vegetation. Analysis of fossil pollen is the most important approach to the reconstruction of
past flora, vegetation and environment.The ability to identify plants on the basis of their pollen
morphology has enabled Palaeobotanists and Ecologists to reconstruct past assemblages of plants
and identify periods of environmental change.
Aeropalynologicalstudies (Pollen in the air) of any locality can be used to get a picture of the local
flora. Pollen grains are studied after Acetolysis (using a mixture of Acetic anhydride and concentrated
H2SO4 in the ratio of 9:1).Pollen morphology has played an important role in identification and
splitting of genera.
Pollen characteristics such as Shape of pollen, Polarity, Symmetry, Size, Number of apertures,
Aggregations and Exine (sporoderm) have been used in taxonomic botanyfor a very long time.
Modern taxonomy has been shown to benefit much from external and internal characteristics of pollen
grains. The taxonomic and evolutionary importance of pollen grains at specific, generic or even higher
levels are significant.
The application of scanning electron microscopy (SEM) has proved to be a useful tool fpr
palynological studies with increased accuracy and precision. This has proved new opportunities for
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better understanding of the exine ornamentation patterns. It enabled application of exine features in
studies involving systematic relationships of microtaxa, particularly subspecies, varieties, cultivars,
cytotypes, bioforms, etc.
The size and shape of pollen, pollen characters such as number and position of furrows, number
and position of apertures and details of sculpturing (The external features of the pollen grain wall
are termed as Pollen sculpturing) of the exine are of taxonomic value.
The form, number, distribution and position of apertures are important palynological criteria in
assessing relationships and phylogeny of plants.
According to the position, the aperture may be proximal, distal, and zonal. In terms of evolution,
the proximal position of aperture is most primitive and zonal position of aperture is most
advanced.
Pollen polarity is the spatial orientation of the pollen grains leading to the formation two distinct poles.
1. Polarity: It is expressed in terms of their arrangement in the pollen – tetrads. The end directed
towards the centre of the tetrad is the Proximal end orPoleand that towards the outside, is the
Distal pole. The hypothetical line connecting the two poles is the polar axis and the axis
perpendicular to it, is the Equatorial end.
2. Apertures: Pollen lacking apertures are called Inaperturate, while with apertures as
Aperturate. Most pollen grains are aperturate. Apertures vary in form, number, position and
distribution on exine surface. Apertures when alongated or elliptical are called Colpate, when
circular, Porate. The margins of apertures may be thin (Tenuimarginate) or thick
(Crassimarginate).
3. Pollen size and Shape: In consideringthe size, the polar diameter (P) and the equatorial
diameter (E) of pollen are taken into consideration, shape is expressed in terms of their
respective sizes. Shape classes have been decided, on the basis of the formula P/E x 100.
50 to 75 Oblate-sphaeroidal
76 to 100 Oblate
101 to 113 Prolate – sphaeroidal
\ 114 to 133 Subprolate
134 to 200 Prolate
>200 Perprolate
In addition to this, there may be Bilateral pollen grains which can be Planoconvex, Biconvex,
Concavoconvex etc.
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POLLEN CHARACTERS CONSIDERED FOR TAXONOMIC ANALYSIS ARE:
1. Pollen unit type, 2. Pollen grain polarity, 3. Pollen grain shape, 4. Pollen grain
symmetry, 5. Pollen grain nuclear state, 6. Pollen wall architecture, 7. Exine
stratification, 8. Exine structure, 9. Exine sculpture, 10. Aperture type, 11. Aperture
number, 12. Aperture position, 13. Aperture shape, and 14. Aperture structure.
(a). Stenopalynous Taxa: Taxa where pollen show no variation in shape, size and sculpture, e.g.
Poaceae and Brassicaceae.
(b). Euripalynous Taxa: Taxa where more than one type of pollen differing in size, shape and
sculpture are found, e.g., Asteraceae, Acanthaceae, and Convolvulaceae.
At family level Pollen grain morphology is found valuable in the classification of the family
Saxifragaceae. Berenicearguta genus was kept in this family because of floral whorl characters. On
pollen study, it was found that this taxa is more close to Campanulaceae. So, it was transferred to
Campanulaceae.
The family Berberidaceae has been variously circumscribed by different taxonomists. Based on
pollen characters, Podophyllum where the pollen grains remain united, has been removed to a
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separate family Podophyllaceae. The pollen grains are free in other members of the family
Berberidaceae.
Pollen grain characters, such as shape, size, number of spicules, transverse size, colpus and polar
area index, were useful for separating Verbesina barrance and V.crocata, two morphologically
close Mexican species.
Studies on the palynology of Orchidaceae indicate that the pollen and pollinaria provide useful
diagnostic features in supra-generic categories.
Aperture type (pantoporate, tricolpate, tricolporate) and the exine ornamentation were found to be
the most valuable diagnostic characters for distinguishing the genera Polygonum, Persicaria,
Antigonon, Rumex, Fagopyrum and Homalocladum.
Pollen morphology of the Rosa species from Western Himalaya found distinguishable on the basis
of exine ornamentation.
Salix and Populus (Salicaceae) can be distinguished on the basis of pollen shape and apertures.
Salix has long and narrowed 3-furrowed pollen as compared to spherical pollen without distinct
apertures in Populus.
Salix Populus
Pollen long and narrow Pollen spherical, without distinct
with 3 furrows aperture
The three subfamilies of Apocynaceae can be distinguished from each other on the basis of pollen
features:
Apocynoideae Asclepiadoideae Periplocoideae
Single-grained pollen and Pollens are attached Pollen grains united
translators are absent. to wishbone-shaped in tetrads that are
translators consisting carried in spathulate
of caudicles and pollen carriers or
corpusculum. translators.
The tribe Bombaceae of the family Malvaceae has been recognised as a separate family
Bombacaceae on the basis of pollen characters.
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Variations in the aperture and exine ornamentation may help in the identification of various species
of Bauhinia.
The thickening of exine around the pores makes a distinguishing character for different genera of
the Betulaceae. In optical section, it is knob-like in Betula, club-shaped in Corylis,
unexpanded in Caprinus and an arcus is present between adjacent pores of Alnus.
The two genera of the Phytolaccaceae, Phytolacca and Rivinia, can be recognized on the basis of
palynological characters. The pollen of Phytolaccais 3-zonocolpate, whereas that of Rivinia is
pentacolpate.
Phytolacca Rivinia
Pollen 3-colpate Pollen colpate
Pollen size has been useful in distinguishing two species of Malva-M. rotundifolia and M.
sylvestris.
Two species of Halorgis can be differentiated and recognized on the basis of pollen
characteristics.
H. ciliata H. alata
Pollen Sub-oblate, Pollen Oblate,
Eqatorial diameter – 36 Eqatorial diameter – 25
Aperture non-protruding Aperture protruding
Palynological studies have also contributed in the elucidation of phylogenetic relationships. These
studies suggest two distinct phylogenetic stocks in the dicotyledons – monocolpate and tricolpate
represented by Magnoliaceae and Ranunculaceae, respectively. The presence of monocolpate element
in the monocotyledons indicates that they are more closely related to the Magnolian stock.
Furthermore, the monocotyledons and the Magnolia dicots (both have monocolpate elements,
characteristic of the preangiospermous arhcaegoniates) are considered more ancient palynologically
than the Ranalian dicots where monocolpate elements are completely absent and new apertural forms
are present.
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Kuprianova (1948), who studied the morphology of pollen in the monocotyledons considered that the
Helobiae are not related to the other monocotyledons but are specialized Polycarpieae with Ranalian
affinities. She also pointed out that most monocotyledonous families could be considered to have
evolved from Arecaceae or Liliaceae.
It is suggested that only palynology can not be taken to ascertain taxonomic positions.
Another limitation is that a particular character can not be universally applied for all
taxa.
Similar type of pollen are found in diverse taxa.
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