Taxonomic Evidences: Structure

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Block 3 Plant Taxonomy – Tools and Evidences

UNIT 14
TAXONOMIC EVIDENCES

Structure
14.1 Introduction 14.5 Evidence from
Phytochemistry
Objectives
Directly Visible Chemical
14.2 Alpha Taxonomy and Omega
Characters
Taxonomy
Primary Metabolites
14.3 Taxonomic Evidences from
Palynology Secondary Metabolites

14.4 Taxonomic Evidences from Semantides


Cytology
14.6 Evidence from Molecular Data
Chromosome Numbers
14.7 Summary
Chromosome Structure
14.8 Terminal Questions
Chromosome Behaviour
14.9 Answers

14.1 INTRODUCTION
In Taxonomy, evidence is the information used in context for a purpose such
as identification or classification. The foundation of plant taxonomy was laid on
a number of characters (attributes) of plants and plant groups studied from
time to time. Such taxonomic information can be realised in many ways to
prove a hypothesis, solve a problem, characterize a taxon, classify a group of
plants or derive evolutionary relationship among plants. The gross morphology
provides the foundation of plants taxonomy and it is supplemented by the
information from various other fields of botany.

In this Unit you will study about taxonomic information that can be collected
from a few specific fields such as: palynology; cytology; phytochemistry and
molecular biology.

Objectives
62
Unit 14 Taxonomic Evidences
After studying this unit you should be able to:

 know the history of plant taxonomy, with special emphasis on plant


taxonomy in ancient India;

 comprehend the concept of plant taxonomic evidences;

 classify the taxonomic evidences as physical, chemical and biological;

 specifically infer the taxonomic evidences from the fields of palynology,


cytology, phytochemistry and molecular botany;

14.2 ALPHA TAXONOMY AND OMEGA


TAXONOMY
The scope and methods of taxonomy have evolved with times. However, the
principal core of the discipline has remained the same: concepts, classification
of plant and plant groups; and nomenclature. To achieve these objectives the
approaches adopted are broadly classified into two kinds: alpha taxonomy
and Omega taxonomy (Turrill 1935).

On the other hand, taxonomists such as Valentine and Love (1958) and Davis
and Heywood (1963) have suggested that the developments in taxonomy
occurred in 4 phases. These are (i) the pioneer or exploratory phase, (ii)
the systematic or consolidation phase, (iii) the experimental or
biosystematic phase, and (iv) the encyclopedic or holotaxonomic phase.
According to this classification of the developments in taxonomy, the first 2
phases are more or less equivalent to alpha taxonomy while phases (iii) and
(iv) are similar to omega taxonomy.

Alpha Taxonomy

At the level of alpha taxonomy, the organisms are characterized, identified,


classified and named. The pioneer or exploratory phase of taxonomy is
basically analytical. It involves collection of plants for analysis of their
characters. Therefore, in this sense, it is a pioneering effort for understanding
the plants. At the same time it is also considered as an exploration of nature.
Hence this is regarded as the pioneer or exploratory phase in taxonomy
This relates to the basic or preliminary classification based almost entirely on
external morphology. This can be considered as the empirical approach in
taxonomy where the classification is synthesised from the observed facts. The
vast amount of data collected by taxonomists must be evaluated and
consolidated. This leads to the development of systems of classification. Thus,
the pioneer or exploratory phase progresses to a systematic or
consolidation phase. Alpha taxonomy has also been called Classical or
Orthodox or Formal taxonomy. This has been practised since ancient times
and different aspects of this approach are in use even today.

Omega Taxonomy

After synthesising a basic classification, the taxonomist can attempt to improve


upon it. Therefore, the observed facts are interpreted in Omega taxonomy to
provide an interpretive classification. Evolutionary and phylogenetic
approaches are applied to understand taxonomic and evolutionary 63
Block 3 Plant Taxonomy – Tools and Evidences
relationships of plants at all levels. This can be achieved by experimentation or
use of a larger number of features such as cytology and genetics for
evaluating taxonomic relationships. Thus developments in taxonomy involving
detailed cytological and genetical studies became important. This has been
referred to as the experimental or biosystematic phase. It was then realized
that knowing all aspects of the biology and chemistry of plants would provide a
much better understanding of relationships. Thus collecting and collating
information from as many sources as available led to the recognition of the
encyclopedic or holotaxonomic phase in taxonomy. Omega taxonomy has
also been called Beta taxonomy or Neotaxonomy or Modern taxnonomy.

Taxonomic evidences are the characters (attributes) gathered from a variety of


disciplines. Most of these characters are used in describing patterns of
variations at or below the species level. The source(s) of such characters
could be from morphology, anatomy, embryology, palynology, cytology,
biochemistry, sementides, molecular biology, etc.

In this Unit we will describe about the following of taxonomic evidences:

Palynology;

Cytology;

Phytochemistry; and

Molecular biology.

SAQ 1
a) Differentiate between alpha- and omega- taxonomy.

b) Identify the following statements as true or false by putting letter T for


true of F for false in the brackets provided at the end of the statements.

i) Omega taxonomy is also called formal taxonomy. [ ]

ii) Turrill proposed the term alpha taxonomy. [ ]

iii) The observed facts are interpreted in alpha taxonomy. [ ]

iv) Any evidence is an information used in context for a purpose. [ ]

14.3 TAXONOMIC EVIDENCES FROM


PALYNOLOGY
Palynological Evidence

Palynology is the study of pollen and spores. The taxonomic characters of


pollen grains include wall structure, polarity, symmetry, shape and grain size.
In angiosperms pollen grains are of two kinds viz. monocolpate and tricolpate
(Fig: 14.1). Monocolpate pollen grains are boat shaped, have one long
germinal furrow and one germinal aperture. This is characteristic of primitive
dicotyledons, such as Piperaceae and Chloranthaceae, majority of
64 monocotyledons, cycads and seed ferns. According to some palynologists the
Unit 14 Taxonomic Evidences
first flowering plants had monocolpate pollen grains. Tricolpate grains are
globose, symmetrical, have three germinal apertures and are characteristic of
advanced dicotyledons.

Fig: 14.1: Monocolpate and tricolpate pollen grain.

Some of the characters of the pollen grains that constitute taxonomic evidence
are: unit; polarity; shape, symmetry; nuclear state; wall architecture; exine
stratification; exine structure and sculpture (Fig: 14.2). It also includes shape,
structure, position, number and types of apertures.

Let us now learn some examples.

The pollen grains of Magnoliidae are binucleate whereas those of


Caryophyllnidae are trinucleate. In Ericaceae the pollen grains occur as
tetrads while pollinium (pl. pollinia) are characteristics of Asclepiadaceae,
Apocyanaceae) and Orchidaceae.

Within the family Betulaceae the exine of pollen grain in section is knob-like in
Betula; club-shaped in Corylus; unexpanded in Carpinus, arch-like in Alnus.
Within the family Epacridaceae the pollen grains of the genus Brachyloma are
solitary while those of the genus Epacris are in tetrads.

Pollen apertures can be distinct in different species of the same genus. It is


pilate in Bauhinia acuminata, striate in B. krugii and echinulate in B.
malabarica.

The pollen grain size could also be used to identify the species of a genus. For
example, the pollen grain size in Malva rotundifolia is 74-84 µm while it is in
range of 105-126 µm in M. sylvestris.

Fig. 14.2: Pollen grain showing various sculpturing.

65
Block 3 Plant Taxonomy – Tools and Evidences
An interesting example of palynological evidence in taxonomy is of the genus,
Trisyngyne. This plant has separate male and female plants (dioecious). On
the basis of morphological evidences, the genus was placed in the family
Euphorbiaceae. However, when female plant was discovered it was
transferred to the family Fagaceae. Later, when the pollen grains of male
specimen was examined, they were found to be exactly similar to the member
of Fagaceae. So, now Trisyngyne is regarded as a member of the family,
Fagaceae. (Some taxonomists suggest the name Nothofagus for this genus,
and classify it in the Family Nothofagaceae).

14.4 TAXONOMIC EVIDENCES FROM


CYTOLOGY
Cytology is the study of the morphology and physiology of cells. The
information about the chromosome number, shape and pairing at meiosis is
used for classification purpose. Cytotaxonomy refers to the use of
chromosome number, morphology, ploidy level, ploidy type and chromosome
aberrations as data for classification. Cytogenetics includes those studies
dealing with observations of chromosome pairing or behaviour at meiosis.

In the following sub-sections you will study about importance of chromosome


number, structure and behavior at meiosis as taxonomic evidence.

14.4.1 Chromosome Numbers


We are generally aware that the number of chromosomes in each cell of all
individuals of a single species is constant. It is also established that the more
closely related species are likely to have similar chromosome numbers while
the more distantly related ones shall have different numbers. Due to this
relative conservativeness, chromosome number becomes an important and
frequently used taxonomic character. In addition, there is a very wide range of
chromosome numbers in the angiosperms from as low as 2n=4 (in
Haplopappus gracilis) (Asteraceae) to as high as 2n=530 (in Poa litorosa)
(Poaceae). A large number of angiosperms have been analysed for their
chromosome numbers, providing useful taxonomic information (see Table
14.1).

Table 14.1 : Basic chromosome numbers of a few plants.

Haplopappus gracilis n=2

Zea and Sorghum (all species) n = 10

Pinus (all species) n = 12

Quercus (all species) n = 12

Kalanchoe sp. n = 125

Poa litorosa n = 132

Morus nigra n = 154


66
Unit 14 Taxonomic Evidences
Table 14.2 : Chromosome Series - a few examples.
Brassica n = 6, 7, 8, 9, 10

Aster n = 9, 18, 27

Taraxacum n = 8, 12, 14, 20, 24

Vicia n = 5, 6, 7, 12, 14

Festuca 2 n = 14, 28, 42, 56, 70

Many interesting ideas have developed from knowledge of chromosome


numbers. For example, in the genus Festuca, different species have different 2n is equal to the
diploid or somatic
chromosome numbers forming a mathematical series (See Table 14.2). The
chromosome number:
chromosome numbers are 2n = 14, 28, 42, 56, 70 etc. From this information, a
while n is the haploid
generalisation can be made, that different species may have some common
or the gametic
basis. If we assume that these chromosome numbers are based on a common
number
denominator called x (and x = 7), then we can consider the different species to
have multiples of this number. This denominator or base number (x = 7) can
be considered as the basic set of genetic information carried by a plant, and
due to the multiplication of this basic genetic set, the evolution of different
species has occurred. Such a series is said to be polyploid in which the basic
number (x) is equivalent to the haploid number of chromosomes in a diploid
species (i.e. x = n = 7). The other species would then be tetraploid (4x = 2n),
hexaploid (6x = 2n), octaploid (8x = 2n), decaploid (10x = 2n), respectively.

14.4.2 Chromosome Structure


Cytologists have studied chromosome morphology and have pointed out that
the most interesting feature about the chromosome structure is the position of
the centromere. The centromere or constriction in the length of the
chromosome provides information about the relationship of the 2 arms of the
chromosomes. Thus, depending on the position of the centromere,
chromosomes are described as metacentric, acrocentric, and telocentric (Fig.
14.3) Metacentric chromosomes are considered to be more advanced than the
acrocentric chromosomes.

(a) (b) (c)

Fig.14.3: Chromosome structure- a) acrocentric, b) metacentric, c) telocentric.

The appearance of the basic chromosome set in a dividing cell is known as


the karyotype of the cell. This can be analysed to provide information not only
of the chromosome number, but also about the chromosome size,
chromosome volume, and type of chromosomes in the cell. This information is
used by taxonomists for identifying plants and understanding relationships.
The karyotype can be represented diagrammatically as an idiogram or 67
Block 3 Plant Taxonomy – Tools and Evidences
karyogram, and these diagrams can be compared for taxonomic purposes.
Another interesting observation is that the absolute size of the chromosomes
of a karyotype is fairly constant since it is controlled by the genotype.
Taxonomists have found that monocots generally have larger chromosomes
than dicots, and that smaller chromosomes are found in hardwood plants in
comparison to their herbaceous relatives.

Let us now describe some examples where chromosome structures have


been successfully used as taxonomic evidence. Paeonia was earlier placed in
the family Ranunculaceae. However, on the basis of embryological and other
evidences it is suggested that it be placed in an independent family,
Paeoniaceae. Cytologically, it is found that Paeonia possesses large
chromosomes as opposed to smaller chromosomes that are characteristic of
members of the family Ranunculaceae.

Plants of Polygonum growing in three different habitats exhibited


morphological differences and were hence considered belonging to different
species. Interestingly, it was found that all of them were not only interfertile but
had similar chromosome morphology. Hence, they are now clubbed together
as Polygonum amphibium.

Chromosome karyotype studies have suggested placing the genus Butomus in


a separate family Butomaceae while retaining Limnocharis, Hydrocharis and
Tenagocharis in the family Alismataceae. Closer affinities of the families
Cyperaceae and Juncaceae have been established due to the presence of
holocentric chromosomes in their members.

Karyotype studies have confirmed the separation of the family Agavaceae


from Amaryllidaceae. Presence of 5 large and 25 small chromosomes in their
cells have helped transfer of Yucca from the family Liliaceae to the family
Agavaceae.

14.4.3 Chromosome Behaviour


When we study meiosis, we not only observe the regularity of pairing which is
important for the fertility of the plants, we are also able to make a chromosome
to chromosome comparison. This provides valuable information about the role
of chromosomes in heredity. Taxonomists use this information to understand
relationships amongst different species of plants. We can also determine the
nature of the genome to find out if it is homozygous or heterozygous. Genome
analysis in plant taxonomy has been particularly useful in understanding
polyploidy and for establishing the parentage of polyploids. A very significant
study in this regard is the case of the common hexaploid bread-wheat,
Triticum aestivum. The genome of this economically important plant has been
designated as AABBDD with 2n=42 chromosomes. Detailed genome analyses
have established that 3 diploid species have contributed to the evolution of
this hexaploid wheat. The A genome is from a wild grass Aegilops speltoides
(2n=14), and the D genome has been contributed by Aegilops squarrosa
(2n=14). Many geneticists have suggested different grasses as the source of
68 the B genome.
Unit 14 Taxonomic Evidences
Finally, we must remember that as with any other character, the value of
cytotaxonmic data depends upon the group or category under investigation. A
combination of cytological information with data from other disciplines will
provide a more useful tool to the taxonomist.

SAQ 2
a) Fill in the blank spaces with appropriate word(s):
i) Monocolpate pollen grains are ……………..………shaped.
ii) Union of 3 or more chromosome complements of the same
species result in ……………….. .
iii) The basic number of chromosome (x) is equivalent to the
………………. number of chromosomes in a diploid plant.
iv) The karyotype can be represented diagrammatically as an
………………….
v) Diploid chromosome number in Haplopappus gracilis is
……………… .
b) Write a note on “pollen grains as taxonomic evidence”.

14.5 EVIDENCE FROM PHYTOCHEMISTRY


Chemotaxonomy is a science which uses chemical information as a character
for taxonomic purposes. Before we analyse the basis of this modern trend in
plant taxonomy, let us for a moment think about the different kinds of plants in
our daily lives. When we drink tea or coffee, we appreciate the flavour or
aroma and differentiate the two by this character. Similarly, when we eat fruits
such as the mango, the banana or an apple, we find that they taste differently.
These differences are due to the chemical constituents of these foods and this
forms the basis of chemotaxonomy where the chemical features or chemical
constituents serve as the evidence for taxonomy. The potential importance of
chemical evidence in plant taxonomy has been suggested by both botanists
and chemists and this has become an important recent trend especially
because newer techniques for quick analysis of plant material have been
developed. Chemotaxonomists suggest that chemical characters have a
particularly high taxonomic value because they are i) stable, ii) unambiguous,
and iii) not easily (if at all) changeable. Further, chemical characters will show
chemical relationships amongst plants in the same way as morphological
characters show morphological relationships.

Although chemotaxonomy is considered to be a relatively recent development


in modem taxonomy, its origin can be traced to very early classical taxonomy.
You will recall that the spice plants were identified on the basis of their
aromatic properties, or the medicinal plants by their curative value. These
aromatic properties or the curative value was largely based on the chemical
constituents of the plants and taxonomists have classified them since ancient
times using these chemical features along with morphological characters.
However, it is only in recent years that chemotaxonomy as an important field
of study has been established. 69
Block 3 Plant Taxonomy – Tools and Evidences
A review of the large amount of literature published in this field reveals that
chemical data may be obtained from any part of the plant. Secondly,
depending on the purpose of the investigation, the chemical information may
be used for description or identification of plants, or for establishing
relationships. This evidence assumes greater significance when it is used to
sort out differences in taxonomic relationships when 2 or more possibilities are
suggested on the basis of morphological characters.

Although theoretically, all chemical constituents of a plant are potentially


valuable to a taxonomist, in practice some kinds of molecules are more useful
than others. Thus we can use directly visible chemical constituents such as
crystals, raphides, or starch grains occurring in different plants as chemical
characters.

Alternatively, we can chemically analyse plant material for different chemical


constituents and use this information for taxonomic purposes. Most
chemotaxonomists recognise three broad categories of chemical compounds,
primary metabolites, secondary metabolites, and semantides, as important
taxonomically.

14.5.1 Directly Visible Chemical Characters


Very few chemical substances in plants can be observed directly, but the few
substances such as starch grains which are present in most green plants as
food reserves have been used for chemotaxonomic purposes (fig. 14.4). The
types of starch grains present in different plants are very specific and this
information can be used without any ambiguity. Reichert (1913) examined the
starch grains in 350 species of plants and established their differentiation and
specificity of occurrence in relation to genera and species, thus providing
chemotaxonomic information. Tateoka (1962) reviewed the starch grain form
in the grass family (Poaceae) and used this as additional information to
classify the family into tribes. For example, in the Tribe Hordeae, the typical
members such as Hordeum have compound starch grains, while other genera
like Lolium, Nardus, and Papapholis have simple starch grains.

Fig. 14.4: Morphology of starch grains in some plants.

Raphides are the crystals of calcium oxalate which are present in large cells in
different plant tissues and can be observed directly. They are long needle
shaped crystals, pointed at both ends and usually occur in bundles, thus being
easily identified. They occur very commonly in some aquatic plants such as
70 Pistia and Eichhornia and found in the members of Araceae.
Unit 14 Taxonomic Evidences
The presence or absence of raphides is a key character, in natural
classification of family Rubiaceae, along with some other ones. All those
Rubiaceae members that possess raphides are placed in the tribe Rubioideae.

Another example where raphides are important taxonomically is the sub-


division of the Order Scitamineae. The raphides are present in the families
Heliconiaceae, Musaceae, Sterlitziaceae and Lowiaceae. Further,
classification within these families is based on other taxonomic features. On
the other hand, within order Scitamineae, in the families Costaceae,
Marantaceae, Zingiberaceae, and Cannaceae the raphides are absent.

Another kind of crystal, made-up of calcium oxalate are called druses and
provide good taxonomic evidence. A druse is an aggregate or conglomerate of
a large number of crystals. They are characteristic of the families Caricaceae,
Apocyanaceae, etc.

Cystoliths are the crystals made up of calcium carbonate and are usually very
common in leaves of various species of Ficus.Presence of crystals of gypsum
(a soft mineral) made of Calcium sulphate (CaSO4.2H2O) is characteristic of
family Tamaricaeae(Fig 14.5).

(a) (b) (c)

Fig. 14.5: Crystals of various types: a) Raphide; b) Druses; c) Cystolith.

14.5.2 Primary Metabolites


As the name indicates, primary metabolites are molecules involved in vital
metabolic pathways. They are of universal occurrence and not very significant
in chemotaxonomy. However, these molecules become useful as
chemotaxonomic features when the quantity of such molecules varies
considerably between taxa. For example, the sugar ‘sedoheptulose’ is stored
in large quantities as a reserve food in the genus Sedum. Thus members of
this genus can be easily identified by the presence of this primary metabolite.
Interestingly, sedoheptulose biphosphate is a part of the photosynthetic
carbon cycle and in a majority of the plants sedoheptulose does not
accumulate at all. In the same way, the 22 amino acids are of universal
occurrence. They serve as the building blocks of proteins. They can provide
useful macromolecular data for chemotaxonomy. The amino acid sequence of
different proteins can be investigated and the degree of similarity is
presumably proportional to the degree of genetic relationship. However, only a
few out of about 3 lakh species of angiosperms have been analysed for amino
acid sequences. For example, the amino acid data on wheat and barley
confirms the relationship of these genera as suggested by classical
taxonomists. 71
Block 3 Plant Taxonomy – Tools and Evidences
14.5.3 Secondary Metabolites
Secondary metabolites or secondary plant products are those macromolecules
that lack nitrogen and are of restricted occurrence and therefore, of greater
taxonomic importance than primary metabolites. This group includes different
kinds of compounds such as flavonoids, terpenes, iridoids, alkaloids,
anthocyanins, glucosinolates, cyanogenic glycosides, polyacetelenes etc.
They are usually not involved in vital functions and are largely storage
products or pigments.

FLAVONOIDS

Amongst the secondary metabolites, flavonoids, which are the commonest


phenolic compounds of leaves, have been very useful for chemotaxonomic
purposes. Both monocots and dicots have been extensively surveyed for
these compounds which show structural variability and chemical stability
besides widespread distribution. They can be rapidly and easily identified and
provide important chemical characters for taxonomic purposes. For example,
80 species of plants from the family Ulmaceae were investigated for their
flavonoid chemistry by Giannasi (1978). Majority of the species contain
flavonols, but a few species have glyco-flavonols and these two types of
flavonoid compounds are never present together in any species. Interestingly,
enough, in most classical systems of classification, the family Ulmaceae is
divided into two subfamilies called Ulmoideae and Celtoideae which are also
distinguishable by the flavonoid chemistry. Therefore, morphological criteria
combined with flavonoid dichotomy can be used to divide the family Ulmaceae
Kinds of Terpenes into two distinct families: family Ulmaceae is characterised by the presence of
C5 = Isoprenes flavonols, and family Celtaceae is characterised by the presence of
C10 = Monoterpenes glucoflavonols.

C15 = Sesquiterpenes Several other studies have used flavonoid chemistry for taxonomic purposes
C20 = Diterpenes in families such as Arilidaceae, Cornaceae, Labiatae (Lamiaceae),
Leguminosae (Fabaceae), Orchidaceae, Rutaceae, Lemnaceae and others.
C30 = Triterpenes
C40 = Tetraterpenes Although useful in assessing relationship among closely related species
(infraspecific variations), flavonoids are occasionally useful in assessing
Cn = Polyterpenes
phylogenetic relationship at higher levels.

TERPENES

A second group of secondary metabolites commonly examined by


chemotaxonomists are the terpenes. Volatile terpenes are major components
of essential (ethereal) oils which are characteristics of the order Magnoliales,
Austrobaileales, Piperales and Laurales. They are also reported in families
Myrtaceae, Rutaceae, Apiaceae, Lamiaceae, Asteraceae. Chemically
speaking, these compounds can be classified on the basis of their molecular
structure into monoterpenes, diterpenes, triterpenes, sesquiterpenes, etc., and
each group can be used for taxonomic purposes. For example, in the genus
Salvia, 19 species could be distinctly identified and classified on the basis of
their monoterpenes. The terpene composition was as useful as the
morphological characteristics in the analysis of introgression and hybridisation
within the genus. Similarly, triterpenes and sesquiterpenes have been
particularly important and useful in the classification of the families
72 Cucurbitaceae and Compositae (Asteraceae) respectively.
Unit 14 Taxonomic Evidences
IRIDOIDS

Iridoids are 9-10 carbon monoterpenoid derivatives. They are commonly found
in plants grouped as Asterids.

ALKALOIDS

Alkaloids are structurally diverse chemicals that are derived from amino acids
or from mevalonic acid. Cocaine, morphine, codeine, atropine, colchicines,
quinines are some important alkaloid plant products. The plant groups, where
they are biosynthesized represent a vast diversity, hence of little systematic
interest.

BETALAINS AND ANTHOCYANINS

Betalains are nitrogenous red and yellow pigments are restricted to the Order
Caryophyllales except the families Caryophyllaceae and Molluginaceae.

Anthocyanins, in contrast, are red, yellow, blue or purple pigments of most


other plants.

Betalins and anthocyanins are mutually exclusive; as they have never been
found together in the same species.

GLUCOSINOLATES

The glucosinolates are also known as mustard oil glucosides. They are
hydrolysed by the enzymes known as myrosinases to yield pungent mustard
oil. They are characteristic of Brassicaceae, Resedaceae and Tovariaceae.

CYANOGENIC GLYCOSIDES

The cyanogenic glycosides are defensive compounds that are hydrolysed by


various enzymes to release hydrogen cyanide. The process cyanogenesis, is
widespread in angiosperms. Leucine derived cyanogenic glycosides is
common in the families Rosaceae, Fabaceae, Sapindaceae. In the Orders
Magnoliales and Laurales, these compounds are derived from the amino acid,
tyrosine.

POLYACETYLENES

Polyacetylenes are a large group of non-nitrogenous secondary metabolites.


They are formed by linking of acetate units via fatty acids. Taxonomically, they
are important because of their presence in related groups of Asteroid families:
Asteraceae; Apiaceae; Campanulaceae; Caprifoliaceae etc.

14.5.4 SEMANTIDES
The information carrying molecules in plants are called semantides, and they
have been recognised to be of 3 kinds; deoxyribonucleic acid or DNA (primary
semantide), ribonucleic acid or RNA (secondary semantide) and proteins
(tertiary semantide) following the sequential transfer of the genetic code. Of
these, the proteins are the most favoured molecules for chemotaxonomic
purposes. Plant proteins can be studied by different methods; by
electrophoresis or by serological methods, and both processes have been
used for obtaining information about the protein chemistry of different plants. 73
Block 3 Plant Taxonomy – Tools and Evidences
In the common breadwheat, Triticum aestivum, the storage proteins were
analysed by electrophoresis. For comparative purposes, the storage proteins
of the tetraploid wheat, Triticum dicoccum and the diploid grass Aegilops
squarrosa were also analysed electrophoretically. This study confirmed the
conclusion that the hexaploid wheat did contain a sum of the proteins
possessed by the diploid species which have contributed to the evolution of
the hexaploid wheat. This study supports the observations based on
morphology and cytological evidence.

Serological analysis of proteins is based on the immunological reaction shown


by mammals when a foreign protein is introduced into the system. In other
words, this is based on the antibody-antigen reaction wherein the antibodies
being specific to an antigen bringing about coagulation. This information can
then be analysed to understand the relationships of the different plants on the
basis of the serological evaluation of the plant proteins. Serology has proved a
useful taxonomic tool at different levels of classification. J. G. Hawkes (1960)
and his co-workers studied several tuber-producing species of Solanum to
understand the evolution of the cultivated potato Solanum tuberosum and
determine the species of Solanum which could be established as the
ancestors of the common cultivated potato, Similarly, in the family
Ranunculaceae, serological studies supported cytological data for the
classification of the family into tribes and genera. Fairbrothers (1959) and his
co-workers have studied several plant groups serologically particularly the
members of grass family. A general conclusion from such studies is that the
different amount of serological activity in members of different plant families
may be interpreted as a reflection of the evolutionary differences in the primary
structure of the proteins due to which serological differences can be
recognised between members of different families.

14.6 EVIDENCE FROM MOLECULAR DATA


In the preceding section you learnt that the nucleic acids and proteins
constitute semantides. However, only nucleic acids (DNA and RNA) contribute
towards significant analysis of phylogenetic relationships among plants.
Traditionally the taxonomists use tools of use taxonomy to construct
phylogeny of plants or plant groups. However, it has now been realized that
the molecular data based on variations in DNA and RNA of different plants
provide more reliable inputs. Such an assumption is based on the hypothesis
that the phylogenetically closer species should possess similar genetic
material and distant species should exhibit variations in their genetic
constitution. In recent times, DNA fingerprinting has become a very useful
source of information in taxonomy. It allows comparison of different species of
a genus enabling understanding of phylogenetic relationship.

From where do the scientists obtain genetic material for studies?

Our understanding of cell biology reveals that such molecular data lie within
the cell organelles, nucleus, mitochondria and chloroplast of a cell. All of them
possess DNA and are referred to as: nuclear-DNA (Nucleus); mt DNA
(mitochondrion), and cp DNA (in chloroplast) respectively. A nuclear DNA of a
74 cell generally consists of a million to many billions kilo base pairs (kbp) and is
Unit 14 Taxonomic Evidences
biparental in origin (a combination of male and female genome). Though
stable, it evolves rapidly and is more difficult to analyse. However, it is a
source of data that could throw light on final evolutionary processes.

Both mitochondrial and chloroplast DNA are uniparental in origin.

Based on the observations that mt DNA (made up of 200-2500 kbp) undergo


a lot of rearrangement as well as occurrence of many forms of it within a
mitochondrion, it is regarded relatively as of little significance in the
phylogenetic systematics.

The most suitable genetic material as taxonomic evidence of green plants is


cpDNA. In higher plants the size of a cpDNA in a chloroplast is only 120-140
kbp. Thus, it is smallest among all the three sources of genetic material within
a cell. The advantages of cpDNA in taxonomic studies could be summarised
as:

 relatively smaller size;

 large number of chloroplasts per cell;

 relatively stable not only in an organism but also in a species;

 easy to isolate and analyse;

 not much altered by evolutionary processes; and

 all cpDNA molecules carry basically same set of genes, however,


arrangement varies in different species.

Molecular data is being used to construct phylogenetic trees of green plants,


at almost all ranks. It principally helps in establishing affinities and
relationships within and outside the ranks. For example : monophyly of
families Apiaceae, Caprifoliaceae, Plantaginaceae, Scrophulariaceae,
Apocyanaccae, Saxifragaceae etc is well supported by molecular data
analysis. It supports the union of families Apocyaceae and Asclepiadaceae;
Braliaceae and Apiaceae; Brassicaceae and Capparaceae.

However, the importance of evidences from other, rather classical taxonomy,


should not be brushed aside. All sources of evidences should be incorporated
and analysed to arrive at a phylogenetic understanding of living organisms.

SAQ 3
a) Choose the correct alternative from the two provided in the parantheses.

i) Hordeum vulgare possesses (simple/compound) starch grains.

ii) Chemically raphides are (Calcium sulphate/Calcium oxalate).

iii) Betalains are (nitrogenous/non-nitrogenous) pigments.

iv) C10 - terpenes are called (diterpenes/monoterpenes).

v) cpDNA is (not/very) much altered by evolutionary processes.

75
Block 3 Plant Taxonomy – Tools and Evidences
b) What are semantides? How do they differ from secondary metabolites.

c) Match the chemical items in column I with the choices provided in


column II.

Column I Column II

i) Volatile terpenes a) Anthocyanins

ii) Purple pigments b) Alkaloids

iii) Cyanogenic glycosides c) Ethereal oils

iv) Colchicine d) Chloroplast

v) Rubisco e) Rosaceae

14.7 SUMMARY
 The scope and methods of plant taxonomy have evolved through times.
Broadly, they are classified as: Alpha and Omega taxonomy.

 Alpha taxonomy is analytical and empirical. At this level an organisms is


identified ,characterised and named on the basis of morphology. It is
also referred to as: classical, orthodox or formal taxonomy.

 Omega taxonomy provides intrepretive classification. It helps understand


evolutionary and phylogenetical relationships among organisms. It is
also referred to as: beta-; neo-, and modern taxonomy.

 The characters/character-states/attributes provide useful inputs to


identify, classify a plant. All of them together are called taxonomic
evidences. Such evidences could be physical, chemical or biological.

 Taxonomic evidences from morphology, anatomy, embryology,


palynology, cytology etc. constitute physical taxonomic evidences.
Primary and secondary metabolites constitute chemical taxonomic
evidences. Information-carrying biomacromolecules such as nucleic
acids and proteins are called biological taxonomic evidences. They are
also called semantides.

 Molecular data obtained from the DNA of nuclear, mitochondrial and


chloroplast origin provides important insight into phylogenetic
systematics.

 Study of structure, polarity, symmetry, shape and size of pollen grains is


called palynology. Palynological evidences from both extanct as well as
extinct plants are very important physical taxonomic evidences.

 Chromosome morphology, karyotype studies (idiograms), position of


centromere in a chromosome and behavior of chromosomes at meiosis,
all together are also very informative taxonomic evidences.

 Directly visible chemicals, within a cell, in the form of starch grains and
76 crystals, are also helpful in identification, and determination of
Unit 14 Taxonomic Evidences
relationships in plants. Chemicals such as flavonoids, iridioids, alkaloids,
betalains, anthocyanins, glucosinates, polyacetylenes, cyanogeneic
glycosides, terpenes etc. are termed secondary metabolic taxonomic
evidences.

 For proper characterisation, identification, classification of plants,


taxonomic evidences from a variety of sources should be gathered prior
to any definite interpretations regarding phylogenetic relationships
amongst various plants or plant groups.

14.8 TERMINAL QUESTIONS


1. Differentiate between the following pairs. Provide one example of each.

a) Semantide and Secondary Metabolites.

b) Alkaloids and Flavonoids.

c) Autopolyploidy and Allopolyploidy.

2. Mention briefly the importance of molecular data in modern plant


systematics.

3. List the characteristics of the pollen grains that are useful as taxonomic
evidences.

4. Write a note: “Cytology as taxonomic evidence”.

5. Discuss the significance of secondary metabolites in taxonomy.

14.9 ANSWERS
Self-Assessment Questions
1. a) Refer section 14.2

b) i) F; ii) T; iii) F; iv) T

2. a) i) boat

ii) autopolyploidy

iii) haploid

iv) ideogram

v) 4

b) Refer Subsection 14.3

3. a) i) compound

ii) Ca-oxalate

iii) nitrogenous

iv) monoterpenes

v) not 77
Block 3 Plant Taxonomy – Tools and Evidences
b) Refer Subsection 14.5.4 and 14.5.3

c) i) c.; ii) a.; iii) e.; iv) b.; v) d

Terminal Questions
1. a) Refer to Sections : 14.5.4 and 14.5.3

b) Refer to Section: 14.6

c) Refer to Section: 14.4.1

2. Refer to section : 14.6

3. Refer to section : 14.3

4. Refer to section 14.4

5. Refer to section 14.5.3

78

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