64

International Seminar on "Multidisciplinary Approaches in Angiosperm Systematics"

INVESTIGATION ON SEEDLING MORPHOLOGY IN TAXONOMIC STUDIES

OFANGIOSPERMS
Avinash Mundhra, Biswajit Roy and N. D. Paria
Abstract
The study of seedling morphology, a less explored field in flowering plants, particularly in angiosperms,
has been emerged as an essential discipline for taxonomic research. It is well accepted that the taxonomic
research should not be restricted to the study of morphological characters of mature specimens only,
but the characters of juvenile specimens, particularly the seedling, should be considered also, as because
such characters are valued as primary genetic characters. There has been a considerable development in
the study of seedling morphology in different countries abroad, but little progress has been made in
India, in spite of its rich biodiversity of plants. The well-knit terminology, glossary and standard
scheme for describing seedling plants are available in seedling literature. Based on them, Paria and his
associates (1986-2010) have made significant contributions in taxonomic studies of angiosperms, some
of which include a number of families and other taxa, e.g. Anacardiaceae, Combretaceae, Euphorbiaceae,
Leguminosae, Rubiaceae, Bauhinia spp., Jatropha spp., etc. The progress made so far in the field of
seedling morphology in relation to taxonomy as above has been promising and significant in the context
of Indian flora. The data obtained in this study also correspond to some other data, e.g. pollen
morphology, cytology, phytochemistry, general morphology, etc. More works are needed to be done
from this field to prepare a sizeable and comprehensive account for seedling flora of India.
Kew words: Seedling morphology, Germination types, Taxonomic markers, Systematics.

Introduction
The seedling stage is arguably the busiest phase in a plant's lifetime (Farnsworth, 2008).
Once germination has occurred, the seedling depends on its own morphological and physiological
characteristics to cope with the various factors threatening its survival. Comparative seedling
morphology has, however, been of very little interest to botanists since Lubbock's (1896) classic
work on the subject. In fossil botany, it has been ignored, possibly due to failure of delicate parts of
seedlings to be preserved. Even evolutionists have not appreciated to study this vital stage in the
life cycle of flowering plants. Moreover, the seedlings of many taxa, particularly herbs, are so
small as to escape attention in their native habitats. But juvenile stages of plants, particularly of
trees, are often so strikingly different from the adult stages, that even with good field knowledge of
plants it is difficult to correlate the seedling plant with an adult shoot of the same species. It is now
emphasized that seedling morphology should be thoroughly investigated for a better comprehension
of germination, establishment and juvenile growth during the natural regeneration of vegetation
(Troup, 1921; Jackson, 1974).
In this direction, considerable literature have been built up dealing with seedlings based
on various investigations (Duke, 1969; Burger, 1972; Vogel, 1980), encouraging to undertake the
study of seedling morphology. However, it is unfortunate that, in spite of considerable taxonomic
potentiality and prospects of seedling morphology, very little attention has been paid to this field,
particularly in the context of Indian flora. The present article is intended as a brief review based on
the works carried out in the laboratory of the present authors.
Review of Literature
Earlier several workers have made use of data from seedlings in taxonomic research.
As quoted by Bernhardi (1832), Caesalpino provided a description of seedling morphology in 1583.
Bernhardi {I.e.) gave an interpretation of the different parts of embryos and seedlings. The nature
of rootlets, hypocotyl, collet and cotyledons as well as seedling parts was discussed by A. P. de
Candolle (1825). Studies on seedling characters of a large number of taxa and their importance in
taxonomy were made by Duke (1965, 1969).
Systematic studies on seedling morphology were carried out at different taxonomic levels.
Different families have been investigated for seedling morphology, e.g. Sapotaceae (Bokdam,

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1977), Annonaceae (Garwood, 1995), Droseraceae (Conran et ah, 1997), Casuarinaceae (Hwang
and Conran, 2000), Sterculiaceae (Wilkins and Chappill, 2002), etc. At generic level, the most
promising work was carried out by Canne (1983). He examined the seedlings of 18 species of
Agalinis (Scrophulariaceae) and showed several seedling features distinctive at the species level
which correlates well with data from other sources with regard to sectional and subsectional
placement of species within the genus. Different workers have also carried out investigations on
seedling morphology at the level of genus with reference to their taxonomic importance, e.g.
Erythrina (Flores and Dora, 1984), Eucalyptus (Lodiges et ah, 1984), Cicer (Nozzolillo, 1985),
Nothofagus (Raffaele et ah, 1998). In addition to the above, Fernando et ah (1999) established
the importance of the phenetic analysis in characterization of woody Medicago species using 20
morphometric characters of seedlings and provided keys for identification of species.
Seedling morphology has also been utilized in the preparation of seedling flora. Lubbock's
Seedling (1892), Burger's (1972) 'Seedlings of shrubs and trees of South East Asia', Muller's
(1978) 'Flora of Seedlings for the 'North-Western European lowland', and Vogel's (1980)
Seedlings of Dicotyledons are some of the noteworthy contributions in this regard.
A lot of work has been done outside India, but very little work has been done in the
context of seedling morphology in India in spite of its rich floristic resources. The usefulness of the
cotyledonary leaves as characteristics in the taxonomy of Indian Convolvulaceae was highlighted
by Sampathkumar (1982) which is one of the pioneer works in this field in the context of Indian
flora. Seedlings of 50 commercially important tree species of Kerala forest were studied by
Balasubramanyan and Swarupanandan(1986). Augustine (1993,2004a-b) investigated the seedling
morphology of some species of Tephrosia, Acacia holosericea, and Cassytha filiformis.
Bandopadhyay (2003) studied the seedling morphology of Bauhinia. The seed and seedling
morphology of Catharanthus roseus was investigated by Singh et ah (2008).
Paria and his associates have investigated the seedling morphology of angiosperms and
established their taxonomic significance base on their work for more than two decades. Paria and
his associates (1986-2010) have so far studied seedling morphology of good number of taxa at
family, genus and species level. At family level, a good number of taxa of Anacardiaceae,
Asteraceae, Boraginaceae, Bombacaceae, Combretaceae, Euphorbiaceae, Lamiaceae,
Leguminosae, Malvaceae, Pedaliaceae, Polygonaceae, etc. have been examined. At generic level,
investigations have been carried out on Bauhinia, Jatropha, Knoxia, etc. Further, seedling
morphology of sixteen true mangrove species under ten families of Sunderban area were also
investigated (Das et ah, 2001). The application of phenetic analyses using seedling characters
were also employed by Mundhra and Paria (2008) in determining affinities between the taxa of
Jatropha for the first time in the context of Indian flora.
Methodology
Seedlings of different stages of growth and development representing possible inter- or
intra-specific diversity were directly collected from the field. In case of confusion for correct
identity, the natural seedlings were compared with those of seedlings raised from identified seeds
in the experimental garden of the Botany Department, University of Calcutta. The different stages
of seedlings were photographed, dried and preserved in the form of herbarium sheets. The
morphology of seedlings was described following the scheme of description of Vogel (1980) and
terminology as proposed by Duke (1965), Burger (1972), Hickey (1973), Dilcher (1974) and Vogel
(1980) was used. For minor details in morphology (venation, hairs, etc), the seedlings were observed
both in transmitted and incident light by using binocular microscope (Wild M-3 Stereomicroscope)
using different magnifications. All parts of seedlings were described in the sequence of germination
type, taproot, hypocotyl, (para)cotyledons, internodes, first two leaves and subsequent leaves. The
seedling herbarium sheets of the investigated taxa were deposited in the Calcutta University
Herbarium (CUH).

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International Seminar on "Multidisciplinary Approaches in Angiosperm Systematics"

Seedling Characters of Taxonomic Importance

In taxonomy as well as in seedling literature occasionally remarks are found concerning
the importance of seedlings for taxonomic considerations. Differences in seedling morphology
have sometimes been basis for rearrangement of taxa in classification. The taxonomic significance
of the seedlings derives principally from its morphology and all its parts contribute to information
(Fig. 1). These seedling parts exhibit a panorama of diverse characters which are discussed
below.
Germination: On germination the majority of dicotyledons conform to two of several patterns of
developments. The cotyledons are either withdrawn from the testa to serve as photosynthetic
organs or they remain enclosed. When the cotyledons emerge from the testa, the seedlings are
described as phanerocotylar, and when they have not, as cryptocotylar (Duke, 1965). An
alternative terminology often applied to seedlings reflects the position of the cotyledons with respect
to the soil surface and assumes the seed to have been buried. When the cotyledons are raised
above the soil surface, the germination is described as epigeal, and when they remain in the soil as
hypogeal. Seedlings in which only the blades of the cotyledons appear above the soil surface,
either enclosed or free of the testa, are described as exhibiting geal germination (Jackson, 1974).
The majority of seedlings with hypogeal or geal germination are usually cryptocotylar, and seedlings
with epigeal germination are generally phanerocotylar. However, during release irom the testa, the
foliaceous cotyledons (paracotyledons) expand strongly and fleshy cotyledons hardly or not at all.
In the investigated members of angiosperms, six different conditions of germination are encountered
considering the two basic types (cryptocotylar and phanerocotylar) of seedling development and
also epigeal, geal and hypogeal situations of cotyledons with respect to whether the hypocotyl
extends or not. These conditions are cryptocotylar-epigeal (e.g. Jatropha multifield), cryptocotylargeal (e.g. Polyalthia longifolia), cryptocotylar-hypogeal (e.g. Terminalia bellirica),
phanerocotylar-epigeal (e.g. Solarium nigrum), phanerocotylar-geal (e.g. Manihot glaziovii) and
phanerocotylar-hypogeal (Cleistanthus collinus). However, the phanerocotylar seedlings exhibiting
epigeal germination outnumber the other types in angiosperms.
During germination, paracotyledons spread out on the stem axis and persist up to several
leaves stages which vary from species to species. The terminal bud develops into a shoot with
leaves. The first organ which emerges from the testa is the radicle. This later enters the soil,
anchoring the embryo. The hypocotyl is next produced in the shape of a bent knee, but soon
stretches and become erect, pulling the cotyledons out of the surrounding endosperm and testa, or
if adherent, lifting the both.
The Root: The root is the descending part of the seedling axis. It is different from the ascending
part of the axis in having the collet (collet is the junction between the descending root part and the
ascending stem part). In hypogeal species, the radicle consists usually of the axis below the
cotyledons. A short to very short hypocotyl is present, comprising of the transitional zone between
the radicle and cotyledonary node. The establishment of young plant is effected through the
development of sturdy tap root. Depending on measurement tap roots may be reduced (i.e. <2 cm,
e.g. Ageratum conyzoides), shortly elongating (i.e. 2-4 cm, e.g. Knoxia roxburghii), elongating
(i.e. >4 cm, e.g. Knoxiaplantaginea). Lateral roots in some seedlings normally appear when the
primary root has developed into a fair-sized tap root. Their number may be small or large, they may
appear rather soon or relatively late, and they have often a specific arrangement. Lateral roots
also may be branched to various degrees. In Erythrina variegata, Terminalia myriocarpa, etc.,
the side roots (lateral roots) are numerous and profusely branched. Tap roots may have different
colour i.e. white or grayish-white, brownish, creamish white, etc. Root characters are not often
used for identification of seedlings, but sometimes they have diagnostic values.
The Hypocotyl: Hypocotyl is the basal portion of ascending axis and consists of one internode
between the paracotyledonary node and the collet. In species germinating in an epigeal manner the

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hypocotyl is already pronounced in the embryonic stage, and on germination, will develop into
elongated hypocotyl. The characters of hypocotyl like measurement, colour, and forms are
sometimes diagnostic and useful in the delimitation of genera and species. It is usually terete or
may be angular-square in Bauhinia malabarica, Bischofia javanica, etc. It may be hairy (e.g.
Hyptis suaveolens, Sesamum indicum, etc.) or glabrous (e.g. Combretum roxburghii, Knoxia
sumatrensis, etc.). Regarding length it may be reduced (i.e. <1 cm, e.g. Terminalia bellirica,
Mangifera indiaca, etc.), shortly elongating (i.e. 1-2 cm, e.g. Acalypha lanceolata, Eupatorium
odoratum, etc.) and elongating (i.e. >2 cm, e.g. Terminalia catappa, Trewia nodiflora, etc.).
The texture of hypocotyl varies from soft to moderately soft or moderately woody to woody.
Hypocotyl may have different colours like light brown, greenish, etc.
The Cotyledons : Seed-leaves, seed lobes or cotyledons, in the broad classical meaning, are the
first leaves of the juvenile plant (de Candolle, 1825). In the embryonic stage they are already
present in a rather definite shape. According to Vogel (1980), cotyledon occurs in three different
manifestations. They may be either food-storing (fleshy), haustorial or photosynthetic at the moment
of germination (Fig. 2). He used the term 'paracotyledons' for the photosynthetic cotyledons.
Real cotyledons are usually food storing or haustorial in nature, massive, succulent and light in
colour. Cotyledons have been observed both in cryptocotylar species and also in phanerocotylar
species such as Schleichera oleosa, Erythrina variegata, Pongamia glabra, etc.
The position of cotyledons is on the top of hypocotyl, inserted at the cotyledonary node.
In phanerocotylar seedlings, the cotyledons are of opposite phyllotaxy (e.g. Erythrina variegata).
However, in cryptocotylar species (e.g. Jatropha multifida, Madhuca indicd), the cotyledonary
blades are often borne lateral on the stem (secund) although their place of attachment is opposite,
or they are opposite on either side of the stem. Among the investigated angiosperms, cotyledons
are exstipulate and sessile or shortly petiolate. The duration of persistence of cotyledons depends
on the seedling types and their functions. The shape of cotyledons does not vary greatly as in
paracotyledons and foliage leaves. They may be suborbicular, ellipsoidal, ovate or obovate. The
morphology of bases of cotyledons may be cuneate, cordate, rounded, etc. and apices may be
truncate, obtuse, rounded, etc. Margin is almost entire in the investigated angiosperms. The venation
pattern of cotyledons in cryptocotylar species is hardly determined.
The Paracotyledons: The paracotyledons are, in general, the exposed, green, leaf-like assimilating
seed leaves. They are relatively thin, but usually 2-7 times thicker than the subsequent foliar leaves
(Marshall and Kozlowski, 1976,1977). In most of the angiosperms, photosynthetic cotyledon or
paracotyledon has been observed. The number of paracotyledons is always two and opposite in
arrangement.
The paracotyledons are petiolate or sub-sessile or sessile. In Ricinus communis, petiole
is 1-2 cm long. The petiole may be hairy or glabrous, terete or angular, etc. The paracotyledons are
exstipulate i.e. there is no stipule at the base of paracotyledon. The paracotyledon persists for
variable duration and stages. The long duration of paracotyledon supports their photosynthetic
nature.
The blades, as a rule, of fully developed assimilating paracotyledons have different shape
compared to foliar leaves or first few leaves of the seedlings. Paracotyledon exhibits wide diversity
in shapes ranging from ovate, oblanceolate, oblong, obovate, suborbicular, lanceolate, linear and
elliptic. Morphological differences are also noted in the form of bases of paracotyledons which
include the forms, e.g. subrounded, rounded, acute, cuneate, attenuate, obtuse, etc. The morphology
of apices of paracotyledons include retuse, obtuse, rounded, acute, mucronate forms, etc. Margin
of the paracotyledons is usually entire. Venation pattern in paracotyledons is mostly distinctly
visible. The primary veins may be one or more. In the latter case, they may be either of pinnate or
palmate type. The secondary vein forming different patterns are also seen in different taxa. Venation
patterns are acrodromous, brochidodromous, cladodromous, eucamptodromous, simple

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International Seminar on "Multidisciplinary Approaches in Angiosperm Systematics"

craspedodromous and semicraspedodromous. The texture of paracotyledons is herbaceous,

membranous, or fleshy. The colour is almost always green, but sometimes reddish colour is observed
specially on ventral surface. The surface is usually glabrous, sometimes hairy (e.g. Eclipta
prostrata). Paracotyledons are very important for seedling identification. Their nature, shape,
dimension, persistency and colouration may be specific, and give clues for the identification of
taxa.
The Epicotyl and subsequent Internodes: The first internode of the seedlings above the
hypocotyl is called epicotyl. In literature, epicotyl is also used for the entire embryonic axis consisting
of several internodes above the cotyledonary node (Vogel, 1980). The development of seedling
axis is variable in different taxa and accordingly the elongation of epicotyl takes place. The length
of epicotyl and subsequent internodes vary in different species. In several taxa, the successive
internodes are irregular in length. In some phanerocotylar seedlings, usually the first internode i.e.
epicotyl is comparatively longer than the subsequent internodes, e.g. Jatropha multifield, Ageratum
conyzoides, etc. The epicotyl and internodes are generally terete, sometimes angled. They may
be glabrous or hairy. Different colours like green, yellowish green, brown, pale purple, etc. also
occur in epicotyl and internodes.
The Eophylls: Eophylls are the juvenile leaves (including first two and few subsequent leaves)
with green expanded blades developed by the seedlings. Subsequent leaves include leaves on main
axis excluding first two leaves. In the present study, emphasis has been given on the first two
leaves and subsequent leaves i.e. from third leaf onwards. These are usually different from the
preceding paracotyledons in shape as well as in texture, although the resemblance of leaves with
paracotyledons may still be rather striking.
The texture of the juvenile leaves is variable as that of adult leaves i.e. herbaceous,
membranous and coriaceous. Eophylls may be stipulate or exstipulate. Eophylls are usually petiolate
but may be sessile or subsessile. The petiole is generally terete, sometimes angular, channelled, or
concave above and convex below. Petioles may be glabrous or hairy.
The phyllotaxy is an important characteristic feature. It may be specific in seedling both
for the first two and subsequent leaves. Two types of phyllotaxy have been found in the investigated
angiosperms - opposite and alternate. In some species, first two leaves are opposite and subsequent
leaves alternate (e.g. Parthenium hysterophorus). Alternate phyllotaxy of eophylls is found in
several taxa (e.g. Briedelia retusa). Opposite phyllotaxy of eophylls are found in Chrozophora
plicata including others.
The shape of the lamina is various. It may be ovate, lanceolate, obovate, oblanceolate,
orbicular, linear and oblong. Bases of the juvenile leaves are of different types such as rounded,
cuneate, acute, truncate, obtuse, attenuate, decurrent and cordate, etc. The morphology of apices
of juvenile leaves include forms like obtuse, acute, acuminate and mucronate, etc. and serves as
important morphological character in distinguishing species. Base balance is mostly symmetric,
sometimes asymmetric. Margin of the lamina of the juvenile leaves may be crenate, entire, serrate,
lobed, toothed, dentate, pinnatifid, undulate, etc. The lamina of eophylls display different venation
patterns. The number of primary veins varies from one to many. The gross venation exhibits
different patterns like acrodromous, brochidodromous, cladodromous, eucamptodromous, simple
craspedodromous and semicraspedodromous. The veins may be prominent or depressed within
the lamina. The surface of the blade may be glabrous or hairy.
Heteroblastic development of leaves:
During seedling morphological studies, it is often observed that the mature leaves characters
of adult shoots differ from first few leaves characters of seedling stages. There is a gradual
transition from juvenile traits, present soon after germination, to stable adult traits acquired by the
time flowering commences, called heteroblastic development. The changes in morphology of leaves

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are most conspicuous feature of heteroblastic development in angiosperms. In the simplest case
there is merely an increase in leaf size without considerable change in form. However, in Phyllanthns
urinaria, apart from size changes, there are progressive changes in leaf shape from node to node
(Mundhra and Paria, 2009).
In addition to changes in leaf shape, the ontogeny of a plant frequently involves changes
in the phyllotaxy and nature of leaves. Heteroblastic development associated with phyllotaxy of
leaves is generally noticed in cryptocotylar species where first two leaves are opposite and
subsequent leaves are alternate, e.g. Madhuca indica (Mundhra and Paria, 2009). The
developmental changes in nature of leaves, that is, from simple to compound have been noticed in
a large number of taxa. For example, in Erythrina variegata (Paria et a\, 2006) and Bombax
malabarica (Paria et al., 1990), the first-formed leaves are simple and follow the compound
leaves either pinnate or palmate respectively. This heteroblastic development of leaves serves as
marker character and will help in identification of the taxon from closely related species.
Correlation with other Botanical Disciplines:
It is generally accepted that palynology, cytology, phytochemistry, embryology, general
anatomy, morphology, etc. are some of the disciplines which are generally considered by taxonomists
to offer comprehensive taxonomic and phylogenetic trends in a taxon (Chanda and Mukherjee,
1978). It is remarkable to mention that different members of the investigated angiosperms have
been studied from different botanical disciplines, and it is of interest to compare the results of the
seedling morphological investigation with those of the other disciplines. However, for this purpose
cytology, phytochemistry and pollen morphology have been taken into consideration.
A perusal of chromosome number and seedling morphological features of some members
of Malvales reveals an apparent relationship among the taxa. Bombax ceiba of Bombacaceae
related to Hibiscus subdariffa of Malvaceae in having same chromosome number namely 2n =
72 (Fedorov, 1969) once was included under the Malvaceae by Bentham and Hooker (1862) while
in morphological features of the seedlings, B. ceiba differs from H. subdariffa in having simple to
compound eophyls against simple eophylls of latter. The members of the Malvaceae and the
Sterculiaceae represent closer affinity between the two except there are 3-9 main veins in eophylls
in the former and 5 in the latter, though they do not show any affinity in relation to chromosome
number. At intra- and inter-generic as well as interspecific level of taxa such as Sida cordata,
Sida rhombifolia and Urena lobata, where the chromosome number is constant i.e. 2n = 28
(Fedorov I.e.), certain seedling features are of value in taxonomic delimitation (Paria et al., 1990).
Seedling morphology of three genera- Martynia, Pedalium and Sesamum from Indian
Pedaliaceae s.l. has been studied (Mukhopadhyay and Paria, 1994). Some salient features of
these taxa are found useful in distinguishing their identities. Due to differences in placentation
characters, pollen morphology (Pal and Paria, 1986) and distributional pattern of phenolic acids in
leaves and fruits (Parvati and Narayana, 1978), it is recommended to place the genus Martynia in
a separate family Martyniaceae. The discriminating features of seedlings lend support towards
arriving at a decision as to separate the genus Martynia having large oblong paracotyledons and
cladodromous type of venation pattern from the Pedaliaceae s.l. and placed in an independent
family Martyniaceae. In sharp contrast, Pedalium and Sesamum have paracotyledons of similar
shape i.e. ovate-oblong and similar venation pattern, i.e. eucamptodromous type.
Lamiaceae is composed of two natural subfamilies: Lamioideae with tricolpate pollen
grains and Nepetioideae with hexacolpate pollen grains. Seedling morphological investigation reveals
that Lamioideae (Anisomeles, Leonurus) are characterized by orbicular paracotyledons and
Nepetoideae {Ocimum, Hyptis) have deltoid paracotyledons with retuse apex. This suggests that
this segregation of Lamiaceae into two subfamilies on palynological ground get support from seedling
studies also (Das et al, 2009).

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International Seminar on "Multidisciplinary Approaches in Angiosperm Systematics"

Concluding Remarks
Seedling taxonomy, therefore offers possibilities for separating taxa. The seedling
morphological characters are as important, reliable and conservative characters as that of floral
ones, and should be used in the delimitation of different taxonomic groups. Although, the organs
and characters of seedlings for survey and observation are limited in number, yet their diversity in
form, shape, size, etc. are so large, that a specific combination of such characters serves the
purpose of delimitation as well as identification. Further, Seedlings also represent the final stage in
the process of regeneration from seeds. Thus, understanding the relationship between the morphology
and function of seedlings can also provide an insight into the regenerative strategies of species in
different vegetation types and assist in constructing appropriate models for biodiversity management.
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24(1): 243-250.
Augustine, K.T. 2004a. Seedling morphology of Acacia holosericea A.Cunn. ex GDon. J. Econ. Taxon. Bot.
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Augustine, K.T. 2004b. Seedling morphology of Cassytha filiformis L. (Lauraceae) from Thumba,
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Farnsworth, E. J. 2008. Physiological and morphological changes during early seedling growth: roles of
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International Seminar on "Multidisciplinary Approaches in Angiosperm Systematics"

Subsequent leaves

First Two Leaves

p
H
Y
L
L

Epicotyl
Paracdtyledon
* Hypocotyl

Fig. 1: Parts of a Seedling Plant

(XaMtkum strumarium)

t-^. fj-WM* *

\ *f

(a)
Fig. 2: Different forms of cotyledons (ct), scale bar: 1 cm
a- houstorial (Teminaha bellmca),
b- fleshy (Schleichera oleosa),
c- foliaceous/paracotyledon (Scholium \iride)