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Gymnosperms Gnetopsida - 1 - 1. Details of Module and its Structure Module Detail Subject Name Gymnosperms Paper Name Systematics and Evolution- II: Pteridophytes and Gymnosperms Section B: Gymnosperms Module Name/Title Gnetopsida –I: Morphology Module Id Bot/Syst II/Gnetopsida-I: Morphology 30 Pre-requisites Gymnosperms, basic knowledge of cell and structural biology Objectives To provide Morpho-anatomical knowledge about Gnetopsida Keywords Gnetopsida, Ephedra, Gnetum, Welwitschia Structure of Module / Syllabus of a module (Define Topic / Sub-topic of module) 1. Leaf (Ephedra) 1.1. Morphology 1.2. Anatomy 2. Stem (Ephedra) 2.1 Morphology 2.2. Anatomy 3. Leaf (Gnetum) 3.1. Morphology 3.2. Anatomy 4. Stem (Gnetum) 4.1 Morphology 4.2. Anatomy 5. Leaf (Welwitschia) 5.1. Morphology 5.2. Anatomy 6. Stem (Welwitschia) 6.1 Morphology 6.2. Anatomy 7. Summary 8. References Systematics and Evolution- II: Pteridophytes and Botany Gymnosperms-Section B: Gymnosperms Gnetopsida - 2 - 2. Development Team Role Name Affiliation National Coordinator Prof A. K. Bakshi Vice-chancellor of Allahabad based Uttar Pradesh Rajarshi Tandon Open University Subject Coordinator Prof Sujata Bhargava Deaprtment of Botany, University of Pune. 09604079253 [email protected] Paper Coordinator Dr. P. L. Uniyal Deaprtment of Botany, University of Delhi. 09968279822 [email protected] Content Writer/Author (CW) Dr. Prabha Sharma Deaprtment of Botany, University of Delhi. 08010575034 [email protected] Content Reviewer (CR) Prof. Dinesh Kumar Deaprtment of Botany, University of Lucknow. 09415014059 [email protected] Language Editor (LE) <LE Name> Systematics and Evolution- II: Pteridophytes and Botany Gymnosperms-Section B: Gymnosperms Gnetopsida - 3 - TABLE OF CONTENTS (for textual content) Introduction Ephedra 1. Leaf 1.1. Morphology 1.2. Anatomy 2. Stem 2.1. Morphology 2.2. Anatomy Gnetum 3. Leaf 3.1. Morphology 3.2. Anatomy 4. Stem 4.1. Morphology 4.2. Anatomy Welwitschia 5. Leaf 5.1. Morphology 5.2. Anatomy 6. Stem 6.1. Morphology 6.2. Anatomy 7. Summary 8. References Systematics and Evolution- II: Pteridophytes and Botany Gymnosperms-Section B: Gymnosperms Gnetopsida - 4 - e-Text Ephedra L. Introduction The plant division Gnetophyta of Gymnosperms consists of three genera Ephedra, Gnetum and Welwitschia. Gentophytes differs from other gymnosperms (cycads, conifers and Ginkgo) in having vessel elements (water transport system within the plant) which are predominantly found in flowering plants. The gnetophytes evolutionary relationships remain unclear: in some classifications, all three genera are placed in a single order (Gnetales), while in other treatments they are distributed among three orders, each containing single family with a genus. Most morphological and molecular studies confirm that Gnetum and Welwitschia diverged from each other more recently than from Ephedra (Crane et. al. 2004; Ickert-bond et. al. 2009). Present module gives an insight into the plant morphology and anatomical details of stem and leaf of Ephedra, Gnetum and Welwitschia. Ephedra The genus Ephedra L. comprises 50–60 species that are distributed in the arid regions of New and Old Worlds from the Mediterranean and Black sea shores to China including northern and north-eastern India. (Stevenson, 1993; Price, 1996). The plant exhibits extreme xerophytic characters. It is mostly shrubby, generally having a height of less than 2m. However, some species are lianas, others spread by underground rhizomes and E. triandra is a small tree with stem diameter of about 30cm. E. campylopoda bears pendulous branches. Eleven species have been reported in India, including E. foliata Boiss, E. gerardiana Wall. ex Stapf, E. intermedia Schr. & Meyer, E. nebrodensis Tineo, E. pachyclada Boiss, E. saxatilis (Stapf) Royle ex Florin, E. regeliana Florin, E. przewalskii Stapf, E. sumlingensis Sharma & Uniyal, E. kardangensis Sharma & Uniyal, E. khurikensis Sharma & Uniyal. These species are distributed in the hot desert area of Arawali as well the cold deserts of Western Himalaya (Sharma et. al. 2010). 1. Leaf 1.1. Morphology Fig. 1, 1. Habit Fig. 2. Cuticle of the upper surface. The lower side shows 2,3. Stem with papillate epidermal cells. opposite decussate leaves. The leaves are small, thin scaly, connate and fused into a basal sheath. They are green when young but turn brown at maturity and eventually shed. Accessory buds arise below and at the base of axillary shoots. Papillae are found in some of the cells of the upper (adaxial) epidermis of leaf, which are more prominent in the leaf base (Fig. 2). Papillae of the foliar epidermis are usually directed upwards towards the tip of the leaf. They are normally inserted at or near the distal end of each epidermal cell. Stomata occur on both sides of the leaf but their number decreases steadily towards the base where there are practically none on the upper side. Stomata occur almost up to the leaf apex. Foliar stomata are longitudinally orientated over the lamina but near the leaf base they tend to be irregularly placed (Pant and Mehra 1964). 1.2. Anatomy Fig. 2a. T.S. Leaf Ephedra sp. Transverse section the leaf exhibist an oval outline (Fig. 2a). The epidermis bears a thick cuticle. A few deeply situated or sunken stomata interrupt the epidermis. The epidermis is followed by three layers of chlorophyllose palisade cells. Remaining space is filled by the spongy cells, which are devoid of chloroplasts. Two small vascular traces are embedded in the spongy parenchyma (Bhatnagar and Moitra 1996). Systematics and Evolution- II: Pteridophytes and Botany Gymnosperms-Section B: Gymnosperms Gnetopsida - 5 - 2. Stem 2.1. Morphology In the stem of Ephedra the internodes are longitudinally ridged, and also the ridges of successive internodes alternate. Similarly, the leaves on successive internodes also alternate. Ephedra has two types of branches - indeterminate and determinate. The indeterminate branches at each node have leaves in whorls of threes and sometimes in fours. The determinate branches are borne in the axil of some of the leaves of indeterminate branches, and these bear opposite and decussate leaves (Fig. 11-3). Fig. 3, 1-4 Stem cuticle details. 1. Ridges showing papillations and furrows showing stomata 2. A portion enlarged as in 1 3. Magnified view of stomata showing polar and lateral lamellae 4. Magnified view of stomatal pore. The epidermis of mature internodes is ridged and consists of longitudinally elongated epidermal cells (Fig. 3 1-4) which tend to be in rows parallel to the long axis of the stem. The size of the epidermal cells varies considerably but those of the nodal region tend to be shorter and are often wider than long. The epidermal bands above the ridges are non-stomatiferous and are of somewhat larger cells with papillae on their surface. The form and size of the papillae is very variable, e.g. in the ridges of the internodes of the plant a papilla may usually appear as a short local bulge of a cell where its cuticle is thickened but just above a node, usually on its side which faces a leaf, the papillae tend to be more elongated. Cauline stomata are usually confined to the furrows. In the internodes the stomata are typically placed in longitudinal rows and those of adjacent rows are often alternating (Pant and Mehra 1964). 2.2. Anatomy The longitudinal ridges on the stem impart it a wavy outline. In a transverse section it shows the presence of ridges and furrows (Fig. 4A.2a). The epidermis has a thick cuticle which is interrupted in the furrows where stomata are present. The stomata are sunken, and each consists of two guard cells with a prominent substomatal cavity (see Fig. 3, Fig. 4A.2b). The hypodermis is present only below the ridges and is composed of elongated sclerenchyma which provides mechanical strength to the stem. The cortex is differentiated into an outer single layer of compactly arranged, elongated palisade cells containing abundant chloroplasts. The inner zone is made up of loosely-arranged cells containing fewer chloroplasts. The cortex is delimited by a single-layered endodermis. Vascular bundles are open and endarch. The centre of the pith consists of thick-walled tanniniferous cells. Xylem consists of tracheids, vessels and parenchyma. The tracheids exhibit annular and spiral thickenings and bordered pits, whereas the vessels have only bordered pits. The phloem consists of sieve cells, parenchyma and albuminous cells. The internodes of the stem are quite elongated due to the activity of an intercalary meristem or meristematic plate which is present just above each node (see Fig. 4A.3a, b). After the active phase, the intercalary meristem either forms an abscission layer or matures into a band of sclerified parenchyma. The intercalary meristem is also known as the nodal diaphragm present at the base of each internode makes the stem easily separable at the nodes (Bhatnagar & Moitra, 1996). Fig.4A-3: Ephedra foliata a) A part ofstem at the nodal region in longitudinal section showing the rneristematic plate. b) The intercalary region magnified. (from Bhatnagar & Moitra, 1996). The stem undergoes secondary growth in the usual manner. The beginning of secondary growth is marked by the differentiation of interfascicular cambium. This joins with the intrafascicular cambium to form a ring. The cambial ring cuts off secondary xylem towards, inner side and secondary phloem towards outer side. In the cambial cells, ray initials cut off xylem and phloem rays, whereas the fusiform initials give rise to vascular elements. The annual growth rings are visible with each year's secondary growth (Fig. 4A.4a, b). In a young stem the medullary rays are uniseriate, but in mature stems they become multiseriate (Fig, 4A.4d) by longitudinal division of the ray cells or by fusion of uniseriate rays. The tracheid show bordered pits (Fig. 4A.4C). Presence of vessels is the most prominent feature of the stem (Fig. 4A.4a-c). The secondary phloem Systematics and Evolution- II: Pteridophytes and Botany Gymnosperms-Section B: Gymnosperms Gnetopsida - 6 - consists of sieve cells, phloem parenchyma, albuminous cells and rays.
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