i Yi, _§P Bhatnagar ~~ Alok MoCopyright © 1996, New Age International (P) Ltd., Publishers Reprint 2004 NEW AGE INTERNATIONAL (P) LIMITED, PUBLISHERS 4835/24, Ansari Road, Daryaganj, New Delhi - 110 002 Offices at: Bangalore, Chennai, Cochin, Guwahati, Hyderabad, Jalandhar, Kolkata, Lucknow, Mumbai and Ranchi This book or any part thereof may not be reproduced in any form without the written permission of the publisher. This book cannot be sold outside the country to which it is consigned by the publisher without the prior permission of the publisher. Rs. 250.00 ISBN : 81-224-0792-7 5678910 11 12 Published by New Age International (P) Ltd. 4835/24, Ansari Road, Daryaganj, New Delhi-110 002 and printed in India at Print Perfect, Mayapuri, New Delhi-110 064.CONTENTS 2, PROGYMNOSPERMS AND THE ORIGIN or GYMNosPERMS 3.___PTERIDOSPERMALES | 5. CayTONTALES 7. CYCADEOIDALES Susyect INDEX$7 teow BPR RATEZ ES ee a me Co it We CS ALAC Aes LAO of the Discovery of Spermatozoids ibe) - Ginkgo biloba (Maidenhair tree) growing at the Botanical Gardens, University of Tokyo ‘Campus from which Professor Hirase made the significant discovery of spermatozo- ids in 1896. Tablet below commemorates the 60th anniversary of the discovery (courtesy: Dr Masahiro Kato).CHAPTER 1 INTRODUCTION Gymnosperms include mainly evergreen trees and shrubs which are extremely captivating because of their graceful habit and attractive shapes. The conifers are greatly valued by garden lovers. Their wide range of shapes, colours, textures, and simple cultivation, make them pre-eminent specimens offering an all the year-round appeal (Pls 1.1-1.5). They form the major component of any temperate forest of the world (Pls 1.6, 1.7). Economically, gymnosperms are highly important particularly in for- estry and horticulture, yielding timber, resins, essential oils, drugs and edible nuts. Conifers are used as Christmas trees. Ginkgo is worshipped and is found growing in compounds of Buddhist temples in China and Japan. The newspaper industry almost wholly depends on the conifer wood for its paper requirements. To a great extent, gymnosperms are responsible for our fossil fuel such as coal. The group has a long history which goes back to at least 2 or 3 hundred million years. Its antiquity goes so far back into the geological times that its origin is lost in the distant past. The first glimpse of this group shows that there were two distinct lines, namely, the Cycadophytes and the Coniferophytes (Fig. 1.1A, B), and these were easily distinguishable by simple characters. We still do not know whether these two lines had a common origin, but what we certainly know is that these two lines are as sharply divided today as they were in the past (see Chamberlain, 1935). The seed plants are generally divided into two groups, gymnosperms and angiosperms. Theophrastus, one of Aristotle’s pupils, is credited with the use of the word ‘gymnosperm’. Gymnosperms, as the name suggests, include plants with naked seeds. At the time of pollination, the ovules are exposed in gymnosperms (gytinos=naked, sperma=seed), unlike in angio- sperms (angios=closed, sperna=seed) where they are enclosed in an ovary. tion. Insect pollination, which is common in angiosperms, is nearly absent in_ gymnosperms. Another very important difference is the absence of double fertilization in gymnosperms, a feature which is unique to angio- sperms. We now have an exception in Ephedra where double fertilization hasbeen reported (see Khan, 1943; Friedman, 1990a,b, 1991) and the fusion product of the male nucleus and the ventral canal nucleus produces supernumerary embryos (Friedman, 1992 a, b). The female gametophyteFig. 1.1:Diagrammatic representation of the habit and comparative size of a cycad (A) and a conifer (B) (after Chamberlain, 1935). GYMNOSPERMS or endosperm in post-fertilization stages remains haploid in gymno- ears In angiosperms, on the: other ind, it is usually triploid. Oftheliving orextantgymnosperms, Cycadales and Ginkgoales are very ancient with a long fossil history and, for this reason and some other characters, their members are aptly called ‘living fossils’. Both the orders were very well represented and had a wide distribution in the past, but now Cycadales include 11 genera (the latest being Chigua) and Ginkgoales are monotypic represented by a single species, Ginkgo biloba. They now show limited distribution in the tropical and subtropical parts of the world. Presently, conifers (Coniferales) are the most conspicuous group of gymnosperms and grow in nearly all parts of the world. Araucariaceae and most of the Podocarpaceae are restricted tothe Southern Hemisphere. In contrast, the majority of Pinaceae are found inthe Northern Hemisphere. The two large families, Taxodiaceae and Cupressaceae, occur in both Northern and Southern Hemispheres. Taxaceae, which is sometimes not considered a true conifer, isdistributed in North America, Europe and Asia. Conifers are represented by nearly 50 genera, notable among them are: Pinus, Cedrus, Abies, Picea, Larix, Tsuga, Pseudotsuga, Taxus, Juniperus, Cupressus, Chamaecyparis, - Biota, Taxodium, Cryptomeria, Sequoia, Sequoiadendron, Podocarpus, Dacrydium, Agathis and Araucaria. Ephedrales, Gnetales and Welwitschialesconstitutea remarkable group among gymnosperms not only because of their restricted distribution but also because of their interesting morphology and reproductivebiology.Introduction 3 Ephedra is found in arid regions of both Eastern and Western Hemispheres. Gnetum, on the other hand, grows in the tropical rain forests of Asia, South America, Africa and in some Pacific islands. Welwitschia is most peculiar and occurs in a narrow strip in south west Africa. The gymnosperm component of Indian flora comprises 14 genera and more than 40 species (see Raizada & Sahni, 1960). These include, besides conifers, Cycas, Ephedra and Gnetum. Members belonging to Coniferales are: Abies, Cedrus, Larix, Picea, Pinus, Tsuga, Cephalotaxus, Cupressus, Juniperus, Podocarpus, and Taxus. Among exotics, which have established fairly well, are Cryptomeria and Callitris. Whereas Callitris is found in South India, Cryptomeria growsextensively in the Eastern Himalayas. Most of thenative conifers grow in the Western Himalayas. Some of the gymnosperms which are commonly found under cultivation are Cycas, Ginkgo, Biota, Thuja, Agathis, Araucaria, Cunninghamia and Taxodium. The gymnosperms comprise woody plants which may be large or small trees, or shrubs. A few may be lianas or climbers. Whereas most of the gymnosperms are evergreen, some are deciduous, such as Larix and Taxodium. The plants are mostly xerophytic. Gymnosperms have some of the tallest trees known in the plant king- dom. Sequoia sempervirens (California or Coast redwood; P1.1.8) is probably the tallest living tree reaching a height of nearly 112 m into the sky and attaining a girth of 15 m; it is as tall as a 36 storey building (see Laetsch, 1979). Perhaps the smallest gymnosperm is a cycad, Zamia pygmaea whose fronds are only 4 or 5 cm long. The most massive and among the oldest, however, is Sequoiadendron giganteum (Big tree; Pl. 1.9). The plants have a well developed tap root system. Coralloid roots as in Cycas and mycorrhizal roots as in Pinus and other conifers also occur. The aerial trunk is branched as in conifers or unbranched as in cycads. In Ginkgo andsomeconifers like Pinus, there are two types of branches orshoots, long shoots and dwarf shoots. The dwarf shoots bear green leaves /needles at their apices. Collectively the whole structure is called spur as in Pinus. There is a great diversity in both form and arrangement of leaves. In addition to foliage leaves, scale leaves may also be present. The leaves may be small (microphyllous) or large (megaphyllous). Both simple and com- pound leaves are known, and their size varies froma minute scale toleaves as long as 2m as in some cycads. Besides size, there is variation in their shapes too. The leaves may be pinnate or fern-like as in cycads or acicular ie. needle-like as in conifers. In Ginkgo the leaves are fan-shaped. The leaves of Gnetum superficially resemblethose of dicotyledons. In Welwitschia the leaves are persistent and strap-shaped (PI. 1.10). Leaves are generally spirally arranged. They may, however, be whorled asin Cedrus or opposite and decussate as in Cupressaceae, Welwitschia and Gnetum. In the stem the vascular cylinder is made up of open, collateral, endarch bundles arranged in a ring. Secondary growth is quite pronounced. An4 GYMNOSPERMS important feature of secondary xylem is the absence of vessels. It comprises mainly tracheids. The tracheids show uni- or multiseriate bordered pits. There may be fiber-tracheids in the late wood, but libriform fibres are absent. It is for this reason the gymnosperm wood is also referred to as softwood, in contrast to hardwood present in dicotyledons. There are two types of wood, namely, manoxylic and pycnoxylic. In the manoxylic type, the wood is sparse with wide parenchymatous rays, large pith and cortex. The pycnoxylic wood, on the other hand, is dense or compact with small xylem rays, and small amount of cortex and pith. In the pycnoxylic type, the wood forms the main bulk of the trunk. Because of its small amount , the manoxylic wood is not important commercially. It is the pycnoxylic wood that provides the timber of commerce. The well defined differences between the two types of wood are of taxonomic significance. It has been shown that there is a correlation between the wood structure, leaf form and seed symmetry. The manoxylic wood is related to megaphyllous leaves and radial symmetry of seeds.On the other hand, the pycnoxylic wood is associated with microphyllous leaves and bilateral symmetry of seeds (see Sporne, 1974). In roots the xylem is di- to polyarch. Tracheids constitute the major part of xylem. Phloem lacks companion cells. An important feature of the leaf of gymnosperms is the transfusion tissue. Although discovered by Frank (1864) in Taxus, the term “transfu- sion tissue” was first used by Mohl in 1871 (see also Hu & Yao, 1981). Stomata are sunken and may occur on the abaxial or both the surfaces. The mesophyll may be differentiated into palisade and spongy parenchyma (Cycas, Gnetum) or undifferentiated (Pinus). In Podocarpus both dorsiventral and isobilateral leaves occur. Venation may be parallel as in Agathis and Welwitschia, reticulate as in Gnetum, dichotomous as in Ginkgo, or there may bea single vein as in most genera. Resin canals occur in the leaves of almost all conifers, except Taxus. Cycads and Ginkgo shows mucilage ducts. In Gnetum latex tubes are present. The vascularsupply to the petiole is quite variable. In Cycas the bundles are arrangedina horse-shoe shaped manner or look like an inverted omega @). These bundles are diploxylic possessing both centripetal and centrifu- gal xylem. In Ginkgo the petiole receives only two bundles and each half of the lamina is supplied by one bundle which, later, dichotomizes. The petiole of Gnetum shows an arc-shaped arrangement of vascular bundles. Vegetative methods of reproduction are rare in gymnosperms. Cycads do propagate through bulbils. Except for Cycas which has ovules borne on loose megasporophylls, in all other gymnosperms, the reproductive structures are borne in cones or strobili that are either staminate (male) or ovulate (female).Introduction 5 Microsporangia are borne on the abaxial or lower surface of the micros- porophyil. The development of the microsporangium in gymnosperms follows mainly two set patterns. It may develop from a meristem that differentiates into epidermis and archesporium as in most gymnosperms or from a meristem with exposed initials as in Cedrus, Larix, Pinus and Pseudotsuga (Fagerlind, 1961, 1971). The exposed initials divide to give rise toepidermisand archesporium. The archesporial tissue divides periclinally to form an outer primary parietal layer and an inner primary sporogenous layer. The primary parietal layer undergoes periclinal divisions to form a multilayered microsporangial wall, the innermost layer of which func- tions as the tapetum. The epidermal cells of the microsporangium develop fibrous thickenings, forming an exothecium. The sole exception is Ginkgo, where fibrous thickenings develop in the subdermal layers, forming an endothecium as in angiosperms. The middle layers are ephemeral and become crushed during maturation. The primary sporogenous cells, on repeated divisions, form a mass of sporogenous cells that mature into microspore mother cells. The microspore mother cells divide meiotically giving rise to tetrads of microspores. Ovules are generally borne on the adaxial or upper surface of the megasporophyll or the ovuliferous scale. The ovule or the megasporan- gium is generally orthotropous. The young gymnospermous ovule con- sists of a central body or the nucellus which is normally surrounded by a single envelope or integument. Exceptionally, the number of envélopes may be 2 or 3as in Ephedra, Welwitschia and Gnetum. A narrow passage left above the nucellusis the micropyle. The chalazais well developed. In some gymnosperms, besides integument, there are other structures surround- ing the ovule. These are the collar as seen in Ginkgo, aril as in Phyllocladus and taxads, and epimatium as in most Podocarpaceae. Another cover- ing reported in some fossil groups such as Lyginopteridaceae and Caytoniales is the cupule. A general pattern of development of a gymnospermous ovule with few representative stagesis shown in Fig. 1.2. There areone to several hypoder- mal archesporial cells. The archesporial cell undergoes periclinal division and gives rise to the primary parietal cell and the primary sporogenous cell. The former by further periclinal divisions forms the parietal tissue and, in this process, pushes the sporogenous cell deep into the nucellus. The sporogenous cell functions as the megaspore mother cell. A densely staining mass below the nucleus in megaspore mother cell in many gymnosperms was seen early in the century (see Coker, 1904; Ottley, 1909). With improvement and advancement in optics, this mass was found to consist of amyloplasts in Pinus. In Encephalartos (De Sloover, 1961) and Ginkgo (Stewart & Gifford, 1967) both plastids and mitochondria were polarised this way.This mass has been referred to in:the literature as the kinoplasmic body. The megaspore mother cell after meiosis forms a tetradGYMNOSPERMS Fig. 1.2: Diagrammatic representation of major stages during the growth of a gymnosperm ovule (arch, archegonium; en, egg nucleus; fg, female gameto- phyte; fm, functional megaspore; fng, free nuclear gametophyte; int, integu- ment; mic, micropyle; mn, male nucleus; ne, neck cell/s; nu, nucellus; pe, proembryo; pg, polien grain; pt, pollen tube; st, spongy tissue; ven, ventral canal nucleus). A.Ovuleatpre-pollination stage. The megaspore tetrad is deep-seated in the nucellus. The lowermost megaspore (larger in size) is functional and the upper three are non-functional and degenerating. The micropyle is open. B. Post-pollination ovule. Micropyle is closed and two pollen grains are seen germinating on nucellus. The nucleus of the functional megaspore after re- peated divisions has given rise to a free nuclear female gametophyte. The nucellar cells adjacent to the female gametophyte are nutritive in nature and constitute the spongy tissue. Degenerated remnants of non-functional megas- pores are visible. C. Same, showing cellular female gametophyte formed through alveoli with two archegonia. The one on left shows neck cells, ventral canal nucleus and egg nucleus and is ready to receive the male gamete. The archegonium on right shows proembryonal cells at its base, arranged in U:S:E pattem.Introduction 7 or triad of megaspores of which the lower one functions and forms the female gametophyte (Fig. 1.2A). The nucellar cells, adjoining the sporogenous cells, become densely cytoplasmic and differentiate into spongy tissue. Some of the non-func- tional sporogenous cells also contribute to the spongy tissue. The latter functions as a nurse tissue to the female gametophyte. By the time the female gametophyte becomes cellular, the spongy tissue gets disorga- nized. In several cycads, after the degeneration of spongy tissue, an endosperm jacket is formed. This develops when the female gametophyte has become cellular and consists of asingle layer of large and multinucleate cells. Fig. 1.3: A, Pinus; B, Larix (al, alveolus reaching up to centre of the female gametophyte;ci, cytoplasmic inclusions; en, egg nucleus; mi, mitochondria; pa, precociously closed alveolus; ta, transversely cut alveolus). A. LS female ga- metophyte just after completion of cellularization through alveoli formation. Whereas most alveoli (al) have reached up to the centre of the gametophyte, some have precociously closed (pa). In the micropylar end, few alveoli are transversely cut (ta). B. Diagrammatic representation of the large and small inclusions (ci) in the archegonial cytoplasm. (A, after Sokolowa, 1890; B, after Camefort, 1968 a). The development of the female gametophyte is monosporic (formed from a single functional megaspore) in gymnosperms, except Welwitschia and Gnetum where it is tetrasporic. There is a fairly uniform pattern of development of female gametophyte. The functional megaspore nucleus8 GYMNOSPERMS divides mitotically to form the free nuclear gametophyte (Fig. 1.2B). The number of nuclei in a gametophyte is more or less fixed for a species. The free nuclear gametophyte becomes cellular through the formation of alveoli. The female gametophyte, therefore, appears to consist of radiating rowsofcells (see Sokolowa, 1890; Figs 1.2C;1.3A).In Gnetumand Welwitschia walls are laid down by free cell formation. Pennell & Bell (1987) claim a similar ontogeny for Taxus. The pollen grains of gymnosperms are wind-borne and it has been aptly remarked that in this group there is hit-a-few and miss-a-lot affair during pollination. The group owes itssuccess to sheer extravagance or abundant pollen production. This becomes obvious if you happen to be in a pine forest at the time when male cones dehisce and pollen grainsare shed. The pollen cover almost everything in gold , and that is why they are referred to as “sulphur showers”. Though generally the gymnosperms are wind- pollinated, some cycads are insect-pollinated (see Tang, 1987a; Wilson, 1993). Entomophily has also been claimed for Ephedra, Gnetum and Welwitschia (see Dogra, 1964; Singh, 1978; Van Jaarsveld, 1990, Kato & Inoue, 1994). The micropyle where the pollination drop is secreted, is variously modified in several taxa. Many conifers show winged pollen. The exact role of wings is still not clear, They either aid in wind dispersal or help pollen float in pollination drop ina definite orientation. A germinal furrow in the pollen grain has been reported in cycads, Ginkgoand some conifers. The furrow closes when conditions are dry, but becomes wide open under high humidity and also when in contact with the pollination drop. The pollen tube emerges through this furrow. In some gymnosperms, such as Cupressaceae, Taxodiaceae, Cephalotaxus, Gnetum and Ephedra, the exine is usually cast off because of the swelling of the pollen grain before germination. Animportant difference between gymnosperms and angiosperms is the absence of pollenkitt in the former, although both entomophily and anemophily are known in the two groups. The pollenkitt production is restricted to angiosperms, and this provides an important proof for the hypothesis that angiosperms are of monophyletic origin (see Hesse, 1980, 1984). The terminology to describe the different cells of the male gametophyte is quite varied. The general course of development of male gametophyte is given on p. 9. Microspore is the first cell of the male gametophyte. In those taxa where prothallial cell or cells are produced, the microspore nucleus divides to form the central cell and the prothallial cell. The central cell divides again producing the antheridial initial and the second prothallial cell. Incycads, where only one prothallial cell is cut off, the antheridial initial is formed after the first division of the microspore nucleus. In taxa lacking prothallial cells, the microspore directly functions as the antheridial initial. ThePlate 1.1: Cyeas revoluta growing at Delhi University Campus.Plate 1.2: Pinus roxburghii growing in natural surroundings (courtesy: Forest Research Institute, Dehra Dun).Plate 1.3: Araucaria bidwilli growing at Lalbagh, Bangalore.Plate 1.4: Biota orientalis growing at Delhi University Campus.“ANSIOATUN TTA JO WapreD [eoUEIOg ayy ye SupMosB wuyof mupaydy :g°1 eld*(ungq eryaq ‘amyysuy ypzeasey ysas0g :Asaymoo) aBues aisnoyyeg ut ys010y aarasay youyureg ul SuLmorS j18sngxod surg Jo Mata onMRIOURd Vy 19° eTPlate 1.7: A luxuriant stand of Cedrus deodara (courtesy: Forest Research Institute, Dehra Dun).Plate 1.8: Sequoia sempervirens (Coast Redwood).Plate 1.9: Sequoiadendron giganteum (Big Tree).sumasnyy Ar0}STH [eNIEN OFeoTYD Je UOHINAySUODAY ‘SIPIQHAIU DIYISPIONIM OUT BELAPlate 1.11: Cycas revoluta growing in front of the Museum of Kagoshima Prefecture in Kagoshima city of Japan. The explanatory plate in the picture means that “This tree of Cycas revoluta is a memorial one from which Professor Ikeno (1866-1943) discovered spermatozoids in 1898, so we must keep it carefully” (courtesy: Professor Tetsuo Nakajima)Plate 1.12: Ephedra foliata. A pair of mature ovules showing prominent pollination drops (courtesy: Dr M.N. Singh).Introduction 9 Microspore central cell prothallial cell antheridial prothallial initial cell antheridial tube cell cell spermatogenous stalk cell cell male male gamete gamete Development of male gametophyte antheridial initial divides to form the antheridial cell and the tube cell. The antheridial cell, in turn, gives rise to a sterile cell or the stalk cell towards the pollen wall, and the fertilespermatogenous (body) cell towards inside. The spermatogenous cell, on division, forms 2 equal or unequal male gametes. In the male gametophyte with regard to the terms stalk cell and body cell, referred to as sterile cell and spermatogenous cell, respectively by Sterling (1948), we tend to agree with him only partially. Since there are other sterile cells like prothallial cells and tube cell in the pollen grain, to label stalk cell as sterile cell would be inappropriate. However, body cell should now be called spermatogenous cell because it tells us about the future derivatives, the sperms or the male gametes, from this cell. Male cells are formed in Cupressaceae, Taxodiaceae, Araucariaceae and Gnetum, whereas male nuclei are produced in Pinaceae, Cephalotaxaceae and Ephedra. Both situations are met within Podocarpaceae and Taxaceae. In cycads and Ginkgo the male gametes are motile i.e. they bear cilia or flagella which is reminiscent of pteridophytes. Also, the pollen tubes in these plants donot carry sperms, but are haustorial invading thenucellus. Multiple male gametes occur in Microcycas and in a few species of Cupressus and infrequently in Juniperus. Hirase’s (1896a) discovery of motile sperms in Ginkgo was a landmark in the history of embryological study of gymnosperms which evoked a lot of interest. This was followed by similar discoveries in Cycas by Ikeno10 GYMNOSPERMS (1898) and in other cycads by Lang (1900) and Weber (1901). Frontispiece and Pl. 1.11 show photographs of plants of Ginkgo biloba and Cycas revoluta from which Professor Hirase (1896a) and Professor Ikeno (1898), respec- tively discovered spermatozoids. The histochemistry and ultrastructure of the male gametes have been studied in some gymnosperms such as Ginkgo, Zamia, and Biota. These studies have provided information with regard to the formation of blepha- roplast and cilia in the motile sperms of Ginkgo, Microcycas and Zamia, and zonation of male cytoplasm in Biota. At the time of shedding, normally, the pollen grains are multicelled containing prothallial cell(s), a tube nucleus and an antheridial cell. In Cupressaceae, Taxodiaceae and Taxaceae, the pollen are shed at the uninucleate stage. A prominent megaspore wall is seen in many fossil seeds, cycads and Ginkgo. Its presence can, however, be demonstrated in almost all the gymnosperms. Ultrastructural studies have shown that the megaspore wall is ontogenetically similar to the pollen wall. It can be compared with the exine and intine of the pollen grains (see Pettitt, 1977; Singh, 1978). The archegonia in the female gametophyte may occur singly or in archegonial complexes. The archegonia are quite large and elongated, and lack neck canal cells (the report of their presence in Taxus baccata by Pennell & Bell, 1987, needs reinvestigation). In Welwitschia and Gnetum there are no archegonia. Instead the former shows prothallial tubes or embryo sac tubes, and the latter shows egg nuclei. Except for Gnetum and Welwitschia, in almostall other gymnosperms the gametophytic cells immediately around the archegonium develop into a nutritive layer or the jacket. Its cells are densely cytoplasmic with promi- nent nuclei. Proteid vacuoles reported in the egg cytoplasm are actually islets of cytoplasm or cytoplasmic inclusions of various types (Fig. 1.3B). The egg, nucleus is fairly large, but stains feebly for chromatin. In post-fertilization stages, the cells of the female gametophyte get filled with reserve food material, giving rise to the ‘endosperm’, which is a haploid tissue. At the time of pollination a ‘pollination drop’ is secreted by the ovule in almost all the gymnosperms (PI. 1.12). This is asugary exudate seen at the micropyle. Some of the exceptions are Abies, Cedrus, Larix, Araucaria and Agathis. The exudate not only receives the pollen, but also transports them to the nucellus. Opinions differ as to how do the pollen grains come to lie on the nucellus- whether they are sucked in or transported through active absorption. Four types of situations have been found to exist with regard to the transportation of pollen to the nucellus. The first situation with erect ovules and wingless pollen is met with in cycads, Ginkgo, Ephedra, Gnetum, and Taxodiaceae. The second situation with inverted or obliquely placed ovulesand wingless pollengrains occurs in Athrotaxis, Taxus and in severalIntroduction 11 members of Cupressaceae. The third situation, typified by Podocarpaceae, Pinaceae and some fossil conifers, shows inverted ovules and winged pollen. The last situation with erect ovules and winged pollen is seen in Phyllocladus, some fossil conifers and Cordaitales (see Singh, 1978). In gymnosperms, usually a long time gap exists between pollination and fertilization. Interestingly, the ovule, during this interval, shows some significant changes. The first noticeable change is the closure of micropyle. This is usually brought aboutby elongation of the subepidermal cells lining the micropyle. The pollinated ovule undergoes considerable enlargement both in the micropylar and the chalazal regions, being much more massive in the chalazal region as in cycads and Cephalotaxus (Fig. 1.4A). In some fossil groups, and in Callitris, Sequoia and Athrotaxis, however, the enlarge- ment is more in the micropylar region than the chalazal region (Fig. 1.4B). Fig. 14: A, Cephalotaxus drupacea; B, Callitris robusta. Post-pollination enlarge- ment of ovule. Note that enlargement is more in the chalazal region in A and in micropylar region in B. (A, after Singh, 1961; B after Baird, 1953). The pollen grains after landing on the nucellus germinate immediaiely or after a resting period. The pollen tube reaches the female gametophyte which may beat different stages of development. Detailsof fertilization are12 GYMNOSPERMS ‘available bothat the light microscope and ultrastructural levels forseveral gymnosperms. The pollen tube after growing through the nucellus and crushing its cells on the way, finally makes contact with the neck of the archegonium. The neck cells degenerate and the contents of the pollentube are released in the archegonium. The functional male nucleus reaches the egg nucleus, sinks into it, and eventually fuses with it, giving rise to the zygote. The cytoplasm of the zygote is contributed either by the male cytoplasm or female cytoplasm, or jointly by both male and female cytoplasm. This new cytoplasm is called “neocytoplasm”. Unlike angiosperms where the division of the zygoteis followed by wall formation, in gymnosperms there is a free-nuclear phase in the develop- mentofembryo. Theexceptionsare Sequoia, Gnetum and Welwitschia where the division in the zygote is followed by wall formation. Similarly in angiosperms, Paeonia is an exception showing a free-nuclear phasein early embryogeny (see Bhojwani & Bhatnagar, 1992). There are three distinct phases in the development of embryo: (i) proembryogeny comprising stages prior to suspensor elongation, (i) early embryogeny including stages after the elongation of suspensor and before the establishment of root meristem, and (iii) late embryogeny involving establishment of the root and shoot meristems, and subsequent development of embryo. Proembryogeny in gymnosperms is quite heterogeneous, and the de- velopment of proembryo can be classified into 4 types: (1) Cycad and Ginkgo type, (2) Conifer type, (3) Ephedra and Sequoia type, and (4) Gnetum and Welwitschia type (for details, see Singh, 1978; Sugihara, 1992). A brief description of conifer type of proembryogeny is given here. The other types will be discussed under respective genera. The conifer type of proembryo development has been termed as the basal plan (Doyle, 1963; Dogra, 1978). The young proembryo is freenuclear because the first and subsequent divisions are not followed by wall formation. The free nuclei lie in the chalazal part of the zygote containing neocytoplasm. The number of free nuclei, before wall formation, varies in different genera being 4 in Athrotaxis, 8 in Pinus, 16 in Cephalotaxus, 32 in Podocarpus, and 64 in Agathis. Prior to wall formation, the free nuclei (Fig. 1.5 A, B) show a definite orientation, arranging themselves in two groups. Whereas the nuclei in the upper group are in one plane, those in the lower group show variable arrangement (Fig. 1.5C). Walls are now laid down resulting in a lower group of variously arranged cells, the primary embryonal tier/group/ cells (pE) and an upper group in one layer, the primary upper tier (pU) (Fig. 1.5D). Because there are no walls on the upper side of the primary upper tier, it is open (Fig. 15D). This is the primary proembryo which denotes the proembryo at wall formation having primary upper and primary embryonal tiers. All the cells of the primary proembryo undergo internal divisionIntroduction 13 (Fig. 1.5E), so called because this is the first division taking place inside the cells, earlier divisions being free nuclear. The internal division results in the doubling of the number of cells. The lower tier (pE) is now called embryonal tier/group (E). The pU tier which undergoes transverse divi- sion gives rise to an upper tier (U) and a lower suspensor tier (S). Thus the Fig. 1.5: A-F. Basal plan of proembryo development in conifers. (E, embryonal group; pE, primary embryonal group; pU, primary upper tier;S, suspensor tier; U, upper tier). A-C. Free nuclear proembryo. D. Primary proembryo showing primary upper tier (pU) and primary embryonal group (pE). E. Cells of primary proembryo undergoing internal division. F. Secondary proembryo with US:E arrangement (for details, see text). primary proembryoafter internal division forms the secondary proembryo with U:S:E arrangement (Fig. 1.5F). Divisions in the pE and pU tiers may not be synchronous, the pU frequently dividing earlier than the pE cells. Cells of the suspensor tier elongate and those of embryonal tier divide forming a mass of cells (e). Since U and S are sister tiers, they contain a similar number of cells. The cells of the U tier and the maternal cytoplasm above it usually degenerate forming a plug. A common feature in the embryogeny of gymnosperms is polyem- bryony, with young seeds having more than one embryo. This could be due to fertilization of more than one egg and developement of many zygotes. This is simple or archegonial polyembryony. More frequently, a single zygote gives rise to multiple embryos usually due to cleavage or splitting of the embryonal tier. This is cleavage polyembryony. Ina mature seed, normally only one embryo is present. There are few reports claiming the occurrence of apomixis in gym nosperms (see Dogra, 1966a). Dogra, basing his observations on Abies pindrow has rightly stated that fusions of gametophytic (prothallus) nuclei, fusion of archegonia, fusion of ventral canal cell nucleus and egg nucleusi4 GYMNOSPERMS or even parthenogenetic division of egg nucleus are no indications of agamospermy. Although there may be a tendency towards initiation of embryo development, maturity is never reached. Such ovules are always sterile. Only through experiments preventing pollination can the occur- rence (or otherwise) of apomixis be proved in gymnosperms. Asa regular phenomenon apomixis seems to be absent in this group of plants. The seed coat or testa in gymnosperms is usually vascularized, conifers being an exception. Among conifers, however, Cedrus and Cephalotaxus show vascular bundlesin the integument. The simplest and most common form of seed vasculature is seen in pteridosperms, Ginkgo, taxads and Ephedra where a ring of two to several bundles traverses the entire length of testa. In cycads and Weltwitschia there are two ringsof vascular bundles. The outer system in cycads comprises several unbranched traces travers- ing the sarcotesta. In the inner system there is a much larger number of traces traversing the endotesta. These traces may fuse with each other forming a network. Whereas the bundles of the outer ring reach almost up to the tip of the integument, those of the inner system stop at the junction of nucellus and integument. In Welwitschia the outer ring, traversing the outer envelope, consists of two unbranched traces. The inner ring has eight traces moving in the inner part of seed up to the base of the inner envelope or integument. In Cephalotaxus the seed coat is supplied by twoinverted bundles giving out two horizontal branches in the middle region towards inner side. In Ephedra, of the two envelopes, only the outer envelope receives a ring of three vascular bundles and from these a few tracheids also move up to the base of nucellus. In Gnetum, however, all the three envelopes of the seed show vasculature. In some fossil forms such as members of Medullosaceae and Cordaitaceae, the nucellus is vascularized. Nucellar vasculature has also been reported in some cycads (see Pant, 1973). A tracheal plate (also called nucellar disc, tracheidal cup, pad or plat- form) has been reported in several taxa.In Welwitschia it is present in the form of basal cup of spirally thickened tracheids and parenchyma in the chalazal region of the ovule. However, unlike other taxa, it is not connected with the vascular supply toovule. Martens (1974) considered this tracheidal cupule as a relict nucellar vasculature. There is a general cytological stability in gymnosperms. The available data follow taxonomic groupings. Except for Cycadaceae and Podocarpaceae, other families show a single basic number. In Cycadaceae the basic number varies from 8 to 13 (x=8,9,11,13); Cycas has x=11 Podocarpaceae show more variation (x=9-13,15,17-19) and in Podocarpus alone the numbers vary from 10 to 19. Ginkgoaceae, Pinaceae, Taxaceae, and Cephalotaxaceae show x=12, whereas Taxodiaceae and Cupressaceae have x=11, Sciadopityaceae shows x=10. The basic number inIntroduction 15 Araucariaceae is 13. Monogeneric families Ephedraceae, Welwitschiaceae and Gnetaceae show basic number as 7,21 and 22, respectively (Khoshoo, 1962). The familial distinctions in Coniferophyta are apparently based on gain or loss of chromosomes. If we regard 12 as the basic number of Coniferales, all the foregoing examples of change in the basic numbers can be attributed to reduction or addition in chromosome number due to unequal translocation and fragmentation process. The system of unequal translocation can alter a karyotype numerically (dysploidy ?). The nature of chromatin present in the proximal portion of the chromosomes deter- mines the direction of alteration. The number would increase if the chromatin present in the proximal part is genetically important. On the other hand, it would decrease if the material is genetically inert (see Khoshoo, 1962). . Polyploidy is rare in gymnosperms having been reported i Jimiperus (partly), Sequoia and Ephedra (partly) where gameticnumberis the multiple of the basic number of the families. Sequoia is the solitary gymnosperm which is hexaploid. Ephedra is the only genus where polyploidy is com- mon. It has been suggested that the rarity of polyploidy in gymnosperms could be attributed to the woody habit and ecospecific differentiation of species (see Khoshoo, 1959, 1962). Gymnosperms, most of which are temperate plants, show very little, if any, activity in the development of their reproductive structures during winter. The activity, however, is renewed in the spring. Whereas the male cones initiate in summer, the female cones do so in autumn. Witha couple of exceptions, the pollination of the young ovules takes place in spring. There is thus a seasonal development in the reproductive cycle of temper- ate gymnosperms. Contrarily, in gymnosperms growing in the tropics and subtropics such as cycads, Gnetum, and Ephedra, the development of ovule is a continuous process. Since the development of the male cone and the ovule is more or less synchronous on a tree or trees ina stand, the time of pollination or fertilization is almost the same year after year, and one can, therefore, be almost certain of the stage one is collecting. A non-synchro- nous behaviour can, however, occur in such plants which are under cultivation as has been observed in some species of Callitris and in Ephedra foliata. The time lag between the initiation of ovule and maturation of seed is long in gymnosperms. Ephedra, exceptionally, shows a reproductive cycle of 3 or 4 months. In most gymnosperms, however, the reproductive cycle may be of 1-year, 2-year or 3-year type. The number of winter rests followed by renewed growth in spring that an ovule passes through determines the type of reproductive cycle. A very convenient method to indicate the various stages in the repro- ductive cycle of gymnosperms with regard to the time of the year using certain symbols (different types of lines; Fig. 1.6) has been given by16 GYMNOSPERMS Favre-Duchartre (1962). In the reproductive cyclediagrams (Figs. 1.7-1.10), the numbers represent the months. Since the seasons ofa year donot corre- spond to the same months in the Northern and Southern Hemispheres, these have also been indicated in these diagrams. 2 5 £ : i ¢ ——______---3 young ‘sporogenous g hod 3 sporangia tissue gameu > embryogenesis 3 $a g Fig. 1.6: Major events/stages in the reproductive cycle of a gymnosperm represented by different types of lines (wavy, straight, broken and parallel). These lines have been used in the subsequent four diagrams of life cycles in this chapter (after Singh, 1978). winter winter 4 J CA ‘spring summer ‘summer Fig. 1.7: One-year type life cycle represented by Cedrus deodara (A)and Gnetum ula (B) as occurring in Western Himalayas and Westen Ghats of India, respec- tively. The numbers represent the months of the year. Pollination and develop- ment of gametophytes in Cedrus take place in autumn. The seeds in Gnetum are shed in summer but the embryogeny continues (after Singh, 1978). Briefly, the 1-year reproductive cycle is represented by most Pinaceae, several members of Cupressaceae, Taxodiaceae and Podocarpaceae and Gnetum (Fig. 1.7A, B). The 2-year reproductive cycle is shown by Araucariaceae, several members of Podocarpaceae, some Taxodiaceze, most species of Pinus (Fig. 1.9A), Cephalotaxus and Ginkgo (Fig. 1.8A, B). The 3-year type of reproductive cycle is exemplified by 3 species of Pinus (Fig. 1.9B) and one species of Juniperus (Fig. 1.10A). Since several cycads complete their reproductive cycle in more than two years, they have also been included in the 3-year type (Fig. 1.10B). For details the reader is referred to Singh (1978).Introduction winter winter autumn summer A summer Fig. 1.8: A, Cephalotarus drupacea; B, Ginkgo biloba. Two-year type life cycle. In Cephalotaxus, both pollination and fertilization take place in spring but the latter occurs after one year of pollination. Seeds are shed in late summer of second year. In Ginkgo, on the other hand, the pollination takes place in spring and fertilization in early autumn of thesame year. Interestingly the development of embryo continues even after seed shedding (after Singh, 1978). winter Fig. 1.9: Two- and Three-years type life cycles as exhibited by most species of Pinus (A) and Pinus pinea (B), respectively. A. Pollination and fertilization take place in the late spring of first and second year, respectively. The ovules over winter twice. In Pinus pinea (B), the ovules are pollinated in spring of first year and get fertilized in the spring of third year. Thus the time lapse is of two years. Seed shedding occurs in autumn (after Singh, 1978). 17 Another interesting feature of conifers is their cyclic bearing. There may be abundant cone crops after certain number of years. In Pseudotsuga, it18 GYMNOSPERMS winter spring summer B Fig. 1.10: A, Juniperus communis; B, Cycas circinalis. Three-year type life cycle. Pollination in Juniperus is followed by fertilization after a lapse of about 13 months. Seeds are shed in the following summer after a gap of another 12 months. Cycasshows pollination in first year winter, fertilization in second year summer i., after five months, and seed shedding in third year summer. Note that embryogeny continues even after the seeds are shed (after Singh, 1978). commonly occurs every five years. It has been possible, with the methods available, to predict very poor or very good seed crops years. The gymnosperms are considered less advanced than angiosperms because their ovules and seeds are unprotected. They are thus thought to occupy an intermediate position between pteridophytes and angiosperms. Besides unprotected seeds, there are several other features which put gymnosperms in adisadvantageous positionas compared to angiosperms. The gymnospermous plants fail to grow in varied habitats; grow very slowly; they do not freely propagate vegetatively; xylem lacks vessels except for a couple of genera; there is no self-pollination because cones are unisexual; and seed dispersal is by wind and man, or by birds as in Taxaceae. In several species of Pinus cones open only when stimulated by fire. In their long evolutionary history, the gymnosperms have had many diverse lines of evolution. During this period many groups reached their zenith, but later became extinct. Of these many lines of evolution, Cycadofilicales (pteridosperms or seed-ferns) were once very prominent. The plants had fern-like foliage that bore seeds. Two other lines, also extinct, were the Cordaitales and Bennettitales. Cordaitales belonged to the late Palaeozoic and Mesozoic, and were considered to be the ancestors of conifers and Gnetales. Bennettitales were prevalent during early Meso- zoicand resembled living cycads. They bore bisporangiate cones and were considered by some as precursors of angiosperms.Introduction 19 Earlier studies on the reproductive biology were routine investigations of the life history of different taxa. With the advent of sophisticated techniques such as scanning and transmission electron microscopy, inter- est developed in the subcellular state or fine structure of various reproduc- tive structures. From this flowed new information on the structure and development of male and female gametophytes and fertilization (see Konar & Moitra, 1980; Moitra & Bhatnagar, 1982). Presently, in addition to the histochemical and ultrastuctural studies, the emphasis is on the in vitro propagation or micropropagation of gymnosperms, especially conifers. This is gaining momentum because there is urgent need to rehabilitate the depleting forests, and this is one quick method to achieve this goal. Classification of gymnosperms: Being an important group of plants, gymnosperms have been variously classified by different workers. Some of these are being briefly considered below: Till many years after the recognition of gymnosperms by Robert Brown in 1827, they were placed alongwith angiosperms. Even Bentham & Hooker (1883) in their Genera Plantarum, placed them between monocotyledons and dicotyledons. It was Van Tieghem (1898) who first recognised gymnosperms as one of the two major divisions of Spermatophyta : the Gymnosperms (Astigmatae) and Angiosperms (Stigmatae). The entire group of gymnosperms was straightway divided into orders (Cycadofilicales, Bennettitales, Cycadales, Cordaitales, Ginkgoales, Coniferales and Gnetales) by Coulter & Chamberlain (1917). It was later modified by Chamberlain (1935) who divided the group into two classes: Cycadophytes (including Cycadofilicales, Bennettitales and Cycadales) and Coniferophytes (including Cordaitales, Ginkgoales, Coniferales and Gnetales). Sahni (1920a) laid stress on the origin of ovules (axial or foliar) and divided gymnosperms into (a) Stachyospermae (Cordaitales, Ginkgoales, Coniferales and Taxales), and (b) Phyllospermae (Pteridosperms and Cycadales and probably Bennettitales). The class Cycadophytes of Cham- berlain is similar to the Phyllospermae of Sahni. Also Stachyospermae of Sahni is similar to Coniferophytes of Chamberlain except that former recognized Taxales as a separate order and later included Gnetales as a separate order in the class. Arnold (1948) for the first time brought in evidences from fossil records into his system of classification. He recognized three separate classes viz., (a) Cycadophyta (Pteridospermae, Cycadeoidales and Cycadales); (}) Coniferophyta (Cordaitales, Ginkgoales, Taxales and Coniferales), and (c) Chlamydospermophyta (Ephedrales and Gnetales). According to him the seed habit arose independently in Cycadophyta and Coniferophyta: the Gymnospermae, thus, representing a polyphyletic group. Pant (1957) suggested a classification which is essentially a modification of Arnold’s classification. He raised these classes to three Divisions viz.,20 GYMNOSPERMS Division 1 : Cycadophyta (a) Class Pteridospermopsida (Orders Lyginopteridales, Medullosales, Glossopteridales, Peltaspermales, Corystospermales, Caytoniales). (b) Class Cycadopsida (Order Cycadales) (0) Class Pentoxylopsida (Order Pentoxylales) (d) Class Bennettitopsida (Order Bennettitales) Division 2: Chlamydospermophyta Class Gnetopsida (Orders Gnetales, Welwitschiales) Division 3: Coniferophyta (a) Class Coniferopsida (Orders Codaitales, Coniferales, Ginkgoales) (b) Class Ephedropsida (Order Ephedrales) (c) Class Czekanowskiopsida (Order Czekanowskiales) (d) Class Taxopsida (Order Taxales) Pant’s classification of gymnosperms into three Divisions was followed by Andrews (1961) classification where he created six divisions. These are: Pteridospermophyta (Pteridospermales), Cycadophyta (Cycadales and Bennettitales), Ginkgophyta (Ginkgoales), Coniferophyta (Coniferales and Cordaitales), Gnetophyta (Gnetales) and Gymnosperms of uncertain origin (Pentoxylales and Vojnovskiales). Sporne (1974) has by and large adopted the classification of Pilger & Melchior (1954). He recognized three classes viz., Cycadopsida (Pteridospermales, Bennettitales, Pentoxylales and Cycadales); Coniferopsida (Cordaitales, Coniferales, Taxales and Ginkgoales), and Gnetopsida (Gnetales). Stewart (1983) placed Progymnospermopsida, Gymnospermopsidaand Gnetopsida as distinct classes under the Division Trachaeophyta (vascular plants) of Kingdom Plantae. He included orders Aneurophytales, Archaeopteridales and Protopityales under Progymnospermopsida, and twelve orders viz., Pteridospermales, Cycadales, Cycadeoidales, Caytoniales,Glossopteridales, Pentoxylales, Czekanowskiales, Ginkgoales, Cordaitales, Voltziales, Coniferales and Taxales in Gymnospermopsida. The class Gnetopsida included Gnetales, Ephedrales and Welwitschiales. Meyen (1984) adopted a system of classification of gymnosperms with the single Division, Pinophyta (=Gymnospermae). He included three classes namely Ginkgoopsida, Cycadopsida and Pinopsida. Under Ginkgoopsida he placed nine Orders which included besides Ginkgoales, such orders as Caytoniales, Glossopteridales, Pentoxylales, Ephedrales, etc. Cycadopsida had six orders including Gnetales and Welwitschiales. Class Pinopsida had two orders namely Cordaitales and Pinales, the latter with 13 families both extinct and extant. On the face of it, Meyen’s system is a good survey but the placement of Ephedrales, Gnetales and Welwitschiales appears highly unsatisfactory. Gifford & Foster (1989), in the process of establishing major taxa, have raised the important groups to the rank of Divisions. They have recognizedIntroduction 21 seven Divisions namely Progymnospermophyta, Pteridospermophyta, Cycadophyta, Cycadeoidophyta, Ginkgophyta, Coniferophyta and Gnetophyta In their treatise, Kramer & Green (see Kubitzki, 1990) have divided Gymnosperms into two subdivisions : Coniferophytina and Cycadophytina. Under Coniferophytina, they have included two Classes namely Ginkgoatae and Pinatae. Ginkgoatae has a single family, Ginkgoaceae and Pinatae comprises nine families viz., Taxaceae, Cephalotaxaceae, Phyllocladaceae, Podocarpaceae, Araucariaceae, Sciadopityaceae, Taxodiaceae, Cupressaceae and Pinaceae. Interestingly under Cycadophytina, Kramer & Green grouped two Classes namely Cycadatae and Gnetatae. Cycadatae has a single order Cycadales with four families as recognized by Stevenson (1985). These are Stangeriaceae, Boweniaceae, Cycadaceae and Zamiaceae. Like Cycadatae, Gnetatae also contains a single order Gnetales with three families, Ephedraceae, Gnetaceae and Welwitschiaceae. Without going into the merits and demerits of their classification, to us there seems hardly any justification to club Gnetales with Cycadales. In our opinion the different classifications which are in vogue, differ widely in their contents and placement of various orders and families. The classification adopted in this book is a synthesis of views prevalent today. The plan followed in this book is as follows: Gymnosperms 1Class : Progymnospermopsida Order : Aneurophytales Family : Aneurophytaceae Order : Archaeopteridales Family : Archaeopteridaceae Order : Protopityales Family : Protopityaceae Il Class : Cycadopsida Order : Pteridospermales Families : Calamopityaceae, Lyginopteridaceae, Medullosaceae, Callistophytaceae Order ‘ Glossopteridales Family : Glossopteridaceae Order : Caytoniales Families: —_Caytoniaceae, Corystospermaceae, Peltaspermaceae Order : Cycadales Families : Cycadaceae Order z Cycadeoidales (=Bennettitales) Families : Williamsoniaceae, Wielandiellaceae, Cycadeoidaceae22 Order Family Ii Class Order Family Order Family Order Family Order Family Order Families IV Class Order Family Order Family Order Fami GYMNOSPERMS Pentoxylales Pentoxylaceae Coniferopsida Ginkgoales Ginkgoaceae Czekanowskiales Czekanowskiaceae Cordaitales Cordaitaceae Voltziales. Voltziaceae Coniferales Pinaceae, Taxodiaceae, Cupressaceae, Podocarpaceae, Araucariaceae, Cephalotaxaceae, Taxaceae. Gnetopsida Ephedrales Ephedraceae Gnetales Gnetaceae Welwitschiales WelwitschiaceaeCHAPTER 2 PROGYMNOSPERMS AND THE ORIGIN OF GYMNOSPERMS Fossils: Before we study some of the important fossil types belonging to the group gymnosperms and attempt to trace the evolutionary tendencies within this group, it would be worthwhile to acquaint the reader with various types of fossils, their reconstruction and nomenclature, and meth- ods used to study them. Since frequent references will be made to the geological history of gymnosperms in subsequent chapters, a summary of the geologic time scale is depicted in a tabular form in this chapter. Fossils are the vestiges or traces of plants and animals of the past. It is derived from the Latin word ‘fossilis’ which literally means “to dig up’. Thus, to begin with, large number of things dug out of earth’s crust were called fossils. Besides plant and animal remains, these included minerals and rocks also. Gradually, however, the word was used in a much re- stricted sense, limited to plants and animals only. It may bea shell, abone, aleaf imprint or a piece of petrified wood. The matrix in which the fossils are preserved is usually hardened into rocks. Fossils have attracted the attention of man since long. As early as 450 B.C. Greek historian Herodotus concluded correctly that Mediterranean Sea extended across Africa by observing the presence of seashells in Northern Africa. Fossils are of great importance in thestudy of present day plant and animal lifeas they furnish evidence of the prehistoric lifeand also provide missing links in the evolutionary chain. The study of the fossil remains is anestablished science now. In India, palaeobotany (the study of fossil plants) has generated a lot of interest among the workers. India has the distinction of having a full-fledged research institute (Birbal Sahni Institute of Palaeobotany, Lucknow), the only one of its kind which is entirely devoted to the study of fossil plants. Conditions for fossilization: The basic requirement for fossilization is that physical part of any organism (plant or animal) must be entombed within a protective, usually sedimentary matrix in the earth's crust. This kind of sedimentary environment can be - stream, lake, inland sea, estuarine or continental-shelf deposits. Sometimes diatom frustules may also be incorporated in deposits of deep sea basins. Rarely, the sedimentary environmentis subaqueous (volcanic deposits). Subsequently, the portions preserved in sediments usually become lithified or stony.4 GYMNOSPERMS Sedimentary rocks are those formed by the accumulation of rock particles ranging from colloids to boulders. These particles are derived from the Relative duration fa Period Epocn Duration in Began ot major mrltons of Fons of geologe intervals vous years ave Condes Recent ast 5000 ws Poerstocene 7s ‘Phocene 45 ae Coneroe fos ——-—s—} vn Tertiary [Oigocene 12 = are Eocene 16 a Paleocene W “a 6 Cretaceous n Mesczo1c 136 Jurassic 54 190 Triassic 35 226 Povmian cy | = | Pennsytvanon 45 28) Precambrian Mississippian 20 Paleczoic 248 Devonian 50 398) Siluran 35 430 Ordoncian 70 5007 Cambrian 70 $70. = Precambrian 4030 soo—! The geological column (after Banks, 1970; redrawn from Taylor, 1981) weathering and mechanical abrasion of existing rocks. The breakdown of rocks into smaller units may also occur by chemical weathering, flooding, etc., with rock component released into solution, which resolidifies later. As sediments accumulate, water is squeezed out of them so that they become much more compact and solidify in due course of time. Actually there is a vast range of sediment size and structure, with plant parts even preserved in conglomerates (rocks with various sized particles). Anaero-Progymnosperms and the Origin of Gymnosperms vi} bic conditions and low pHalso favour fossilization. Fossils can sometimes be preserved after forest fires in the form of fossil charcoal or the so called fusain. The sedimentary material such as silicates, carbonates or iron salts should be abundantly present in soluble or precipitate form. Plant parts thatresistdecay and maceration such as walls ofdiatomsand pollen grains provide a good source for fossilization. Among the plant organs usually fossilized leaves are the most commonly preserved part. Types of fossils: A wide range of plant materials is known which have been fossilized under diverse environmental conditions and by different modes of preservation in time and space. We have selected only those examples which reveal the fine preservation that is generally encountered. The large majority of plant materials are preserved as fossils in sedimen- tary rocks. The well known example of sedimentary rock which we come across everyday is coal. In coal, the sediments are of plant origin that are crushed by overlying pressure. Thus the present coal belts represent dense forests of the past. Interestingly, more details of the plant material could be observed in less metamorphosed coal. Thus, lignites which represent the early stage of coal formation, carry plants (or their parts) which are less crushed and could be studied easily. As the degree of metamorphosis increases from lignites to bituminous to anthracite coal, the plant parts are excessively crushed. By special techniques (through sectioning or macera- tion by chemicals), we can study the plant material in coal such as wood fragments, cuticle, bark, algal colonies, cyanobacteria, and the most com- monly occurring (well preserved too) pollen grains. Bituminous coal is, thus, of great importance to the study of palynology. Rarely, coalis formed consisting only of cuticle fragments and amorphic organic materials. The cuticle could be peeled off in thin layers. Such special type is known as paper coal. Other interesting examples of plant fossils are amber and diatomite. Amberis the fossilized plant resin mostly secreted by conifer trees that grew in the geological past ranging from Carboniferous to Pleistocene. Here fungal spores, pollen grains, insects, etc. are trapped before fossilization and they become engulfed init. Finally the resin fossilizes into amber and inside this are left fossilized plant parts or insects. Amber, thus, represents fossils within fossils. It has helped researchers to have an insight into evolution of insects and the plants they pollinated. Diatomite is the sedi- mentary rock formed by the remains of diatoms (atype of unicellular algae whose walls have silicon deposits). These diatoms, frequently found in lakes and sea, get deposited at the base and form the rock in course of time. Pseudofossils are a wide variety of wholly inorganic structures that may resemble plantorgans. They are formed due to mineral depositions caused by the percolation or seepage of water in rock crevices, superficially resembling fern leaves. A careful examination reveals an absence of the26 GYMNOSPERMS characteristic regularity of form of a plant organ. These are also known as Dendrites. Some fossils have also been formed in ice-frozen environments in the polar regions (Mummification). The moisture of the tissue gets converted to micro-crystals of ice. The process is similar to deep freezing, e.g, Pleisto- cene preserved Rhino. Another spectacular example is of mammoths trapped in snow and ice fields during the Late Pleistocene (about 2m years) in which unaltered remains have been unfrozen and edible meat obtained. There is a very interesting type of fossil called biochemical fossil which consists of chemical substances like flavonoids, amino acids, chlorophyll, aromatic acids, steroids and branched hydrocarbons. These may be found in the rocks or in the fossilized remains of organisms (Niklas & Giannasi, 1977; Niklas & Brown, 1981). Substances related to cellulose, lignin, cutin and sporopollenin have also been studied (Niklas, 1981). Depending upon the type of fossilization, the fossils have been vari- ously named. Some of these are briefly introduced here. Impressions: In this type of fossils, the rock upon splitting shows negative replica and all the surface of the plant part involved viz., leaf shape and venation in the case of leaf impression. No cellular details can be seen on an impression because there is no organic matter involved during fossilization. Molds, as the name implies, are formed by three dimensional structures, dissolved by percolating ground water, leavinga hollow cavity in the rock. These hollow cavitiesare similar to the original organ in shape and size and are helpful particularly in studying the external features of a plant in three dimensions like leaf bases on a stem or ornamentation of seed and fruits. Molds are formed in circumstances when, before the plant part is crushed, the sediment surrounding it hardens. After the internal plant material dissolves, the hollow of the sediment is the mold. The process is similar to modern day sculpture making. The sculptor makes the original with wood or wax and prepares the mold. When the mold is prepared, the original statue of wood or wax is removed from inside. The molten bronze is then poured into the mold to form the bronze statue (see Taylor, 1981). Casts: Tree trunks, hard seeds and fruits are frequently fossilized as casts, A cast is formed when a hollow (formed by the decay of the tissue) gets filled with sediments (mineral matter). This produces an exact replica of the original plant material and again no actual partof the plantis involved. Both molds and casts are important in the study of external form of plant parts in a three dimensional plane. Compressions: Coalified compression fossils are those in which bulk of plant material gets compressed in layers of sediment. As additionalProgymnosperms and the Origin of Gymnosperms 27 sediments are laid from above, water is squeezed out of them and the plant parts become more compact and flattened. All that remains is the thin carbonaceous film corresponding to original outline of the plant but unlike an impression in which little or no cell structure remains, we find in compression fossils the cuticularized epidermal cells of various plant organs, spores, epidermal hairs and occasionally other cellular details. Thus very valuable diagnostic data may be accumulated from such type of fossils, Rarely, when compressions are formed in low heat and pressure, intercellular details such as microfibrillar organization of cell wall, chloro- plasts with grana, nuclei with chromatin, and plasmodesmata have been observed using TEM (Niklas et al., 1978). A split along the bedding plane, generally yields one part bearing the compression and the other bearing its impression. In compression much of the organic matter of the plant is preserved. During the formation of a compression, a sort of maceration of tissues takes place in nature resulting in the loss of soluble material and gas. These structures, due to heavy pressure, consolidate into dark black coal. While discussing compressions, it may be of interest to know about the artificial compressions made by Walton (1936). He actually used a hydraulic press to compress plants after embedding them in mud and sand. He observed that while vertical dimension of the organ got reduced, the horizontal dimension remained unaffected. Also, the upper face of the fossil was distorted, whereas the lower face remained unaffected. Compressions are mainly formed in a deltaic system. They may also be formed in lagoons, swamps, ponds or in other such situations. The most well preserved plants are found in clay or shale. Sometimes compressions are also found in consolidated volcanic ashes. Petrifaction: This is the first and, for palaeobotany at least, the most informative mode of preservation. A large number of specimensare found in petrified or cellular permineralized (Schopf, 1975) state. Petrifactions are formed when plant parts are completely submerged in water bodies containing dissolved minerals. During their course of formation, various soluble minerals (silicates, carbonates and iron compounds) infiltrate the cells and intercellular spaces replacing the organic molecules and water. The precipitation of these minerals surrounding the cellular remains forms the rock matrix. This precipitation of the soluble salts takes place due to gradual evaporation of moisture from the swamps leading to their super- saturation. Precipitation occurs when the saturation point is reached. A petrified plant fossil is, thus , a mass of plant tissue infiltrated with hardened mineral substances so that most of the internal structure is preserved. Petrifaction is readily produced by freshly ejected voleanicash. A process similar to petrifaction takes place when a plant material is embedded in paraffin wax in the laboratory. The impregnating material, whether mineralsin case of fossils or waxin case of living material, replaces air and water in tissues and cells. Petrifactions are of great value because28 GYMNOSPERMS when examined as thin sections they reveal perfect details and, exception- ally, even protoplasmic structures are preserved. Of the different types of petrifactions, ‘coal bails’ are the greatest source of information on ancient plants and their structure. These are found as nodules in the coal beds of Carboniferous age and are detected during mining operations. The coal balls are found scattered in coal seams like raisins in a cake. Their size ranges from a few centimeters in diameter to very heavy specimens. These are aggregates of plant materials. The coal balls basically comprise of calcium carbonate as a petrifying mineral which prevents plant debris being altered to coal. It is obvious that perfect preservation is extremely rare. Some degree of rotting always takes place. Softer tissues such as parenchyma get distorted first. More resistant and harder tissues are better preserved. This is one of the reasons why we have more information about xylem and bark of petrified plants than of other parts. The cellular structure is much better preserved in a petrifaction than in a compression. Wood, which is a resistant structure, is commonly seenasa petrifaction. One does, however, come across delicate plant materials, nuclei and chloroplasts. Sometimes, thin walled structures such as fleshy seed coats and root hairs are better preserved than tougher parts and this is because of quick impregnation by water rich in minerals. Study of fossils: An impression or mold is generally studied by preparing artificial casts (also called positives). A compression, on the other hand, can bestudied by the transfer technique or by maceration in Schulze’s solution. A compression could also be studied by softening it and subjecting to microtomy to obtain thin sections. For sectioning, the material is embed- ded in plastic. Peel section method is used for petrified material. Earlier, palaeobotanists had to labour very hard to study the petrified fossils. The technique consisted of cutting thin slices of rock containing, fossils by a circular toothless saw with its periphery having diamond particles. The cut surface was then ground smooth using carborundum and cemented toa slide with the help of resin. A second cut was made on this piece leaving a thin slice on the slide. The slice was then ground till it became thin and transparent so that it could be examined under the microscope. Since this was a cumbersome method involving lot of time and effort, palaeobotanists evolved some more methods that were less time consuming, easier and gave better results. This was achieved through etching technique or peel method. In this method the polished surface of the rock or the coal ball is first etched by an appropriate acid, usually nitric acid or sulphuricacid. This results in the dissolution of matrix and only cell walls remain standing in relief above the surface. The surface is flooded with acetone and a cellulose acetate film is rolled down without entrap- ping any air bubble. When dry, the film is removed from the surface of the specimen. The strip carries a thin and clear impression of the specimen thatProgymnosperms and the Origin of Gymnosperms 29 can be made permanent and examined under the microscope. Time of etching is very critical because the thickness of the peel depends on it. A series of peels obtained through this method are comparable to serial microtome sections. Modern day palaeobotanists regularly use transmission and scanning electron microscope, interference and phase contrast microscopy and X- ray analysis to study their material. Pollen grains and spores are specially observed under scanning electron microscope as it shows surface in great details due to range of magnification and depth of focus. X-ray analysi: useful in studying specimens covered by matrix. The X-ray analysis provides details that are impossible to obtain with regular methods but excellent reconstructions could be made by using stereoscopic X-ray analysis. Reconstruction and nomenclature: Entire plants have rarely been found preserved as fossils. The most frequent form is an organ or even a part thereof. These plant parts undergo distortion and fragmentation toa large extent. The biggest problem a palaeobotanist faces is to synthesize the knowledge in reconstructing the entire plant. A detailed study of the vegetative and reproductive parts, during all developmental stages, is needed to accomplish this. Since this is well-nigh impossible, direct or indirect evidences are used in assigning an organ toa particular plant. The discovery of an organic connection between two parts is used as a direct evidence of these parts belonging to the same plant. An indirect evidence is the one in which either structural similarity or the constant association in the same area or presence of similar pollen grains in the ovulesand other such factors are taken into consideration. The following example would bring out the point clearly. Small capitate glands on the surface of the cupule of Lagenostoma lomaxi were observed by Oliver & Scott in 1904. Similar glands were already found on the stem and leaves of Lyginopteris oldhamia (see also Chapter 3). This was considered as an indirect evidence to prove the association of these two organs. Much later (about 1930), a compressed foliage specimen of Sphenopteris hoeninghausi was discovered which had glandular cupules in actual organic connection. The leaves of the above specimen were identical to those borne on Lyginopteris stem. This, then, became the direct evidence in reconstructing the plant. Pollination is also a strong tool in assigning relationships. The associa- tion of Caytonanthus to Caytonia (Harris, 1940, 1941) and the pollen-bearing organs of Glossopteris to its seed-bearing organs (Pant, 1958, 1977) have been established in this way. Thedetached organs or fragments, which arestudied by palaeobotanists, are described under separate generic names, called form genera. These follow the same binomial system of nomenclature under the International Rules of Botanical Nomenclature as for the present day living plants. Let us consider the various organ genera of the Pentoxylales. The stem has30 GYMNOSPERMS beennamed under organ genera Pentoxylon sahnii and Nipanioxylon guptai, leaf under Nipaniophyllut raoi, male fructification under Salitia nipaniensis and female fructifications under Carnoconites laxum and C. compactum. While reconstructing, using these organ genera (except Sahnia nipaniensis which was not known then), Sahni (1948) proposed the Type genus Pentoxylon for the complete plant. Determination of age: The age of various fossils is ascertained by determining the age of the rock in which they are found using radiometric dating techniques. Radioactive isotopes of Uranium **U, *°U), Thorium @*U) and Potassium ("K) are used for determining the age of rocks and strata. These radioactive isotopes are also called ‘geological clocks’. The decay of radioactive isotopes leads to stable isotopes, releasing energy in the process. The rate at which any isotope decays, and gives rise to the stable isotope, is constant. Thus, by measuring relative quantities of radioactive isotopeand the stable isotope the age of the rock and the plants present can be calculated. Use of radioactive carbon, or carbon dating as it is called, is frequently used in estimating the age of biological specimens back to about 60,000 years, since half-life of carbon is very short (5,568 +30 yrs). Developed by Libby (1955), the technique takes into account the natural bombarding of nitrogen (#N) in upper atmosphere by high energy cosmic rays to form radioactive "'C. ¥C, thus formed, is utilized by plants during photosynthesis alongwith the stable isotope of carbon, #C or BC. With the death of the plant, the incorporation of “C also ceases. The amountof “C which remains in the plant material can be estimated and its age determined by using high energy accelerators. Progymnosperms The gymnosperms are believed tohave appeared during Upper Devonian and Lower Carboniferous in the Palaeozoic era i.e., appoximately 350 million years ago. They were preceded by pteridophytes of the psilophytic stock in Upper Silurian when vascular plants are believed to have first originated. This antiquity led some workers (Oliver, 1906; Arber, 1906; Scott, 1906) to propose that ferns , through pteridosperms, gave rise to gymnosperms. A different view was held by Sahni (1920a) and Chamber- lain (1935) who considered that origin of gymnosperms is polyphyletic. However, there was an element of uncertainity which was due to the lack of a connecting link between the groups. This gap was filled, to a great extent, by Beck’s discovery in 1960 of an organic connection between Archaeopteris (with fern like characters) and Callixylon(with gymnosper- mous wood) from a 80 cm long specimen recovered from the Upper Devonian beds of New York State. This discovery led to the establishment of a new class of vascular plants - the Progymnospermopsida (see also Pant, 1984).Progymnosperms and the Origin of Gymnosperms 31 The Class Progymnospermopsida consists of three orders, viz., Aneurophytales (Ancurophyton, Protopteridium, Tetraxylopteris, Triloboxylon and Proteokalon), Archaeopteridales (Archacopteris) and Protopityales (Protopitys). Here we have described the main genus Archacopteris in some detail. Prior to the discovery by Beck in 1960, Archaeopteris and Callixylon were considered as separate genera occurring in the same beds of Upper Devonian. Archaeopteris looked like a fern having large bipinnate fronds where fertile and sterile pinnae were borne on a rachis. These pinnae, in turn, bore sterile and fertile pinnules. Two rows of adaxial fusiform sporangia were present on fertile pinnules. Callixylon, on the other hand, was represented by fragments of branches, twigs and trunks. The wood was pyenoxylic which is characteristic of conifers. Besides other similari- ties, the tracheids exhibited circular bordered pits on the radial walls. Because of close similarity between the wood of Callixylon and that of conifers, Callixylon was considered as an early representative of conifer clan. Beck had, thus, found a plant with leaves of a ‘free sporing’ fern attached to the stem of gymnospermous type. The plant was eventually named Archaeopteris by him. He reconstructed it as a tree, 18 m high, with a crown of spirally arranged bipinnate fronds (Fig. 2.1) The leaves of Archacopteris were considered similar to the bipinnate ‘frond’ of ferns. However, later studies by Carluccio et al. (1966) showed that in A, macilenta the ultimate appendages or ‘pinnules’ are actually spirally arranged simple leaves. They could also demonstrate that leaf traces arise from branches of Archacopteris in a spiral manner. Later work by Beck (1971) on the same species (A. macilenta) showed that the penultimate branches bear both leaves and ultimate branches in a spiral. Since the ultimate branches and the leavesare in the same spiral, the former are, thus not axillary in origin, a situation much different from seed plants. A large variation in leaf morphology has been observed in different species of Archaeopteris. The stelar anatomy of Archaeopteris macilenta (Fig. 2.2B) shows that the mesarch primary xylemis nine ribbed. Branch and leaf traces diverge from these nine ribs - two of them give rise to branch traces and the remaining seven to leaf traces. All the traces are spirally arranged. Before entering a leaf, a trace divides into two and further divisions take place after it enters the leaf to give rise to typical dichotomous venation of Archaeopteris leaf. Secondary xylem is abundantly produced. As stated earlier secondary xylem is compact and, in appearance, it is typically coniferophytic. It comprises only tracheids and vascular rays. There may be clear growth rings or these may be inconspicuous or absent. Pits are confined to radial walls of tracheids. In the late wood pits occur on tangential walls also. Ray tracheids as seen in conifers are also present. Vascular rays, depending upon species, may be multi-, bi-, or uniseriate32 GYMNOSPERMS Fig. 2.1: Archaeopteris tree, a reconstruction (after Beck, 1962).Progymnosperms and the Origin of Gymnosperms 33 with different heights. The most characteristic feature of the wood is its pycnoxylic nature and groups of circular bordered pits present on radial walls of tracheids. The available evidence clearly indicates that all the major, important features of gymnosperm secondary xylem had evolved in the progymnosperms (see Beck, 1970). Fig. 2.2: A,C-E, Archaeopteris halliana; B, A. macilenta (btr, branch trace; Ib, leaf base; Itr, leaf trace). A. Reconstruction of a branch bearing sterile and fertile leaves. Note the leaves are spirally arranged and four ranked. B. TS stem showing prominent leaf bages, leaf traces and branch traces. C. Fertile leaves bearing micro- and megasporangia. D, E. Cluster of microspores and megas- pores, respectively. (A, after Phillips et al., 1972;B, redrawn from Stewart, 1983; CE, after Arnold, 1939).34 GYMNOSPERMS The fertile leaves which occur with sterile leaves on ultimate branches (Fig. 2.2A) are also spirally arranged and bear one or two rows of fusiform sporangia on adaxial side (Fig. 2.2C). Arnold (1939) for the first time showed heterospory in A. halliana (Fig. 2.2D, E). Other interesting observations on Progymnospermopsida relate to the first evidence of cambial activity producing secondary xylem in Protopteridium (Aneurophytales), fossil records of which appear in the Middle Devonian, more than 370 million years ago. Some other genera of this class such as Tetraxylopteris, Triloboxylon and Proteokalon show cam- bium producing phellogen (Scheckler & Banks 1971 a,b). These plants date back to nearly 360 million years. Beck (1971) put forth the concept that Archaeopteris may be the ancestor of conifers because of the pycnoxylic wood and simple decurrent leaves on its branches. Thedeterminate axis of Archaeopteris is believed to have given rise to long and dwarf shoots of conifers with their simple leaves (see Banks, 1970). Itis believed that both Cycadophytes and Coniferophytes have evolved from progymnosperms. The Aneurophytales have given rise to the Archaeopteridales and Cycadophytes, and Archaeopteridales, in turn, gave rise to Coniferophytes. Aneurophytes are considered older because they are protostelic throughout with three-dimensional dichotomous branching and terminal sporangia. The protostele has given rise to eustele as seen in Archaeopteridales, Cycadophytes and Coniferophytes. A study by Scheckler (1978) has revealed that branches of Archaeopteris are protostelic at the base and become eustelic distally. Aconsensus has emerged in recent years that gymnosperms could be of monophyletic origin (Scheckler & Banks, 1971a) having a common ances- try derived from a plexus of ancient plants showing gymnospermous wood and pteridophytic reproduction (Stewart, 1981). Origin of gymnosperm stele: The origin of the gymnospermous stele was earlier believed (Jeffrey 1899, 1902, 1917) to be from a basic protostelic pattern which gave rise to siphonostele by enclosing extrastelar parenchy- matous cortical tissue (as pith) and simultaneous appearance of leaf gaps (ie., formation of parenchymatous tissue in the wall of tubular stele above the point of departure of leaf trace). Jeffrey considered that all other stelar forms are derivatives of the siphonostele by gradual reduction and loss of inner phloem and endodermis and subsequent loss of tubular nature of stele by overlapping leaf gaps. The presence of leaf gaps was considered by Jeffrey (1917) as one of the characteristic features of the siphonostele in Filicales, gymnosperms and angiosperms. He even established a class Pteropsida comprising these three groups. The above concept of the origin of gymnosperm stele changed consi- derably after the study of stele in progymnosperms. Ina series of publica- tions Beck (1964) and Namboodiri & Beck (1968a, b, c) established thataa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.36 GYMNOSPERMS divides tangentially to form two radially aligned strands placed opposite to each other (i.e., these two strands are oriented along the same radius). One of the strands gives rise to a leaf trace and the other one continues as a reparatory strand (Fig. 2.3 B, C). Such a situation is observed in Archaeopteris, Stenomyelon, Calamopitys and some other taxa from Upper Devonian to Lower Mississipian era (c)Change in the orientation in the division of the two strands (leaf trace and reparatory) from radially oriented to tangentially oriented, alternat- ing with each other or placed side by side (i.e., oriented along the same circumference). The nature of the strand remains the same (Fig. 2.3D, E). This situation is found in Lyginopteris, Callistophyton, Poroxylon, etc. and in the present day conifers. Gd) Figure 2.3E shows, in addition to the leaf trace and the sympodial segment, the undulating path taken by the latter during its upward course, a situation common in primary vasculature of living conifers. The reticu- late pattern in some conifers with opposite leaves involves increase in the number of traces supplying a leaf from one to two and change in the direction of divergence of trace are modifications of this scheme. The primary vasculature of Abies concolor described by Namboodiri & Beck (1968b) will reveal the similarity in nature. Figure 2.4 shows vascular cylinder of Abies split openand drawn in two dimension. There are thirteen sympodia following an undulating path. The sympodia, at any stage, do not fuse with each other. Leaf traces have been shown ending in a triangle. It is quite clear that successive leaf traces from any sympodium arise at intervals of 13 nodes e.g., leaf trace number 19 arises above leaf trace number 32 and so on. The primary vascular system of progymnosperms and extinct gymnosperms resembles closely that of extant gymnosperms.’ Leaf traces in progymnosperms and gymnosperms arise directly from sympodial bundles of the eustele without the formation of leaf gaps. According to Namboodiri & Beck (1968c) this would mean that gymnosperm stele evolved from a ‘dissected’ protostele. They suggest that the ‘leaf gap’ in gymnosperms is not homologous with that of siphonostelic ferns. If we at all wish to apply the term leaf gap for the parenchymatous interfascicular region opposite to leaf trace in gymnosperms, it ‘should be used ina purely descriptive sense’, Since in the evolution of stele in progymnosperms and gymnosperms leaf gaps were never formed, the origin of gymnosperms from ferns no longer appears valid, thus refuting Jeffrey's view (see also Gifford & Foster, 1989). Pteropsida comprising ferns, gymnosperms and angiosperms is now considered an unnatural group. Namboodiri & Beck’s (1968 a-c) study supports the view that gymnosperms have evolved from progymnosperms rather than from ferns, and ferns and gymnosperms should be included in separate subdivisions or divisions.Progymnosperms and the Origin of Gymnosperms 37 1 on mW vMovm vinx x xixn xm Fig. 2.4: Abies concolor. Primary vasculature split open to show sympodia and leaf traces. The leaf traces are ending in a triangle. Sympodia are thirteen in number (numbered using Roman numerals). Leaf traces have been numbered according to their ontogenetic sequence (after Beck, 1970). Homospory, Anisospory, Incipient Heterospory and Heterospory Before discussing the origin of ovule and integument, it will be worth- while to relate the steps through which plant groups ultimately attained heterospory from homospory. Unlike some vascular land plants belong- ing to Pteridophyta (Lycopsida, Sphenopsida and Filicopsida) and all Spermatophyta, which seem to have developed heterospory by parallel evolution, none of the bryophytes, which too are land plants show heteros- pory. Nevertheless, afew mossesand hepatics of the family Orthotrichaceae and Targioniaceae, respectively (Ernst-Schwarzenbach, 1936; Ramsay, 1979; Pant &Singh, 1989) show anisospory whichis an allied phenomenon. All thesteps towards heterospory are basically involved around thesize and production of spores within the same or different sporangia. Homos- pory or isospory can be defined by the occurrence of equal sized spores within the same sporangium, which are morphologically similar but genetically they may be similar or different. Most of the bryophytes and pteridophytes show homospory of one kind or the other.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Progymnosperms and the Origin of Gymnosperms 41 A. Fig, 2.6: Archaeosperma arnoldii (cu, cupule; ov, ovule). A. Reconstruction of the cupule complex. Each cupule has two ovules which it covers partially. B. Ovule showing tetrahedral arrangement of megaspore tetrad and lobes of integument (after Pettitt and Beck, 1968, redrawn from Pettitt, 1970). megaspores as seen in Bothrodendrostrobus where only four equal sized megaspores are present (Fig. 2.7B). This is followed by a condition, seen in Bensonites, where two of the four megaspores are larger than the other two (Fig. 2.7C). Archaeosperma, as described above, shows four megaspores and, of these, three are greatly reduced in size (non-functional megaspores?) (Fig. 2.7D). A further reduction in the size of the three non- functional megaspores in a tetrahedral tetrad is seen in Cardiocarpus (Fig. 2.7E). The last stage is a linear tetrad as in Cycas (and in most present day gymnosperms) where the functional megaspore is much enlarged,aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Progymnosperms and the Origin of Gymnosperms 45 off layer, and in Eurystoma it is bowl-shaped. In Salpingostoma dasu, once the pollen are inside the long micropylar tube, they are directed towards pollen chamber by lining of the hairs and the trumpet-shaped salpinx (Fig. 2.10C). A 8 Fig. 2.10: A, Hydraspermia tenuis; B, Eurystoma angulare; C, Salpingostoma dasu (cc, central column; Ig, lagenostome; pe, pollen chamber; sp, salpinx). Median longitudinal sections through seeds to show modification of the tip of the nucellus to form pollen chamber and salpinx (after Andrews, 1963). Geminitheca has a long tube-like salpinx and in Genomosperma kidstoni it is represented by flared out blown off layer. According to Long (1960b), the barrel-shaped structure usually labelled as lagenostome may actually be salpinx in Conostoma, Long, (1961a) emphasizes that evolution of a well developed salpinx may have been one of the earliest modifications of apex of megasporangium or nucellar beak following the establishment of heterospory. Development of salpinx has preceded the establishment of integumentary micropyle in the evolution of ovule.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Pteridospermales 49 Fig. 3.1: A,B, Lyginopteris oldhamia; C, Lyginorachis, D, Kaloxylon hookeri (gl, gland; Itr, leaf trace; oc, outer cortex; pi, pith; pxy, primary xylem; sn, scleren- chymatous nests; sxy, secondary xylem). A. TS stem showing characteristic fibrous outer cortex, massive secondary xylem, prominent double leaf traces and pith with sclerotic nests. B. TLS outer cortex to show anastomosing fibrous strands with ‘windows’ of parenchymatous cells. C. TS petiole; note the capitate glands and trough shaped vascular bundle. D. Same, root with two distinct zones in cortex. (A, C, redrawn from Stewart, 1983; B, redrawn from Andrews, 1961; D, after Scott, 1923). structure is parenchymatous and some cells of the head are secretory in nature. It is believed that these cells secreted an oily or waxy substance of a protective nature. The capitate glands are mere appendages as they are not supplied by any vasculature (Fig. 3.2).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Pteridospermales 53 correspond to the protoxylem points or strands in the vascular segment and not to vascular segments themselves. In Medullosa noei, for instance, Fig. 3.3: A-C. Hypothetical stages in the evolution of Medullosan stems. A. Three ribbed protostele showing mesarch primary xylem with five protoxy- lem points (small white circles). B. Radiating plates of parenchyma (stippled) have dissected the protostele into three segments. C. Each vascular segment is then invested with secondary xylem (hatched). D,E. TS stem Medullosa noei (D) and M. primaeva (E); note the vascular segments with ‘mixed’ protostele. Protoxylem points vary in number. (A-C, After Stewart, 1976; D,E,after Basinger et al., 1974), there may be five or six sympodia (axial bundles) for the two or three vascular segments (Fig. 3.3D). Similarly vascular cylinder of M. primaeva shows 4-5 vascular segments and 13 sympodia (Fig. 3.3E). Figure 3.3 represents hypothetically the steps in the evolution of the Medullosa type eustele (see also Stidd, 1981; Gifford & Foster, 1989). Apart from this interesting feature in the stem, some of the other characteristics of the group are: fronds with bifurcate rachis; large petioles with many scattered traces; large radiospermic, non-cupulate ovules with integument free from nucellus except at the base; simple pollen chamber; large synangiate male reproductive structures with tubular sporangia and monolete pollen grains. Based ona study of a large number of specimens, Stewart & Delevoryas (1956) could reconstruct the Medullosa plant (M. noei; Fig. 3.4). It resembles a tree fern, 3.5 to 4.5 m tall with a crown of pinnately compound leaves. Massive spirally arranged leaf bases are a prominent feature of the stem.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Pteridospermales 57 in association with leaves of Alethopteris. This issupported by thesimilarity of the leaf base structure. The petioles are mostly oval or round in outline and may vary greatly in diameter from a few centimeters at the base to few millimeters in the upper region. In general, the petiole shows a well developed epidermis surrounding strands of radially elongated scleren- chymatous cells. The ground tissue contains numerous scattered vascular segments and gum canals. Mycloxylon lanatum, which is the petiole of Medutllosa noei, shows a little different pattern. Here, the vascular bundles are arranged in a ring followed by an inner ring of gum canals. The sclerenchymatous strands are irregular. The individual vascular bundle of Myeloxylon exhibits a compact xylem tissue which is made up of scalari- form or spirally thickened tracheids, separated from phloem by a narrow parenchymatous patch. The bundle is covered by a sclerenchymatous sheath. Medullosa shows adventitious roots which may be triarch (M. anglica) or tetrarch (M. noei). The prominent secondary wood is divided by broad rays which develop against each protoxylem strand into wedge-shaped segments. Male fructifications: There had been a lot of confusion regarding the identity of the pollen-bearing organs of Medullosaceae. Only rarely have these been found in association with the stem. Also, most of the fructifica- tions have been studied as impressions. Halle (1933) found that some of the previously described pteridosperm “seeds” were in fact pollen organs containing microspores. The earliest studied fructification is that of Whittleseya (Fig. 3.6A) from Ohio, USA. Halle (1933) could prepare mic- rotome sections of the carbonised material. These are cup-shaped struc- tures formed by the fusion of a large number of sporangia side by side with a hollow cavity in the centre and a wide opening. In the compressed state, they look more like a leaf with parallel venation. The individual “veins” actually are sporangia. They were, for a long time, wrongly associated with Ginkgoales. The pollen grains are large (0.2 to 0.25 mm in length) and elliptical when flattened. Some other pollen organ genera have been described which exhibit similar structures. Goldenbergia represents about 8mm long synangium with twelve to sixteen elongated stalked sporangia fused together. Dolerotheca is the most complex and one of the largest pollen organs of Medullosaceae. It consists of four independent radial synangia (each homologous to a single Whittleseya type of structure) which are fused together to form a compound structure. The campanulum of D. formosa (Fig. 3.6B) is hemispherical, up to 4 cm in diameter and with an eccentri- cally placed pedicel. The dehiscing surfaces of opposite sporangia face one another. Each microsporangium is supplied by a vascular trace and dehisces longitudinally (Dennis & Eggert, 1978). Based on Dufek’s (1980)aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Pteridospermales 61 between the seed forms and the corresponding stem forms was found. Most of the inferences are drawn from structural similarities and other evidence of association. CALLISTOPHYTACEAE Established by Stidd & Hall (1970b), the family Callistophytaceae consists of only one genus Callistophyton which was mainly described from the Fig. 3.7: Habit sketch of Callistophyton plant; note the scrambling stem with roots arising at nodes, pinnately compound leaves, and circinate vernation in young leaves (after Rothwell, 1975, redrawn from Rothwell, 1981). Upper and Middle Pennsylvanian sediments of North America. It is a small, shrubby plant with scrambling stem up to 3.cm in diameter. The bipinnately to quadripinnately compound leaves (up to 30 cm long) exhibit circinate vernation when young. Each leaf bearsa bud in its axil; theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Caytoniales 7 Sagenopteris have been compared to angiosperms by Thomas (1925) on the basis of epidermal structure. Harris (1951), too, compared them with dicotyledonary leaves by referring to the formation of abscission layer during the shedding of the leaves. | Fig. 5.1: A,B, Sagenopteris phillipsii; C,D, Caytonanthus kochi; E,l, Caytonanthus; , G, Caytonia nathorstil; H, C. thomasii. A. Palmately compound leaf. B. Portion of leaflet showing reticulate venation. C. Microsporophyll with large number of anther like synangia. D. Two synangia cut open; one with four and other with three sporangia. E. Bisaccate pollen grains. F. Megasporophylll bearing two rows of cupules. G. Young cupule; note the position of ‘lip’ or ‘flange’. H. LS cupuleto show the arrangement of ovules. I. Caytonanthus type pollen in the micropyle of Caytonia ovule. (A, B, F, after Thomas, 1925; C, D, after Harris, 1937; E,L, after Harris, 1951; G, after Harris, 1964; H, after Harris, 1933). Male fructifications: The microsporangiate fructifications (pollen-bearing organs) named Caytonanthus were studied as early as 1829 by Phillips in Yorkshire when these were described as leaves. According to Harris (1964) it was a modified, pinnate microsporophyll. The rachis or the central axis is dorsiventral (the upper and the lower epidermis differing from each other) bearing opposite pinnae. The pinnae branch irregularly and each branchlet bears a single terminal, pendant, tubular, quadrilocular synangium (Fig. 5.1C). The four pollen sacs (Fig. 5.1D) or sporangia, on dehiscence, separate out except at the tip thus releasing the pollen grains (Vitreisporites). The pollen of Caytonanthus are small, nearly 25-35 um,aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Caytoniales 79 PELTASPERMACEAE The family Peltaspermaceae was erected by Thomas in 1933 on the basis ofhis collections made from Natal (South Africa). Similar collections were made by Harris (1932) in Greenland and by Antevs (1914) in Sweden. The family belongs to Upper Permian age. It occurred in Argentina, Australia, Madagascar and China. Only leaves, male and female fructifications are known. Since stems, béaring these organs, have so far not been reported, it is not possible to know the general growth habit of the complete plant. y ( Voc a E n o Fig. 5.2: A. Dicroidium lancifolia pinna. B. Microsporophyll and sporangia of Pteruchus africanus. C. Umkomasia macleani; megasporophyll with cupules. DE, Pilophorosperma cupules with bifurcate micropyle. F. Part of Lepidopteris frond. G. L. otionis, restoration of cupulate disc. Three seeds are still intact. (A-E, after Thomas, 1933; F, after Thomas, 1955; G, after Harris, 1932). The fronds belong to the genus Lepidopteris (Fig. 5.2F). These are up to30 cm long, lanceolate, bi- or tripinnate with a single unforked rachis. Theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Cycadales 83 2. C. pectinata Griff. It grows in the sal forests of Sikkimand Assam, Khasi hillsand Manipur. 3. C. circinalis Linn. This is the most abundant, natu- rally occurring species. It grows in the deciduous forests of Western Ghats and on eastern side as far as Orissa. 4. C. rumphii Miq. It is distributed on the beach for- ests of Andaman and Nicobar Islands Apart from the above mentioned wild species, C. revoluta Thunb. and C. siamensis Mig. are commonly cultivated in the gardens. Morphology: The plants present a palm-like appearance in possessing columnar aerial trunk and a crown of pinnately compound leaves. Some forms have underground short tuberous stems as in C. siamensis (as also in Zamia). The stem is normally unbranched but, sometimes, older trees do exhibit branching. The branch develops as a small bulbil which is actually an adventitious bud. These bulbils arise from the lower fleshy region of the leaf bases. Initially bulbils show only scale leaves but, with further growth, they start producing a crown of leaves, so typical of the main trunk. Later they crush back the leaf bases from which they originated and start appearing as branches. The bulbils also serve as means for vegetative propagation of the plant both artificially and in nature. The branching is usually enhanced by physicalinjury and onecomes across highly branched specimens in parks as against unbranched or poorly branched ones in their native habitat which remains undisturbed (see Pls 1.1 and 1.12). A thick armour of alternating bands of large and small rhomboidal leaf bases covers the aerial stem in a spiral manner (Pl. 6.1A). The large leaf bases belong to the foliage leaves and the smaller ones to the scale leaves (in male plants) and sporophylls (in female plants). The armour is persis- tent but in aged trees the old leaf bases tend to abscise thereby exposing the corky stem surface. This accounts for the trunk being thinner in the lower region than in the upper portion. The age of Cyaas plantscan be determined by counting the number of leaf bases on the trunk and dividing their number by half the number of leaves in the crown (the two seasons’ growth being present at any time). Two crowns of leaves are produced annually, one in the spring and the other during monsoon, coinciding with the favourable seasons of growth. Cycas produces two types of leaves, ie. it exhibits dimorphism. There are large green foliage leaves and smaller scale leaves or cataphylls. Crowns of foliage and scale leaves alternate regularly. The scale leaves areaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Cycadales 87 Each megasporophyll (up to 30cm or more in length) bears two rows of opposite or sub-opposite, 2-12 orthotropous and shortly stalked ovules Fig. 63: A,B, Cycas circinalis; C, C. revoluta (ov, ovule; sc, scale leaves). A. Crown of megasporophylls; note sterile scales at the base. B. Megasporo- phyll showing five pairs of ovules. The upper part of megasporophy!l is sterile and dissected. C. Megasporophyll is pinnatifid and leaf-like bearing ovules in two rows below the pinnae. (A.B, after Maheshwari, 1960; C, redrawn from Gifford & Foster, 1989). (Fig. 6.3 B, C). Cycas produces the largest ovules in the plant kingdom (6 cm long in C. circinalis and 7 cm in C. thourasii). The mature seeds are elliptical or egg shaped, slightly flattened laterally and bilobate. They are fleshy, and bright orange or red. The apex forms a short micropylar tube. The ovule surface may be smooth or hairy, hairs being shed off at maturity. Root: The primary root of Cycas is short-lived and is soon replaced by adventitious roots arising from the stem. It is diarch at the tip but becomes tetrarch or even more higher up at the point of attachment. The xylemaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.90 GYMNOSPERMS full of starch, which is commercially exploited in some species as ‘sago’. The vascular bundles are radially elongated, collateral and open. The protoxylem elements are endarchand show spiral thickenings. The metaxy- lem elements consist of tracheids mostly with scalariform thickenings but sometimes with multiseriate pits. The phloem comprises sieve cells, phloem parenchyma and fibres. Fig. 6.5: A, Cycas circinalis; B, Zamia floridana (cor, cortex; b, leaf base; gr, girdling leaf trace; md, mucilage duct; pi, pith). A. TS stem showing polyxylic condition, armour of leaf bases and a large cortex and pith. B. Same, Zamia showing prominent girdling leaf traces and manoxylic wood. Both cortex and pith are full of mucilage ducts. (A, after Maheshwari, 1960; B, after Chamberlain, 1935). Secondary growth takes place very early in the life of the plant. The interfascicular cambium develops in the parenchymatous cells between the vascular bundles and joins the intrafascicular cambium. The cambium then cuts off radially seriated secondary xylem towards the pith and secondary phloem towards outside. The secondary xylem shows tracheids with bordered pits and scalariform thickenings. The pitting is multiseriate, there being as many as four or five rows of bordered pits which may be oval or round (Fig. 6.6A, C). The cross fields are rarely distinct and show 8 to 12 bordered pits per cross field (Fig. 6.6B). The vascular rays are conspicuous and of three kinds viz., (2) uniseriate rays which are 1-10 cells high, (b) multiseriate rays which are 2-5 cells in width and much higher and (©) foliar multiseriate rays which are continuous from the pith to the phloem. These have a foliar bundle or leaf trace above which is a mucilage duct, usually situated on the phloem side. The ray cells and cortex are full of starch. Calcium oxalate crystals also occur frequently. The phloem is very well developed in the stem and the petiole. It sometimes exceeds the xylem in bulk. In a longitudinal section the indi- vidual sieveelements show numerous sieve areas on their radial walls. Theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.94 GYMNOSPERMS rachis show only a few centrifugal xylem tracheids. Thus, at higher level (distal end), the centripetal xylem is more abundant than the centrifugal xylem (Fig. 6.8B). Such vascular bundles are termed as pseudomesarch or diploxylic bundles as they show the presence of both centripetal and centrifugal xylem (Fig. 6.8C). As the bundles move up into the pinnae, they become exarch with no trace of centrifugal xylem. This condition is, however, rare. The phloemin thebundle is on the side of the protoxylem. Each bundle is surrounded by a bundle sheath. Leaflet: The structure of the leaflet of Cycas is strongly xerophytic (Fig. 6.9A). It is dorsiventral with an epidermis of thick-walled, highly cuticularized cells showing simple pits. The hypodermis is 1- or 2-layered except for the midriband sometimes at margins, where it is3 or 4 cells thick. The hypodermal cells are highly cutinized and lignified. The mesophyll is. differentiated into palisade and spongy parenchyma. A single layer of lignified palisade cells is presentbelow the hypodermis. These cellsexhibit long fibrous thickenings, which make them resistant to the outside radial and longitudinal pressures. The cells of the spongy parenchyma areplaced in away that connects them to the ends of two adjoining palisade cells. The vascular bundle is diploxylic and isolated thick-walled cells extend from it to the hypodermal cells above and below. Some tracheid-like cells with reticulate thickenings or bordered pits on their walls are present on either side of the centripetal metaxylem. These constitute the transfusion tissue. The cells of the transfusion tissue are connected with the surrounding parenchymatous cells. Additionally, between the palisade and spongy parenchyma lie a few layers of empty thick-walled, tracheid-like cells with bordered pits. These cells are at right angles to the longitudinal axis of leaf and run from either side of the midrib to the margins of the lamina forming the accessory transfusion tissue. It probably functions as lateral conduct- ing tissue in the leaflet which lacks veins. The centripetal xylem is sepa- rated from centrifugal xylem by a few parenchymatous cells. A small arc of cambium lies outside the protoxylem towards phloem and probably cuts off some secondary xylem. Towards the phloem, the cambium pro- duces a few layers of secondary phloem. A fibrous sheath surrounds the bundle (Fig. 6.9B). The stomata are sunken (Fig. 6.9A-C) and confined wholly to the underside. They possess a crater-like appearance. The two guard cells lie at the bottom of the crater and between them is theslit-like pore. The guard cells are bordered by 8-10 subsidiary cells arranged in a ring-like manner (Fig. 6.9 D). In later stages these are not easily distinguishable from the rest of the epidermal cells. Since the subsidiary cells and the guard cells do not have a common origin, the stomata are haplocheilic. In haplocheilic stomata the guard cells and subsidiary cells are not related ontogeneti- cally. The stomatal mothercell gives rise to guard cells, and epidermal cellsaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.98 GYMNOSPERMS The wall layers of the sporangium become disorganized and only the epidermis is conspicuous which forms the exothecium (Fig. 6.10B). The cells at the upper corners of the microsporangium are very much elon- gated, gradually becoming smaller at the region of dehiscence which comprises two rows of thick-walled cells. All the cells develop heavy thickenings on the radial and the inner tangential walls and remain thin on the outer tangential side. At the time of dehiscence, the sporangium breaks open because of difference in the turgidity of cells. The formation of male gamete: The antheridial cell divides into the stalk and the spermatogenous cells. The latter enlarges considerably in the longitudinal axis of the pollen tube. Two blepharoplasts arise as darkly- Fig. 6.12: Cycas circinalis (b1, blepharoplast; nu, nucellus; pe, prothallial cell; pt, pollen tube; sm, spermatozoid; sc, spermatogenous cell; st, stalk cell; tn, tube nucleus). A. Several pollen tubes growing in the nucellar tissue; pollen tubes do not carry sperms but arehaustorial. B. Grain end showing largespermatogenous cell,a small tube nucleus, a prothallial cell and a stalk cell; note two prominent blepharoplasts in the spermatogenous cell. C. Same, at a later stage with two large spermatozoids formed asa result of division of spermatogenous cell (after Maheshwari, 1960).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.102 GYMNOSPERMS movement (called amoeboid by Webber, 1901) by spermatozoids, or (ii) functioning as cytoskeleton and supporting the flagellated band along, with the vierergruppe. Norstog (1990) has made an ultrastructural study of the spermatozoids of Microcycas calocoma. The spermatozoids are almost identical with those of Zamia except that they are very small. An important feature is that the first-formed spermatozoids are more developed and more active than the last-formed ones which are less developed and less active thus showing a reductional gradient. Microcycas is considered as the most primitive member of the Zamiaceae. Megasporangium: The ovules are initiated as marginal protuberances formed by rapid, localized divisions in the surface layer cells of the sporophyll.In a young ovulea single hypodermalarchesporial cell divides periclinally to form a primary parietal cell and a primary sporogenous cell. The primary parietal cell, by repeated divisions, gives rise to a massive nucellus. The integument soon appears as a ring-like outgrowth from the base of the nucellus. It is fused with the nucellus for most part, but is free in the upper one third portion (Fig. 6.14D, E). It extends beyond the nucellus to form the micropylar tube. The deep-seated sporogenous cell functions as the megaspore mother cell (Fig. 6.14A) which divides meioti- cally to forma row of three cells representing an undivided upper dyad cell and two megaspores (Fig. 6.14B). Of the two megaspores, the lower one functions (Fig. 6.14C). Some cells of the nucellus, surrounding the develop- ing megaspore, are small, radially elongated and densely cytoplasmic. They divide rapidly keeping pace with the enlarging gametophyte and are probably nutritive in function. These cells constitute the spongy tissue. As the female gametophyte develops, the nucellar beak dissolves anda well defined pollen chamber is formed. The integument becomes differen- tiated into three zones, viz., the outer zone with mucilage canals and tanniniferous célls, the riddle zone with elongated cells and few scalari- form tracheids, and the inner zone of thin walled cells. The middle zone later develops into the stony layer of the mature testa (Fig. 6.14E). Female gametophyte: The nucleus of the functional megaspore divides many times forming nearly a thousand nuclei unaccompanied by wall formation. The megaspore, in the meantime, enlarges rapidly and devel- opsa central vacuole in its cytoplasm. The nuclei come to lie ina thin film of cytoplasm around the big central vacuole (Fig. 6.14D). Walls are laid down in the free nuclear gametophyte in a centripetal fashion, from periphery inwards, till each nucleus is separated by walls (Fig. 6.14E). The peripheral cells are small, isodiametric and without any food reserves, whereas the inner cells are large, irregular, and packed with starch.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.106 GYMNOSPERMS The pollen grains germinate immediately with the rupture of exine at the tube cell end. The tube nucleus passes into the pollen tube. A noteworthy feature of the pollen tube is that it does not grow towards the archegonium. Instead, it penetrates the nucellar tissue and becomes highly branched (Figs 6.12A; 6.15). Light microscopic and ultrastructural studies made by MH oo nie nee TA A La Fp Fig. 6.15: Dioon edule (arch, archegonium; dpt, discharged pollen tube; en, egg nucleus; fg, female gametophyte; ne, neck cell; nu, nucellus; mw, megaspore wall; pt, pollen tube; sm, spermatozoid). Reconstruction of the upper part of an ovule at fertilization stage. Of the five pollen tubes seen, two have already discharged theirsperms which are seen in the archegonium on the left and above it on the right in thick liquid. Of the remaining three pollen tubes, one shows a spermatogenous cell, the second with two sperms and the third shows the divided spermatogenous cell, prothallial and stalk cells. Two pollen tubes are seen penetrating the nucellus indicating their haustorial nature (after Chamberlain, 1935).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Cycadeoidales 119 distributed from Upper Jurassic to Upper Cretaceous, comprising about thirty species. The majority of the species are found as petrified trunks from America and a few from Europe, and India. The cycadeoidean members have ovoid or short columnar trunks which are unbranched or Fig. 7.4: Cycadeoidea. Reconstruction of the plant (after Delevoryas, 1971; redrawn from Stewart, 1983). apparently branched. The trunks are massive, not more than 1 m in length and about 60 cm in diameter. The surface of the stem is covered with prominent rhomboidal leaf bases and multicellular hairs in between them. The short stout trunks are much different from the members of Williamsoniaceae and Wielandiellaceae (Delevoryas, 1971). The anatomical details have been well worked out in Cycadeoidea. A cross-section of the trunk shows large parenchymatous pith (Fig. 7.5A) surrounded by a thin zone of primary wood with endarch protoxylem and secondary wood with scalariform tracheids (circular brodered pits haveaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Cycadeoidales 123 Relationships with other groups: Cycadeoidales, as a group, reflects similarities (and differences too) with many groups but the reproductive structures specially those having stalked ovules are unique. Some of them are discussed below: Cycadeoidales and Cycadales: Both the groups exhibit similar habit and morphologically similar fronds. Histologically the stems are similar hav- ing manoxylic wood. They have monocolpate pollen grains, orthotropous ovules and dicotyledonous embryo. Some of the major differences between the two groups are: different anatomical details of the frond; haplocheilic stomata in Cycadales and syndetocheilic in Cycadeoidales; girdling leaf traces which are promi- nently displayed inacycad stemare absentin the members of Cycadeoidales. Cones are monosporangiate in Cycadales, whereas in Cycadeoidales they are bisporangiate also. The microsporophylls are arranged in whorls in Cycadeoidales, but spirally in Cycadales. The arrangement of stalked ovules alongwith interseminal scales is a characteristic feature of the Cycadeoidales which is not found in Cycadales. Based on these differ- ences, Andrews (1961) opined thatboth the groups have evolved along two different and independent lines. Cycadeoidales and Pteridospermales: The two groups share a large num- ber of characters such as presence of ramental hairs, similar internal structure, syndetocheilic stomata, direct leaf traces, presence of leafy microsporophyll bearing synangia and the presence of cupule. The bisporangiate ‘flower’ of Cycadeoidales could be compared tobisporangiate fronds of Cycadofilicales (Pteridosperms). According to some authors thereare two lines in the course of evolution from Cycadofilicales. Of these one gave rise to monosporangiate forms like cycads, and the other to Cycadeoidales which has both mono- and bisporangiate forms. The major differencesare in the organization of ovules as they aresmaller and stalked in Cycadeoidales (much different in Medullosaceae) and exhibit promi- nent vascular supply to the nucellar tissue (absent in pteridosperms). Cycadeoidales and angiosperms: The similarity in ‘strobili’ or ‘flower’ of Cycadeoidales and the flowers of some angiospermous groups (such a Ranales and Magnoliales which have primitive flowers with numerous floral parts arranged spirally) lead a few morphologists (Bessey, Hallier, Hutchinson, etc.) to believe that the angiosperms might have originated from mesozoic Cycadeoidales. Some of the other similar characters are the spirally arranged hairy bracts around reproductive organs in Cycadeoidales which could becompared to the perianth of angiosperm; the mesoparacytic stomata of Magnoliales and the syndetocheilic stomata of Cycadeoidales; homoxylous (without vessels) wood comparable to few angiospermsaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Pentoxylales 127 The secondary wood is compact (pycnoxylic) like conifers, with well marked growth rings and xylem rays. The radial walls of the tracheids show uniseriate bordered pits. On the broader tracheids, the pits may be multiseriate, alternate and hexagonal like that of Araucariaceae and Cordaitales. The rays are uniseriate, up to seven cells high. Fig. 8: A.B,D, Pentoxylon salmii; C, Sahnia nipaniensis; E, Carnoconites (int, integument; nu, nucellus). A. Reconstruction of foliage-bearing branch showing long and short shoots. The leaves are borne on short shoots B. The stelar system in long shoot consisting of five distinct vascular segments. C.Shortshoot bearing male ‘flower’. Note rhomboidal leaf bases. D. Short shoot bearing ovulate cones which are mulberry-like. E. LS ovule. (A,B,D,E, after Sahni, 1948; C, after Vishnu-Mittre, 1953). Leaves: The leaves are represented by the organ genus Nipaniophyllum raoi (named after the village Nipania from where these were discovered). Theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.CHAPTER 9 GINKGOALES GINKGOACEAE The order Ginkgoales is a remarkably old group, represented by a single family Ginkgoaceae, which came into existence in the Permian. It was very well represented and had an almost world wide distribution in the Jurassic. The order consisted of sixteen genera and many species. From Cretaceous-onwards, the members of this order gradually started fading, so much so that now it is represented by a single monotypic genus, Ginkgo biloba. For many years Ginkgo was included in the family Taxaceae under the Coniferales till the discovery of motile sperms (Hirase, 1896a). Of the fossil ginkgophytes, Trichopitys is probably the oldest reported from Lower Permian. It has spirally arranged dichotomously branched leaves which have no lamina. Fertile ovule-bearing structures are borne in theaxil of some leaves. This appears similar to Ginkgo biloba. Another fossil representativeis Sphenobaierahaving wedge-shaped, slender dichotomous leaves. Leaves of Baiera are without a distinct petiole. They are bilobed, wedge-shaped and each lobe is repeatedly dichotomous. Ginkgoites, an- other leaf genus, hasa distinct petiole and semicircular lamina. Variations in leaf morphology are similar to those found in the modern Ginkgo. The genus Ginkgo is known in rocks as old as Triassic and the species G. biloba probably goes back as far. This may well be the oldest living seed plant. Its peculiar leaves have been identified with certainty in the Permian and with probability in the Carboniferous. It is, therefore, aptly called the “Living Fossil”. Professor Seward of Cambridge wrote in 1938 that hesaw G. biloba “asan emblem of changelessness, a heritage from worlds ofanage, too remote for our human intelligence to grasp, a tree which has in its keeping the secretsof an immeasurable past”. G. biloba is not as genetically stagnated as its apparent antiquity might lead one to suspect. It shows variation from specimen to specimen in frequency ofcertain leaf forms and of anamolous reproductive structures and in hardiness of the entire plant, of reproductive structures that overwinter within the buds, and of incom- pletely developed seeds after they drop in autumn. Such obvious points of variability are inconsistent with a species that has been in existence, relatively unchanged, for as long as G. biloba is reputed to have been. Itaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.140 GYMNOSPERMS resulting four strands fork repeatedly forming the dichotomous system of veins which are ‘open’. However, Arnott (1959), who made an extensive Fig. 9.7: Ginkgo biloba. A. Leaf showing dichotomous venation. Of the two bundles that enter the petioleone each supplies its respective half of the lamina. B, Section of a leaf cut at right angles to a vein. Note the large mucilage cavity and presence of stomata on the lower surface. Thesubsidiary cells have a finger like projection covering guard cells. The protoxylem elements in the vascular tissue show distinct lignification. C. Portion of leaf showing veins alternating, with mucilage ducts; note numerous crystals near veins. D. TS petioleshowing two distinct vascular bundles and several mucilage cavities and crystals in cortical tissue. (A,B, after Chamberlain, 1935; C,D, after Sprecher, 1907). study of Ginkgo leaves, observed vein anastomoses and in approximately 10% of the leaves there were one or more vein unions (see also Gifford & Foster, 1989). Interestingly, each petiole strand forms the venation of its respective half of the leaf (Fig. 9.7A). The bundles are usually all endarch, interspersed with some indistinct centripetal xylem in the form of transfu- sion tracheids. Surrounding the bundles there is a sheath of thick-walled cells. The cortex shows several large and conspicuous mucilage ducts (Fig. 9.7D).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.156 GYMNOSPERMS Embryogeny: The zygote nucleus divides in situ unaccompanied by wall formation (Fig. 9.14C). Subsequent divisions give rise to 256 nuclei before walls are laid down. The free nuclei become evenly distributed in the cytoplasm of the developing proembryo. After wall formation the newly formed proembryo fills the entire archegonial cavity (Fig. 9.14D). This situation is in contrast to cycads where, initially, the wall formation is restricted to the chalazal end only, the upper portion remaining free nuclear. The cells at the micropylar end of the proembryo elongate very rapidly to form a massive suspensor (Fig. 9.14E). Thoseat thechalazal end develop into embryonal cells with distinct meristem. The shoot apex and the two cotyledons differentiate at this end, and the root apex develops at the opposite end. The mature embryo may, rarely, show three cotyledons. There is a weak coleorrhiza. Two mesarch bundles are present in each cotyledon. Seed: Usually only one out of a pair of ovules develops to maturity (Fig. 9.16A); the other aborts at an early stage. The seed is bilaterally symmetrical and the longer diameter is marked by two prominent ridges on thetesta. In theripe seed, embryo develops several foliar primordia that along with the shoot apex constitute the first apical end of the seedling (Fig. 9.16B). The germination is hypogeal. In Ginkgo, a noteworthy feature is the development of the embryo which is acontinuous process from the time of fertilization until the germination of seed. In fact the major part of the growth of embryo takes place when the seed is detached from the tree and is lying on the ground (see also Dogra, 1992). The seeds are shed in autumn before or just after fertilization. This takes place seven months before the embryos attain the average size of 10.5 mm, and are ready to germinate. Ginkgo follows the two-year reproductive cycle (see Fig. 1.8B). Excised embryos are, however, able togerminate even when they are only 3 mm in length. This suggests that the delayed germunation of the seeds is not due to any immaturity of the embryos but, perhaps, the endosperm requires an after ripening process.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.164 GYMNOSPERMS which are fused at the base (Fig. 11.2 B). The primary axis of Cordaianthus is a flattened, bilaterally symmetrical axillary structure. It contains an endarchmedullated stele which gives off traces departing alternately from two areas situated at the opposite end of the stele to the four ranked bracts and secondary shoots. The single trace which enters the secondary shoot divides so that each branch enters one scale. The mesarch vascular strands supplying the scales divide dichotomously near their tip so that each microsporangium is vascularized (Fig. 11.2B). Fig. 11.2: (br, bract;fca, fertile scale; mic, micropyle; ms, microsporangium; nu, nucellus; pe, pollen chamber; pg, pollen grain; sca, sterile scale; sn, stony layer; vb, vascular bundle; w, wing). A. Reconstruction of Cordaianthus concinnus primary shoot bearing two dwarf shoots in the axilof thebract. Each dwarfshoot consists of spirally arranged sterile and fertile scales. The fertile scales terminate in ‘pollensacs’ or microsporangia. B. Fertile scale enlarged to show six ‘micros- porangia’. C. Primary shoot of Cardiocarpus cordei bearing ovulate secondary shoots with pendant ovules. D, E. LS and TS of Mitrospermum compressum. Micropyle and pollen chamber are distinctly seen in D. The seed coat shows inner stony layer (sclerotesta) and outer (sarcotesta) wing like extension (D). Spine like projections of sclerotesta extend into sarcotesta (E). The two lateral strands divide into several bundles in sarcotesta (D). F. Cordaianthus Grand d’ Euryi, part of nucellar beak with two pollen grains. The lower pollen grain exhibits part of the markings on exine and a part of internal cellular structure. (A, after Delevoryas, 1953; B, after Florin, 1944, redrawn from Andrews, 1961;C-E, after Taylor & Stewart, 1964, redrawn from Stewart, 1983; F, redrawn from Chamberlain, 1935).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.168 GYMNOSPERMS Ernestiodendron possess a long slender monopodial stem on which five to six branches arise in a whorl. The branches, in turn, have spirally arranged, acicular leaves (few millimeter long, needle-like or scale-like with a pointed tip). The leaves borne on the trunk and main branches, however, have a bifid tip. The leaves are amphistomatic with stomata being present in bands or isolated rows. The stomata are of haplocheilic type; having few papillate subsidiary cells. The leaves of Ernestiodendron are borne at right angles to the stem, whereas those of Lebachia are adpressed or slightly spreading (Fig. 12.1A, B). Moreover, in Ernestiodendron, the leaf scars on stem reveal that leaves abscissed from the branches. According to Florin (1951) the stem shows a large pith and eustelic arrangement of vascular bundles with endarch primary xylem. The secondary growth is abundant with tracheids showing one to three rows of closely arranged alternating bordered pits, uniseriate rays and absence of wood parenchyma. Rothwell (1982) noted resin ducts and leaf traces originating from eustele. Fructifications: The plants are monoecious. The male and female monosporangiate cones are borne singly at the tip of the ultimate branches. Female cones, in some species (e.g. Lebachia hypnoides), were borne in penultimate branches too. Whereas, the female cones were upright, the male cones were pendulous. The male cones are simple strobili with the main axis bearing spirally arranged flattened microsporophyils like the present day conifers. Each sporophyll had an upturned leaflike distal segment that partially covered the uppersporophyll. The proximalend, on the other hand, was narrow stalk-like that bore two sprorangia abaxially. The sporangia dehised longitudinally. The situation is quite similar to present day conifers. Florin (1951) interpreted the entire cone as simple as. compared to compound ovulate cone. His conclusion is based on the absence of bracts subtending a secondary shoot in male cones. The pollen grains had a balloon-like reticulately ornamented air sac (saccus) sur- rounding the entire gametophyte except at the distal end Ultrastructural studies of afew Voltzialean pollen, suchas Willsiostrobus, Sertostrobus and Darneya have been made. The pollen are bisaccate and germinate from the furrow on the distal surface. The sexine in Sertostrobus and Darneya is columellate with distal ends forming a tectum. In Willsiostrobus the sexine is thin on both proximal and distal surfaces adpressed to nexine. Whereas nexine is thick and lamellate in Sertostrobus, in Willsiostrobus it shows numerous lamellae. The pollen are protosaccate with sporopollenin threads of variable thickness. The corpus is thick (see Taylor & Taylor, 1987). The female cone of Lebachia (Fig. 12.2A) is a much complex structure. Each cone is ellipsoidal to cylindrical with the axis bearing spirallyaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.CHAPTER 13 CONIFERALES Coniferales are a large group comprising a sizeable number of plants distributed in temperate regions and high altitudes of tropics. The group, in general, is composed of woody perennial plants. Their growth habit varies from tall lofty trees as in Sequoia to miniature forms of Dacrydium which are only some centimeters high. In between are forms which are spreading trees and shrubs. One of the Juniperus species, J. horizontalis, is completely prostrate The living conifers show a wide range of geographical distribution Whereas some genera are widespread, many have a markedly restricted distribution and a few are endemic. This indicates great antiquity, a fact substantiated by fossil records also. Conifers were differentiated by the Jurassic period of the Mesozoic era. The families Pinaceae and Araucariaceae are slightly older than the Jurassic period. On the other hand, Cephalotaxaceae and Cupressaceae are slightly younger, having appeared in the Upper Jurassic or Lower Cretaceous. Conifers, however, reached their climax in mid-Mesozoic when extensive forests were present in Northern Europe. Comparative studies on the fossil groups and the extant plants made by Florin and others reveal that different families of the order Coniferales have evolved from a group of extinct plants which are included in the family Voltziaceae and are called ‘transition conifers’. Some of the general characters of the order are: (a) Conifers are woody perennials. Some exhibit two types of branches viz., (i) long shoots or shoots of unlimited growth, and (ii) dwarf shoots or shoots of limited growth. (b) Dimorphism of leaves is shown by the presence of scales and foliar leaves. The foliar leaves may be needle-like, linear or broad. () The tap root system of conifers is mostly associated with mycor- rhiza which may be ectotrophic or endotrophic. The endotrophic mycorrhizal association is found only in the members of the family Araucariaceae, : (d) The wood is pycnoxylic. Secondary xylem consists of tracheids, ray and xylem parenchyma. Fibres are absent. Xylem parenchyma is absent in Araucariaceae, scarcely present in Pinaceae and abun- dant in Podocarpaceae, Taxodiaceae and Cupressaceae.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.176 GYMNOSPERMS the axil of scale leaves (Fig. 13.1 A-C). Each dwarf shoot bears two opposite scaly leaves, called prophylls followed by 5-13, spirally arranged scaly cataphylls in 2/5 phyllotaxy. The leaves on the dwarf shoots are of two types: 2-5, long, needle-like foliage leaves and the scale leaves of protective nature. The needle number is constant for a species and is used as a taxonomic character, e.g. P. monophylla has one, P. sylvestris has two and P. roxburghii has three needles (Fig. 13.1C), and P. wallichiana has five needles (Fig. 13.1D). The primary (deciduous) leaves, which are always single, appear soon after germination and function as foliage leaves. The secondary (perma- nent) leaves, or the needles, are borne on short shoots or brachyblasts in fascicles in the axil of primary leaves. When a pine plant is 2-3 years old, all the primary needles fall off leaving scarious bracts on the shoot. These bracts on the lower part of the shoot are without any axillary bud, whereas those on the upper part of the shoot sport an axillary bud that later gives rise to secondary needles. . The tree is monoecious, but the male and female cones are borne on separate branches. The male cones (see Figs 13.2 A-D; 13.3A) arise on the lower and the female cones on the upper branches. The male cones, which replace the dwarf shoots, occur in clusters. The number of male cones per cluster varies considerably from 15 (P. wallichiana) to about 140(P. roxburghii). Each male cone consists ofa central axis on which the microsporophylls are spirally arranged (Fig. 13.4). Each microsporophyll bears two sporangia on the lower or the abaxial surface (Figs 13.2E; 13.3B,C). The cone axis elongates exposing microsporangia which dehisce longitudinally. The female cones (Figs 13.2F-H; 13.3D-F) replace thelong shoots and are generally borne in pairs, but thenumber may go up tosix. Each femalecone consists of a central axis on which 80-90 megasporophylls (ovuliferous scales or ovule - bearing scales), axillary to scale leaves or bract scale (Fig. 13.5), are arranged in aspiral fashion. The bract and the ovuliferous scales together forma seed-scale-complex (Fig. 13.6).Each megasporophyll bears two ovules on the upper or the adaxial surface (Fig. 13.3 G). The ovules are inverted with the micropyle facing the axis of the cone. The ovuliferous scale is initially much smaller than the bract scale, but in post pollination stages it overgrows the latter. The exposed part of the ovuliferous scale (also called cone scale in a mature cone) is known as ‘apophysis’. The tip of the apophysis becomes the ‘umbo’ in the mature cone. The umbo of all diploxylon pines is dorsal, whereas it is terminal or dorsal in haploxylon pines. Sporophylls at the apex and the base of a cone are generally sterile. Seed crops are known in Pinus. These are not annual but occur at intervalsof twoorevenmore years. As the tree grows old, the seed-bearing capacity declines but doesnot stop entirely. Pineskeep on producing seeds till their death. There isa great variation in the longevity of pines; P. radiata lives up to 140 years and P. lambertiana up to 600 years. A tree of P. aristata,aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.180 GYMNOSPERMS Fig, 13.5: Pinus roxburghii (bs, bract scale;ca, cone axis; ov, ovule, ovs,ovuliferous scale; rd, resin duct; vt, vascular trace). Longisection of a young female cone showing seed-scale complexes. At this stage, the bract scale is longer than the ovuliferous scale. The latter bears ovules on its upper or adaxial surface (after Maheshwari & Konar, 1971). Fig. 13.6: Pinus banksiana (bs, bract scale; ov, ovule; ovs, ovuliferous scale; pg, pollen grain; rd, resin duct). Longisection of bract, ovuliferous scales and ovule to show their relative positions. The ovule is in post-pollination stage (after ‘Chamberlain, 1935).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.184 GYMNOSPERMS rod-shaped horizontal thickenings in the middle lamella between the radial walls of secondary wood tracheids. Recent electron microscopic studies have shown them to be mere refraction patterns (see Mauseth, 1988). The secondary phloem consists of sieve elements which, according to Srivastava (1963), arise directly by transformation of the phloem initials. Rarely, the phloem initial divides once to give rise to two sieve cells. It is interesting to note that the sieve cells cutoff during coolerand moistseason show wider lumen and thinner walls. The sieve cells (or tubes) are arranged in rows. The most characteristic feature of the sieve cell is the occurrence of sieve plate (or the sieve area) on the radial walls throughout the length of the sieve cell. Sometimes two or more sieve areas may be grouped together. The sieve plates consist ofnumerous narrow channels (0.1-0.5 pm in diam.) lined with callose. In a good preparation, these sieve areas look like nuclei and the entire sieve cell as a multinucleate cell. The vascular rays are initiated by the cambium and, once formed, they continue to increase in radial length indefinitely. The cells are always elongated in the direction of the long axis of the ray. The rays are of two types: (i) uniseriate rays, and (ii) multiseriate or fusiform rays (Pl. 13.6.A,B). Most of the rays are of the uniseriate type and are seen in a transverse section as uniseriate files of cells which are radially elongated. Theray may be likened to a brick wall in which each brick corresponds to a single cell. In height, the ray varies from 1 to 12 cells. The multiseriate ray is always more than one cell wide and several cells in height. It is associated with a centrally situated resin canal (Pl. 13.5B). The detailed structure of a ray can be best understood by examining sections cut in all the three planes. The transeverse section (TS) shows the width and the length of the ray. The longitudinal sections should be cut in the radial (RLS) as well as in the tangential (TLS) planes. If one began cutting tangential sections and continued to cut inwards, sections passing through the pith would be radial (see Fig. 13.8A). It should be understood clearly that aradial section cuts along the radius and also along the vascular rays while the tangential section cuts at right angles to vascular ray. The TLS shows the height and width of the ray (Pls 13.4; 13.6A, B). The RLS depicts the length and height of the ray (Pls 13.4; 13.6C). A radial longitudinal section (RLS) shows that the ray is composed of two kinds of cells (Pl. 13.6C). The starch filled central cells are living. The peripheral cells are, however, dead and are called the marginal ray tracheids or tracheidal cells. These have no protoplasmic contents and are further distinguished from the central cells or ray parenchyma in having bordered instead of simple pits. In the phloem region, the ray consists of the usual central cells, which are living and store starch, and the marginal cells which are also living but store only albuminous matter instead of starch.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Plate 13.9: Pinus banksiana (sa, saccus). A. Early stage of saccus (wing) develop- ment. The region under saci is PAS positive (arrows). B. Microspore within the tetrad showing nexine (arrows) separating the microspore cytoplasm from the saccus (after Dickinson & Bell, 1970).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.188 GYMNOSPERMS cells are devoid of Casparian strips. The pericycle is parenchymatous interspersed with transfusion tracheids. The transfusion tissue is com- posed of parenchymatous cells, albuminous cells and tracheidal cells with bordered pits on their tangential and transverse walls. A sclerenchyma- tous sheath is present around the vascular bundle. There may be one or two vascular bundles. When there is only one vascular bundle (as in P. wallichiana) it is medianly placed. In those species where there are two vascular bundles (as in P. roxbirrghii) they are placed at an angle to each other (Figs 13.9B; 13.10). The vascular bundles consist of protoxylem elements (scalariform pitted tracheids) and metaxylem elements (helically thickened tracheids with bordered pits). The tracheidal elements are in radial rows alternating with rows of parenchymatous cells and albumin- ous cells. The protoxylem gets crushed in mature leaves. Phloem consists of sieve cells and parenchyma. A cambium is present in the vascular bundle. It cuts off Secondary phloem and little ornosecondary xylem. One or two layers of secondary phloem are formed every year in P. aristata. The stomata are sunken. The walls of the subsidiary cells and the guard cells are partly lignified. The stomata may be located on all surfaces of the needle as in diploxylon pines or they may be absent on the outer surface of the needle as in some of the haploxylon pines. The stomata are situated in longitudinal rows. When the needles are examined with a hand lens, the cavities leading’to the stomata appear as white dots arranged in longitu- dinal series from the base to theapex. Edges of pine needles may be smooth or serrated. Microsporangium: One or more hypodermal cells differentiate into arches- porial cells which divide repeatedly to form an archesporial tissue (Fig. 13.11A). The peripheral cells of the archesporium divide periclinally to give rise to a primary parietal layer and primary sporogenous cells. The former divides both periclinally and anticlinally forming a wall of 3 or 4 layers, the inner most of which differentiates into tapetum. The remaining 2or3layersare the middle layers, which degenerate with the maturation of the sporangium. There is no trace of middle layers at the time of shedding of pollen. The primary sporogenous cells, on the other hand, divide in all planes producing a mass of sporogenous tissue (Fig. 13.11B), the last cell generation of which is known as microspore mother cell stage. The epidermal cells, in later stages, develop fibrous thickenings except for a few cells inarow, which mark the suture of dehiscence. The outer surface of the epidermis acquires a coating of cuticle (Fig. 13.11C-E). The newly formed tapetal cells cannot be differentiated from the other wall layers. During maturation they acquire dense cytoplasm and may become bi- or multinucleate. Broad cytoplasmic channels connect tapetal cells to each other (Dickinson, 1970). The cell walls of the tapetal and sporogenous tissue are largely pectinaceous as compared to other layersaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.192 GYMNOSPERMS future region of the wall and gradually fuse with each other. Later, a very thin line of fine fibrillar material, which appears in the centre of these fusing, vesicles, is replaced by highly electron transparent callose. During inter- phase, initially, an aggregation of amyloplasts is observed at the equatorial plate (Pl. 13.7). In addition to Golgi activity, the microspore mother cell cytoplasm also shows packets of rough endoplasmic reticulum (RER) during zygotene-pachytene stage. The RER changes to smooth endoplas- mic reticulum (SER) during pachytene but reappears when tetrads mature. A quantitative change in the polysomal population hasalso been observed. Lipid complexes appear till zygotene and after telophase II. It is felt that Fig. 13.12: Pinus wallichiana (ae, antheridial cell; ai, antheridial initial; cc, central cell; pe, prothallial cell, tc, tube cell; w, wing). A. Microspore mother cell. B,C. Meiosis I and II, respectively. D. Microspore tetrad. E. Uninucleate pollen grain. F,G. Stages inthedevelopmentof male gametophyte. In G, both prothallial cells have been cut off and the antheridial initial is undergoing division. H. Mature, four-celled (two degenerating prothallial cells, an antheridial cell and a tube cell) pollen grain at shedding stage (after Konar & Ramchandani, 1958). whereas polysomes are related to callose and pollen wall formation, lipids play a role in the formation of sporopollenin. The mitochondria and plastids remain constant during meiosis. However, some increase in the population of vesicles has been observed at late tetrad stage. It is at this stage vacuoles develop in microspores. At the end of the division, the callose sheath is dissolved by enzymatic action and the young haploid microspores are set free. Before the nuclear division, the microspore undergoes a maturation phase during which it builds DNA, RNA and polysaccharides (Moitra & Bhatnagar, 1982b).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.196 GYMNOSPERMS autofluorescent in P. roxburghii showing thereby the presence of sporopollenin (Moitra, 1980). Megasporangium and megasporogenesis: The ovulesof Pinus are unitegmic and crassinucellate. The integument is free from the nucellus for most part of its length, except at the chalazal end. It forms a fairly broad micropylar tube, whichis curved-in during pre-pollination stages, but is curved-outat the time of pollination. A hypodermal archesporial cell (rarely more) is formed at the micropy- lar end of the broad nucellus. It divides periclinally to form a primary parietal cell (towards micropylar end) and a primary sporogenous cell ({towardschalazalend). The former divides both anticlinally and periclinally so that the primary sporogenous cell is pushed deep in the nucellus. The sporogenous cell functions as the megaspore mother cell (Fig. 13.13A). It forms a linear tetrad of megaspores after meiosis (Fig. 13.13B, C). A polar migration of starch grains towards the chalazal end takes place in the mother cell before meiosis so that after meiosis it is the lowermost cell whichreceives most of thestarch and actsas the functional megaspore. The upper three megaspores degenerate. Callose is formed around the cyto- plasm of the megaspore mother cell preceding meiosis. As in the case of pollen formation, callose separates the individual megaspores of a tetrad but disappears during the maturation of the functional megaspore. The nucellar cells in the immediate vicinity of the developing mega- spore are rich in their cytoplasmic content and are nutritive in function. These cells constitute the spongy tissue. The cells of spongy tissue (also known as tapetal cells) often become compressed between the female gametophyte and the outer tissues of the ovule. Female gametophyte: The functional megaspore enlarges considerably; its nucleus divides mitotically to give rise to the free nuclear gametophyte (Fig. 13.13D-F). During early stages, the nuclear divisions are synchronous. With the enlargement of the megaspore, a vacuole develops in the centre gradually pushing the cytoplasm alongwith the nuclei towards the periphery. Thecytoplasm at this stage hasabundant mitochondria, plastids and dictyosomes. The number of nuclei ina free nuclear gametophyte is constant fora particularspecies; in P. roxburghiiand P. wallichiana itis about 2500 (Fig. 13.13E, F) and approx. 2000 in P. gerardiana. At the end of last mitosis, every nucleus gets connected to six neighbouring nuclei by the formaton of secondary spindles. Wail formation takes place through alveoli. The newly synthesized walls are unevenly thick and grow centripetally as open tubesor pipes from megaspore cell wall till they reach the centre of the vacuole. Each alveolus has a nucleus at its mouth and directs its growth. Cytoplasmic organelles surround the nucleus. Now,aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.200 GYMNOSPERMS of the grain contacts the drop. Another advantage of such an orientaion is that after being sucked inside, the germinal pore faces the nucellus, thus avoiding the twisting or curving of the pollen tube after germination and entry into the nucellus. This claim has, however, been contested by McWilliam. According to him, the pollen grains move because of active absorption of the drop, and there is no definite orientation of pollen on the nucellus. Moreover, winged pollen occur in plants such as Abies, Cedrus and Tsuga where the pollination drop is absent. The pollen tube arises from the inner pectic-cellulosic layer of intine or internal intine (Martens & Waterkeyn, 1962). While moving through the nucellus the pollen tube secretes enzymes stich as pectinase and cellulase dissolving the nucellar cell walls (Willemse & Linskens, 1969). As the tube emerges, the tube nucleus is the first to move into it followed by spermatogenous and stalk cells (Fig. 13.15A). The spermatogenous cell eventually divides into two unequal male nuclei (Fig. 13.15 B, C). Fig, 13.15: Pinus gerardiana (arch, archegonium;ma, malenucleus; nu, nucellus; pt, pollen tube; sc, spermatogenous cell; st, stalk cell, tn, tube nucleus; w, wing). A. LS nucellar tip with three germinating pollen grains. In one of the pollen grains, stalk and spermatogenous cells are separating prior totheir entry into the pollen tube. B. Same, part of ovule showing a fertilized (left) and an unfertilized (right) archegonium. A portion of the pollen tube is seen in the nucellar tissue. C. Pollen tube from B enlarged to show the two unequal male gametes, thestalk cell and the tube nucleus (after Konar, 1962). Fertilization: Fertilization is the union of the male and the female gametes that results in the formation of zygote. The pollen tube enters theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.204 GYMNOSPERMS i Hi Hi rea I Py Fig. 13.18: A, Pinus sp; B-I, P. roxburghii (cot, cotyledon; dS, disfunctional suspensor; e, embryonal mass; Es, and Es,, first and second crop of embryonal suspensor; hy, hypocotyl; U, upper tier). Progressive stages in the development of embryo. (Contd.)aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.208 GYMNOSPERMS Cytology: The diploid number of chromsomes in Pinus is 24; all the chromosomes have a median constriction (see Mehra & Khoshoo, 1956a; Natarajan et al., 1961). Of the 12 pairs of chromsomes, 2 pairs are long with satellites and 2 pairs are short without satellites. The remaining 8 pairs are medium-sized and of these 3 pairs bear satellites. Of the 5 pairs of chromosomes with satellites, four pairs show seconday constriction, whereas the fifth pair shows distinct, separated satellite heads. Moreover, in this fifth pair, one chromosome has a longer short-arm and shorter satellite head, and the other has a shorter short-arm and longer satellite head. All the satellited chromosomes are capable of organizing nucleoli. The number and position of centromeres vary in different species. Most species of Pinus show two-year type of reproductive cycle, except three species (P. pinea, P. leiophylla and P. torreyana) which show three-year type of reproductive cycle (see Fig. 1.9). CEDRUS Cedrus also belongs to the family Pinaceae. The wood of the Indian species, C. deodara (the ‘deodar’), produces an invaluable timber. Deodar wood is the strongest coniferous wood and is weight for weight as strong as teak. Because of its great economical importance, C. deodara is described in this chapter to represent the genus Cedrus. The name Cedrus is derived from the Greek word Kedros, meaning conifer. It is a small genus, being represented by four species: C. atlantica (Loud.) Manetti is the Atlas or Algerian cedar found on the Atlas Moun- tains in Algeria and Morocco; C. brevifolia (Hook.f.) Henry is native to Cyprus and is called the Cyprus cedar; C. libani Loud. is the Lebanon cedar indigenous to Lebanonrange. ang. Asia minor; and C. deodara (Roxb.) Loud. is the Himalayan cedar or deodar'spread over Western Himalayas (from Afghanistan to Garhwal), Mediterranean and Morocco. The specific name deodara indicates the tree's connection with objects of worship and is derived from Sanskrit words deva meaning deity and dara meaning wood. Cedrus deodara is distributed throughout the Western Himalayas from Afghanistan toGarhwal. Itis predominant between 1800-2600 m elevation above sea level, though the tree extends from 1200-3000 m. Low tempera- ture restricts its distribution beyond this elevation. The deodar belt, as the deodar dominated zone is called, varies in different localities. This belt is usually higher in the Southern thanon the Northern slopes. Best specimens are found in places where rainfall varies from 100-175 cm. Though the plants occur in regions with lower rainfall (75cm or even less provided the soil is deep and good), they fail to grow in areas with higher rainfall. This explains the absence of deodar forests from the eastern range. The best growth is attained on deep, fairly porous, well drained fertile soil and inaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.212 GYMNOSPERMS are thick-walled, the rest being uniformly thin. Few tannin-filled cells are present in the cortex. The root may be tetrarch or pentarch. The protoxylem is exarch and alternates with the phloem. In a young root, small cavities could be seen in the cortex just outside each phloem group. Their origin is not very clear. The endodermis and pericycle are not well demarcated. Resin ducts are absent. The pith is large and parenchymatous. In an old root, the secondary growth takes place normally. With the formation of secondary tissue, primary phloem and pith are crushed. The cork cambium cuts off cork cells towards outside resulting in the disinte- gration of major portion of cortex and the whole of epidermis. Resin ducts, which were absent in young roots, appear in older ones. The mycorrhizal association is purely ectotrophic in Cedrus. The fungal mantle over the infested roots consists of 8-12 cell layers and is up to25 yum thick. The hyphae grow in the intercellular spacesin the outer 2 or 3 layers of cortical cells. The mycorrhizal roots are coralloid in nature. They are very small and bear unbranched roots laterally. Stem: A study of TS of young stem at seedling stage shows a wavy outline with ridges and furrows bound by a thick-walled epidermis. Some of the epidermal cells elongate to form unicellular hairs. The cortex is made up of 6-8 layers of cells. The cells are thick-walled below the ridge and thin- walled below the furrow. The cortical cells are green (at the seedling stage) filled with starch grains and interspersed with resin ducts. The xylem is endarch. The pith is parenchymatous and contains starch. Secondary growth takes place very early in the development. The long shoot is covered by a layer of cork made up of parenchymatous cells filled with some tanniniferous material. Itis followed by thin-walled cortical cells, some containing starch and others containing tannin. The cortical cells are interrupted by large resin ducts which are arranged in a ring. These ducts are bound by epithelial cells and are oval in outline. The cortex also exhibits the presence of abundant sclereids (both single and in groups) and occasional leaf traces. The xylem is pycnoxlic. The pithis small and parenchymatous. The dwarf shoot in a TS shows a circular outline with persistent leaf bases. A cork cambium differentiates in the cortex which is parenchyma- tous with a few tanniniferous cells. Resin ducts are morenumerous than in the long shoot and are lined by a double layer of epithelial cells. A number of sclereids are present in the cortex and pith. Leaf traces are seen in the cortex. Vascular cylinder is in the central portion of the shoot. The tracheids show uniseriate, sometimes biseriate, bordered pits on their radial walls. Each bordered pit is circular in outline and the margin of the torus is ornamented with rounded projections. The xylem rays are usually 20 cells or more high. Ina TLS both uni- and multiseriate rays areaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.222 GYMNOSPERMS because of the colour of bark and heartwood, reaches an average height of 100m, the highest being 112 m, anda diameter of 5 m. The tree is often clear of branches for nearly one-third to half its length. A second species, S. gigantea, renamed as Sequoiadendron giganteum, or the ‘Big Tree’, is some- what shorter (rarely reaching 100 m) but more massive (up to 11 m in diameter). Big Trees happens to bea small station 3 to 4 hoursjourney from San Francisco. From China, a new genus, Metasequoia or the ‘Dawn Red- wood’ was discovered. This was known only in the fossil state earlier. In 1944, T. Wang (see Chaney, 1948), a Chinese forester, made the discovery of three living trees in a remote valley of central China. The discovery was amazing because the foliage and the cones of this tree resembled the fossil specimens of Metasequoia. The tree that became extinct 20 million years before was suddenly found ina living state. One could indeed say thathere was a fossil come to life. This can well be considered as one of the most remarkable and dramatic discoveries. According to Stebbins (1948), Metasequoia seems to be closely allied to Sequoia. It has only one species, M. glyptostroboides. Cryptomeria or the Japanese Cedar is a native of China and Japan but has got naturalised in Eastern and Western Himalayas. Darjeeling has exten- sive forests of Cryptomeria. In the Western Himalayas, it grows at an altitude of 1800-2400 m. The genus is represented by only one species, C. japonica. The name is derived from Greek Kryptos (=hidden) and Meroe (=part), meaning obscure The genus Taxodium, after which the family is named, is commonly knownas ‘cypress’. It derivesits name from Taxus and the Greek word vides (=like) meaning that is it is Taxus-like (the resemblance is in the leaf arrangement). The genus is a native of north America and Mexico and is peculiar in having deciduous leaves. There are three species, namely T. ascendens (the pond cypress), T. distichum (the bald cypress), and T. mucronatum (the Montezuma yew cypress). According to Sargent (1933, 1947) there are only two species, the pond cypress being a variety of the bald cypress. T. ascendens is common in wet grounds from Florida to Mexico. One specimen of T. mucronatum, that stands in a church yard in Tule, about 400 km south of the old city of Mexico, is a giant among its Kind. Itiscalled the ‘Big tree of Tule’ (PI. 13.12). Itsurpasses even the largest sequoias of America. The trunk is 16 m in diameter and the tree has been estimated by some botanists to be 5,000 years old, probably the oldest living thing in the world. The tree is most remarkable and does not seem to possess a single dead twig, It is massive but not tall, being about 50 m high. Unlike the swamp cypress, which is deciduous, it is evergreen (see Chamberlain, 1921, 1935). Taxodium is described as the Type genus of the family. The following description is mainly based on T. mucronatumt (see Vasil & Sahni, 1964) which is growing at the Forest Research Institute, Dehra Dun. It is notaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Plate 13.11: Cedrus deodara. A. TS wood showing annual rings and resin ducts. B, Same, magnified.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Plate 13.19: A and C, Agathis robusta; B and D, Araucaria cunninghamii. A, B. Twigs bearing male cones. C, D. Twigs bearing female cones (after Kaur, 1979).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.226 GYMNOSPERMS cone axis (Fig. 13.27E). The seeds are dark brown, somewhat triangular or rhombodial with a rudimentary wing. Fig. 13.27: Cryptomeria japonica (br, bract; fc, female cone; mco, male cone; ms, microsporangium; msp, microsporophylll; ov, ovule; ps, process). A Branchlet bearing male and female cones. B. A single male cone alongwith the bract enlarged. C. Abaxial view of microsporophyll bearing sporangia. D. Female cone ready for pollination. E. Bract scale and seed scale complex; thebract at the tip is seen as a recurved process. Ovules are borne adaxially with micropyles pointing away from cone axis. F. Abnormal female cone where its axis has grown into a vegetative shoot bearing a cluster of male cones (after Singh & Chatterjee, 1963) Stem: The stem of Taxodium is circular in outline. A young stem shows an epidermis on the outside. The cortex is four or five-layered consisting of isodiametric parenchymatous cells with intercellular spaces. The old stem shows a well developed periderm which is dark brown in colour and is formed quite early. Its cells are rectangular and are arranged in rows without intercellular spaces. Groups of stone cells are seen in the cortex. In a young stem, the xylem elements are represented by narrow, elongated tracheids with annular or spiral thickenings. Annual rings are not distinct in young stems but become prominent in the mature wood (Pl. 13.13A, B). The late wood tracheids have thick walls with a narrowaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.238 GYMNOSPERMS Embryogeny: The zygote nucleus divides to form two daughter nuclei which divide twice to give rise to eight free nuclei lying embedded in the dense neocytoplasm. The free nuclei organize into two groups: the nuclei of the upper group are arranged in one plane, and those of the lower group show variable arrangement. Also, the number of nuclei in the two groups is not always equal. Wall formation takes place to form the primary upper tier (pU) and the primary embryonal cells (pE) (Fig. 13.37A). The cells of Fig. 13.37: Taxodium mucronatum (e, embryonal mass; E, embryonal tier; et, embryonal tubes; pE, primary embryonal tier; pU, primary upper tier; S, suspensor tier; U, upper tier). A. Primary proembryo showing primary upper (pU) and primary embryonal (pE) tiers. B. Same, showing internal division in pU; pEis still undivided. C. Secondary proembryo with U:S:E arrangement; a non-functional nucleusis visible in the upper part of archegonium. D. Embryo- nal clump showing three dividing embryonal units. E. Developing embryonal tubes (et) from embryonal mass (A, B, D, after Dogra, 1966b; C,E, after Vasil & Sahni, 1964). the primary upper tier do not have walls on the upper side and are thus in continuity with the egg cytoplasm. This is the primary proembryo. At thisaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.278 GYMNOSPERMS traversing the nucellus, the pollen tube reaches the free nuclear female gametophyte; sometimes several pollen tubes establish contact with the female gametophyte. The pollen tube expands and sends out many branches which grow on all sides of the female gametophyte (Fig. 13.63B, C). Fig.13.63: Podocarpus gracilior (fg, female gametophyte; mn. male nuclei: pg. pollen grain; pt, pollen tube; sc, spermatogenous cell;stn, sterile nucleus). A. LS upper part of ovuleshowing one of the pollen grains germinating atnucellar tip. The pollen on the right belongs to Pinus. B. Cellular gametophyte showing few growing pollen tubes overt. C. Dissected whole mount of pollen tube showing its highly branched nature; note the spermatogenous cell and sterile nuclei. D. Portion of pollen tube to show two unequal male gametes. The non-functional male gamete is being extruded out (after Konar & Oberoi, 1969 a). The spermatogenous cell enlarges, shows dense mass of cytoplasm and a large nucleus which is eccentrically placed. It is usually surrounded by 2 or 3 sterile nuclei. When the female gametophyte starts becoming cellular, the spermatogenous cell nucleus divides. Initially, both the nuclei are of the same size but, subsequently, the inner nucleus enlarges and moves towards the centre of thecell. The smaller nucleus, whichis towards the outside, is eventually extruded from the cell (Fig. 13.63D). Thus, there is only one functional male nucleus.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Coniferales 287 vascular bundle. The mesophyll consists of only parenchymatous cells with intercellular spaces in type a. Only in the two longer arms of leaf, a geegete 2) Leis sys deeh. LS "Fig. 13.67: VS part of leaf of Agathis robusta showing well developed hypoder- mis, mesophyll and fibres (after Chamberlain, 1935). palisade layer is visible. On the other hand, in type b, a single layer of well developed palisade and many layered spongy tissue are present. There are abundant branched sclereids. A single median (type a) or several parallel vascular bundles (type b) are present with completeaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.296 GYMNOSPERMS a column and the peripheral zone. The distal portion gives rise to cotyle- dons and the epicotyl. The pith, the vascular bundles, and the cortex differentiate in the central portion of the hypocotyl. Both the genera show two cotyledons (Fig. 13.72) except that in Araucaria the cotyledons are deeply cleft and appear as four. Seed coat: The seed coat consists of two zones. The outer zone is composed of 4 or 5 layers of thick-walled cells; the cells of the outermost layer or epidermis are also thickened and crescent-shaped resulting in undulation of the surface. The inner zone is of 5 or 6 layers of loosely arranged cells which remain thin-walled up to very advanced stage of embryo develop- ment. The cells become thick-walled at a much laterstage. Ina mature seed, the inner 2 or 3 layers of inner zone get disorganized (Fig. 13.73A-E). The Fig.13.73: Agathis robusta. A-E. Stages in the development of seed coat (after Kaur, 1979) wing in this family seems to arise from the entire bract scale. The germination of the seed is epigeal in both the plants (Fig. 13.74 A-F).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.300 GYMNOSPERMS secondary growth are similar to those of stem. The cork develops quite early. Singh (1961) found the secondary xylem to be eccentric in the roots examined by him. Stem: A young stem has a fluted outline because of the long leaf bases. The epidermis bears a thick cuticle. There is a narrow cortex of thin-walled cells. An interesting feature is the presence of a resin duct in the centre in young stages (Pl. 13.22A). The secondary growth is normal. In the second- ary phloem sieve cells show circular sieve areas along their radial walls. The phloem fibres occur in single-layered rings. The secondary xylem shows distinct growth rings (P1. 13.22B). Tracheids are long with uniseriate bordered pits both on radial and tangential walls (PI. 13.22C). Like taxads, the tracheids are characterised by the presence of tertiary spiral thicken- ings. There is abundant xylem parenchyma which is diffuse and resinous. Xylem rays are uniseriate and 1-8 cells high (P1. 13.22C, D). Leaf: The leaf is dorsiventral and its margins are slightly curved towards inside. In the midrib region there is a keel-like structure on the upper or adaxial surface. The upper epidermis is covered witha thick cuticle. There is no hypodermis. The mesophyll is differentiated into thick-walled pali- sade and thin-walled spongy parenchyma. The endodermis is ill defined. Rarely sclereids may be present in the mesophyll. Stomata occur in numerous lines (stomatal lines) as two broad silvery bands, one on either side of the midrib, on the lower surface. A single unbrached bundle traverses the leaf. A resin duct is present below the phloem. The transfu- sion tissue is not well developed and consists of reticulately thickened tracheids confined to the lateral sides of the bundle (Fig.13.76). Fig.13.76: Cephalotaxus (rd, resin duct; tt, transfusion tissue; vb, vascular bundle). VS leaf (diagrammatic). The margins are curved inside and a single resin ductis present below the vascular bundle. Transfusion tissue isseen on the lateral sides of the bundle. Microsporangium and microsporogenesis: A plate of hypodermal arches- porial cells differentiates on the abaxial surface near the base of the microsporophyll (Fig-13.77A). Ineach microsporangium archesporial cellsaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.GYMNOSPERMS os'er Styaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.312 GYMNOSPERMS reproductive structures. The male cones occur singly as stalked globose heads in the axil of the foliage leaves (Fig. 13.82A) on the undersides of branchlets of the preceding year and are more numerous in the distal region of certain branchlets. The cone axis bears at its base a few decussate, sessile, decurrent, imbricate scales. Each male cone consists of 6-14 radially symmetrical, perisporangiate microsporophylls which are arranged spi- rally (Fig. 13.82A). The sporophylishave the same peltate appearance as in Equisetum. Each sporophyll bears 6-8 microsporangia which are pendent and elongated. They, however, do not hang freely from the underside of the disc, but are fused with the stalk on their inner side (Fig. 13.82B). Unlike other conifers, the female reproductive structures in Taxus are not organized into a typical cone. Ona female tree, an axillary shoot bud develops on a branch which is protected by a number of scales. The ovule is initiated by the transformation of the apical meristem. The apical initials of the shoot apex give rise to the nucellus, the flank meristem to the integument, and the chalazal portion of the ovuleis contributed by the pith meristem, flank meristem and the subapical initials. The ovule is, thus, strictly terminal and cauline in origin and unrelated to any of the scales which cover it. These scales probably represent sterile megasporophylls which have lost their fertile nature in the course of evolution and the cone axis itself gives rise to the fertile ovule. The ovule is unitegmic, crassinucellate, orthotropous and sessile. The single integument is free from the nucellus exceptat the base. Ina young ovulea ringlike outgrowth, the aril, develops at the base of the integument which, later, covers the entire ovule. At first it is a green saucer-shaped structure but eventually it takes the form of a large red fleshy cup enclosing the hard bony seed (Fig. 13.83). Root: TS of the root shows an epidermis which, in young stages, exhibits long unicellular hairs. The epidermis is followed by a large parenchyma- tous cortex with intercellular spaces. Beneath the cortexis a single-layered endodermisand amultilayered pericycle. Resin canals are absent incortex, but resin parenchyma is present. The root of Taxus is usually diarch. The metaxylem elements have uniseriate bordered pits in addition to tertiary spirals. Phloem bundlesalternate withxylem. Secondary growthis normal and resembles that of Pinus. Stem: The young stem is covered by long ridge-like leaf cushions which become thinner as they go downwards (Fig. 13.84A). They scale off in old branches (Fig. 13.84B). The epidermis is parenchymatousand cuticularized, and is followed by a broad parenchymatous cortex. A few cells of the epidermis are tannin filled. Resin canals or resin parenchyma are absentaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Coniferates 319 (non-functional megaspores) are without them (Pennell & Bell, 1987). In some nucelli all the four megaspores may grow resulting in the formation of more than one female gametophy te. The spongy tissue is poorly devel- oped in Taxus. Female gametophyte: The nucleus of the functional megaspore divides mitotically (Fig. 13.87B) unaccompanied by wall formation, thereby result- ing ina free nuclear gametophyte. Approximately eight successive divi- sions produce 256 nuclei which arrange themselves in a thin peripheral layer of cytoplasm around a central vacuole (Figs 13.87F; 13.88A). The walls are laid down by alveoli formation to give rise to the cellular gametophyte (Fig. 13.88A). In the developing megasporangium, the free nuclear gametophyte is pear-shaped, the apical beak representing the original position of the megaspore. In the relatively mature ovule, a tent pole is present. According to Dupler (1917; see also Chamberlain , 1935; Maheshwari & Singh, 1967) in T. canadensis the cellularization of free nuclear gametophyte was presumably effected through alveoli (see Fig. 13.88A). However, Pennell & Bell (1987) found no evidence of secondary spindles in the free nuclear gametophyte of T. baccata. The cytoplasm and nuclei became enclosed directly by growing walls. This was accomplished by the cister- nae of endoplasmic reticulum. Looking at the illustrations of Dupler, itcan be said, beyond doubt, that wall formation is through alveoli, a feature common in gymnosperms. Subsequent divisions in the cellular gametophyte occur, first at the micropylar end and later at the chalazal end. As the gametophyte enlarges, it expands at the upper end so that the original pear-shaped outline becomes more or less cordate with a tapering basal end. A few cells at the micropylar end of the female gametophyte enlarge to. function as the archegonial initials (Fig. 13.87C). The first division of the archegonial initial gives rise to a neck initial and a central cell (Fig. 13.87 D). The former divides to form a neck of 2-4 cells arranged in a single tier. The central cell, on the other hand, enlarges rapidly so much so that its cytoplasm is unable to keep pace with it. This results in the formation ofa lot of vacuoles in the central cell (also knownas foam stage). The central cell nucleus directly functions as the egg (Dupler, 1917; Sterling, 1948; Favre- Duchartre, 1958) without cutting off any ventral canal cell or nucleus (Fig. 13.87E). This is an important feature in which Taxus and other members of the Taxaceae differ from the preceding gymnosperms. Pennell & Bell (1987) made the most unusual observation of the presence of neck canal cell in the archegonium of T. baccata. These authors are certainly in error and this cell may be, if at all it is formed, the ventral canal cell, since neck canal cell is absent in gymnosperms (see Chamberlain, 1935; Mahashwari & Sanwal, 1963; Maheshwari & Singh, 1967). Theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Coniferates 323 ing in the formation of the zygote. The fusion of the two gametes usually takes place near the base of the archegonium. Embryogeny: The zygote nucleus divides mitotically unaccompanied by wall formation to give rise to a 2-nucleate proembryo. Three more such divisions take place so that a 16-nucleate proembryo is formed. Walls are laid down at this stage. Rarely, wall formation takes place after 32 nuclei have been produced. After wall formation the proembryo showsa primary upper tier (pU) of 9-13 cells and a primary embryonal group (pE) of 3-7 cells. The cells of the pU tier have no walls on the upper side and are, therefore, continuous with the general cytoplasm. Internal division in pU gives rise to the upper tier (U) and the lower suspensor tier (S). The cells of the pE, also undergo internal division forming the embryonal tier/ group (E; Fig. 13.89A, B). The cells of the E tier divide rapidly to give rise to the embryonal mass (e). Some of the proximal cells of the latter elongate forming embryonal tubes (et; Fig. 13.89 C). The cells of the U tier degenerate to form a plug. Cleavage of suspensor (S) cells occurs in Taxus (Sterling, 1948). The suspensors separate from each other and each may carry one or more embryonal units. In some cases, groups of meristematic cells are seen at the base of the suspensor cells. These are called the ‘rosette embryos’. They do not, however, grow further. As the suspensor elongates, it pushes the growing embryonal mass deeper in the gametophyte or endosperm. A group of actively dividing (anticlinally and periclinally) cells is seen at the apex. Later, the tip of the embryo shows periclinal divisions whereas anticlinical divisions continue a little below. The anticlinal walls are visualized as concentric arcs radiat- ing from the apex and accounting for the successive growth increment in the apical development. The shoot apex of Taxus differs from that of Pinus in not exhibiting prominent cytohistological zonations. The meristematic activity now shifts to a point slightly in concentric circles and it appears that this meristematic zone is at the focus of this circle (hence named focal zone by Sterling, 1949 a). The concentric dividing cells at the lateral boundary of the focal zone constitute the procambial cylinder. At the base of the procambial cylinder is the root generative meristem which is constituted by larger cells with prominent nuclei. The root cap is marked by the transeversely dividing cells below the generative meristem. The embryonal cortex originates from the peripheral regions of the hypocotyl /shoot axis. It is continuous with the peripheral region of the rootcap-the number of cells increasing from cortex to the rootcap. In Taxus the hypodermal layers, alongwith the epidermis, are continuous from cotyledons to the suspensor. It has been attributed to the indistinct column in the root cap. The cotyledons, usually two in number, are relatively short. Thestriking feature is the relatively minor developmentof theroot capand suspensor region.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Ephedrales 327 the lowest pair which is usually sterile. Each flower consists of an unbranched sporangiophore, bearing at its tip 2-6 bilobed, sessile, microsporangia which dehisce apically. The sporangiophore develops as a small protuberance in the axil of the bract scale. The protuberance, to begin with, has a tunica-corpus organization which soon gets disorganized. The sporangiophore is enclosed in a pair of perianth (hyaline appendages) leaves and it grows out of the latter during maturation of the sporangia (Fig. 14.1B) The female strobilus consists of four to seven pairs of opposite and decussate green bracts, fused at the base forming a cup-like structure. In the axil of each of the uppermost pair of bracts there is an ovule (Figs 14.1C, D; 14.5A). Each ovule consists of two envelopes. The inner envelope, or the integument, is thin and fused with the nucellus except in the upper region which protrudes to form a long micropylar tube. The outer envelope is thicker and completely free from the integument (Fig. 14.5G). In Ephedra the apical meristem transforms into an ovule. The initiation of the ovule is marked by periclinal divisions in the outermost layer of the lateral shoot meristem. The outer envelope, also called perianth or a pair of connate bracteoles by Eames (1952), Fagerlind (1971) and Martens (1971), arises as a dorsal ridge but becomes annular. The outer envelope is formed by anticlinal and periclinal divisions in the epidermis. Immedi- ately above the outer envelope, the integument arises as a protuberance. After initiation the integument becomes annular, but later shows one- sided growth or becomes asymmetric on the dorsal side. The integument remains asymmetric throughout and thisis also reflected in the micropyle. Both the envelopes are inserted at almost the same level in a young ovule. Later, however, the integument (inner envelope) is seen at a much higher level than the outer envelope. This is probably because of growth in the region below the nucellus and integument. The outer envelope grows by an independent meristem. Theepidermal cells of the nucellus undergo periclinal divisions forming a nucellar cap. According to Fagerlind (1971) the massive nucellus arises as a result of periclinal divisions in the epidermis; a parietal tissue is thus absent. However, Singh & Maheshwari (1962) have shown that in E. gerardiana, the nucellus is of dual origin having been contributed by the nucellar epidermis and parietal tissue. After discussing Fagerlind’s obser- vations, Singh (1978) opined thata parietal tissue is presentin some species of Ephedra and absent in others. Ephedra, thus, shows both crassinucellate and tenuinucellate ovules. Root: The root of Ephedra exhibits typical structure from outside within. Theoutermost layer, epidermis, is followed by a well developed parenchy- matous cortex which is generally differentiated into an outer compact portion of thick-walled cells and aninner portion of comparatively loosely arranged thin-walled cells.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.GYMNOSPERMS Fig.14.3: A, Ephedra foliata; B-D, E. trifurca. A.TS part of old stem showing periderm and secondary xylem with vessels. B.Same, showing early wood with (Contd.)aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.338 GYMNOSPERMS Later, Golgi body becomes active and vesicles form a thick wall across plasma membrane (cf Cunninghamia). The cell wall becomes prominent after the degeneration of cytoplasm and organelles. The long, columnar neck appears similar to the transmitting tissue in the style of angiosperms (see Singh, 1978). The central cell is highly vacuolate to begin with. Its nucleus divides to form an ephemeral ventral canal nucleus and an egg nucleus (Fig.14.6B). In some species the ventral canal nucleus is known to persist and remain in the upper part of archegonium. The egg nucleus enlarges and comes to lie in the centre of archegonial cytoplasm. The archegonium, in the meanwhile, acquires a lot of cytoplasm and becomes granular. The archegonium of Ephedra contains only small inclusions and is rich in protein but poor in polysaccharides and RNA. The mature egg is Feulgen negative. The gametophytic cells in the immediate vicinity of the developing archegonium differentiate to form a single or a double-layered jacket. The jacket cells are elongated, densely cytoplasmic and uni- or multinucleate. Generally, each archegonium has a separate jacket but, sometimes, 2 or 3 arhegonia are enclosed in a common jacket layer (Fig. 14.6B). The jacket cells are rich in RNA and total proteins indicating their highly metabolic state (Moitra, 1980). The jacket nuclei regularly migrate to egg cytoplasm and many undergo divisions. Thearchegonium of E. nevadensis is surrounded by a multilayered jacket of bi- or multinucleate densely cytoplasmic cells. There is a high frequency of migration of jacket nuclei into the central cell or egg. The egg nucleus moves down the egg into ‘comet-like’, ‘cytoplasmic dense zone’, having rich amount of organelle DNA (Friedman, 1990b). In another species, E. distachya, Moussel (1977, 1978a) reported numerous plastids and mitochondria in the cytoplasmic dense zone. In the micropylar part, where ventral canal nucleus is located, most of these organelles are absent. Pollination: The pollen grains of Ephedra are elliptic, generally inaperturate, wingless and are airborne. However, Steeves & Barghoorn (1959) observed a colpus at the base of each furrow, i.e. pollen are polycolpate (see also Bharadwaj, 1963) or polyplicate according to Kedves (1987). This is op- posed to monocolpate condition in Welwitschia. Pollen are caught in the pollination drop whichis present on the micropylar tip and are sucked in. The pollen grains thus come to lie on the nucellar tip. The pollination drop contains several amino acids, peptides, malic and citric acids (Ziegler, 1959). Inorganic phosphates and sugar also occur. The concentration of sucrose is very high, nearly 25%. This is supported through artificial cultures where pollen of Ephedra have been found to germinate in a high concentration of sugar. Whereas wind pollinationis important, entomoph- ily too has been reported in E. aphylla and E. campylopoda where ants and other insects visit cones. Drops of nectar rich in sugar occur in both maleaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.342 GYMNOSPERMS (tubular outgrowth) on one side in each of the eight units which grows towards the archegonial base (Fig. 14.9C). The nucleus then leaves the spherical bulb-like portion and comes to lie in the tube. It divides here by Fig.14.9: Proembryogeny in Ephedra (e, embryonal cell; emb, embryo; Es, embryonal suspensor). A. Eight-nucleate proembryo, four (produced from fusion of ventral canal nucleus and second male nucleus) in upper and four (produced from zygote) in lower region of the archegonium. Cell walls have started separating the upper four nuclei. B. Same, at a later stage showing eight proembryonal units; each nucleus is surrounded by radiating cytoplasm. C, Elongation of the embryonal units, nuclei in three of them are dividing Few jacket nuclei are also seen (9). D. Same, ata later stage with an elongated embryonal suspensor and an embryonal cell. E. Embryonal cell has divided into three cells. F-H. Later stages in embryo development. (A-B,D, after Lehmann- Bearts, 1967; C, E-F, after Khan, 1943; G, H, after Narang, 1956).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.348 GYMNOSPERMS is composed of uniform cells. The xylem ray consists of thin-walled, starch- containing parenchymatous cells. The xylem elements produced after secondary growth are larger than those of the stem. In older roots some xylem elements contain starch. Stem apex: As far as the presence of a discrete surface layer is concerned, the shoot apex of Gnetum has assumed an angiospermous character. It exhibits a typical tunica-corpus organisation. The tunica is the outermost single layer of cells extending from the first pair of leaf primordia over the shoot apex. The corpus consists of subapical initials, central mother cell zone, flanking layers and pith rib meristem. The flank zone of the corpus (2 or 3 layers) is derived from the subapical initials and peripheral cells of the central mother cell zone. The latter also gives rise to the pith rib meristem. The flanks contribute to leaf primordia. £ ™s~ Fig.15.3: Gnetum gnemon. Transection of young stem (diagrammatic). Stem: A young stem of G. ula and G. gnemon shows a single layered epidermis of rectangular cells with a thick coating of cuticle and sunken stomata. The cortex consists of 12-16 layers of parenchymatous cells; some cells in the inner zone become fibrous with narrow lumen. In older stems, a ring of parenchymatous cells becomes sclerenchymatous in the inner cortex, and it is referred to as the ring of spicular cells. The sclerotic cells show branched as well as unbranched pit canals. Endodermis and peri- cycle are not distinct. There are 20-24 collateral and endarch vascular bundles, arranged ina ring (Fig. 15.3). The xylem consists of tracheids and few vessels. The phloem is composed of sieve cells and phloem paren- chyma. Medullary rays between vascular bundles are broad and high. Theaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Gnetales 355 G. africanum (Waterkeyn, 1959), the orbicules are present on all the faces of tapetal cells. The epidermis develops thickenings and is the only layer that persists in the mature sporangium which dehisces by a median longitudi- nal slit. The epidermal cells lining the slit are thin-walled. The sporogenous cells (Fig. 15.9B) divide and increase in number, the last cell generation of which differentiates into microspore mother cells (Fig. 15.9D). Broad cytoplasmic channels interconnect microspore mother cells or meiocytes forming a syncytium (Waterkeyn, 1959). A space is formed between the plasmalemma and the mother cell wall. The latter is gradually dissolved away followed by the rounding off of the protoplasm. As the mother cell enters meiosis, it is surrounded by a thick layer of callose. Meiosis results in the formation of decussate, tetrahedral or isobilateral tetrads still embedded in the callose cover (Fig. 15.9E). This covering is soon absorbed releasing individual haploid microspores. Male gametophyte: The microspore nucleus divides to form a small lens- shaped prothallial cell and a large antheridial initial (Fig. 15.9F). The prothallial cell rounds up and does not undergo any further division. The antheridial initial divides forming a antheridial cell and a tube cell. Since a stalk cell is not formed in Gnetum, the antheridial cell directly functions as spermatogenous cell. At thé three-celled stage (Fig, 15.9G), thepollenare shed. Whereas the tube nucleus is hyaline with a large nucleolus, the spermatogenous cell nucleus is rich in chromatin with a cytoplasmic sheath. The prothallial cell eventually degenerates. Anentirely differentand anovel mode of developmentof male gameto- phyte in G. ula was reported by Swamy (1974). According to him the microspore nucleus divides into a small cell and a large cell. The small cell (Swamy calls it generative cell) has its own sheath of cytoplasm and lies free in the large cell. It moves into the pollen tube and gives rise to two male gametes. The nucleus of the large cell divides into two daughter nuclei (the morphology of these two nuclei is not known, because Swamy unfortu- nately could not discuss it). The pollen grain wall consists of an outer thick exine with minute spines and an inner thin intine. Double pollen grains have occasionally been observed in several species. They probably arise due to the non-separation of two or more cells of a tetrad. Like Ephedra and Welwitschia, pollenkitt is lacking in Gnetum (Hesse, 1980). Megasporangium: Nearly four to ten ovular primordia differentiate from an annular meristem or rim below each collar of the female strobilus. The ovular primordium lies on a cushion. The three envelopes arise in a centripetal manner. Whereas in G. ula the ovule is stalked (Vasil, 1959), it is sessile or subsessile in some other species. Of the three envelopes (Fig. 15.10A), only the inner one is the integu- ment. The outer envelope differentiates first. Often called perianth, thisaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Gnetales 365 area called axial tissue. The upper part is gradually consumed by down- ward growing suspensors, and later becomes compressed and crushed. The endosperm is rich in starch and oil droplets and provides nutrition to the undifferentiated embryo which continues to grow after the seed has fallen to the ground. The endosperm cell walls show simple pits. Embryogeny: The early development of the zygote in different species of Gnetum shows variation (Maheshwari & Vasil, 1961a; Martens, 1971). In G. gnemon (Madhulata, 1960) zygotes usually occur in pairs. The zygote may give rise to a small protuberance into which the nucleus moves (Fig. 15.15A) (Lotsy, 1899). It may even divide into two cells and bothor one of the cells (Fig. 15.15 B) may give out the tube. Alternatively, it gives rise to a branched tube, and the nucleus passes into one of the branches (Fig. 15.15C). The tubes have variously been designated as proembryonal tubes, suspensor tubes, or primary suspensor tubes (Fig. 15.15C, D). We prefer to call them primary suspensor tubes to differentiate them from the secondary suspensor which will be discussed later. The tubes become septate and much elongated and coiled, and penetrate the female gameto- phyte or endosperm. These primary suspensor tubes always move down- wards i.e. towards chalazal end (Fig. 15.15E). Some of them may be seen growing over the periphery of the gametophyte (Fig. 15.15F); still others come out of the gametophyte and enter nucellar tissue (Fig. 15.15G). Because of their tortuous course, it is difficult to trace them in a section. However, mass of tubes can easily be dissected out from the centre of the endosperm. Embryo development starts at the tips of some of these primary suspensor tubes where a small cell is cut off at the tip. This cell divides transversely and longitudinally, thus producing a quartet (Fig. 15.18A). Further divisions in this quartet of cells result in a globular embryo (Fig. 15.18B). The cells towards the primary suspensor tube end divide and elongate considerably to form long, coiled secondary suspen- sor which pushes developing embryo deeper in the endosperm. In G. africanum the zygote and its derivatives produce a row of cells and éach of these forms a primary suspensor tube. There is some controversy with regard to the early development of the embryo in G. ula (see Vasil; 1959; Maheshwari & Vasil, 1961a; Swamy, 1973). According to Vasil, the zygote divides into 2 cells which grow into tubular structures. These tubular structures undergo divisions. The daughter cells thus produced elongate forming a bunch of uninucleate primary suspensor tubes growing in different directions in the female gametophyte. The nucleus of the primary suspensor tube (Fig. 15.16A) endospermcells are also seen. D. Branched cellular primary suspensor tube. E,F. Young and old endosperms. G. Lower portion of ovule in LS showing many primary suspensor tubes growing beyond endosperm in the nucellar tissue (A-C, after Sanwal, 1962; D, after Madhulata, 1960; E-G, after Vasil, 1959).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Gnetales 369 The root tip is differentiated at the opposite end. The root cap cells are confluent with the massive secondary suspensor. After the differentiation of the shoot and the root apices, a small protrusion appears in the region between the two apices. This is is the beginning of the so-called ‘feeder’. It shows differentiation into epidermis, cortex, vascular bundles and pith. The feeder, which is very prominent in the mature embryo, is generally longer than the hypocotyl (Fig. 15.18C, D) Thereare several ways by which polyembryony occurs in Gnetum. Each of the primary suspensor tube may develop an embryo at its tip resulting in the formation of many embryos. The embryonal mass at the tip of the secondary suspensor may proliferate to give rise to additional embryos (Fig. 15.18 E). Sometimes the cells of the secondary suspensor may become meristematic producing numerous embryos. Ina primary suspensor tube, instead of one, two or more groups of cells may be produced forming many embryos at the tip (see Maheshwari & Vasil, 1961a). Seed: The seed in most species of Gnetum is elongated. It is oval in G. gnemon. The colour ranges from green to red. The seeds are shed at a stage when the embryo is not fully formed. The nucellus represents a thin strip at the apex; the endosperm is massive, surrounded by a three-layered seed coat. The seed coat of G. africanum and G. ula has been studied in detail by Thoday (1911). The outer envelope (sarcotesta) is green and succulent and is free for most of its length. It consists of mostly parenchymatous cells interspersed with numerous branched fibres or sclereids. Stomata occur on the outer epidermis and as on the leaves they are haplocheilit. Laticifers are formed in this layer. The inner envelope (sclerotesta) is highly complex and could be divided into three separate zones, viz., the outer zone of dark coloured cells with indurated walls, middle palisade layer and inner fibrous lignified zone. Towards the micropylar end of the seed, the thickness of the fibrous zone increases, whereas the strands of indurated tissue diminish. The integument (endotesta) is fused with the nucellus for major part of its length. Its free portion encircles the nucellar cap and projects for some distance beyond the nucellus. In this region several elongated sclereids occur in a few species. All the three layers of the seed coat are supplied with vasculature. A ring of well developed bundles enters the base of the seed. Each bundle bifurcates, the outer series supplying the sarcotesta. The inner series again bifurcates, the outer supplying the sclerotesta and the inner entering the endotesta (see Maheshwari & Vasil, 1961a; Martens, 1971). As mentioned earlier, the embryo is immature when the seed is shed. There is always a time lag between seed shedding and seed germination. InG. gnemon seeds are shed in April and germinate in September, whereas in G. ula they take one year to germinate. The germination is epigeal (Fig. 15.19A-G).aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.CHAPTER 16 WELWITSCHIALES WELWITSCHIACEAE WELWITSCHIA The monogeneric family Welwitschiaceae has only one genus Welwitschia which has been considered as the most bizarre of all gymnosperms, being unique among living and fossil plants. It was discovered, growing near Cape Negro, Angola, more than a century ago in 1859 by an Austrian physician, explorer and botanist, Friedrich Welwitsch. Thomas Baines, a British painter, independently reported Welwitschia in the same year (1859) as abulbous plant with 4 leaves. His name was even suggested asa specific name of this plant (W. bainesii). Exploration since its discovery had dis- closed that Welwitschia covers an area of nearly 1200 km extending along Atlantic coast of Africa from Angola to South-West Africa (Namibia). A large number (5000-6000) of specimens grow in the Welwitschia Flats which is about 45 km east of Swakopmound. One of the oldest specimens, which is about 2000 years old, grows here. The estimated lifespan of Welwitschia is 400-1500 years (Herre, 1961). The rainfall in these regions is exceedingly low, varying between 0 and 100 mm a year. In the coastal regions, the plants depend for their water supply on night fog or dew. According to von Willert (1985), however, itis stilla matter of controversy. The generic name Welwitschia was given by Hooker (1863) in honour of its discoverer, Welwitsch. The latter, however, in a letter to Hooker suggested Tumboa as the generic name. This he based on ‘Tumbo’, the native name of the plant. This name somehow did not find favour with botanists (see Rodin, 1953), who thought that this word had no precise meaning. Welwitschia isa monotypic genus, with only one species, W. mirabilis. The plant resembles a giant turnip or an enormous woody carrot, or giant octopus (PI. 1.10; Fig, 16.1A). It reaches a diameter of more than one meter with two large opposite, strap-shaped leaves. The part of the stem above the leaves has been referred to as the crown and the region between the crown and root as the stock (Fig. 16.1A, B; see Hooker, 1863). A periderm covers the whole plant. A notable event in the life of this plant is that its shoot tip aborts or dies. Although the cause of death is unknown,aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.378 GYMNOSPERMS fibres are not lignified. Gum canals are occasionally found in the leaves of mature plants. These occur between vascular bundles on the lower side near the phloem. Stomata occur on both the surfaces, being more on the upper side. Both the guard cells and the subsidiary cells arise from a single epidermal cell, ie. the stomata are syndetocheilic. The stomata are sunken and are nor- mally seen in areas between the bundles of hypodermal fibres. The leaf is traversed by numerous parallel longitudinal veins that become connected with smaller, obliquely oriented veins. Because of the radial arrangement of cells in xylem and phloem tissues of vascular bundles, the later formed vascular tissues are considered as secondary. Rodin (1958) could not decisively demonstrate the presence of a fascicular cambium in the vascular bundle, so hebelieves that there are no secondary tissues and growth is only primary. Both the tracheids and vessels bear large uni- or biseriate bordered pits. There is a single perforation on each end wall of vessels. Unlike the xylem, the phloem of Welwvitschia is gymno- spermous comprising sieve cells, parenchyma and albuminous cells, the latter shown for the first time. Fine structural studies have revealed intact nuclei (withenvelope) in the mature sieve cell. This, according to Bornman (1972), is the first demonstration of its kind in either gymnosperms or angiosperms, because the nucleus, after becoming necrotic, eventually disappears from a developing sieve element. In some gymnosperms, the nucleus is seen in the mature sieve cell, but it is without its membrane. Strands of fibres are located as caps on both xylem and phloem sides of vascular bundles. Transfusion tissue forms almost a complete sheath around these fibres. It consists of cells having reticulate or scalariform thickenings with small bordered or simple pits. Reproductive structures: Welwitschia is dioecious. ‘Inflorescences’ arise in the leaf axil that are hidden in the groove. Occasionally an inflorescence may be found below the groove. The ovulate (Fig. 16.4A) as well as the staminate (Fig. 16.4B) cones or strobili are compound and are borne in large numbers. The stalk is dichotomously branched several times, and the strobili are borne at the tips of these branched stalks. Extensive work on the embryology has been doneby Martens and his pupil Waterkeyn (Martens, 1959, 1961, 1963, 1971, 1974, 1975, 1977a, b; Martens & Waterkeyn, 1962, 1964, 1974). Male cone: The male cone consists of opposite and decussate cone scales whosearrangementis beautifully geometrical. Figure 16.5C represents the floral diagram of a male flower with associated bracts and cone axis. Each male flower is remarkably bisporangiate and is subtended by a large cone scale. It comprises two lateral bracts and a perianth of two parts, alsoaa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.Welwitschiales 383 -B c Fig.16.7: Welwitschia mirabilis (br, bract; fg, female gametophyte; int, integu- ment; nu, nucellus; oe, outer envelope; ov, ovule; pe, perianth). A. Floral diagram of female flower. B. LS female flower showing nucellus, integument and outer envelope. C. Female flower showing lateral bracts, fused perianth forming wings, and the ovule with a long micropylar tube formed by integu- ment. D. LS female flower at free nuclear female gametophyte stage. (A,C, after Martens, 1959; B, after Chamberlain, 1935; D, after Martens, 1975). 2-8 nuclei, and the lower segmented ‘chalazal’ or ‘sterile’ zone comprises densely cytoplasmic cells with 6-12 or more nuclei (Fig. 16.9D). The upper and central regions of the chalazal zone are vacuolate, and the central and peripheral regions remain meristematic for a considerable time. Several cells of the micropylar zone give out cluster of ‘embryo sac tubes’, or ‘endosperm tubes’, or, more appropriately, ‘prothallial tubes’.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book.GYMNOSPERMS Gymnosperms, as a group, enjoy a ‘unique position in the world flora. The present book is an attempt to include: important aspects of living (extant) and fossil (extinct) gymnosperms. It encompasses distribution, morphology, anatomy, reproductive biology, seed biology, ultrastructure and histochemical studies. The book gives a family-wise description of the group based on therepresentative genus with relevant information on other taxa for better and fuller understanding. Latest research findings have been included to make the text highly usefull and meaningful. Chapters on forest biotechnology and economic importance area special feature of the book. ft The book, besides being profusely illustrated, provides an extensive bibliography. Students of undergraduate and postgraduate classes would find information on gymnosperms under one cover. * S.P. Bhatnagar is Professor in the Department of Botany, University of Delhi. He was a CAS Senior Visitor of the British Council and worked at the Botany School, Cambridge, UK. He has visited several laboratories in Europe, Middle East, Singapore, Thailand, Japan, USA and ‘Canada, He has considerable research and teaching experience in the field of developmental and experimental reproductive biology of angiosperms and gymnosperms. He has participated in several national and international symposia held in India and abroad. Professor Bhatnagar has published nearly 80 research papers in both Indian and foreign journals, including a monograph on Loranthaceae. He is also a co-author of the book The Embryology of Angiosperms. He has contributed review articles in books published from India and abroad. He has also compiled a Bibliography on the Embryology of Angiosperms. Alok Moitra did his Ph.D. on Reproductive Biology of Gymnosperms from University of Delhi under Professor Bhatnagar, the senior author. He has taught gynmosperms to the undergraduate students: and has contributed original research papers and reviews in the subject. He is presently employed in Indian National Science Academy as Officer on Special Duty. Cover pictures: Taxus baccata, Courtesy : N.N. Bhandari Ginkgo biloba, Courtesy : H.Y. Mohan Ram Pinus roxburghii, Courtesy : R.N. Konar Cyeas circinalis, Courtesy : H.Y. Mohan Ram Juniperus virginiana, Courtesy : R.N. Konar NEW AGE INTERNATIONAL (P) LIMITED, PUBLISHERS New Delhi» Bangalore * Chennai * Guwahati * Hyderabad. Kolkata * Lucknow * Mumbai ISBN 81-224-0792-7