I M.Sc., BOTANY PLANT DIVERSITY II UNIT: I, II &V Unit :I General topics in pteridophytes Introduction of pteridophytes Pteridophytes (ferns and lycophytes) are free-sporing vascular plants that have a life cycle with alternating, free- living gametophyte and sporophyte phases that are independent at maturity. The body of the sporophyte is well differentiated into roots, stem and leaves. The root system is always adventitious. The stem is either underground or aerial. The leaves may be microphylls or megaphylls. Their other common characteristics include vascular plant apomorphies (e.g., vascular tissue) and land plant plesiomorphies (e.g., spore dispersal and the absence of seeds. General characters of pteridophytes: Pteridophyta (gr, pteron = feather, phyton = plant), the name was originally given to those groups of plants which have well developed pinnate or frond like leaves. Pteridophytes are cryptogams (gr. Kruptos = hidden, and gamos = wedded) which have well developed vascular tissue. Therefore, these plants are also known as vascular cryptogams or snakes of plant kingdom. They are represented by about 400 living and fossil genera and some 10,500 species. Palaeobotanical studies reveal that these plants were dominant on the earth during the devonian period and they were originated about 400 million years ago in the silurian period of the palaeozoic era. (i) Majority of the living pteridophytes are terrestrial and prefer to grow in cool, moist and shady places e.g., ferns. Some members are aquatic (e.g., marsileci, azolla), xerophytic (e.g., selaginella rupestris, equisetum) or epiphytic (e.g., lycopodium squarrosum) (fig. 1). (ii) Majority of the pteridophytes are herbaceous but a few are perennial and tree like (e.g., angiopteris). Smallest pteridophyte is azolla (an aquatic fern) and largest is cyathea (tree fern). (iii) Plant body is sporophytic and can be differentiated into root, stem and leaves. (iv) Roots are adventitious in nature with monopodial or dichotomous branching. Internally usually they are diarch. (v) Stem is usually branched. Branching is monopodial or dichotomous. Branches do not arise in the axil of the leaves. In many pteridophytes stem is represented by rhizome. (vi) Leaves may be small, thin, scaly (microphyllous e.g., equisetum), simple and sessile (e.g., selaginella) or large and pinnately compound (megaphyllous e.g., dryopteris, adiantum). (vii) Vascular tissue is present in stem and root. It consists of xylem and phloem. Xylem consists of tracheids only and phloem has only sieve tubes. (viii) The steel is protostele (e.g., rhynia, lycopodium), siphonostele (e.g., equisetum), dictyostele adiantum) or polycyclic (e.g., angiopteris). (ix) Cambium is absent; hence, they do not show secondary growth. 3. Reproduction in pteridophytes: (i) Reproduction takes place by means of spores which are produced inside sporangia. (ii) The development of the sporangium may be leptosporangiate (sporangium originates from a single cell) or eusporangiate (sporangium develops from a group of cells). (iii) Sporangia may be borne either on stem or leaves. On the stem they may be terminal (e.g., rhynia) or lateral (e.g., lycopodium). On the leaves (sporophylls) they may be ventral, marginal (pteris, adiantum) or dorsal (e.g., polypodiceae). In equisetum the sporangia are borne on special structures called sporangiophores which constitute a cone. In marsilea, azolla, salvinia sporangia are produced in sporocarps. (iv) Spores on germination give rise to multicellular gametophytic bodies called prothalli (sing. Prothallus). (v) In homosporous pteridophytes prothalli are monoecious (antheridia and archegonia develop on the same prothallus). In heterosporous species prothalli are always dioecious. Microspores on germination give rise to male prothalli and megaspores to the female prothalli. (vi) Antheridia and archegonia are developed on prothalli. (vii) Antheridium is surrounded by a single layered sterile jacket. (viii) Archegonium consists of four vertical rows of neck cells, 1-2 neck canal cells, ventral canal cell and egg. (ix) Antherozoids are unicellular, biflagellate (e.g., selaginella) or multiflagellate (e.g., equisetum and ferns) and motile. (x) Antherozoids are attracted towards the neck of the archegonium chemotactically by certain substances like malic acid) present in the mucilaginous substance formed by the degeneration of neck canal cells and venter canal cell. (xi) Water is essential for fertilization (zooidogamous). Therefore, pteridophytes are also known as amphibians of the plant kingdom. (xii) Fertilization results in the formation of zygote or oospore, which ultimately develops into well-developed sporophyte. (xiii) The fertilized egg divides transversely or vertically. Another cross wall forms a quadrant stage producing stem, leaf, foot and root. (xiv) Plants show heteromorphic alternation of generation. The main plant body is sporophytic and forms a dominant phase in the life cycle. Classification of pteridophytes (parihar 1996): the modern classification of pteridophytes have been suggested by many scientist according to the Sporne(1975) pteridophytes have been classified into six classes. They are psilopsida, psilotopsida, lycopsida, sphenopsida, pteropsida and progymnospermopsida. This classification is a modification is a modification of Reimer’s (1954) system. According to the modern system of classification by Parihar in 1996; pteridophytes have been divided into seven divisions. They are rhyniophyta, trimerophytophyta, zosterophyllophyta, psilotophyta, lycopodiophyta, equisetophyta and polypodiophyta. Bold (1973) differentiated psilophytes into living and fossil groups. The living psilophytes were placed under the division psilotophyta and extinct psilophytes were divided into three division. They are rhyniophyta, trimerophytophyta and zosterophyllophyta Types of stele in pteridophytes: Stelar system of plant: definition and types (with diagrams) Definition of stelar system: According to the older botanists, the vascular bundle is the fundamental unit in the vascular system of pteridophytes and higher plants. Van tieghem and douliot (1886) interpreted the plant body of vascular plant in the different way. According to them, the fundamental parts of a shoot are the cortex and a central cylinder, is known as stele. Thus the stele is defined as a central vascular cylinder, with or without pith and delimited the cortex by endodermis. The term stele has been derived from a Greek word meaning pillar. Van Tieghem and Douliot (1886) recognized only three types of steles. They also thought that the monostelic shoot were rare in comparison of polystelic shoots. It is an established fact that all shoots are monostelic and polystelic condition rarely occurs. The stele of the stem remains connected with that of leaf by a vascular connection known as the leaf supply. Types of steles: 1. Protostele: Jeffrey (1898), for the first time pointed out the stelar theory from the point of view of the phylogeny. According to him, the primitive type of stele is protostele. In protostele, the vascular tissue is a solid mass and the central core of the xylem is completely surrounded by the strand of phloem. This is the most primitive and simplest type of stele. There are several forms of protostele: (a) Haplostele: This is the most primitive type of protostele. Here the central solid smooth core of xylem remains surrounded by phloem (e.g., in selaginella spp.). (b) Actinostele: This is the modification of the haplostele and somewhat more advanced in having the central xylem core with radiating ribs (e.g., in psilotum spp.). (c) Plectostele: This is the most advanced type of protostele. Here the central core of xylem is divided into number of plates arranged parallel to each other. The phloem alternates the xylem (e.g., in lycopodium). (d) Mixed-pith stele: Here the xylem elements (i.e., tracheids) are mixed with the parenchymatous cells of the pith. This type is found in primitive fossils and living ferns. They are treated to be the transitional types in between true protosteles on the one hand and siphonosteles on the other (e.g., in gleichenia spp. And osmunda spp.). 2. Siphonostele: This is the modification of protostele. A stele in which the protostele is medullated is known as siphonostele. Such stele contains a tubular vascular region and a parenchymatous central region. Jeffrey (1898) interpreted that the vascular portion of siphonostele possesses a parenchymatous area known as a gap immediately above the branch traces only or immediately above leaf and branch traces. On the basis of these branch and leaf gaps Jeffrey (1910), distinguished two types of siphonosteles. In one type, however, the leaf gaps are not found and they are known as cladosiphonic siphonosteles. In the other type both leaf and branch gaps are present and they are known as phyllosiphonic siphonosteles. Jeffrey (1902, 1910, and 1917) interpreted the evolution of the siphonostele from the protostele as follows. He supported that the parenchyma found internal to the phloem and xylem has been originated from the cortex. The supporters of this theory believe that the inner endodermis found to the inner face of the vascular tissue and the parenchyma encircled by this endodermis have been originated from the cortex. According to Jeffrey and other supporters of this theory the siphonosteles with internal endodermis are more primitive than those without an internal endodermis. The siphonosteles which do not possess the inner endodermis are