Syllabus topics (official RTMNU)
- Gymnosperms: General characteristics, Classification (Stewart, 1982) and Economic Importance
- Fossil Gymnosperms: Cycadeoidea (Morphology and Reproductive structure)
- Life cycle of: Cycas (Morphology, Anatomy and Reproductive structures)
Gymnosperms: General characteristics, Classification (Stewart, 1982) and Economic Importance
Definition: Gymnosperms are seed plants whose ovules are not enclosed in an ovary; they lie exposed on megasporophylls or cone scales. Hence the seeds are "naked". They were dominant in the Mesozoic era.
General characteristics
- Mostly woody, perennial, evergreen trees or shrubs (tallest: Sequoia).
- Plant body is the diploid sporophyte, differentiated into root, stem and leaves. Tap root system; coralloid roots in Cycas (with cyanobacteria) and mycorrhizal roots in Pinus.
- Vascular bundles are conjoint, collateral, open; secondary growth present. Xylem has tracheids but no vessels (except Gnetales); phloem has sieve cells but no companion cells (albuminous cells instead).
- Plants are heterosporous: microspores (pollen) and megaspores. Sporophylls usually grouped into male and female cones (strobili). Plants monoecious (Pinus) or dioecious (Cycas).
- No flowers, no ovary, no style and stigma; hence no fruit is formed.
- Wind pollination; pollen lands directly on the ovule, caught by a pollination drop.
- Male gametophyte is highly reduced; female gametophyte (endosperm) is haploid and formed before fertilisation. Archegonia are present (except Gnetum, Welwitschia).
- Fertilisation by siphonogamy (pollen tube). Male gametes are motile and ciliated in Cycas and Ginkgo; non-motile in conifers.
- Polyembryony is common. No double fertilisation (true double fertilisation absent).
Classification (Stewart, 1982) โ W.N. Stewart in "Paleobotany and the Evolution of Plants" placed gymnosperms (living and fossil) in Division Gymnospermophyta with three classes:
| Class | Orders | Examples |
|---|---|---|
| 1. Progymnospermopsida (fossil, spore-bearing ancestors with gymnosperm-like wood) | Aneurophytales, Archaeopteridales, Protopityales | Aneurophyton, Archaeopteris |
| 2. Gymnospermopsida (true seed plants) | Pteridospermales (seed ferns), Glossopteridales, Caytoniales, Cycadeoidales (Bennettitales), Cycadales, Pentoxylales, Ginkgoales, Cordaitales, Voltziales, Coniferales, Taxales | Lyginopteris, Glossopteris, Cycadeoidea, Cycas, Ginkgo, Pinus, Taxus |
| 3. Gnetopsida | Ephedrales, Welwitschiales, Gnetales | Ephedra, Welwitschia, Gnetum |
Within class Gymnospermopsida two lines are recognised: the cycad line (seed ferns, Bennettitales, Cycadales) with manoxylic wood (soft, much parenchyma), large pinnate leaves and radiospermic seeds, and the conifer line (Cordaitales, Ginkgoales, Coniferales, Taxales) with pycnoxylic wood (compact), small simple leaves and platyspermic seeds.
Economic importance
- Timber: Cedrus deodara, Pinus, Abies โ furniture, construction, plywood.
- Paper pulp: Pinus, Picea.
- Resin, turpentine from Pinus roxburghii; Canada balsam (mountant) from Abies balsamea.
- Food: sago starch from Cycas pith; chilgoza seeds of Pinus gerardiana.
- Medicine: Ephedrine from Ephedra (asthma, cold); Taxol (anticancer) from Taxus; Ginkgo biloba extract for memory and blood circulation.
- Ornamental: Cycas, Thuja, Araucaria.
Fossil Gymnosperms: Cycadeoidea (Morphology and Reproductive structure)
Systematic position: Class Gymnospermopsida (Cycadopsida), Order Cycadeoidales (Bennettitales), Family Cycadeoidaceae. Genus Cycadeoidea lived from the Triassic to the Cretaceous (Mesozoic). Petrified fossils from the USA were studied in detail by G.R. Wieland.
Morphology
- Stem short, stout, barrel-shaped or globose, up to about 1 m tall.
- The stem surface is covered by an armour of persistent leaf bases with dense ramentum (hairs).
- A crown of large pinnately compound leaves (cycad-like) at the apex.
- Internally: large pith, a ring of bundles with manoxylic wood, large cortex; leaf traces simple (not girdling).
- Stomata are syndetocheilic (subsidiary cells from the same mother cell as guard cells) โ a key difference from cycads (haplocheilic).
Reproductive structure ("flower")
- Cones were borne laterally among the leaf bases, sunk in the armour.
- Cones were bisporangiate (bisexual), resembling an angiosperm flower โ hence called "flowers".
- From base to apex the cone shows: (1) a short peduncle bearing spirally arranged hairy bracts (perianth-like); (2) a whorl of about 10โ20 microsporophylls, pinnate, fused at the base into a cup (like a fern frond), bearing synangia (fused microsporangia) on pinnae; they were incurved in bud; (3) a central conical or dome-shaped receptacle (ovulate region).
- The receptacle bore numerous long-stalked orthotropous ovules intermixed with interseminal scales. The expanded tips of interseminal scales formed a protective layer with openings only for micropyles.
- Each ovule had a single integument extended into a long micropylar tube. Seeds were small, dicotyledonous and exalbuminous (embryo almost fills the seed).
Significance: The bisexual, flower-like cone of Cycadeoidea led to the Bennettitalean (Anthostrobilus) theory of angiosperm origin (Arber and Parkin, 1907).
Life cycle of: Cycas (Morphology, Anatomy and Reproductive structures)
Systematic position: Gymnospermopsida โ Cycadales โ Cycadaceae. Example: Cycas revoluta. Called a "living fossil".
Morphology (sporophyte)
- Palm-like, 1โ3 m tall; stem unbranched, columnar, covered by persistent bases of foliage and scale leaves.
- Roots: tap root; coralloid roots โ apogeotropic, dichotomously branched, swollen, containing the cyanobacteria Anabaena/Nostoc in the cortex (nitrogen fixation).
- Leaves dimorphic: (a) foliage leaves โ large, pinnately compound, crowded at the stem apex; young leaflets show circinate vernation; each leaflet has a single midrib and no lateral veins; (b) scale leaves โ brown, dry, hairy, protective.
Anatomy
- Coralloid root: cortex with a distinct algal zone between outer and inner cortex.
- Stem T.S.: outline irregular due to leaf bases; large parenchymatous cortex with mucilage canals and girdle leaf traces; a ring of conjoint, collateral, open, endarch bundles; large pith; manoxylic wood; secondary growth by successive cambia (polyxylic in old stems).
- Rachis T.S.: omega (ฮฉ)-shaped arrangement of vascular bundles; each bundle diploxylic (centripetal xylem exarch + centrifugal xylem endarch).
- Leaflet T.S.: xerophytic โ thick cuticle, sclerenchymatous hypodermis, palisade only on upper side, sunken stomata on lower side, transfusion tissue (primary beside the midrib and secondary/accessory transfusion tissue running across the mesophyll to the margins). Midrib bundle is diploxylic.
Reproductive structures โ Cycas is dioecious.
- Male cone: large, compact, terminal. Numerous spirally arranged, woody, wedge-shaped microsporophylls, each with a sterile upturned apex (apophysis). Lower (abaxial) surface bears thousands of microsporangia in sori (groups of 2โ6). Microspore mother cells undergo meiosis forming microspores (pollen).
- Female "cone": no true cone; megasporophylls are produced in a loose rosette at the apex, and the stem apex keeps growing through them. Each megasporophyll is leaf-like with a petiole bearing 2โ10 ovules on its margins.
- Ovule: the largest ovules in the plant kingdom (up to 6 cm in C. thouarsii); orthotropous, one thick integument with three layers (outer fleshy, middle stony, inner fleshy), nucellus with a beak, pollen chamber, female gametophyte (endosperm) bearing 2โ8 archegonia at the micropylar end, and an archegonial chamber.
Gametophytes and fertilisation
- Microspore forms 3-celled pollen (prothallial, generative, tube cell). Wind pollination; pollination drop draws pollen into the pollen chamber.
- Generative cell divides into stalk cell and body cell; body cell forms two large top-shaped, multiciliated antherozoids (the largest male gametes, about 300 ยตm, visible to the naked eye).
- Pollen tube is haustorial (absorbs food from nucellus). It bursts and releases gametes into the archegonial chamber; one antherozoid fuses with the egg โ zygote (zooidogamy with siphonogamy).
- Embryo develops with a suspensor; seed has two cotyledons, haploid endosperm and stony seed coat. Polyembryony occurs.
5 likely exam questions
Long answer
- Give an account of general characteristics and economic importance of gymnosperms. Add a note on classification of gymnosperms proposed by Stewart (1982).
- Describe the morphology, anatomy of the leaflet and reproductive structures of Cycas with labelled diagrams.
Short answer / short notes
- Describe the reproductive structure (bisexual cone) of Cycadeoidea.
- Write a note on coralloid roots of Cycas.
- Draw a labelled diagram of L.S. of the ovule of Cycas.