Chapter 19 -- SEEDLESS VASCULAR PLANTS

      Vascular tissue allowed plants to become more complex:  true roots, stems, leaves

      Gametophyte became much reduced in size

      Sporophyte dominates

      Life cycle of seedless plants & nonvascular plants is similar

      Sporangia may be clustered in strobili

I. Evolution of vascular plants (seedless & seed producing):

     A. To prevent desiccation:

            1. spores developed thick, protective  walls, & plants developed a cuticle 

              (waterproof, consists of cutin + waxes)

            2. stomata for gas exchange.

B. Other early adaptations:

            1. conducting systems (xylem & phloem) -- transport water & food

2. incorporation of lignin in cell walls of supporting & water-conducting  cells

            3. true roots, stems, & leaves

            4. reduction of gametophyte stage; sporophyte dominates

            5. seeds (= embryonic sporophyte) -- seed plants only

    C. Direct ancestor -- perhaps a multicellular green alga = Coleochaete       

Oldest known fossil = Cooksonia (~ 420 mya)

    D. Became numerous, diverse by Devonian period (408-360 mya)

     E. 9 phyla (~ 250,000 species)


II. General chs. of seedless vascular plants:

    A. All of above, except seedless

    B. Flagellated sperm -- must have water to reach egg

    C. Sporophyte is dominant, often long lived

    D. Sporophyte & gametophyte are nutritionally independent of each other

     Gametophyte is often short lived --photosynthetic in ferns & horsetails,

     saprophytic in club mosses & whisk ferns

    E. archegonium ---> 1 egg; antheridium ----> several sperm

            Life cycle resembles that of nonvascular plants 


III. Plant body organization (all vascular plants):

    A. Early sporophyte was dichotomously branched (evenly forked) -- lacked leaves & roots; had underground rhizomes

    B. Organ systems later evolved:

            1. root system --anchorage, water & mineral absorption

 2. shoot system --stems & leaves  (photosynthesis, conduction of water & minerals)

    C. 3 tissue systems (continuous in all organs):

            1. dermal = outer, protective covering

2. vascular = conductive tissue (xylem & phloem); embedded in ground tissue

             3. ground tissue (mesophyll, cortex, etc.)

    D. Primary & secondary growth

            1. primary -- occurs near tips of roots & stems; initiated by apical meristems; involved in extension of plant body; primary plant body; occurs in all vascular plants!!!

2. secondary -- thickens stems & roots; produced by lateral meristems (such as vascular cambium);

secondary plant body; occurs in woody plants, such as gymnosperms, woody angiosperms, & ancient seedless vascular plants

Allows plants to be of enormous size

Many trees are almost all secondary plant body 

    E. Conducting cells:

1. tracheid (xylem) = elongate tracheary element with tapering ends; = the first water-conducting cells;  in all vascular plants; lignified walls; dead at maturity

What are the vascular plants? Seedless vascular plants, gymnosperms, and angiosperms

2. vessel element (xylem) = more specialized water-conducting cell of angiosperms; not present in other vascular plants except the gnetophytes     (type of gymnosperm); dead at maturity       

            3. sieve element (phloem) -- soft walled; live at maturity

F. Stele = arrangement of vascular tissues (plus pith, if present) in central   cylinder of stem & root of the primary body; pith = ground tissue

1. protostele -- most primitive type: solid strand of vascular tissue (no   pith);  found in roots of most plants (except some monocots)

Examples?  Found in stems of Psilotum & Lycopodium

2. siphonostele -- has a pith in the center & surrounded by cylinders (rings) of xylem & phloem

Stem may have leaf gaps -- places where vascular strand is missing from the siphonostele; associated with leaf traces -- vascular tissue that enters leaves

Examples? typically found in ferns & woody dicot stems

            3. eustele -- has discrete bundles of vascular tissue; pith

Examples? Typically found in stems of:   horsetails, ferns, some angiosperms (herbaceous dicots & monocots)

An atactostele is a special type of eustele


IV. Origin of roots & leaves

Stem evolved first as axis of the plant

A. Roots are fairly simple & primitive in structure;evolved from underground stems (rhizomes); retained a protostele

B. Leaves = primary lateral appendages of stem

1. arise as protuberances (or primordia) from apical meristem

            2. 2 major kinds of leaves:

a. microphylls -- relatively small; contain a single strand of vascular tissue

Associated with stems having protosteles; found in lycophytes

                    b. megaphylls -- larger; have a complex system of veins

Associated with stems having siphonosteles or eusteles; found in most plants


V. Reproduction in vascular plants

     A. Early vascular plants are homosporous -  produce only one kind of spore ->   typically produce bisexual gametophytes

Examples?  psilophytes, horsetails, some lycophytes, & most ferns

Gametophytes develop outside spore wall

    B. More advanced vascular plants are heterosporous

– produce 2 types of spores (in 2 kinds of sporangia):

     microspores (from microsporangia) -- produce microgametophytes

     megaspores (from megasporangia) -- produce megagametophytes

Gametophytes develop within spore wall

Examples: Selaginella, some ferns, all gymnosperms and angiosperms

   C. Sporangia may be single or aggregate into a cone (= strobilus)

 

VI. 4 phyla of seedless vascular plants:

     A. Phylum Psilotophyta (only 2 genera)

            1. Psilotum (whisk fern) -- occurs in FL, LA, TX, AZ, Hawaii,& Puerto Ric

Only living vascular plant that lacks both leaves and roots!

a. dichotomously branched stem with small outgrowths; underground rhizome with rhizoids; sporangia borne on the ends of short, lateral branches; stem has a protostele

                    b. Has bisexual, subterranean gametophytes

2. Tmesipteris -- epiphyte; occurs in the South Pacific; has leaflike appendages

B. Phylum Lycopodophyta (club mosses)

1. 3 orders (3 families) of living species; all are herbaceous, & sporophyte has microphylls, stems, roots

     2. Lycopodiaceae -- club mosses (e.g., Lycopodium); evergreen

a. sporophyte has branching rhizome from which aerial branches & adventitious roots emerge; stem & roots have protosteles

            b. microphylls are spirally arranged

 c. homosporous, & sporangia are borne on fertile microphylls called sporophylls. In many species, sporophylls are grouped into strobili

3. Selaginellaceae -- Selaginella (only extant genus)

             a. leaves occur in 2 ranks

             b. heterosporous; both kinds ofspores occur in same strobilus

  c. has a ligule (scalelike growth) near base of upper surface of each microphyll & each sporophyll

            d. stem & root have protosteles

            4. Isoetaceae -- Isoetes (only extant genus)

    C. Phylum Equisetophyta -- horsetails or scouring rushes

(Equisetum = only living genus

1. appeared during Devonian; during late Devonian & Carboniferous periods, represented by trees called calamites 18 m tall, 45 cm diameter

2. sporophyte is herbaceous; occurs in moist places, along streams,edges of woods:

a. conspicuously jointed stems, small scalelike leaves, rough texture

b. leaves are whorled at nodes, & branches (if present) occur laterally at nodes, alternating with leaves

c. internodes are ribbed & contain siliceous deposits; adventitious roots arise at nodes of rhizome

                    d. perennial; homosporous

spores ---> sporangia ---> sporangiophore ---> strobilus

                        e. gametophyte is the size of a pinhead, is green & freeliving

D. Phylum Pteridophyta -- ferns (~ 12,000 species; largest group of vascular plants next to flowering plants)

            1. abundant since Carboniferous period (360 mya)

2. tropical & temperate in distribution; greatest diversity in tropics (~ 1/3 of these are epiphytes)

3. sporophyte has megaphylls, stems, adventitious roots, rhizomes; sporangia are borne on fertile leaves or a fertile segment

            4. most are homosporous; 2 groups of water ferns are heterosporous

      5. Order Ophioglossales (eusporangiate)

               grape ferns & adder's tongues. (e.g.,  Botrychium & Ophioglossum)

6. Order Pteridales -- most ferns in this order; leptosporangiate; homosporous; rhizome bears new sets of fronds each year; often these are compound & have leaflets called pinnae;  sporangia -- on lower surface of fronds, on specially modified leaves, or on separate stalks; often in clusters called sori; sorus may or may not be covered by flap of tissue called indusium;

            prothallus = gametophyte; bisexual, flat, membranous, green, heart shaped

            7. Order Marsileales & Salviniales -- water ferns

      Marsilea -- rhizome grows in mud, leaves often float

      Salvinia (duckweed) -- entire plant floats & is tiny

      Azolla -- also tiny and entire plant floats