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Divisions • Three divisions –Hepatophyta (liverworts) –Anthocerophyta () –Bryophyta ()

botit..wisc.edu Beatific

nobuosubaru.blogzine.jp www.hiddenforest.co.nz

Common Characteristics Homospory

• Motile • All bryophytes homosporous • most prominent • Produce 1 kind of generation, not • Spore develops into gametophyte – Thallus = undifferentiated into a • Gametophyte produces both antheridia , stem or (sperm) and archegonia () • Most lack cuticle – For absorption • No true leaves, stems or – General lack of – But see mosses, …

1 Bryophyte Phylogenies But Not a Natural Group! Vascular LiverwortMoss Vascular • Paraphyletic Hornwort Liverwort plants – Some, but not all, descendents of a common ancestor – In other words, what other people thought

was correct was, in fact, wrong Vascular Liverwort Hornwort Moss plants – Based on an old Vascular MossHornwort Liverwort plants

Rhizoids sterile Antheridia jacket • Roots – Absorption – Anchorage fertile – Vascular tissue cells • (sperm) – Anchorage – Minimal or no absorption – No vascular tissue! • – Underground stem – Vascularized, with stem (not root) pattern

2 Archegonia Hornworts •Simplest gametophyte of all bryophytes •Small, flat thallus sporophyte

neck gametophyte

venter

Sporophyte Sporophyte

s • Contains true stomata p o – Gas exchange () r o • Under favorable conditions may outlive p h gametophyte Foot y – Foot acts as root t e – Conducting tissue may develop at Gametophyte sporangium base •Foot supplies nutrients via transfer cells – Unusual condition of free-living •Embedded in gametophyte sporophyte!

3 Hornwort Spermatozoid

Capsule splits (sporangium)

Liverworts Liverworts Using as an example • Gametophyte often ribbon-like – Thallose or leafy • Rhizoids • Sporangia with 4 valves • Capsule with – Specialized, thickened cells – Dispersal • Examples: Riccia & Marchantia • Gametangia buried in deep, lengthwise depressions on upper surface of thallus

4 : Archegoniophores • Marchantia is dioecious (separate male & female plants)

•Form on upper surface. •Contain gemmae (small green disks of tissue) Archegoniophore: structure bearing archegonia •Raindrops break them free of cup, & these can turn into NOT sporophyte tissue!!!!

Marchantia Sporophyte Marchantia Antheridiophores

Antheridiophore: structure bearing antheridia

5 Spermatozoid Marchantia Cycle

Mosses Bryophyta: Mosses

3 classes www.kib.ac.cn sporophyte • Andreaeopsida – Lantern, Granite mosses gametophyte

Andreaea • – Peat mosses

Maire Smith – True mosses

Polytrichum

6 Gametophytes Protonemal Phase • 3 Phases – Protonemal phase • Creeping, filamentous – phase phase – Leafy phase (“phyllids”) • initially dormant – Water + light cause growth – comes out of spore – Growth is directed towards red light

Bud Phase Vascular Plants?

• Some protonemal cells • Some mosses have hydroids & leptoids develop into a mass of • Not the same as xylem & phloem cells called a bud • One bud cell functions as the tip of a caulid, controlling direction of further cell • Each bud develops into a mature gametophyte • Rhizoids grow downward from bud

7 Hydroids Leptoids

• Hydroids have no lignin • Primitive sieve elements? • Very slow translocation of water • Degenerate, inactive nuclei • Limited distribution in plant • Many plasmodesmata in end walls • Mosses can remain dry for a long time, – Plasmodesmata connect cells yet come back to life when wetted • Nearby parenchyma tissue may act as – Up to 20 years in drought-tolerant , companion cells yet come back to life within 4-24 hours! – Details when we cover anatomy – Try it! Get some moss, let it dry for a few – Companion cells (higher plants) help weeks or longer, then wet it load/unload , etc.

Asexual reproduction Sexual Reproduction

cups (splash cups) • Gametangia produced at main caulid apex • Monoecious or dioecious • Clonal • Apparent independent of light – Protonema can produce more – Soil pH, air temp., etc. • Antheridial heads • Phyllid tissue in wet soil – Antheridia elongate may produce protonemal – Outer sterile jacket usually contains strands – Sperm have 2 flagella • Rhizoids can produce buds • Archegonial heads. (sometimes) – Often retain a layer of water (phyllids aid in this) – Any sperm that land can swim on in

8 Moss Sporophyte Moss Sporophyte • Foot, seta and capsule • Mature seta and capsules contain: • Old increases in size, – of cuticle becomes the calyptra – stomata – Acts as temporary – thick-walled steroids (small, thick-walled protective covering over parenchyma cells) sporangium – a cortex region • Operculum covers – central strand of conducting tissue sporangium opening • Can produce 10-50% of • Up to 50 million spores needs per sporangium – Rest from gametophyte

Dispersal: Moss Life Cycle • Calyptra falls off • Operculum shed • Peristome teeth attached to annulus

peristome teeth

annulus

9 What good is it? Sphagnum

• Sphagnum have large, empty, clear cells in epidermis that fill with water (via a pore) & serve as reservoir • Worldwide carbon dioxide buffer

– High CO2 levels, take in more – Low levels, take in less • Environmental indicators • WWI wound dressing What about ecological indicators? – Antiseptic & absorbent • Peat bogs cover ~ 1% earth’s surface – Half size USA!

G.Chandler Various Moss Structures

Sphagnum bog (Tierra del Fuego)

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