Chapter 5 Evolution of Woody and Seed Plants

Michael G. Simpson

Evolution of wood from secondary xylem of vascular cambium enabled plants to grow very tall, as large trees.

Illustrated: Sequoiadendron giganteum, giant sequoia, largest non-clonal organism on earth; most tissue dead secondary xylem (wood).

Divisions of vascular cambium mostly tangential (some radial) 2 xylem > inside, 2 phloem > outside: bifacial growth.

Secondary growth from lateral meristems: vascular cambium & cork cambium.

Vascular cambium: 2 xylem > inside; 2 phloem > outside: = bifacial growth. Tissue to outside (cortex, older 2 phloem) gradually crushed.

In most temperate regions (growth season / winter) and some tropical regions (dry/wet season), seasonal annual rings form. Annual rings can be used to date wood. Dendrochronology study of wood in relation to time and environment/climate.

cork cambium - like vascular cambium bifacial cork > outside, phelloderm > inside

Seed = embryo enclosed by nutritive tissue, surrounded by seed coat

Ovule = immature seed (prior to fertilization/maturation)

Seed Evolution

Heterospory

telomes & lagenostome found in fossil (extinct) taxa, functioned in pollen capture

micropyle hole in integument, where pollen enters (gymnosperms)

pollination droplet secreted from micropyle; pollen grains stick to, are pulled inside as droplet evaporates.

Ginko biloba

Pollen grains = immature endosporic male gametophytes

Pollen grains develop from microspores by mitotic divisions/differentiation.

[When haploid nucleus of microspore starts to divide mitotically, it becomes a pollen grain.]

Pollen tube exosporic, tube-like extension from pollen grain.

Pollen tube haustorial (parasitic, feeding off tissues) in Gymnosperms -In cycads & Ginkgo sperm delivered to fertilization chamber, where sperm swims to archgonium = zooidogamy. -In conifers (incl. Gnetales) tube grows directly to archegonium = siphonogamy.

Ovule development

Adaptive significance of seeds

1) Protection of embryo (seed coat) 2) Nutrition of embryo (nutritive tissue=female gametophyte in Gymnosperms) 3) Dispersal unit (e.g., fleshy, carried by animals; winged, transported by wind) 4) Dormancy mechanisms (seed coat involved in preventing germination except when conditions right)

eustele single ring of vascular bundles

Two fossil (extinct) lignophytes:

Archeopteris large tree, wood like a conifer, leaves like a fern; some heterosporous.

Medullosa a tree fern, bore seeds, but with fern-like foliage.

Gymnospermae (naked seed) seeds not enclosed by carpel; sister to Angiospermae.

Cycadophyta - Cycads

Cycadophyta Cycads
-trunks short (rarely elongate, tree-like)

-leaves pinnate (rarely bipinnate), coriaceous, with circinate vernation (like ferns)

-dioecious (separate male & female individuals)

-sperm motile

Cone = determinate shoot systems, consisting of a single axis with sporophylls = modified leaves with attached sporangia.

Pollen cones (male) bear microsporophylls, with microsporangia.

pollen cone

seed cone

Ovulate / seed cones (female) bear megasporophylls, with ovules/seeds.

Cycadophyta Cycads
Cycadaceae: female plants without cones; seeds born on megasporophylls from stem axis. One genus: Cycas Cycas revoluta sago-palm -starch derived from pith > flour, bread. [C. media -edible seeds]

Cycadophyta Cycads
Zamiaceae: female plants with seed cones; leaves pinnate or bipinnate

Cycadophyta Cycads
Zamiaceae: female plants with seed cones; leaves pinnate or bipinnate

Ginkgophyta Maidenhair Trees One family: Ginkgoaceae

Maidenhair Tree family

One species (monotypic): Ginkgo biloba

Ginkgophyta - Ginkgo biloba

- tree with short shoots - leaves obtriangular - venation dichotomous - dioecious

Ginkgo biloba Pollen cones catkin-like: axis bearing stalk-like microsporophylls, each with two microsporangia.

Ginkgo biloba Ovulate reproductive structures: stalk bearing two, erect ovules, each with basal collar. Seeds fetid (butyric acid).

Coniferae conifers Apomorphy:

pollen tube siphonogamous

-leaves simple Three groups:

Pinopsida Cupressopsida Gnetales

Pinaceae: Pinus

Mystery of the conifer cone: compound

Evolution of the conifer cone:

Compound structure (axis bearing two components): bract homologous with leaf ovuliferous scale homologous with shoot system

Coniferae conifers Pinopsida: One family: Pinaceae

Pinaceae
-pollen cones with 2 microsporangia / microsporophyll -pollen grains of some taxa saccate (with 2 bladders)

Pinaceae
-seeds usu. 2 per ovuliferous scale, inverted, winged.

Pinaceae
-leaves linear to acicular (needle-shaped) - in some taxa, short shoots (e.g., Cedrus) or determinate fascicles (Pinus).

Pinaceae
Include: Abies fir Cedrus cedar Larix larch Pinus pine Pseudotsuga Douglasfir Tsuga - hemlock

Coniferae conifers Cupressopsida

5-6 families We will cover 4: Araucariaceae Podocarpaceae Cupressaceae Taxaceae

Araucariaceae
-leaves broad to acicular -microsporangia numerous (5-20) per microsporophyll -ovule 1 per scale Includes: Araucaria heterophylla Norfolk Island-Pine Araucaria bidwillii bunya-bunya Agathis australis kauri

Cupressaceae
-leaves linear, acicular, or subulate, spiral, opposite, or whorled -branches flattened in some, resembling pinnate leaves in some -seed cones with numerous seeds per scale (2-20) -pollen not saccate Includes: Cupressus cedar Juniperus - juniper Sequoia sempervirens redwood Sequoiadendron giganteum giant sequoia Taxodium bald-cypress

Podocarpaceae
-leaves linear, elliptic, or subulate -seed cones usu. fleshy, often reduced, in some subtended by fleshy receptacle, seed may be enveloped by fleshy epimatium derived from scale Includes: Podocarpus, e.g., P. gracilior Phyllocladus spp.

Taxaceae Yew family

-seed cones usu. reduced to 1 ovule -seed surrounded by fleshy aril

Includes: Taxus yew taxol derived from T. brevifolia, used to treat ovarian cancer

Coniferae conifers Gnetales:

3 families: Ephedraceae Gnetaceae Welwitschiaceae All dioecious

Gnetales:
Apomorphies: 1)Pollen striate 2)Vessels porate

Gnetales Gnetaceae One genus: Gnetum (ca. 30 spp.)

Gnetum tropical vines [trees, shrubs], leaves simple, opposite

Gnetales Ephedraceae
One genus: Ephedra (35-45 spp.) Morman Tea

Ephedra Morman-Tea
Shrubs Deserts of S.W. North America, W. South America, N. Africa, and Eurasia

Ephedra
- pollen cones with stalk-like microsporangiophore, bearing synangia - seeds of seed cones with fleshy, connate bracteoles and micropylar tube

Gnetales
Welwitschiaceae One genus/species: Welwitschia mirabilis -native to deserts of Namibia