vascular transport, mechanical support, storage: plant stems (including vascular pathways, growth...

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Vascular transport, mechanical support, storage: Plant stems (including vascular pathways, growth forms, woody/herbaceous)

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Vascular transport, mechanical support, storage: Plant stems (including vascular

pathways, growth forms, woody/herbaceous)

Questions

• For plants, APweb characters page• Any problems with the wiki?• For posting questions, please put your name

next to your post• Discussion• Questions?

Ferns

Gymnosperms

Angiosperms

A digression into woody vs. herbaceous

Primary vs. Secondary growth

Groover. 2005. What genes make a tree a tree. TREE. 5:210

Plant stems

• What are the functions of a plant stem?

Plant stems

• How do they accomplish these tasks?– Phloem (living cells) – Xylem

• Dead cells: Vessels, tracheids, fibers• Living cells: Parenchyma (axial and ray)

Transport: Water!

Evapotranspiration

Cohesion-tension

Conductivity increases to the fourth power of the radius

Perforation plates

Pits

Tracheid: Diam: 0.01-0.05 mm, L: ~1-5 mm

Vessel elementsDiam: 20-500 um, Vessel L: few mm to few m

Embolisms

• What happens when air gets in?• Freeze- and drought-induced cavitation

Embolisms

• What happens when air gets in?• Drought-induced cavitation: air seeding

Woody and pseudo woody clades Water conducting cells

Pits Support Living cells Wood

Ferns Tracheids, vessels

Homogenous Sclerenchyma bands

? Pseudo woody

Extant gymnosperms

Tracheids, except Gnetales

Torus/margo, with a number of exceptions

Tracheids 4-10% ray, 0% axial parenchyma

Woody, low density

Monocots (bamboos, palms)

Tracheids, vessels

Homogenous Fibers, tracheids

Ground parenchyma

Pseudo woody

Basal angiosperms, Magnolids, Eudicots

Mainly vessels

Mainly homogenous

Fibers, sometimes tracheids

11-30% ray, up to 11% axial parenchyma

Woody, variable density

Evolution of vessels

Evolution of vessels

• Angiosperms: have higher rates of CO2 uptake and transpiration. They have literally transformed their surroundings (more next week!)

• To do this, they must be able to transport efficiently.

Peforation plates

Pits

Tracheid

Vessel elements

Question

• We know that derived angiosperm vessels are more efficient.

• Were basal angiosperms with vessels more efficient than gymnosperms and vesselless angiosperms?

*

*

*

* = vesselless

Herbs or pseudo woody

Vessels, homogenous pits, fibers

Tracheids, heterogeneous pits,

Herbs or pseudo woody

Basal

Derived

* Basal type with long thin vessels

Question

• We know that derived angiosperm vessels are more efficient.

• Were basal angiosperms with vessels more efficient than gymnosperms and vesselless angiosperms? – Answer: At the level of a given unit of stem, NO!

• So then what?

So, then what?

• We know basal angiosperms vessels are more efficient than tracheids at the conduit level. Meaning?

Perhaps…

• Less space needs to be devoted to vessels, so more diverse cell tissues can evolve or different allocation to tissues can arise… (Heteroxyly)

Why did vessels evolve?

Why did vessels evolve?

• Findings: In the basal angiosperm with vessels– Vessels did not confer greater transport efficiency

at the stem level, nor greater photosynthetic ability

– Vessels did allow for “uncoupling” of the hydraulic and mechanical support functions

• Wood was denser and stems were stronger and energy per volume was greater in the species with vessels.

• Also, species with vessels grew taller.

Why did vessels evolve?

• So…– Why vessels were first important (heteroxyly)

appears to differ from why vessels may have led to the rise of the angiosperms (stem level efficiency)

From water to land: Evolution of tetrapod limbs

Elpistostege

http://tolweb.org/Deuterostomia/2466

http://universe-review.ca/R10-19-animals.htm