and directional evolution Greg and Ray and Tim Proteaceae are the - - PowerPoint PPT Presentation
and directional evolution Greg and Ray and Tim Proteaceae are the - - PowerPoint PPT Presentation
Cell size, genome size, plant strategies and directional evolution Greg and Ray and Tim Proteaceae are the best organisms in the world a short stroll through leaf anatomy Palisade Leaf (lamina) Vein thickness Vessels Epidermis see
Proteaceae are the best organisms in the world
a short stroll through leaf anatomy
Palisade Vein Vessels Epidermis see detail… Leaf (lamina) thickness
Detail of an epidermis
Epidermal cells stomate
Epidermis in surface view
small epidermal cells small stomates high stomatal density big epidermal cells big stomates low stomatal density
paradermal section showing vein density
a short stroll through leaf anatomy
Palisade cells Vein Vessels Epidermis with stomata Leaf (lamina) thickness
unified changes in leaf cell size (across 80 million years!)
stomatal size epidermal cell size vessel diameter leaf thickness palisade cell size phenotypic phylogenetic significant correlations
Do we expect directional evolution
* Systematic changes over the Cenozoic (last 65 million years)
Stomatal size affects photosynthetis
packing density and stomata
10 20 30 40 50 60 70 80 90 200 400 600
stomatal length (μm) abaxial stomatal density (mm-2)
hypostomatic species amphistomatic species
stomatal length ~ 1/√stomatal density
small stomata are more efficient
- per stomate conductance scales linearly with
size
- number of stomates scales 1/quadratically
with size
- small stomates lead to high conductance
matching water supply (veins) and demand (stomata)
functional links
1/stomatal size vein density stomatal density 1/vessel size packing constraints
Expected directional evolution
1 CO2 model
– low CO2 = a need for greater conductance – small stomata are more efficient – guard cell size should have decreased
genetic link
- genome size
- other genetic factors
10 100 0.9 9 90 guard cell length (microns) genome size pg
Grevilleoideae Persoonioideae + Bellendena Proteoideae + Symphynematoideae
Do we expect directional evolution
1 CO2 model
– low CO2 = a need for greater conductance – small stomata are more efficient – guard cell size should have decreased
2 genome size model
– genome size drifts up (one-way path to obesity) – Guard cell should have increased
- strong evolutionary association with open
vegetation (versus rainforest)
- leaf thickness
- stomata on both leaf surfaces
Do we expect directional evolution
1 CO2 model
– low CO2 = a need for greater conductance – small stomata are more efficient – guard cell size should have decreased
2 genome size model
– genome size drifts up (one-way path to obesity) – Guard cell should have increased
3 ecological model
– follows changes in habitat – mostly increase - as woodland replaced rainforest
So, what has happened to stomatal size through time?
- ancestral state analyses
- fossils
ancestral state reconstruction
scatter plot of reconstruction versus age for each node
10 100 25 50 75
guard cell length (μm) - log scale Time (millions of years ago)
but fossils say
10 100 25 50 75
guard cell length (μm) - log scale Time (millions of years ago)
ancestral states fossils
and it happens within clades
5 10 15 20 25 30 35 40 20 40 60 80
guard cell l length (μm) Time (millions of years ago
Banksieae
fossil ancestral states
directional evolution
- driver either
– habitat shift with global climate change – systematic trend in genome size
- invalidates the ancestral state reconstructions