Leaf Development And Evolution I: What Is A Leaf Flashcards
Leaf function
1) support
2) photosynthesis
3) defence
4) bearing spores
5) nutrition
6) floatation
Monocot grass leaf
- blade, ligule, sheath
Eudicot simple leaf
Blade, petiole
Seed leaves
Cotyledons
Reduced leaves
- cactus spines
- butcher’s broom cladode
- whisk fern
What is an angiosperm leaf?
- determinate lateral organ associated with a bud
SAMs
- make leaves
- transition from indeterminate to determinate growth via leaf primordia
SAM zones
1) central
2) peripheral
3) rib
SAM Layers
1) L1 (outer)
2) L2 (sub-surface)
3) L3 (inner)
SAM function
- perpetuate stem cell pop
- produce organ dedicates
Perturbed SAM mutants
- wuschel
- clavata1, 2, 3
Wuschel
Homeodomain protein Responsible for proliferating cells
wuschel
- defective (reduced) meristem forms defective organs
- can form axillary meristems
- decreased CLV; less cells in central zone
clavata1, 2, 3
- enlarged meristem
- more cells in central zone
CLAVATA1, 2, 3
Component of receptor-ligand pathway that makes less cells in central zone
WUS/CLV interactions
- WUS promotes CLV expression
- CLV inhibits WUS zone
- feedback inhibition
CLV OX
~ wus (due to strong inhibition)
Expression doesn’t always mean function
Just because the transcript is there, doesn’t mean the protein is
Stem cell maintenance
- kept beneath stem cell zone
- easy to self-regulate upon environmental cue
- WUS = essential for stem cell identity
Periclinal chimaeras
- one layer has a different identity
- generated by grafting
- helps you work out what contributions each layer makes to final structure
- reveal contributions of SAM layers to leaves
Sectorial chimaeras
- for clonal (sector) analysis
- deduce what part of the meristem does what
- reveals no. of leaf founder cells
- shows plastochron
Sector induction to determine cell lineage relationships
- in tobacco
- genetic stock: mid-green double heterozygote
- double chance of sector
- one break = green
- one break = yellow
Lineage analysis
- pattern of division at each plastochrons
- division continues at leaf base
Leaf initiation in tobacco
- 120-180 founder cells (15x2-3x4 in a dome)
- L1, 2 and 3 maintained
- requires co-ordination
L1
Upper and lower epidermis
L2
Palisade upper mesophyll
L3
Inner mesophyll
Vascular tissue
KNOX genes
- knotted1-like homeobox
- Homeodomain proteins bind DNA
- green algae and land plants
Ectopic cell division in KNOX GOF
- 35S constitutive for expression
- necessary and sufficient
Simple angiosperms
Maize, antirrhinum, arabidopsis, pea, tobacco
Simple leaf formation in angiosperms
- KNOX turns off in primordium
- what turns it off?
- need recessive LOF
rs2
- -ve Kn1 regulator (transverse visualisation)
- rough sheath 2
- similar to kn1-0
- ectopic Kn1 accumulation in leaf primordium
RS2 and Kn1
Expressed in mutually exclusive domains / zones
ASl
- inhibits KNOX in leaves
- RS2 ortholog
asl
KNOX expression in leaves
PHANTASTICA
- ARP gene
- turns KNOX off in primordium
ARP genes
Encode Myb TFs
Timing of leaf evolution
- c425Mya; earliest land plants
- c360-409Mya; leaves formed in Devonian
- c130Mya; flowers formed in Cretaceous
Cooksonia
- leafless fossil
- branched axis supporting terminal sporangia
- c400Mya
Leafless fossils
- Rhynia
- Asteroxylon
- Psilophyton
- study absence/arrangement of vasculature
Rhynia
- 410Mya
- lycophyte stem lineage
Asteroxylon
Lycophyte (pre-microphyls)
Psilophyton
- monilophyte stem lineage
- pre-euphyllophyte megaphylls
At the very least, there are
2x Leaf origins, 50My apart
Microphylls
- simple vein
- simple shaped leaf
Megaphylls
Tries to unify many types of leaf
Lycophytes
Microphylls
Telome theory of leaf evolution intro
- explains fossil trajectory
- leaves evolved from flattened branches
Telome theory of leaf evolution
1) branching
2) overtopping (Actinoxylon); dominant branch
3) planation (Archaeopteris); flattening
How does the telome theory of leaf evolution explain M/M divide?
Megaphyll - fusion
Microphyll - reduction
Enation theory of leaf evolution
1) leaves evolved through lateral outgrowth from unbranched axes
2) followed by vascularisation
Sterilisation theory of leaf evolution
- leaves evolved through sterilisation of lateral sporangia
- Zosterophyll
Selaginella kraussiana
- lycophyte (v. hard to transform£
- dichotomous prostrate branching pattern; v. rigorous
- meristems bifurcate
- major and minor branches in specific leaf pairs w/ specified rhizophores
S. kraussiana counting mechanism?
- SkKNOX1
- SkKNOX2
- no 1:1 ortholog
- conserved expression patterns
- in indeterminate equivalent
- where leaf is budding
SkKNOX
- present in meristem
- different to angiosperms!
- conservation of pattern
In order to prove function
you have to KO
35S:SkARP1
- complements asl
- transgene expression matches degree of rescue
Independent recruitment of KNOX pathway
- at least twice
- > 30My apart from
- did it arise de novo?
KNOX expression in bifurcating shoot
When meristem bifurcates, KNOX does too
KNOX hypothesis
- SkARP expression makes KNOX genes switch off in that cleft at the time the meristem needs to branch
- facilitates bifurcation
- can’t prove until KO
Independent recruitment of pathway to enable leaf evolution
1) KNOX promotes cell division in apex of multicellular sporophyte (Bryophyte stem lineage)
2) acquired for branching (Lycophyte stem lineage)
3) KNOX leaves (Lycophytes)
How many times has it happened in euphyllophytes?
Open q!