Plant Development Flashcards
Multicellularity evo in plants?
No common multicellular ancestor w animals
Independent
Development may be different from animals then
In plants - cells joined by cell walls so don’t change neighbours
-no cell migration
-so development potentially simpler?
-done by directional growth instead
Why no central control unit?
land Plants stationary
Also get bigger to compete
Need to harvest nutrients from ground w roots so can’t move
Can’t run away
Perpetually getting eaten partially
So if had central unit and it was eaten whole plant done for
So instead have many stem cell pockets - and the stem cells are more flexible
Higher flexibility plant stem cells?
Animal adults usually have just multipotent progrnitor types that can just make specific tissue
Plant stem cells are much more totipotent
Makes cloning easier too
Plant organ origin?
Apical meristems
Shoots and roots grow from tips
Right in middle of meristem are stem cells
Make organs - eg leaves and flowers
SAM makes?
Basically everything above ground
Germ cells
Leaves
Flowers
Often times parts of roots?
Modularity of plant development
Iterated developmental unit
Primary shoot meristem makes PHYTOMERS - functional units of the plant -can give rise to other shoots??
If bit is taken off and eaten - another dormant meristem is activated and get another healthy plant
Plant germ cell origin differences
In animals - cells are put away v early to become germ cells - strict differentiation
Plant germ cells develop much later
Multicellular diploid embryo evolution
Evolved in land plants
Land plants all evolved from freshwater aquatic algae
Multicellular gen in FW algae is haploid
Fertilisation to make diploid zygote but that immediately undergoes mitosis to make 4 haploid spores
Bryophyte embryo
In land plants - bryophytes
Diploid zygote divides to make multicellular embryo
Benefiti as swimming sperm cannot easily swim on land (too dependent on water)
Then that can produce many haploid spores (many more than 4)
Benefitial as increases odds of successful spore on land
Vascular land plant embryo
Have specialised cells for carrying water and nutrients
So vascular plants can grow bigger
Embryos in vascular plants covered in seed
Seed can store nutrients for development as seed is likely underground when germinating
Helps with dispersal
Plant embryo development within seed
-double fertilisation
2 gametes within female ovule
-haploid egg cell - 1 speed fertulises this to make zygote -> goes on to become embryo
-2 sperm fuse with homoploid central cell - makes endosperm which fills seed and is consumed as embryo grows - is ephemeral and consumed by embryo
Maternal tissue in seed
Integuments
Surround seed
Suspensor
Holds embryo in seed
Nutrients can go up
Undergoes apoptosis during embryo development
Somatic embryogenesis
Possible due to totipotent stem cells being present dispersed around body
Can be activated by hormones or muse pressing certain embryo genes
Can be used for cloning
Occurs naturally in a few species
Kalanchoe (“mother of thousands”) forms embryos around its leaves
Apomixis
eg Dandelion - is triploid
Are sterile due to triploidy - usually cannot make seed
Except they do it with unfertilised flowers - embryo ends up with no paternal info
Often meiosis breaks down giving diploid egg cell - parthenogenesis
Gives clinal propagation by seed - APOMIXIS
The PAR gene confers parthenogenesis
Is expressed in sperm cells - so sworn cell delivers product to mother which kicks off parthenogenesis
Mutant in dandelion which expresses PAR in ovule ends up w no seed in embryo
Embryo Germination
Stem cells at what will be SAM and RAM
Cotyledons - embryonic leaf
Hypocotyl - embryonic stem
So apical basal axis set up in embryo
But many adult structures (seed, fruit, flowers) are not
So development is continuous throughout life(flowers develop in adult plant)
Plant developmental patterning timing
Most occurs post embryonically
Embryogenesis establishes the:
-apical/basal pattern (shoot/root)
-Inside/outside pattern (epidermis/ground tissue/vasculature)
-and stem cells (shoot and root meristems)
Most of plant body plan produced after embryogenesis
Post embryonic patterning
Most of plant body plan produced after embryogenesis
Flowers
Germline
Lateral roots
Branches
Most leaves
Tubers
Continued patterning from groups of stem cells termed meristems
Different to animals where body plan largely formed in embryo
Gives flexibility
Predation problem
Difference to plants - Drosophila embryo patterning
Counter to plants. - patterning of adult stuff set up in embryo
AP and DV axes prefigured in egg
So maternal info in egg determine zygote axes
Localised determinants are localised at piles of egg/zygote (eg bicoid)
Kinase signalling pathways determine terminal elements AP pattern
AP fates determined early on
Localise TF expression (homeodomain proteins) specify regional fate on AP axis
Cell-cell interactions and signals important for segment polarity (wingless etc)
Apical basal pattern appearing in arabidopsis
Egg cell long and thin
The asymmetric cell division giving small apical cell and large basal cell
Apical cell:
Divides longitudinally to give 4 then 8 cells
Then makes layer around to make future epidermis
Apical cell goes on to form embryo
Basal:
Basal cell forms the suspensor and parts of root meristem
Apical basal fate determination
If after zygote 1st division kill apical cell
Basal cell becomes apical in nature
So fate of the cells are due to signaling
If signalling changes then fate of cell changes
Basal cell is capable of producing embryo but is blocked by inhibitory signal from apical cell that inhibits embryo identity/promoted basal fate
Auxin in apical and basal cells
Auxin normally accumulates in apical cell
Important for establishing the vertical/longitudinal division planes and setting off 3D growth
Auxin accumulation promotes vertical rather than horizontal division patterns?
Polar PIN7 efflux protein localisation in basal cell causes this accumulation
Isolating basal cell from both apical and maternal tissue means it does not respectfully like if just apical is ablated
So some +ve signal must be coming from maternal tissues
Basics - how is apical and basal axis and shoot root suspensor fates specified?
Kinase mediate signalling pathways
Localised determinants
Localised TFs
Polar transport of auxin
Zygote polarisation properties
Is long and thin
Nucleus toward one end
Vacuole towards other
Zygote is transiently symmetrical after fertilisation but elongates and becomes polar
Promoter of basal fate in embryo
YODA Activity promoted basal fate
Loss of yoda causes suspensor cells to divide longitudinally like in embryos
YDA gene encodes a MAP3K
Component of signalling pathways
Phosphorylase’s downstream MAP kinases such as MPK3 and MPK6
Constitutive activation of YDA gives opposite effect of loss of function
Hyperactive YODA phenotype
Have defective more suspensor like embryos
Lose YDA - base become embryo like
Hyperactive YDA - apical becomes suspensor like
YDA kinase signalling promotes basal fates and inhibits apical fates in early embryo
Stomata formation
Stomata with 2 guard cells forming mouth like structure
Involves asymmetric divisions in epidermis - smaller daughter remains in stomata lineage - divides again to form 2 daughters becoming guard cells
Bigger daughter becomes elidermis pavement cell
YDA and stomata
YDA inhibits stomata fate
YDA- = more stomata
Hyperactive YDA = all pavement cells
Breaking of asymmetry in stomat lineage BASL
BASL determines asymmetric division in stomatal lineage
Mutant gives lots of nearby stomata instead of spread out
because BASL needed for asymmetric division
No good BASL = symmetric stomatal lineage division -> both go on to guard cell lineage causing close by stomata
BASL localisation
Is in polar localisation on cell periphery
Predicts the asymmetric division plane
Localised to the nucleus and the edge of the cell
Cell will then divide so that the cell was forms distal to the cortical BASL crescent
The larger daughter cell inherits cortical BASL and the smaller daughter enters the stomatal lineage
YDA and BASL
YDA required for BASL polar localisation
YDA mediates BASL phosphorylation and is required for its polar localisation
A similar pathway may control asymmetrical division in zygote
YDA and zygote asymmetrical division
(BASL not present here)
Shirt suspensor (ssp) mutant resembles YDA mutants
Ssp encodes a PM localised pelle-like receptor kinase
Misexoresson of ssp in leaves inhibits stomata development - so confirms intersction wit YDA?
Ssp genetics
Self heterozygote for mutation and WT plants
50% if resusltung embryos give mutant phenotype not expected 25% that would be mutant homo
Cross male WT to homozygous mutant females
All progeny is WT
Cross ssp mutant males with WT females
All embryos are mutants
Suggests that ssp is active in male side (sperm)
Male activity of ssp
Ssp active in male side (sperm)
Pollen expresses ssp RNA
Deposits that RNA in the ovule where it is translated to protein
Explains the odd genetics as maternal is never active
Paternally supplied SSP protein
Localised to one side?
promotes YDA activity
WUS gene (WUSCHEL)
Encodes homeodomain TF
Required for stem cell niche in shoot meristem
Wox homeodomain genes
Patterned on apical basal axis
Due to auxin localisation
Important for apical basal fate specification
WOX2 WOX8
WOX2
WOX8
WOX8 WOX9
WOX9
Homeosis
Assumption by one member of a meristic series of the form or characters proper to other members of the series
Meristic series. -series of repeated homologous units
Homeotic mutations - one member of a repeating series is replaces by another member
Homeotic genes control
Differences between these repeated units
Eve difference between segments in drosophila
Whorl organs
Most flowers have organs in them
Organised in whorls
Whorl number in flowers
1- sepals - form bud
2- petals
3- stamens - male
4- carpel - female
Homeotic mutations in floral organ identity
Class a - affect identity of organs in whorls 1 and 2
Class b- 2 and 3
Class c - 3 and 4
ALWAYS 2 adjacent whorls
Class a mutant example
Arabidopsis
1- sepal - transformed to carpel
2- petals - transformed to stamen
Apetal1
Sepals more leaf like
Petals absent
Class B mutants example
Sepals sepals carpels carpels
Arabidopsis- apetala3 ap3, pistillata pi
Antirhinnum- deficiens def and globosa glo
2 and 3 affected
Class C mutants
Agamous
No sex organs
Whorls 1 and 2 fine
Stamen replaced by extra whorl of petals
Carpel also converted - and unlike WT where stem cells stop and only get 4 whorls - in agamous keep getting more and more whorls of sepals and petals - double flower
Eg plena in antirrhinum
abc model of floral organ identity
3 Homeotic functions
a b and c
-a function in whorl 1 and 2
-b in 2 and 3
-c in 3 and 4
Combinations on Homeotic functions in a whorl specifies the identity of the organs that form there
a =sepal
a+b=petal
b+c=stamen
c=carpel and STOP
Mutual inhubition in a and c
Class c mutations give sepals and petals everywhere even though A function is needed there and is not normally everywhere
Same with C expression in whorls 1 and 2 in A mutants
Lead to idea that A and C are mutually inhibitory
So mutating one expands range of other into the two other whorls
A and C functions are antagonistic
Triple mutant for a b and c
ap2 pi ag triple mutant
Organs are leaf like
Flowers are modified shoots
Floral organs are modified leaves
Like shoots flowers also the compressed internodes, organs in concentric rings
Flowers are determinate
Shoots usually indeterminate
Genes in the abc model
TFs
MADS box mediating DNA binding
C function genes
Agamous RNA expressed where whorls 3 and 4 will be
B function genes
Apetella3 RNA active where whorls 2 and 3 will be not others
A function genes
On where whorl 1 and 2 will be
Are the floral Homeotic genes sufficient for floral identity?
Ectopic expression of them in leaves does not transform them
So not sufficient
SEP genes also needed
SEP genes
SEP1 2 and 3 isolated
Sequence similarity to AGAMOUS
Expressed in whorls 1-4
Single mutants have no phenotype (redundancy)
Triple mutant all floral organs are like sepals
SEP1-3 genes are needed for activity of b and c class Homeotic genes
SEP1-4 quadruple mutant - all organs are leaf mike and have branched trichomes
SEP genes also needed for class a activity too