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nitrogen fixation
conversion of molecular nitrogen to ammonium
can occur through lightning, industrial N fixation and biological N fixation
three types of biological N fixation
symbiosis
associations
free living
symbiosis
energy source: sucrose and metabolites form host plant
ex- rhizobium
fixes 50-400kg
legumes
associations
energy source: root exudates from host plant
fixes 10-200kg
ex- azospirillum
free living
energy source
heterotroph: plant residue
autotroph: photosynth
ex- azobacter
nitrogenase complex
uses reducing power to convert atmospheric nitrogen gas into ammonia
only catalyzed by bacteria/prokaryotes
req a lot of reducing power and atp
plant provides carbon source
N2 is triple bonded, hard to break
3 major steps of symbiotic association formation
signaling
infection
nodule development
Signaling
release of elicitors by plant roots
chemoattractants (attracts rhizobia)
nod genes (produced by bacteria)
confirms compatibility by making interactions specific to certain plants and bacteria
molecules involved in signlaing
flavonoids and isoflavonoids
ex- luteolin (alfalfa, nod gene inducer)
ex- genistem (soybean, nod gene inducer)
betaines and aldonic acids
nod factors
bacterial products that act as morphogens to induce nodulation in the plant
molecules involved, lipochitooligosaccharides (LCOs), lipids and carbs, hormones, nodulation outer proteins
plants recognized very low nod factor concentration
perceived by plants via nod factor receptor kinases
signaling and host specificity in rhizobia
root produces chemoattractants/elicitors into the soil
rhizobia in the rhizosphere recognize elicitors, their nod gene is activated
rhizobia produces and releases nod factors into the soil
plant recognizes nod factors via receptors
infection thread
a tunnel built by the plant for the bacteria to move into
formed within 18-30h of plant recognizing nod factors
determinate nodules
soybeans
nodule develops at roughly the same rate
have a set end and are not meristematic, cannot grow indefinitely
spherical in shape
nodule primordia
in crop legumes, initiated opposite to protoxylem poles
indeterminate → w/in inner root cortex
determinate → w/in outer root cortex
indeterminate nodule
alfalfa and peas
have meristem region and can keep growing
oval/oblong in shape
have 3 regions, 1 with new cells forming, 1 with cells doing the fixation, and 1 with cells senescing
transfer of fixed N into host
nh3 diffused out of bacteroid into the symbiosome and is protonated to NH4+ due to low pH of symbiososome
nh4 transported across peribacteroid membrane and into the cyotplasm of the infected cell, where they can be used
determinate N exporting
nh3 assimilated into glutamine, then xanthine that moves into uninfected cells where ureides are produced and exported via xylem
indeterminate N exporting
nh3 made into glutamine, then asparagine which is used for export
oxygen permeability barrier
layer of empty cells, reduces rates of oxygen penetration into nodule core
leghemoglobin
high oxygen affinity
controls availability of free oxygen in infected cells to limit nitrogenase inhibition
creates the red color of the rhizobia
tissue
a group of similar cells doing the same job
organ
different types of tissue (ex- leaf)
growth
irreversible change
a positive change over a period of time
usually quantitative
differentation
functional changes
qualitative change
development
growth and differentiation
overall changes that a plant undergoes through its life cycle
ex- seed germ, flower formation, fruit formation, etc.
types of growth
cell division and expansion
increase in dry weight
Protoderm
forms epidermis
primary meristems
protoderm
ground meristem
procambium
ground meristem
forms pith, cortex, and mesophyll
cortex
forms cork cambium, which makes periderm
procambium
forms primary phloem and xylem as well as vascular cambium (which then forms secondary xylem and phloem
phytomer
a unit consisting of a leaf, a node, an internode, and an axillary meristem
intercalary meristem
present in grasses
meristem at base of the leaf sheath, leaf blade, and intersection of node and internode
root zones
root apical meristem
elongation zone
maturation zone
cellulose
arranged in microfibrils in cell wall
primary cell wall component
synthesized at the plasma mem
primary cell wall components
cellulose
hemicellulose (ex- xyloglucan, arabinoxylan, etc)
pectin
diffuse growth
entire cell as a whole expands at the same rate
cell tip growth
one side of the cell expands
expansion is unequal
ex- root hairs, pollen tubes
acid induced growth
cell walls extend much faster at an acidic ph rather than a neutral ph
mediated by expansins
creep
time dependent irreversible extension, typically the result of slippage of wall polymers relative to one another
plant cell expansion steps
osmotic uptake of water across the plasma mem is driven by the gradient in water potential
turgor pressure builds because of cell wall rigidity
biochemical wall loosening occurs, allowing the cell to expand in response to turgor pressure
phyllochron
rate of appearance of successive leaves (or phytomers)
when the leaves appear visually, not when they are created
plastochron
rate of initiation of successive leaves
cell death
localized process that culminates in the death of the cell and is often quite rapid
ex - hypersensitive response
senesence
slower, systemic process that includes nutrient remobilization and under most conditions, culminates in the death of cells
ex- deciduous trees shedding leaves in autumn
programmed cell death
a developmental program in many tissues
ex- tracheary cells, arenchyma formation, leaf senesence, sepal and petal senesence
hormones/phytohormones
naturally occurring substance which influences physiological processes at low concentration
factors influencing hormone effects on tissues
biosynthesis
catabolism
compartmentation
transportation
modulation/modification
sensitivity of the tissue to the hromone
auxin major functions
cell elongation
tropism (photo, gravi, and thigmo)
regulation of developmental affects
apical dominance, floral bud development, leaf abscission, lateral root formation, vascular differentiation, fruit development
auxin synthesis
synthesized in meristem and young dividing tissue, especially SAM and young leaves
basipetal
movement from shoot tip to base of the plant (area b/t shoot and root)
acropetal
movement from base to root apex
auxin transportation (cellular level)
transport proteins at the end/base of the cell, not on side or top
establish gradient of auxin movement in vascular tissue
auxin moved into apoplast, depending on pH, apoplast is protonated or not
not protonated → diffuse through membrane
protonated → co transported with protons into the cell
eflux carriers move auxin out of bottom of the cell
largely active transportation
phototropism
growth in response to light
auxin is distributed on the far side of the light source, causing higher rates of cell elongation on one side, forcing the shoot to bend towards the light
gravitropism
growth in response to gravity
auxin is redistributed to cause cell elongation on one side, forcing the plant to grow upright
ex- goosenecking in corn
auxin in roots
inhibits growth/elongation in main axis of root/root tip
moves to root from synthesis in the shoot via phloem
statoliths
starch granules within the root cap
statoliths when root growing straight down
uniform distribution of auxin via the stele
statoliths when root is horizontal
statoliths fall and press on one side of the cells, causing auxin to move to other side of the root, inhibiting elongation and causing the root to bend as elongation is increased on one side vs the other
apical dominance
auxin suppresses growth of axillary/lateral buds
when the terminal bud/apical meristem is removed, the suppression is subsided and the axillary bud will grow
strawberry achenes
achenes in strawberries (seeds) cause the receptacle to swell and form the fruit
when achenes are removed, receptacle does not swell as much
but if supplemented with auxin (achenes removed) the receptacle swells
cytokinin function
promote cell divison in the shoot
regulate auxin action and distribution
delay leaf senesence
promote nutrient movement
involved in formation of N fixing nodules in legumes
cytokinin cell division in sam and ram
promote shoot growth by increasing cell proliferation in the SAM
inhibit root growth by promoting exit of cells from the RAM, keeping the meristem cell number small
cell proliferation
increased production of cells in the meristem
auxin vs cytokinin in the SAM
auxin promotes differentiation into lateral organs/organ initiation
cytokinin increases cell proliferation, promoting stem cell fate
auxin vs cytokinin branching in SAM
auxin inhibits branching
cytokinin promotes branching
auxin vs cytokining root branching
auxin promotes branching
cytokining inhibits branching
cytokinin oxidase
inactivates cytokinin
axillary buds and auxin vs cytokinin
auxin produced in apical bud inhibits axillary bud growth by turning off cytokinin biosynthesis and turns cytokinin oxidase on, reducing cytokinin concentration
if apical bud is removed, no flow of auxin turning off genes
so then cytokinin biosynth gene is turned on and cytokinin oxidase is turned off, leading to axillary bud growth