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Where are plant hormones located?
Move through xylem, cells, or air as a gas
Cell signaling stages
Reception, transduction, response
Reception
Env’t signals are detected by receptors (often proteins embedded in the membrane
Transduction
Environmental signals are converted to a form that amplifies the signal to trigger a response
Response
High enzyme activity and gene expression, secondary messengers are produced
etiolation
morphological adaptation for growing in the dark, lack chlorophyll, allocate all resources to stem elongation
phytochrome
A pigment attached to a photoreceptor protein, is converted into active form during reception stage to initiate signaling
Transduction example
Active phytochrome triggers secondary messenger production (cGMP and Ca2+) that activate protein kinases
kinases
enzymes that phosphorylate other molecules using ATP
Role of Ca2+ in plants
A common signaling molecule (secondary messenger) that triggers changes in pH, transcription, and vacuolar pumops
Response example
Activated kinases influence transcription factors that bind DNA to allow transcription of light-response genes and proteins for development
phytohormones
Synthesis occurs in many cell types simultaneously and may act there or leave, not a linear relationship between concentration, degree, and type of activity
Animal hormones
Synthesis in discrete organs and tissues, transport from site of synthesis to site of action, localized, control processes in conc-dependent manner
What must be true for hormones to create a response?
Present in a sufficient quantity, target tissue is sensitive to hormone, there is a receptor to bind that will change shape
primary natural auxin
IAA, indole-3-acetic acid, derived from AA tryptophan
How were auxins discovered?
Charles and Francis Darwin covered coleoptile from light and it didn’t bend, proving phototropism
Boysen-Jensen proved auxin is a chemical and diffusible through gelatin but not butter
Fritz Went isolated IAA
Function of auxin
Links env’t signals with directional growth (apical dominance and tropic responses), cell elongation, alone induces root formation in a callus
gibberelins
Part of terpene group, GA1 (first discovered in plants), GA3 (fungal gibberellic acid), GA4 (most common plant gibberelin)
How were gibberellins discovered?
Fungus Gibberella secreted gibberellins causing rice to grow tall/spindly and topple at maturity
Gibberellin function
Stem/internode elongation, seed germination, flowering, fruit growth
Cytokinin
Adenine derivatives with isopentenyl side chains, first was kinetin derived from degraded DNA
Function of cytokinin
Stimulate cell division, alone induces shoot formation of callus
How were cytokinins discovered?
Early tissue culture experiments using coconut milk or tobacco with modified forms of adenine
How do auxin and cytokinin regulate tissue formation?
Both are required in a correct ratio to maintain cells of an undifferentiated callus
ethylene
Gaseous hormone, C2H4, rapidly produced in response to drought, flooding, injury, infection
ethylene triple response
Allows seedlings to circumvent obstacles. Inhibits hypocotyl, reduces elongation, thickening of hypocotyl, exaggerated curvature
climacteric fruits have…
High respiration and ethylene so they ripen post-harvest
abscisic acid
A terpenoid derived from carotenoids in mature leaves, levels rise under stress
abscisic acid function
Regulates maturation of embryos and seed germination, levels rise at embryogenesis, can prevent vivipary through dormancy
abscisic acid response to water stress
Accumulates in stressed leaves to inhibit stomatal opening before water potential changes, low soil moisture transfers ABA from roots to shoots
brassinosteroids
First discovered in pollen, functions in sex determination and similar to sex hormones of animals
strigolactones
Stimulates seed germination, mycorrhizal associations, apical dominance
Role of sunlight
Supplies energy for photosynthesis, c-fixation, and biomass accumulation
solar tracking (heliotropism)
Mov’t of leaf throughout the day so its surface is always capturing the sun
phototropism
Altering overall plant growth patterns in response to light
photomorphogenesis
Influence of light on developmental responses (ex. seedlings)
photoperiods
flowering in response to day length, describes hours of light given
photoreceptor
Chromoprotein with a light absorbing group (chromopore) attached to apoprotein with catalytic abilities, absorb photons to initiate a response
How are photoreceptors different from photosynthetic pigments?
They absorb different wavelengths, result in conformational change, create developmental response
phytochromes
Two forms: red (660nm) and far-red (730nm)
Role: In all stages of development
Relationship b/t red and far-red light
Far-red is active form that moves into nucleus to interact with transcriptional regulators. A flash of red light converts red photoreceptors to far-red, flash of far-red results in far-red photoreceptors
phytochrome reversibility
Some seed germination is promoted by red light whereas far-red inhibits
Red and far-red light under vegetative canopies
Red light absorbed by chlorophyll, far red enriched and transmitted
cryptochromes
Absorb: Blue light (400-450nm), induces conformational changes exposing signaling domains
Role: Seedling development, reg. circadian rhythms, anthocyanin biosynthesis
What light induces flowering?
Blue and far-red, red light inhibits
phototropin
PM associated photoreceptors that contain LOV domains
Absorb: UV-A (320-400nm)
Role: Phosphorylation of kinase domain, activation of downstream signaling pathways (stomatal opening, chloroplast mov’t, phototropism)
blue light receptors
Cryptochromes, phototropin, uncharacterized UV receptors. Contain flavin molecules as chromophores
uncharacterized UV-receptors
Absorb: UV-B (280-320nm)
Role: Stress response
How does auxin contribute to phototropism?
Auxin is transferred to the shaded side causing cells to elongate creating curvature towards light source
Where are chloroplasts localized under different light intensities?
Low light→ Along abaxial and adaxial surfaces
High light→ Parallel to light to minimize absorbance
Dark→ Settle at the bottom
How does blue light regulate stomata?
Closes stomata to optimize CO2
Phototropins in guard cells initiate signaling cascades
Activates H+-ATPase pumps
H+ exported out causing (-) charge, K+ accumulates inside
High turgor opens stomata
development
Changes in form from embryo to seedling, not random and can be reversible
embryogenesis
Formation and development of the embryo inside a seed
growth
Irreversible quantitative change in cell number, size, and/or volume
cell growth = increase in size
tissue and organ growth = increase in cell number and size
callus
A mass of cells resulting from those isolated from plants that can de-differentiate
totipotency
Ability of cells too revert to embryonic state without passing through a reproductive stage, most plants
animal development
Establish body plan during embryogenesis, evolved mobility
lineage dependent mechanisms involving transcription factors
homeotic (Hox) genes for proper placement or structures
plant development
Adaptive with more post-embryonic development, evolved flexibility to environment
cell position determines fate
no migration
continuous meristems
2-cell stage (embryogenesis begins)
Zygote forms (1st diploid cell after fertilization), asymmetry and polarity determined. Apical (becomes embryo) and basal (becomes suspensor) cells
suspensor
anchors embryo to maternal tissue to facilitate nutrient/hormone transport
globular stage
Apical cell undergoes series of divisions, primary tissues differentiate into protoderm (establishes radial symmetry), meristem, procambium (early vascular tissues)
polarity
directionality, structural or chemical differences at opposite ends
heart stage
Cotyledon primordia establishes bilateral symmetry, shoot system organization
Torpedo stage
Elongation along the apical-basal axis
Bent-cotyledon stage
After elongation in torpedo stage, cotyledons bend around the hypocotyl and embryo is anatomically complete
Two patterning systems in embryogenesis
Apical-basal axis→ shoots vs roots, longitudinal growth
Radial axis→ tissue organization (dermal, ground, vascular)
Periclinal cell divisions
Produces cells parallel to radial axis for tissue formation, adds cell layers, dermal tissue produced first
Anticlinal divisions
Produces cells perpendicular to radial axis, increase number of cells within a layer to maintain radial integrity
Auxin role in embryo development
Determines position of SAM, cotyledons, RAM
2-cell stage→ auxin asymmetrically distributed along apical-basal axis
globular stage→ auxin redirected to basal region of embryo
heart stage→ complex and directional transport
How is auxin transported?
Protonated IAAH (uncharged, H+ cotransport) form can enter cells, loses H+ in neutral pH of cytoplasm to become charged and incapable of diffusion across PM
chemiosmotic mechanism of auxin transport
pH gradients facilitate entry/exit because cell wall is acidic (H+-ATPase maintains acidity), cytoplasm is basic and IAA- accumulates,
PIN proteins
Mediate exit/entry of IAA/IAAH through the PM, can reorient
ABCB transporters
Actively export auxin using ATP, work with PIN proteins to increase transport efficiency
gurke
lack apical development, polarity fail from disrupted auxin transport
fackel
defective in central (hypocotyl) region from disrupted auxin transport
monopteros
lack proper basal structures (ex. root) from disrupted auxin transport
gnom
lack both apical and basal structures from disrupted auxin transport
Three features of all seeds
embryo, food storage (often 3n endosperm from double fert), testa (seed coat)
endospermic seeds
Endosperm (3n, major storage tissue) is retained at maturity, embryo small, cotyledons thin and specialized for absorption, reserves mobilized to embryo during germination
scutellum (monocots)
Single cotyledon modified to facilitate absorption from endosperm
coleoptile (monocots)
Protective sheath that covers primary leaves of grass seedling
coleorhiza (monocots)
Protective sheath around the radicle
mesocotyl (monocots)
“stem” similar to the hypocotyl
non-endospermic seeds
Endosperm is used during embryogenesis, cotyledons serve as primary food storage tissue, embryo large, typical of dicots, reserves mobilized to cotyledons during germination
exogenous dormancy
Seed coat or other seed structures inhibit germination (water impermeability, gas exchange interference, mechanical constraint, chemical inhibitors)
endogenous dormancy
Embryo dormancy (abscisic acid or embryo size)
photoblasty
Light requirement for seed germination, often small seeds, phytochrome responses
Phase 1 of seed germination
Imbibition, rapid water uptake driven by water potential fuels cell expansion (matric potential low creating a gradient, water potential becomes less negative as water binds to surfaces)
Phase 2 of seed germination
Lag/metabolic activation with little growth, seed volume increases and coat ruptures, transcription/translation occuring
Phase 3 of seed germination
Post germination, radicle emerges, water uptake increases, cells expanding and dividing as starches, lipids, and oils are mobilized
Hormone balance theory
ratio of ABA:GA determines seed dormancy
When is dormancy promoted?
ABA is synthesized and GA degraded, ABA inhibits embryo growth and reserve mobilization
When is germination promoted?
GA synthesis and ABA degradation, GA stimulated embryo growth and reserve mobilization
vivipary
Germination of seeds on the mother plant, allow quick establishment in complex env’ts (ex. tides, salility)
Preharvest sprouting
Seeds germinate on parent plant, dormancy lost before harvest, triggered by wet or humid env’ts
Precocious germination
Seeds germinate during development before entering dormancy due to mutant that disrupts ABA biosynthesis
stratification
Cold treatment (5 degrees C) for a period of time that releases seeds from dormancy, germination increases with duration
How can gibberellins mobilize endosperm reserves?
GA is produced by scutellum
Stimulates production of amylase near aleurone layer to stimulate transcription/translation
Amylose hydrolyzes starch to simple sugars which gets transferred to embryo (ABA can inhibit amylase)