Seed germination Seeds remain dormant until sense appropriate
- Slides: 73
Seed germination Seeds remain dormant until sense appropriate conditions: • Many require light: says that they will soon be able to photosynthesize: often small seeds with few reserves • Hormones can also trigger (or stop) germination • ABA blocks it • GA stimulates it
Seed germination Germination is a two step process • Imbibition is purely physical: seed swells as it absorbs water until testa pops. Even dead seeds do it. • Next embryo must start metabolism and cell elongation • This part is sensitive to the environment, esp T & p. O 2
Vegetative growth Once radicle has emerged, vegetative growth begins • Juvenile plants in light undergo photomorphogenesis • Juvenile plants in dark undergo skotomorphogenesis • Seek light: elongate hypocotyl, don’t unfold cotyledons
Vegetative growth Once radicle has emerged, vegetative growth begins • Add new leaves @ SAM in response to auxin gradients • Roots grow down seeking water & nutrients • 1˚ (taproot) anchors plant • 2˚ roots absorb nutrients • Continue to add cells by divisions @ RAM
Transition to Flowering Adults are competent to flower, but need correct signals Very complex process! Can be affected by: • Daylength • T (esp Cold) • Water stress • Nutrition • Hormones • Age
Light regulation of Plant Development Plants use light as food and information Use information to control development
Light regulation of Plant Development • Germination • Morphogenesis • Sun/shade & shade avoidance • Flowering • Senescence
Light regulation of growth Plants sense 1. Light quantity 2. Light quality (colors) 3. Light duration 4. Direction it comes from Have photoreceptors that sense specific wavelengths
Light regulation of growth Duration = photoperiodism (Garner and Allard, 1920) Maryland Mammoth tobacco flowers in the S but not in N = short-day plant (SDP) Measures night! 30" flashes during night stop flowers LDP plants such as Arabidopsis need long days to flower SDP flower in fall, LDP flower in spring, neutral flower when ready
Light regulation of growth Measures night! 30" flashes during night stop flowers LDP plants such as Arabidopsis need long days to flower SDP flower in fall, LDP flower in spring, neutral flower when ready Next : color matters! Red light works best for flowering
Phytochrome Next : color matters! Red light (666 nm)works best for flowering & for germination of many seeds! But, Darwin showed blue works best for phototropism! Different photoreceptor!
Phytochrome But, Darwin showed blue works best for phototropism! Different photoreceptor! Red light (666 nm) promotes germination Far red light (>700 nm) blocks germination
Phytochrome Red light (666 nm) promotes germination Far red light (>700 nm) blocks germination After alternate R/FR flashes last flash decides outcome
Phytochrome Red light (666 nm) promotes germination Far red light (>700 nm) blocks germination After alternate R/FR flashes last flash decides outcome Seeds don't want to germinate in the shade!
Phytochrome Red light (666 nm) promotes germination Far red light (>700 nm) blocks germination After alternate R/FR color of final flash decides outcome Seeds don't want to germinate in the shade! Pigment is photoreversible
Phytochrome Red light (666 nm) promotes germination Far red light (730 nm) blocks germination After alternate R/FR color of final flash decides outcome Pigment is photoreversible! -> helped purify it! Looked for pigment that absorbs first at 666 nm, then 730
Phytochrome Red light (666 nm) promotes germination Far red light (730 nm) blocks germination After alternate R/FR color of final flash decides outcome Pigment is photoreversible! -> helped purify it! Looked for pigment that absorbs first at 666 nm, then 730
Phytochrome Red light (666 nm) promotes germination Far red light (730 nm) blocks germination After alternate R/FR color of final flash decides outcome Pigment is photoreversible! -> helped purify it! Looked for pigment that absorbs first at 666 nm, then 730 Made as inactive cytoplasmic Pr that absorbs at 666 nm
Phytochrome Made as inactive cytoplasmic Pr that absorbs at 666 nm or in blue Converts to active Pfr that absorbs far red (730 nm)
Phytochrome Made as inactive cytoplasmic Pr that absorbs at 666 nm or in blue Converts to active Pfr that absorbs far red (730 nm) 97% of Pfr is converted back to Pr by far red light
Phytochrome Made as inactive cytoplasmic Pr that absorbs at 666 nm or in blue Converts to active Pfr that absorbs far red (730 nm) 97% of Pfr is converted back to Pr by far red light Also slowly reverts in dark
Phytochrome Made as inactive cytoplasmic Pr that absorbs at 666 nm or in blue Converts to active Pfr that absorbs far red (730 nm) 97% of Pfr is converted back to Pr by far red light Also slowly reverts in dark: how plants sense night length
Types of Phytochrome Responses Two categories based on speed 1. Rapid biochemical events 2. Morphological changes
Types of Phytochrome Responses Two categories based on speed 1. Rapid biochemical events 2. Morphological changes Lag time also varies from minutes to weeks
Types of Phytochrome Responses Two categories based on speed 1. Rapid biochemical events 2. Morphological changes Lag time also varies from minutes to weeks: numbers of steps after Pfr vary
Types of Phytochrome Responses Lag time also varies from minutes to weeks: numbers of steps after Pfr vary "Escape time" until a response can no longer be reversed by FR also varies
Types of Phytochrome Responses Lag time also varies from minutes to weeks: numbers of steps after Pfr vary "Escape time" until a response can no longer be reversed by FR also varies: time taken for Pfr to do its job Conclusions: phytochrome acts on many processes in many ways
Types of Phytochrome Responses Two categories based on speed 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2
Types of Phytochrome Responses Two categories based on speed 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible!
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible! But action spectrum same as Pr
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible! But action spectrum same as Pr • Induced by FR!
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible! But action spectrum same as Pr • Induced by FR! Obey law of reciprocity: 1 nmol/m-2 x 100 s = 100 nmol/m-2 x 1 sec
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible! But action spectrum same as Pr • Induced by FR! Obey law of reciprocity: 1 nmol/m-2 x 100 s = 100 nmol/m-2 x 1 sec 1. Examples: Cab gene 2. induction, oat 3. coleoptile growth
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 • Changes 0. 02% of Pr to Pfr • Are not FR-reversible! But action spectrum same as Pr • Induced by FR! Obey law of reciprocity: 1 nmol/m-2 x 100 s = 100 nmol/m-2 x 1 sec 1. Examples: Cab gene 2. induction, oat 3. coleoptile growth 4. 2. LF: induced by 5. 1 µmol/m-2, saturate @ 6. 1000 µmol/m-2
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 Are FR-reversible!
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 Are FR-reversible! Need > 3% Pfr
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 Are FR-reversible! Need > 3% Pfr Obey law of reciprocity
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 Are FR-reversible! Need > 3% Pfr Obey law of reciprocity Examples : Lettuce seed Germination, mustard photomorphogenesis, inhibits flowering in SDP
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 Are FR-reversible! Need > 3% Pfr Obey law of reciprocity Examples : Lettuce seed Germination, mustard photomorphogenesis, inhibits flowering in SDP 3. HIR: require prolonged exposure to higher fluence
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Effect is proportional to Fluence
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Effect is proportional to Fluence Disobey law of reciprocity Are not FR-reversible!
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Effect is proportional to fluence Disobey law of reciprocity Are not FR-reversible! Some are induced by FR!
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Effect is proportional to fluence Disobey law of reciprocity Are not FR-reversible! Some are induced by FR! Examples: inhibition of hypocotyl elongation in many seedlings, Anthocyanin synthesis
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Effect is proportional to fluence Disobey law of reciprocity Are not FR-reversible! Some are induced by FR! Examples: inhibition of hypocotyl elongation in many seedlings, Anthocyanin synthesis 1. Different responses = 2. Different phytochromes
Types of Phytochrome Responses 3 classes based on fluence (amount of light needed) 1. VLF: induced by 0. 1 nmol/m-2 , saturate @ 50 nmol/m-2 2. LF: induced by 1 µmol/m-2, saturate @ 1000 µmol/m-2 3. HIR: require prolonged exposure to higher fluence Different responses = Different phytochromes: 3 in rice, 5 in Arabidopsis
Types of Phytochrome Responses Different responses = Different phytochromes: 3 in rice, 5 in Arabidopsis 1. PHYA mediates VLF and HIR due to FR
Types of Phytochrome Responses Different responses = Different phytochromes: 3 in rice, 5 in Arabidopsis 1. PHYA mediates VLF and HIR due to FR • Very labile in light
Types of Phytochrome Responses Different responses = Different phytochromes: 3 in rice, 5 in Arabidopsis 1. PHYA mediates VLF and HIR due to FR • Very labile in light 2. PHYB mediates LF and HIR due to R • Stable in light
Types of Phytochrome Responses 1. PHYA mediates VLF and HIR due to FR • Very labile in light 2. PHYB mediates LF and HIR due to R • Stable in light 3. Roles of PHYs C, D & E not so clear
Types of Phytochrome Responses 1. PHYA mediates VLF and HIR due to FR • Very labile in light 2. PHYB mediates LF and HIR due to R • Stable in light 3. Roles of PHYs C, D & E not so clear PHYA & PHYB are often antagonistic.
Types of Phytochrome Responses PHYA & PHYB are often antagonistic. In sunlight PHYB mainly controls development
Types of Phytochrome Responses PHYA & PHYB are often antagonistic. In sunlight PHYB mainly controls development In shade PHYA 1 st controls development, since FR is high
Types of Phytochrome Responses PHYA & PHYB are often antagonistic. In sunlight PHYB mainly controls development In shade PHYA 1 st controls development, since FR is high But PHYA is light-labile; PHYB takes over & stem grows "shade-avoidance"
Phytochrome Pr has cis-chromophore
Phytochrome Pr has cis-chromophore Red converts it to trans = active shape
Phytochrome Pr has cis-chromophore Red converts it to trans = active shape Far-red reverts it to cis
Phytochrome Pfr is a protein kinase: acts by kinasing key proteins • some stays in cytoplasm & activates ion pumps
Phytochrome Pfr is a protein kinase: acts by kinasing key proteins • some stays in cytoplasm & activates ion pumps • Rapid responses are due to changes in ion fluxes
Phytochrome Pfr is a protein kinase: acts by kinasing key proteins • some stays in cytoplasm & activates ion pumps • Rapid responses are due to changes in ion fluxes • Increase growth by activating PM H+ pump
Phytochrome Pfr is a protein kinase: acts by kinasing key proteins • some stay in cytoplasm & activate ion pumps • Rapid responses are due to changes in ion fluxes • most enter nucleus and kinase transcription factors
Phytochrome stay in cytoplasm & activate ion pumps • Rapid responses are due to changes in ion fluxes most enter nucleus and kinase transcription factors • Slow responses are due to changes in gene expression
Phytochrome most enter nucleus and kinase transcription factors • Slow responses are due to changes in gene expression • Many targets of PHY are transcription factors, eg PIF 3
Phytochrome most enter nucleus and kinase transcription factors • Slow responses are due to changes in gene expression • Many targets of PHY are transcription factors, eg PIF 3 • Activate cascades of genes for photomorphogenesis
• • Phytochrome Slow responses are due to changes in gene expression Many targets of PHY are transcription factors, eg PIF 3 Activate cascades of genes for light responses Some overlap, and some are unique to each phy
• • • Phytochrome Slow responses are due to changes in gene expression Many targets of PHY are transcription factors, eg PIF 3 Activate cascades of genes for light responses Some overlap, and some are unique to each phy 20% of genes are light-regulated
Phytochrome • 20% of genes are light-regulated • Protein degradation is important for light regulation
Phytochrome • 20% of genes are light-regulated • Protein degradation is important for light regulation • Cop mutants can’t degrade specific proteins
Phytochrome • Protein degradation is important for light regulation • Cop mutants can’t degrade specific proteins • COP 1/SPA targets specific transcription factors for degradation
Phytochrome • Protein degradation is important for light regulation • Cop mutants can’t degrade specific proteins • COP 1/SPA targets specific TF for degradation • DDA 1/DET 1/COP 10 target other proteins for degradation
Phytochrome • Protein degradation is important for light regulation • Cop mutants can’t degrade specific proteins • COP 1/SPA targets specific TF for degradation • DDA 1/DET 1/COP 10 target other proteins for degradation • Other COPs form part of COP 9 signalosome
• • • Phytochrome Protein degradation is important for light regulation Cop mutants can’t degrade specific proteins COP 1/SPA targets specific TF for degradation DDA 1/DET 1/COP 10 target other proteins Other COPs form part of COP 9 signalosome W/O COPs these TF act in dark
Phytochrome • COPs target specific TF for degradation • W/O COPs they act in dark • In light COP 1 is exported to cytoplasm so TF can act • Tags PHYA by itself on the way out!
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