The Dual Role of Sunlight: Energy and Information

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6 The Dual Role of Sunlight: Energy and Information Lectured by L-F Chien

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Sunlight •Sunlight is a source of energy which drives photosynthesis. •Sunlight is a source of information for plants that drives phenomena such as photomorphogenesis and photoperiodism.

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The principle topics to be covered: •explore the physical nature of light and how light interacts with matter • discuss some of the terminology used in describing light and methods for measuring it • discuss briefly the characteristics of light in the natural environment of plants • reviews the principal pigments and pigment systems found in plants

6.1 The physical nature of light 6.1.1 Light is electromagnetic energy, which exists in two forms •The electromagnetic spectrum (wavelength)

Figure 6.1

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•The electromagnetic spectrum c λv

(Taiz & Zeigher 2006)

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6.1.2 Light can be characterized as a wave phenomenon •Light has properties of both particles and waves. –The wave properties can be characterized in terms of the wavelength and frequency. c v 

–The frequency (n) is the number of wave crests per second. –The wavelength (l) is the distance between crests of the wave. –The speed of light is c.

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Wave nature of light

(electric field) (magnetic field)

Figure 6.1

c v  c E q h hv λ

c, light speed: 3108 m s1 [lambda], wavelength: nanometer (nm) [nu], frequency: Hertz (Hz) or wavenumber (cm1) Eq, energy: Joule (J) or calorie (cal) h,Pl anck’ sconst ant :6. 6261034 J sec

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Quantum energy (Eq) •The energy (Eq) of a particular photon is called a quantum and is related to the wavelength and frequency. c

─red light of 660 nm Eq ( red light) hv h λ 3 10 10 cm  sec -1 (6.626 10 J  sec)( ) 660 nm 10 23 molecules 6.023 -19 -1 301 10 J  ] photon [ N  mol -1 181 kJ  mol 34

─blue light of 435 nm Eq ( blue

hv h light)

c λ

3 10 10 cm  sec -1 (6.626 10 J  sec)( ) 435 nm 10 23 molecules 6.023 -19 -1 301 10 J  ] photon [ N  mol -1 274 kJ  mol 34

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6.1.4 Light energy can interact with matter •Absorbed light can induce photochemical reactions according to the Gotthaus-Draper principle. •A molecule that absorbs light is a pigment.

(Taiz & Zeigher 2006)

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The absorption light by pigment molecules Figure 6.3

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Molecular orbital of H2 H2

(Chang 2000)

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Molecular orbital of CO CO

(Chang 2000)

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T? S?

Molecular electronic state (ES) 1 2( s)

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where s represents electron spin of +½ or ½

(½)

(+½) or

(+½) (½)

ES of Chl =1+2[(+½)+(½)] =1 (Singlet state) 1Chl

S0: singlet ground-state

(+½)

ES of Chl* =1+2[(+½)+(½)] =1 (Singlet state) 1Chl*

S1: singlet first excited-state

(+½) ES of Chl* =1+2[(+½)+(+½)] =3 (Triplet state) 3Chl*

T1: triplet first excited-state

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Light absorption: (i) 1Chl + hv  1Chl* (ii) 1Chl + hv  1Chl*

1Chl*

Blue

ns=10-9

60 Integral area no. of molecules

Thermal deactivation (relax)

1Chl*

Red

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Fluorescence fs=10-15 1Chl

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Fluorescence Phosphorescence

6.1.5 How does one illustrate the efficiency of light absorption ant its physiological effects? •Action spectrum Figure 6.3

leaf photosynthesis

absorption spectrum of Chl a

spectrum, s. spectra, pl.

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6.1.6 Accurate measurement of light is important in photobiology •Three parameters are relevant to the measurement of light. –Light quantity –Light quality, spectral composition, or spectral energy distribution (SED) –Timing and duration of light treatment

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Concepts and units for measuring light

(Taiz & Zeiger 2006)

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Flat and spherical light sensors

Flat

Spherical

(Taiz & Zeiger 2006)

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 Light can be measured omnidirectionally (from all directions) as fluence rate.  Units: (1) energy fluence rate: (a) watts per square meter (W m2) (b) Joules per square meter per second (J m2 s1) [1 W = 1 J s1] (2) photon fluence rate: mol m2 s1

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Light can be measured as energy, and the amount of energy that falls on a flat sensor of known area per unit time is quantified as irradiance. Units: watts per square meter (W m2) PAR: photosynthetically active radiation

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Light can be measured as the number of incident quanta (quantum, s.; quanta, pl.). Unit: (a) moles per square meter per second (mol m2 s1) (b) microEinstein (Einstein, one mole of photons) per square meter per second (E m2 s1) PPFD: photosynthetic photon flux density

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Spectral energy distribution of sunlight (1) Figure 6.5A

紅外光 紫外光

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Spectral energy distribution of sunlight (2) Figure 6.5B

白熾光

6.2 The natural radiation environment

(A) Radiation from the sun (B) Earth reradiates infrared (heat) back (C) Infrared radiation either reflected back to earth or absorbed by atmospheric gases

Figure 6.6

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6.3 Photoreceptors absorb light for use in a physiological process •Photoreceptors are pigment molecules that absorb and process light into a form that can be used by the plant. •Photoreceptors can be chromoproteins, consisting of a pigment (chromophore) and a catalytic protein (apoprotein).

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Photopigments: (1) Chlorophylls:

chlorophylls a, b, c, d bacteriochlorophyll a, b

(2) Phycobilins:

phycoerythrobilin (PE) phycocyanobilin (PC) allophycocyanin (AP)

(3) Carotenoids:

(about 700 compounds) -carotene lutein xanthophyll cycle pigments

6.3.1 Chlorophylls are primarily responsible for harvesting light energy for photosynthesis •Chlorophylls: (a)chlorophylls: chlorophylls a, b, c, d, (b) bacteriochlorophylls: bacteriochlorophyll a, b

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Chemical structure of chlorophylls

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(chlorophyll, Greek: chloros = green + phyllon =leaf)

Figure 6.7 Porphyrin

Phytol (5-carbon isoprene, lipid-soluble)

Phyeophytin: no Mg 2+

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Chlorophylls

Chlorophyll a, c

Chlorophyll d

Chlorophyll b

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Absorption spectra of Chl a and Chl b

Figure 6.8

6.3.2 Phycobilins serve as accessory lightharvesting pigments in red algae and cyanobacteria •Phycobilins: (a) phycobilins: ─phyco, algal origin ─pr ot ei ns

(i) phycoerythrobilin (PE) (ii) phycocyanobilin (PC) (iii) allophycocyanin (AP) (b) phycobilinsomes: ─l ar gemacr omol ecul arcompl ex es

(c) phytochrome: ─Pr ,Pf r

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Open-chain tetrapyrrole chromophore of phycocyanin Figure 6.9

(water-soluble)

Absorption spectra of phycocyanin and phycoerythrin Figure 6.9 phycoerythrin

phycocyanin

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6.3.3 Carotenoids account for the autumn colors •Carotenoids: ─about700compounds ─C40 terpenoids ─accessory pigments ─in chloroplast membrane or chromoplasts

(a) carotenes: a, , g-carotene lutein (b) xanthophyll cycle pigments

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Carotenes and xanthophylls Figure 6.11

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Absorption spectra of a-carotene and -carotene

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b-carotene

a-carotene

Figure 6.12

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Absorption spectra of some photosynthetic pigments Chlb

Chla

PE

BChla

Chla

BChla

-Car Chlb

(Taiz & Zeiger 2006)

6.3.4 Cryptochorome and photoropin are photoreceptors sensitive to blue light and UV-A radiation •Cryptochrome [隱花色素] –Gr., hidden colour

–blue light/UV-A photoreceptor –Arabidopsis CRY1 and CRY 2 genes –CRY1 protein, a flavoprotein containing FAD and pterin, 75kDa –using blue light to stimulate repair of UV-induced damage to microbial DNA

–regulate germination, photoperiodism

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6.3.4 Cryptochorome and photoropin are photoreceptors sensitive to blue light and UV-A radiation •Phototropins (PHOT) –blue light photoreceptor –Arabidopsis PHOT1 and PHOT2 genes –PHOT1 protein, a flavoprotein containing 2 FMN, 120 kDa –In optimizing photosynthetic efficiency

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6.3.6 Flavonoids provide the myriad flower colors and act as a natural sunscreen Phenylpropane

Riboflavin

C6-C3-C6

Figure 6.14

Pterin Figure 6.13

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Absorption spectrum of the anthocyanin, pelargonin

Figure 6.15 pelargonin (天竺葵花紅素)

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Assay Chapter Review: 1, 2, 3, 6 (p-107) Deadline: next Thursday Delay: 100n2