Light Control

of Plant Development
Mochammad Roviq

Photomorphogenesis is the plants

response to light
An obviously integral element of normal
development in autotrophic organisms like plants
We will be looking at the role of phytochrome in
perceiving and translating light signals into changes
in plant shape and function
Light dictates a plants metabolism
Quality (spectrum)
Amount (flux) more light, more photosynthesis, higher
growth rate
Timing (diurnal patterns) important in development

A light perception system is critical for the

meaningful regulation of plant metabolism
Plants are sessile: must deal with environmental
limitations to survive
Plants use photoreceptors to detect
environmental light changes
These act as initiators of signal transduction
cascades that ultimately direct plants response
to light level
Most light responses are controlled by
chromoproteins
They contain a chromophore absorbs light
They are an apoprotein undergoes conformational
change (initiates signal transduction cascade)

THE PHOTOCHEMICAL AND BIOCHEMICAL

PROPERTIES OF PHYTOCHROME

Phytochrome, a blue protein pigment with a molecular mass of about 125

kDa (kilodaltons)
The first clues regarding the role of phytochrome in plant development
came from studies that began in the 1930s on red lightinduced
morphogenic responses, especially seed germination.

Phytochrome was the discovery

that the effects of red light (650
680 nm) on morphogenesis could
be reversed by a subsequent
irradiation with light of longer
wavelengths (710740 nm), called
far-red light.
The germination of lettuce seeds
is stimulated by red light and
inhibited by far-red light.
But the real breakthrough was
made many years later when
lettuce seeds were exposed to
alternating treatments of red and
far-red light. Nearly 100% of the
seeds that received red light as
the final treatment germinated; in
seeds that received far-red light as
the final treatment, however,
germination was strongly inhibited

Phytochrome Is a Dimer Composed of

Two Polypeptides

Native phytochrome is a soluble protein

with a molecular mass of about 250 kDa.
It occurs as a dimer made up of two
equivalent subunits.
Each subunit consists of two components:
a light-absorbing pigment molecule called
the chromophore, and a polypeptide chain
called the apoprotein.
The apoprotein monomer has a molecular
mass of about 125 kDa.
Together, the apoprotein and its
chromophore make up the holoprotein. In
higher plants the chromophore of
phytochrome is a linear tetrapyrrole termed
phytochromobilin.
There is only one chromophore per
monomer of apoprotein, and it is attached
to the protein through a thioether linkage to
a cysteine residue

Phytochrome is the primary plant

chromoprotein
Phytochrome exists in 2 stable states that
absorb light at different wavelengths
Phytochromes
conformation is
photoreversible:
light changes the
proteins shape!

(665 nm)

Phytochrome
protein synthesis

Pr

Pfr

Fig. 17.3
(730 nm)

There are multiple phytochromes in plants

different proteins expressed at different times
during development
These collectively form the photochromic
receptor system

Only the Pfr form of phytochrome is

involved in signal transduction
Pr is the physiologically inactive form
Exposed to red light becomes Pfr active!

You should see this in the lettuce seed experiment

How light interconverts Pr Pfr not clear

Likely affects protein folding and dimerization

Apoprotein
Conversion between Pr and Pfr involves
rotation between rings of the
chromophore

Chromophore (phytochromobilin)

Phytochrome Can Interconvert between Pr

and Pfr Forms

In dark-grown or etiolated plants,

phytochrome is present in a red
lightabsorbing form, referred to
as Pr because it is synthesized
in this form. Pr, which to the
human eye is blue, is converted
by red light to a far-red light
absorbing form called Pfr, which
is blue-green. Pfr, in turn, can be
converted back to Pr by far-red
light.
When Pr molecules are exposed
to red light, most of them absorb
it and are converted to Pfr, but
some of the Pfr also absorbs the
red light and is converted back to
Pr because both Pr and Pfr
absorb red light

Plant neighbors?

Red
absorbed
by other
plants.

Far red reflected

from other plants.

Far red enriched = neighbors

Under other plants?

Red
absorbed
by other
plants.

Far red
reflected
from other
plants or
transmitted.

Far red enriched = understory

Phytochrome has 2 forms

Red-absorbing phytochrome
Pr

Far red absorbing phytochrome

Pfr
Pr
Pfr
Interconverted
Two forms of the same compound
Total amount same

In red light

Prfr

Pr absorbs red light,

changes to Pfr form.

Pfr

Pfr doesnt absorb red

light, stays the same.

In far red light

Pr

Pr doesnt absorb far red

light, stays the same.

Prfr

Pfr absorbs far red light,

changes to Pr form.

Sunlight

Mostly red
A little far red

In sunlight

Pfrr
Pfrr

Pfr
Pfrr

Pfr
Pfr

Pfrr

Pfr

Pr

Pfr

Pfrr
Pfrr

Prfr

Pfr
Pfr

Pfr
Pfrr

Prfr
Pfrr

Pfr

In sunlight most P gets converted to Pfr form.

Start of night
Most P in Pfr form.
Pfrr
Pfrr

Pfr
Pfrr

Pfr
Pfr

Pfrr

Pfr

Pr

Pfr

Pfrr
Pfrr

Prfr

Pfr
Pfr

Pfr
Pfrr

Prfr
Pfrr

Pfr

In the dark
Pfr form changes gradually to Pr form.
Pfrr

Pfr

Prfr
Pfr
Pfr
Pfrr

Prfr

Pfrr

Pr

Pfr

Pfrr
Pfrr

Prfr

Pfr
Pfr

Prfr
Pfrr

Prfr
Pfrr

Prfr

After a short night

Pfr still left.
Pfrr

Pfr

Prfr
Pfr
Pfr
Pfrr

Prfr

Pfrr

Pr

Pfr

Pfrr
Pfrr

Prfr

Pfr
Pfr

Prfr
Pfrr

Prfr
Pfrr

Prfr

LDP = SNP
Needs short night
Needs Pfr still present at end of night
Pfr promotes flowering for LDPs

Later in the night

More Pfr changes to Pr.
Pfrr
Prfr

Pfr

Pfr

Pfr

Pfrr
Pfr

Pfrr

Prfr

Pfrr

Prfr

Pfrr

Pfr
Pfr

Pr
Prfr

Pfrr

Prfr
Pfrr

Prfr

After a long night

All the Pfr is gone.
Prfr

Prfr

Prfr
Prfr
Prfr
Prfr

Prfr

Prfr

Pr

Prfr

Prfr
Prfr

Prfr

Prfr
Prfr

Prfr
Prfr

Prfr
Prfr

Prfr

Day dawns

Pfrr
Pfrr

Pfr
Pfrr

Pfr
Pfr

Pfrr

Pfr

Pr

Pfr

Pfrr
Pfrr

Prfr

Pfr
Pfr

Pfr
Pfrr

Prfr
Pfrr

Pfr

Most P gets converted to Pfr form again.

LOCALIZATION OF PHYTOCHROME IN
TISSUES AND CELLS
Phytochrome is most
heavily concentrated in
the regions where
dramatic developmental
changes are occurring:
the apical meristems of
the epicotyl and root.
Shown here is the
distribution of
phytochrome in an
etiolated pea seedling, as
measured
spectrophotometrically

There are 3 categories of plant

responses to phytochrome
The categories are light-level dependent
1. VLFRs (very low fluence responses)

< 10-3 mmol photons/m2 (converts 0.01% of

phytochrome)

2. HIR (high irradiance responses)

>1000 mmol photons/m2, continuous irradiation,

dependent on actual fluence

3. LFRs (low fluence responses)

1-1000 mmol photons/m2, FR light reversible

We will concentrate on LFRs only

CHARACTERISTICS OF PHYTOCHROMEINDUCED WHOLE-PLANT RESPONSES

Phytochrome Responses Can

Be Distinguished by the
Amount of Light Required
The amount of light is
referred to as the fluence,
which is defined as the
number of photons
impinging on a unit
surface area
responses fall into three
major categories based on
the amount of light
required: very-low-fluence
responses (VLFRs), lowfluence responses (LFRs),
and high-irradiance
responses (HIRs)

Very-Low-Fluence Responses
Some phytochrome responses can be initiated by fluences as low as
0.0001 mol m2 (one-tenth of the amount of light emitted from a
firefly in a single flash), and they saturate (i.e., reach a maximum) at
about 0.05 mol m2.
For example, in dark-grown oat seedlings, red light can stimulate the
growth of the coleoptile and inhibit the growth of the mesocotyl (the
elongated axis between the coleoptile and the root) at such low
fluences.
Arabidopsis seeds can be induced to germinate with red light in the
range of 0.001 to 0.1 mol m2. These remarkable effects of
vanishingly low levels of illumination are called very-low-fluence
responses (VLFRs).

Low-Fluence Responses Are

Photoreversible

Another set of phytochrome

responses cannot be initiated until
the fluence reaches 1.0 mol m2,
and they are saturated at 1000
mol m2.
These responses are referred to as
low-fluence responses (LFRs), and
they include most of the red/far-red
photoreversible responses, such as
the promotion of lettuce seed
germination and the regulation of
leaf movements,
The LFR action spectrum for
Arabidopsis seed germination is
shown in Figure
LFR spectra include a main peak
for stimulation in the red region
(660 nm), and a major peak for
inhibition in the far-red region (720
nm).

Low fluence responses are the most

studied changes induced by
phytochrome
These regulate important plant growth and
development responses including
De-etiolation
the greening of etiolated seedlings
Important as seedlings emerge from the soil and
begin to become autotrophic

Seed germination
breaking of seed coat, start of active metabolism in
new plant
Light can promote or inhibit germination, depending
on the species

De-etiolation is regulated by phytochrome

Etiolated typical seedling response
when dark-grown
Long hypocotyl (stem below the cotyledons)
Not green no chlorophyll, no chloroplasts
Little leaf development or unfolding

Expose to light (de-etiolated):

Hypocotyl stops growing
Chlorophyll and chloroplasts develop
Leaves unfold

De-etiolated
Etiolated

Grown
under
normal
light

Therefore plants need light to

developmentally progress from
heterotrophic autotrophic organism
Fig 17.9

Seeds can be one of two types

Positively photoblastic: germination in light (need high Pfr/Pr)

Negatively photoblastic: germination in light (need low Pfr/Pr)

High-Irradiance Responses Are Proportional to the

Irradiance and the Duration

The reason that these responses are called high-irradiance responses

rather than high-fluence responses is that they are proportional to irradiance
(loosely speaking, the brightness of the light) rather than to fluence.
HIRs saturate at much higher fluences than LFRsat least 100 times
higher.
Because neither continuous exposure to dim light nor transient exposure to
bright light can induce HIRs, HIRs do not obey the law of reciprocity.

The time scales for phytochromemediated effects vary

Usually long (h d) growth effects

Germination
De-etiolation
Circadian clock (daily light effects)

Some short (s min)

Transmembrane potential
Red light depolarizes membranes!
FR repolarizes
Due to ion movements via specific membrane transporters
(electrically gated channels)

The main form of phytochrome (A) has

characteristics suggesting that it is the primary plant
light sensor
Degrades in light
Other, more sensitive forms of phytochrome may monitor
light quality (e.g., phy B)

Phytochrome Enables Plants to Adapt to

Changing Light Conditions
The ratio of red light (R) to far-red
light (FR) varies remarkably in
different environments
Simulated canopy shading (high
levels of far-red light) induced these
plants to allocate more of their
resources to growing taller. This
correlation did not hold for shade
plants, which normally grow in a
shaded environment.
Shade plants showed little or no
reduction in their stem extension
rate as they were exposed to higher
R/FR values

Shade avoidance in
vegetative plants

dense stand

50

height (cm)

Light environment in
leaf canopies

open stand

60

40
30
20
10
0
0

Lysimachia vulgaris
different canopy densities

20

40
60
irradiance (%)

80

100

0.0

0.2

0.4

0.6
R:FR

0.8

1.0

1.2

R:FR effects independent of PAR

Chenopodium a shade avoiding species

Adapted from H. Smith

Phytochrome Regulates the

Sleep Movements of Leaves
The sleep
movements of leaves,
referred to as
nyctinasty,
(circadian rhythm that
is regulated by light)
In nyctinasty, leaves
and/or leaflets extend
horizontally (open) to
face the light during
the day and fold
together vertically
(close) at night

Circadian rhythm

Circadian rhythm
in the diurnal
movements of
Albizia leaves.
The leaves are
elevated in the
morning and
lowered in the
evening. In
parallel with the
raising and
lowering of the
leaves, the
leaflets open and
close.
The rhythm
persists at a
lower amplitude
for a limited time
in total darkness.

Light affects movement

Light also directly affects movement: Blue light
stimulates closed leaflets to open, and red light
followed by darkness causes open leaflets to
close.
The leaflets begin to close within 5 minutes after
being transferred to darkness, and closure is
complete in 30 minutes.
Because the effect of red light can be canceled
by far-red light, phytochrome regulates leaflet
closure.

CIRCADIAN RHYTHMS
Organisms are normally subjected to daily cycles of light
and darkness, and both plants and animals often exhibit
rhythmic behavior in association with these changes.
Examples of such rhythms include leaf and petal
movements (day and night positions), stomatal opening
and closing, growth and sporulation patterns in fungi
(e.g., Pilobolus and Neurospora),
When organisms are transferred from daily lightdark
cycles to continuous darkness (or continuous dim light),
many of these rhythms continue to be expressed, at
least for several days

Control rhythms
Like animals, plants have both exogenous
and endogenous factors that control
rhythms.
Circadian rhythms shown by plants
include: a) opening and closing of stomata
b) sleep movements, for example,
leaves of beans droop at night
Circaanual rhythms include: seed
germination, flowering and leaf fall. These
rhythms are controlled by daylength and
temperature.

 Under such uniform conditions the period of the

rhythm is then close to 24 hours, and
consequently the term circadian rhythm is
applied.
Because they continue in a constant light or dark
environment, these circadian rhythms cannot be
direct responses to the presence or absence of
light but must be based on an internal
pacemaker, often called an endogenous
oscillator.

Circadian Rhythms Exhibit

Characteristic Features
Circadian rhythms arise from cyclic phenomena that are defined by
three parameters :
Period, the time between comparable points in the repeating cycle.
Typically the period is measured as the time between consecutive
maxima (peaks) or minima (troughs)

Phase and amplitude

Phase, any point in the cycle that is recognizable by its relationship
to the rest of the cycle. The most obvious phase points are the peak
and trough positions.
Amplitude, usually considered to be the distance between peak and
trough. The amplitude of a biological rhythm can often vary while the
period remains unchanged

 Under natural conditions, the endogenous oscillator is entrained

(synchronized) to a true 24-hour period by environmental signals,
the most important of which are the light-to-dark transition at dusk
and the dark-to-light transition at dawn

Noon

Midnight

Subjective day and night

When an organism is entrained to a cycle of 12
hours light and 12 hours dark and then allowed
to free-run in darkness, the phase of the rhythm
that coincides with the light period of the
previous entraining cycle is called the
subjective day, and the phase that coincides
with the dark period is called the subjective
night.
If a light pulse is given during the first few hours
of the subjective night, the rhythm is delayed;
the organism interprets the light pulse as the
end of the previous day

PHOTOPERIODISM:
MONITORING DAY LENGTH
The circadian clock enables
organisms to determine the time of
day at which a particular molecular or
biochemical event occurs.
Photoperiodism, or the ability of an
organism to detect day length, makes
it possible for an event to occur at a
particular time of year, thus allowing
for a seasonal response.
Circadian rhythms and
photoperiodism have the common
property of responding to cycles of
light and darkness.

PHOTOPERIODISM
RESPONSES

The two main photoperiodic response categories are short-day plants and
long-day plants:
Short-day plants (SDPs) flower only in short days (qualitative SDPs),
or their flowering is accelerated by short days (quantitative SDPs)
Long-day plants (LDPs) flower only in long days (qualitative LDPs), or
their flowering is accelerated by long days (quantitative LDPs).

The essential distinction between long-day and short day plants is that
flowering in LDPs is promoted only when the day length exceeds a certain
duration, called the critical day length, in every 24-hour cycle, whereas
promotion of flowering in SDPs requires a day length that is less than the
critical day length.

 Long-day plants can effectively

measure the lengthening days of
spring or early summer and delay
flowering until the critical day length
is reached. Many varieties of wheat
(Triticum aestivum) behave in this
way.
SDPs often flower in fall, when the
days shorten below the critical day
length, as in many varieties of
Chrysanthemum morifolium.
However, day length alone is an
ambiguous signal because it cannot
distinguish between spring and fall.

The photoperiodic regulation of flowering.

Effects on SDPs and LDPs

 Other plants avoid seasonal ambiguity by distinguish ing between

shortening and lengthening days. Such dualday length plants fall
into two categories:
Long-short-day plants (LSDPs) flower only after a sequence of
long days followed by short days. LSDPs, such as Bryophyllum,
Kalanchoe, and Cestrum nocturnum (night-blooming jasmine),
flower in the late summer and fall, when the days are shortening.
Short-long-day plants (SLDPs) flower only after a sequence of
short days followed by long days. SLDPs, such as Trifolium
repens (white clover), Campanula medium (Canterbury bells),
and Echeveria harmsii (echeveria), flower in the early spring in
response to lengthening days
Finally, species that flower under any photoperiodic condition are
referred to as day-neutral plants. Day-neutral plants (DNPs)
are insensitive to day length. Flowering in DNPs is typically
under autonomous regulationthat is, internal developmental
control

The Circadian Clock Is Involved in Photoperiodic

Timekeeping

Rhythmic flowering in response

to night breaks. In this
experiment, the SDP soybean
(Glycine max) received cycles of
an 8-hour light period followed
by a 64-hour dark period. A 4hour night break was given at
various times during the long
inductive dark period. The
flowering response, plotted as
the percentage of the maximum,
was then plotted for each night
break given. Note that a night
break given at 26 hours induced
maximum flowering, while no
flowering was obtained when the
night break was given at 40
hours.

Moreover, this experiment demonstrates that the sensitivity to

the effect of the night break shows a circadian rhythm. These
data support a model in which flowering in SDPs is induced
only when dawn (or a night break) occurs after the completion
of the light-sensitive phase. In LDPs the light break must
coincide with the light sensitive phase for flowering to occur.
(Data from Coulter and Hamner 1964.)

Phytochrome Is the Primary Photoreceptor in

Photoperiodism

Night-break experiments are

well suited for studying the
nature of the photoreceptors
involved in the reception of
light signals during the
photoperiodic response. The
inhibition of flowering in
SDPs by night breaks was
one of the first physiological
processes shown to be
under the control of
phytochrome

Blue-Light Responses:
Stomatal Movements
and Morphogenesis

Responses
Some blue-light responses are for example, chloroplast
movement within cells in response to incident photon
fluxes, and sun tracking by leaves.
As with the family of the phytochrome responses, there
are numerous plant responses to blue light.
Besides phototropism, they include inhibition of
hypocotyl elongation, stimulation of chlorophyll and
carotenoid synthesis, activation of gene expression,
stomatal movements, phototaxis (the movement of
motile unicellular organisms such as algae and bacteria
toward or away from light), enhancement of respiration,
and anion uptake in algae (Senger, 1984).

three-finger
Action spectra for blue
lightstimulated
phototropism,stomatal
movements, inhibition of
hypocotyl elongation, and
other key blue-light
responses share a
characteristic three-finger
fine structure in the 400 to
500 nm region (Figure).

THE PHOTOPHYSIOLOGY OF
BLUE-LIGHT RESPONSES
Directional growth toward (or
in special circumstances
away from) the light, is called
phototropism. It can be
observed in fungi, ferns, and
higher plants.
Phototropism is a
photomorphogenetic
response that is particularly
dramatic in dark-grown
seedlings of both monocots
and dicots

 Blue Light Rapidly Inhibits

Stem Elongation
Blue Light Regulates Gene
Expression : Most of the
studies on light-activated genes
show sensitivity to both blue and
red light, as well as red/far-red
reversibility, implicating both
phytochrome and specific bluelight responses.

 Blue Light Stimulates Stomatal Opening

The stomatal response to blue light is rapid and
reversible, and it is localized in a single cell type, the
guard cell.
The stomatal response to blue light regulates
stomatal movements throughout the life of the plant.
This is unlike phototropism or hypocotyl
elongation,which are functionally important at early
stages of development.
The signal transduction cascade that links the
perception of blue light with the opening of stomata is
understood in considerable detail.

Blue-light under red-light

background

The response of stomata to blue light

under a red-light background. Stomata
from detached epidermis of Commelina
communis (common dayflower) were
treated with saturating photon fluxes of
red light (red trace).
In a parallel treatment, stomata
illuminated with red light were also
illuminated with blue light, as indicated by
the arrow (blue trace).
The increase in stomatal opening above
the level reached in the presence of
saturating red light indicates that a
different photoreceptor system, stimulated
by blue light, is mediating the additional
increases in opening. (From Schwartz and
Zeiger 1984

Gerakan Tanaman
Gerakan tanaman dapat dilaksanakan dengan 2 cara:
Satu sisi organ tumbuh lebih cepat daripada sisi yang lain terjadi
pada bagian tanaman yang sedang tumbuh
Perubahan turgor dari sel-sel tertentu yang menyebabkan
perubahan posisi organ yang bereaksi Gerakan tidur daun
Samanea saman & Albizia julibrissin
Secara konvensional gerakan tanaman dapat dibagi menjadi 2
golongan berdasarkan arah gerakan dan arah dari mana
rangsangan datang:
TROPISME Arah gerakan ditentukan oleh arah asal
rangsangan
NASTI Arah gerakan tidak ada hubungannya dengan arah
rangsangan, melainkan tergantung pada struktur organ yang
bereaksi

Tropisme

Geotropisme / Gravitropism diinduksi oleh gravitasi:

Geotropisme positip pertumbuhan akar primer dan pertumbuhan
tangkai bunga kacang tanah (Arachis hypogea) setelah penyerbukan
Geotropisme positip pertumbuhan batang
Plagiogeotropism akar sekunder dan cabang lateral
Diageotropism mendatar : Rhizome
Phototropism diinduksi oleh cahaya:
Phototropism positip pertumbuhan batang
Phototropism negatip pertumbuhan akar
Diaphototropism sikap daun mendatar / tegak lurus cahaya
Chemotropism diinduksi oleh bahan kimia
Hydrotropism diinduksi oleh air akar respon terdap aplikasi air
Thigmotropism diinduksi oleh singgungan / sentuhan pertumbuhan
sulur

Nasti
Nictinasti Gerakan tidur daun saat gelap dikontrol oleh
interaksi antara lingkungan dan phytochrome
Thigmonasti diinduksi oleh sentuhan Mimosa sp.
Epinasti / Hyponasti respon pertumbuhan sebagian
Daun tumbuh melengkung ke atas karena bagian bawah tumbuh
lebih cepat Epinasti
Daun tumbuh melengkung ke bawah karena bagian atas tumbuh
lebih cepat Hyponasti
Hydronasti / Hygronasti daun melipat /menggulung yang terjadi
karena adanya cekaman air untuk mengurangi transpirasi
Gerakan ini terjadi karena hilangnya turgor sel-sel bulliform (sel
motor yang berdinding tipis, tidak mempunyai cuticula sehingga
hilangnya air transpirasi lebih cepat daripada sel-sel epidermis)
Daun melipat

 TERIMA KASIH