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Chapter 23.

Tropisms and
Nastic
Movement:
Orienting
Plants in
Space

BY: REENA CARLOS

TROPISM
the

turning of all or
part of an organism in
a particular direction
in response to an
external stimulus.
These tropic
responses may either
be positive or
negative.

Phototropism
is

often defined
as a response to
unilateral light
but can also
occur in plants
that receive light
from all sides.

Phototropism
The

magnitude of a light gradient


across an organ such as a coleoptile is
dependent on optical properties of the
tissue as well as differences in incident
light.

coleoptiles
Organs

such as
coleoptiles appear to
function as light
pipes. This means
that light applied to
the tip, for example,
will be transmitted
through the coleoptile
to cells further down
the organ.

Phototropism: A Bluelight
Response
1930s
Probability of the chromoprotein
being a photoreceptor.
1940s
Suggestions that photoreceptors
are Flavin molecules.
Phototropina flavo-protein
pigment
responsible
for

Phototropism orients a
plant for optimal
The singular
impact of phototropism is that
photosynthesis

it
orients growth and leaf angle toward incident
light in order to maximize light interception for
photosynthesis.

Plants

also use blue light to control stomatal


opening and facilitate gas exchange as well as
to relocate chloroplasts within the cell.

Plants also
use blue
light to
control the
high-light
avoidance
response of
chloroplast
s in the
mesophyll
cells.

FLUENCE
CURVES
Fluence

RESPONSE

response curves are generally


obtained by monitoring the response
of the organ to different total amounts
of light (fluence), usually by using a
single fluence rate but varying the
presentation time.

Initial rise
toin
Another
rise
first peak;
response to light;
restricted to
extends
more
the apex of
toward
the basal
coleoptiles
region of the
coleoptile

Reduced positive
response: bending
away from the light

Bunsen-Roscoe
Reciprocity
Law
States that the product of a photochemical
reaction is determined by the total amount of
energy presented, regardless of fluence rate or
presentation time.
The

complexities of second positive curvature


are due to subsequent events in the signal
transduction chain.

THE PHOTOTROPIC
RESPONSE IS ATTRIBUTED
TO A LATERAL
REDISTRIBUTION OF
DIFFUSIBLE AUXIN

Cholodny-Went
hypothesis
Formulated in the late 1920s as a
combination of the ideas of N. Cholodny
and F. W. Went in an attempt to explain
phototropism
States that unilateral illumination induces
a lateral redistribution of endogenous
auxin near the apex of the organ.

Systematic Reevaluation
of the Cholodny-Went
hypothesis by W. R.
Briggs

REGULATION BY A FAMILY
OF BLUE-SENSITIVE
FLAVOPROTEINS

Phototropin
Photoreceptor for phototropism.
120kDa

plasma membrane protein is


phosphorylated by blue light and contained
in the same area most responsive to light.

Extensive

characterization proved that the


protein is a kinase that autophosphorylates
in blue light.

NPH 1 or the Non-phototropic


hypocotyl 1
Encodes

the 120kDa protein,


Was a mutant isolated in Arabidopsis with
failure to exhibit phototropism and lacking
the 120kDa protein.
The NPH1 holoprotein was subsequently
renamed phototropin 1 (phot1) because of
its functional role in phototropism.

Phot 1 has two flavin


mononucleotide (FMN)
chromophores
Phot 1 photosensory domains
has distinctive domains
called LOV (light, oxygen and
voltage)
2 LOV domains are the
sites that bind to FMN to
make Phototropin
responsive to light

Phot 1
First positive
curvature
Second positive
curvature
Stomatal
opening

Phot 2

Second positive
curvature
Stomatal
opening
Avoidance movt
of chloroplasts
under high light
intensities

PHOTOTROPIN ACTIVITY AND


SIGNAL CHAIN
1. Autophosphorylation

of phototropin plays
a significant role in the phototropic
response, probably by initiating a
phosphorylation cascade.

2. Phototropins

regulation.

may be involved in gene

GRAVITROPISM
is the directional
growth of a plant
organ in response
to a gravitational
field where roots
grow downwards
and shoots grow
upwards.

Gravity
Omnipresent

and nonvarying
Constant magnitude
can be detected only by the movement
of some structure or structures within
the cells

GRAVITROPISM IS MORE
THAN SIMPLY UP AND
DOWN
Orthogravitropic

diagravitropic

root and shoot of


the primary plant
axis align parallel
with the direction
of
gravitational
pull.
Organs which grow at

plagiogravitropic

agravitropic

Organs oriented
at some
intermediate
angle (between
0 and 90 to the
vertical)
Organs that

Gravitational
Stimulation
Gravitational stimulation (stimulus quantity or dose)
is the product of the intensity of the stimulus and
the time over which the stimulus is applied:
d

=ta

where a is the acceleration of mass due to gravity


(in g)
t is the time (in seconds) over which the stimulus is
applied.

Threshold Dose
minimum

dose required to induce


gravitropic curvature.

Threshold

dose will vary depending on


the organism or experimental conditions.

Presentation Time
Is

the minimum duration of stimulation required to


induce a curvature that is just detectable.

Reaction Time
is

the interval between the presentation of the


stimulus and the actual development of curvature.

Threshold Intensity.
Is

the minimum stimulus intensity


required to induce a response.

Threshold

intensities have been


determined for a variety of plant organs
under different experimental conditions.

occurring between
1 and 10 seconds
following
occurs
within
perhaps
one
second
reorientation,
occurs between 10
of orienting a root
off the
vertical
involves
the
seconds and 10
and involves biophysical
conversion of the
minutes of
mechanisms
(e.g.,
pressure)
for
biophysical
single
reorientation and
sensing
the
direction
of
to
a
biochemical
involves a
gravitational
pull.
signal.
redistribution of auxin
within the root tip.

4 Phases of Root
Gravitropism

Columella
the

gravity detecting
portion of the root cap.

consists

of cells rich in
dense
amyloplasts:
organelles that are filled
with starch grains.

Gravity Perception
Starch-Statolith

hypothesis
G. Haberlandt
and E.
Nemec, 1900
Statolithsstarch grain

Amyoplasts
Are

a group of statoliths (starch grain)


contained within a membrane.

In

vertically oriented roots the


amyloplasts reside at the lower end of
each columella cell, toward the root tip.

Amyoplast as the
gravity-sensing
1. Gravitropism
is generally absent in plant
mechanism
species that have no starch grains or
amyloplasts.

2.

There is a strong correlation between the rate


of starch sedimentation and presentation time.

Amyoplast as the
gravity-sensing
3. Loss ofmechanism
starch by hormone treatment or
mutation is accompanied by a loss of
graviresponse.
4. Amyloplasts can be displaced by a highgradient magnetic field in place of gravity.

Redistribution of
Auxin
horizontal orientation of the shoot or roots
induces a lateral translocation of auxin toward the
lower side of the organ.
Auxin

redistribution would bias the growth rate in


favor of the lower side such that negatively
gravitropic organs (e.g., coleoptiles and shoots)
would turn upward.

Redistribution of
Auxin
the

higher auxin
content on the lower
side of the root
inhibits elongation
relative to the upper
side and the root
curves downward.

Changes in membrane
potential
Roots

at a horizontal position, the the


columella cells on the lower side of the
root depolarize and those on the upper
side hyperpolarize.
stretch-activated ion channels would be
responsible for the observed changes in
membrane potential in the columella cells,
which in turn would lead to the asymmetric

pH changes
it

has been shown that pH of the root cap


apoplast decreases from pH 5.5 to 4.5
within 2 minutes of gravistimulation.

These

pH changes in the root cap


precede auxin-related pH changes in the
elongation zone by about 10 minutes.

Calciu
m

PIN proteins
PIN protein
1

Location
Importance
Apical end of responsible
cells in the for delivering
stele
the auxin
stream to the
root apex

PIN proteins
PIN protein
2

Location
root cap
Epidermal
and cortical
cell

Importance
mediates the
basipetal
stream of
auxin toward
the cell
elongation
zone.

PIN proteins
PIN protein
3

Location
Lateral wall
of the
columella
cells

Importance
diverts the
flow of
auxin
laterally, or
centrifugally,
toward the
peripheral

NASTIC MOVEMENTS
Nastic

movements are plant movements that


occur in response to environmental stimuli but
unlike tropic movements, the direction of the
response is not dependent on the direction
of the stimulus.

Some

of the most spectacular plant movements


are nastic movements.

Epinasty
is

the downward
bending of an organ,
commonly petioles
and leaves whose
tips are inclined
toward the ground.

Hyponasty
Reverse

response of epinasty,
which is said to be induced by
gibberellins.

Theupwardbendingofaleafo

rotherplantpart,resultingfro
mgreatergrowthofthelower
sidethanoftheupperside.

Thermonasty
Is

a nastic movement
that is associated with
changes in temperature
thermonastic movements
are permanent and result
from alternating
differential growth on the
two surfaces of the
petals.

Turgor Movements
1.
2.
3.

leisurely rhythmic leaf movements in


nyctinastic plants,
very rapid seismonastic movements in a
limited number of species, and
thigmonastic or thigmotropic curling of
threadlike appendages in climbing plants
and vines.

All

NYCTINASTIC
MOVEMENTS

nyctinastic
responses
night
closure
depend on reversible
turgor changes in the
pulvinus.
The extensor region is
formed by motor cells
that lose turgor during
the bending movement,

ION FLUXES AND


RESULTING OSMOTIC
motor cell
volume changes are due to
RESPONSES

osmotic water uptake (or loss) as a result of


ion accumulation (or loss) across the cell
membrane.

Swollen

extensor cells are characterized by


high protoplasmic K+ and low apoplastic K+.

Role of Potassium
K+

exchange across the plasma membrane


occurs through K+ channels and that these
channels can be regulated by changing the
membrane polarity.

Depolarization

of the membrane opens the


channels and allows K+ to move out of the cell
down its electrochemical gradient.

SEISMONASTY
a

nastic movement in
response to a mechanical
shock rapid folding of the
leaflets of the sensitive plant
due to changes in turgor
pressure caused by vibration

three essential
characteristics of the
seismonastic
1. The
rapidity of the response
response.
It follows the all-or-none principle, which
means that there is no obvious relationship
between the intensity of the stimulus and
the extent of the response.
3. Excitation is propagated from the place of
stimulation.
2.

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