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Plant Responses to Internal and

External Signals

The Discovery of Plant


Hormones
Any growth response
That results in curvatures of whole plant organs
toward or away from a stimulus is called a tropism
Is often caused by hormones

Charles Darwin and his son Francis


Conducted some of the earliest experiments on
phototropism, a plants response to light, in the late
19th century

EXPERIMENT In 1880, Charles Darwin and his son Francis designed an experiment to determine
what part of the coleoptile senses light. In 1913, Peter Boysen-Jensen conducted an experiment to
determine how the signal for phototropism is transmitted.

RESULTS
Control

Boysen-Jensen (1913)

Darwin and Darwin (1880)


Shaded
side of
coleoptile

Light

Light

Light

Illuminated
side of
coleoptile

Tip
removed

Tip covered
by opaque
cap

Tip
covered
by transparent
cap

Base covered
by opaque
shield

Tip separated
by gelatin
block

Tip separated
by mica

CONCLUSION In the Darwins experiment, a phototropic response occurred only when light could
reach the tip of coleoptile. Therefore, they concluded that only the tip senses light. Boysen-Jensen
observed that a phototropic response occurred if the tip was separated by a permeable barrier (gelatin)
but not if separated by an impermeable solid barrier (a mineral called mica). These results suggested
that the signal is a light-activated mobile chemical.

Figure 39.5

EXPERIMENT

In 1926, Frits Wents experiment identified how a growth-promoting chemical


causes a coleoptile to grow toward light. He placed coleoptiles in the dark and removed their tips,
putting some tips on agar blocks that he predicted would absorb the chemical. On a control
coleoptile, he placed a block that lacked the chemical. On others,
he placed blocks containing the chemical, either centered on top of the coleoptile to distribute the
chemical evenly or offset to increase the concentration on one side.

In 1926, Frits
Went

RESULTS
The coleoptile grew straight if the chemical was distributed evenly.
If the chemical was distributed unevenly, the coleoptile curved away from the side with
the block, as if growing toward light, even though it was grown in the dark.

Extracted the
chemical
messenger for
phototropism,
auxin, by
modifying
earlier
experiments

Figure

Excised tip placed


on agar block
Growth-promoting
chemical diffuses
into agar block

Control

Control
(agar block
lacking
chemical)
has no
effect

Agar block
with chemical
stimulates growth
Offset blocks
cause curvature

CONCLUSION
Went concluded that a coleoptile curved toward light because its dark
39.6 side had a higher concentration of the growth-promoting chemical, which he named auxin.

A Survey of Plant Hormones

In general, hormones control plant growth and


development
By affecting the division, elongation, and
differentiation of cells

Plant hormones are produced in very low


concentrations
But a minute amount can have a profound effect on
the growth and development of a plant organ

Auxin
The term auxin
Is used for any chemical substance that promotes
cell elongation in different target tissues

Auxin
Is involved in the formation and branching of roots
(Lateral and Adventitious)

An overdose of auxins
Can kill eudicots (Auxins as Herbicides)

Auxin affects secondary growth


By inducing cell division in the vascular cambium
and influencing differentiation of secondary xylem

Cytokinins
Cytokinins
Stimulate cell division

Cytokinins
Are produced in actively growing tissues such as
roots, embryos, and fruits
Work together with auxin (Control of Cell Division
and Differentiation)

Cytokinins, auxin, and other factors interact in the


control of apical dominance
The ability of a terminal bud to suppress development of
axillary buds

If the terminal bud is removed


Plants become bushier

Axillary buds

Stump after
removal of
apical bud

Lateral branches

Cytokinins retard the aging of some plant


organs
By inhibiting protein breakdown, stimulating RNA
and protein synthesis, and mobilizing nutrients from
surrounding tissues

Gibberellins
Gibberellins have a variety of effects
Such as stem elongation, fruit growth, and seed
germination

Gibberellins stimulate growth of both leaves


and stems
In stems
Gibberellins stimulate cell elongation and cell
division

In many plants
Both auxin and gibberellins must be present for fruit
to set

Gibberellins are used commercially


In the spraying of Thompson seedless grapes

Figure 39.10

Abscisic Acid
Two of the many effects of abscisic acid (ABA)
are
Seed dormancy
Drought tolerance

Seed dormancy has great survival value


Because it ensures that the seed will germinate only
when there are optimal conditions

ABA is the primary internal signal


That enables plants to withstand drought

Ethylene
Plants produce ethylene
In response to stresses such as drought, flooding,
mechanical pressure, injury, and infection

A burst of ethylene
Is associated with the programmed destruction of cells,
organs, or whole plants
Apoptosis: Programmed Cell Death

Leaf Abscission
A change in the balance of auxin and ethylene
controls leaf abscission
The process that occurs in autumn when a leaf falls

A burst of ethylene production in the fruit


Triggers the ripening process

A phytochrome
Is the photoreceptor responsible for the opposing effects of red and farred light

Phytochromes exist in two photoreversible states


With conversion of Pr to Pfr triggering many developmental responses

Pr

Pfr
Red light
Responses:
seed germination,
control of
flowering, etc.

Synthesis

Far-red
light

Slow conversion
in darkness
(some plants)

Enzymatic
destruction

Biological Clocks and Circadian


Rhythms

Many plant processes

Oscillate during the day

Many legumes
Lower their leaves in the evening and raise them in
the morning

Cyclical responses to environmental stimuli are


called circadian rhythms
And are approximately 24 hours long
Can be entrained to exactly 24 hours by the
day/night cycle

Photoperiodism and Responses to


Seasons
Photoperiod, the relative lengths of night and
day
Is the environmental stimulus plants use most
often to detect the time of year

Photoperiodism
Is a physiological response to photoperiod

Some developmental processes, including


flowering in many species
Requires a certain photoperiod

Critical Night Length

In the 1940s, researchers discovered that


flowering and other responses to photoperiod
Are actually controlled by night length, not day length
EXPERIMENT
During the 1940s, researchers conducted experiments in which periods of darkness were
interrupted with brief exposure to light to test how the light and dark portions of a photoperiod affected flowering
in short-day and long-day plants.
RESULTS

24 hours

Darkness
Flash of
light
Critical
dark
period
Light

(a) Short-day plants


flowered only if a period of
continuous darkness was
longer than a critical dark
period for that particular
species (13 hours in this
example). A period of
darkness can be ended by a
brief exposure to light.

Figure 39.22

(b) Long-day plants


flowered only if a
period of continuous
darkness was shorter
than a critical dark
period for that
particular species (13
hours in this example).

CONCLUSION
The experiments indicated that flowering of each species was determined by a critical period of
darkness (critical night length) for that species, not by a specific period of light. Therefore, short-day plants are
more properly called long-night plants, and long-day plants are really short-night plants.

Action spectra and photoreversibility


experiments
Show that phytochrome is the pigment that receives
red light, which can interrupt the nighttime portion of
the photoperiod
EXPERIMENT

A unique characteristic of phytochrome is reversibility in response to red and


far-red light. To test whether phytochrome is the pigment measuring interruption of dark periods,
researchers observed how flashes of red light and far-red light affected flowering in short-day
and long-day plants.
24
20
R

FR
R

Hours

16

R
FR
R

FR
R
FR
R

Critical dark period

RESULTS

12
8
4
0

Short-day (long-night) plant

Long-day (short-night) plant

CONCLUSION

Figure 39.23

A flash of red light shortened the dark period. A subsequent flash of far-red
light canceled the red lights effect. If a red flash followed a far-red flash, the effect of the far-red
light was canceled. This reversibility indicated that it is phytochrome that measures the interruption
of dark periods.

Plants respond to a wide variety of stimuli other


than light
Because of their immobility
Plants must adjust to a wide range of environmental
circumstances through developmental and
physiological mechanisms

GRAVITY
Response to gravity
Is known as gravitropism

Roots show positive gravitropism


Stems show negative gravitropism

Mechanical Stimuli
The term thigmomorphogenesis
Refers to the changes in form that result from
mechanical perturbation

Rubbing the stems of young plants a couple of


times daily
Results in plants that are shorter than controls

Growth in response to touch


Is called thigmotropism
Occurs in vines and other climbing plants

Tropisms
Gravitropism: growth response to direction of
gravity.
Phototropism: growth in response to light.
Hydrotropism: growth in response to the
presence of water.
Heliotropism: ability of plants/leaves to follow the
suns movement across the sky.
Chemotropism: growth in response to certain
chemical stimulus.
Thigmotropism: growth in response to a
mechanical stimulus.

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