Chapter 39

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Transcript Chapter 39

LECTURE PRESENTATIONS

For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson

Chapter 39 Plant Responses to Internal and External Signals

Lectures by Erin Barley Kathleen Fitzpatrick

© 2011 Pearson Education, Inc.

Overview: Stimuli and a Stationary Life

• • Linnaeus noted that flowers of different species opened at different times of day and could be used as a

horologium florae

, or floral clock Plants, being rooted to the ground, must respond to environmental changes that come their way – For example, the bending of a seedling toward light begins with sensing the direction, quantity, and color of the light © 2011 Pearson Education, Inc.

Figure 39.1

Concept 39.1: Signal transduction pathways link signal reception to response

• • • A potato left growing in darkness produces shoots that look unhealthy, and it lacks elongated roots These are morphological adaptations for growing in darkness, collectively called

etiolation

After exposure to light, a potato undergoes changes called

de-etiolation

, in which shoots and roots grow normally © 2011 Pearson Education, Inc.

Figure 39.2

(a) Before exposure to light (b) After a week’s exposure to natural daylight

Figure 39.2a

(a) Before exposure to light

Figure 39.2b

(b) After a week’s exposure to natural daylight

• • A potato’s response to light is an example of cell-signal processing The stages are reception, transduction, and response © 2011 Pearson Education, Inc.

Figure 39.3

CELL WALL 1 Reception CYTOPLASM 2 Transduction Relay proteins and second messengers Receptor Hormone or environmental stimulus Plasma membrane 3 Response Activation of cellular responses

Reception

• • Internal and external signals are detected by receptors, proteins that change in response to specific stimuli In de-etiolation, the receptor is a phytochrome capable of detecting light © 2011 Pearson Education, Inc.

Transduction

• • •

Second messengers

transfer and amplify signals from receptors to proteins that cause responses Two types of second messengers play an important role in de-etiolation: Ca 2+ ions and cyclic GMP (cGMP) The phytochrome receptor responds to light by – Opening Ca 2+ channels, which increases Ca 2+ levels in the cytosol – Activating an enzyme that produces cGMP © 2011 Pearson Education, Inc.

Figure 39.4-1

Cell wall 1 Reception CYTOPLASM Plasma membrane Phytochrome Light

Figure 39.4-2

Cell wall 1 Reception CYTOPLASM Plasma membrane Phytochrome Light 2 Transduction cGMP Second messenger Protein kinase 1 Protein kinase 2 Ca 2

channel Ca 2

Figure 39.4-3

Cell wall 1 Reception CYTOPLASM Plasma membrane Phytochrome Light 2 Transduction cGMP Second messenger Protein kinase 1 Protein kinase 2 Ca 2

channel 3 Response Transcription factor 1 NUCLEUS P Transcription factor 2 P Transcription Translation De-etiolation (greening) response proteins Ca 2

Response

• • • A signal transduction pathway leads to regulation of one or more cellular activities In most cases, these responses to stimulation involve increased activity of enzymes This can occur by transcriptional regulation or post-translational modification © 2011 Pearson Education, Inc.

Post-Translational Modification of Preexisting Proteins

• • • Post-translational modification involves modification of existing proteins in the signal response Modification often involves the phosphorylation of specific amino acids The second messengers cGMP and Ca 2+ activate protein kinases directly © 2011 Pearson Education, Inc.

Transcriptional Regulation

• • • Specific transcription factors bind directly to specific regions of DNA and control transcription of genes Some transcription factors are activators that increase the transcription of specific genes Other transcription factors are repressors that decrease the transcription of specific genes © 2011 Pearson Education, Inc.

De-Etiolation (“Greening”) Proteins

• De-etiolation activates enzymes that – Function in photosynthesis directly – Supply the chemical precursors for chlorophyll production – Affect the levels of plant hormones that regulate growth © 2011 Pearson Education, Inc.

Concept 39.2: Plant hormones help coordinate growth, development, and responses to stimuli

• Plant

hormones

are chemical signals that modify or control one or more specific physiological processes within a plant © 2011 Pearson Education, Inc.

The Discovery of Plant Hormones

• • • Any response resulting in curvature of organs toward or away from a stimulus is called a

tropism

In the late 1800s, Charles Darwin and his son Francis conducted experiments on

phototropism

, a plant’s response to light They observed that a grass seedling could bend toward light only if the tip of the coleoptile was present © 2011 Pearson Education, Inc.

• They postulated that a signal was transmitted from the tip to the elongating region © 2011 Pearson Education, Inc.

© 2011 Pearson Education, Inc.

Video: Phototropism

Figure 39.5

RESULTS Control Light Shaded side Illuminated side Boysen-Jensen Light Darwin and Darwin Light Tip removed Opaque cap Trans parent cap Opaque shield over curvature Gelatin (permeable) Mica (impermeable)

• In 1913, Peter Boysen-Jensen demonstrated that the signal was a mobile chemical substance © 2011 Pearson Education, Inc.

• In 1926, Frits Went extracted the chemical messenger for phototropism, auxin, by modifying earlier experiments © 2011 Pearson Education, Inc.

Figure 39.6

RESULTS Excised tip on agar cube Growth-promoting chemical diffuses into agar cube Control Control (agar cube lacking chemical) Offset cubes

A Survey of Plant Hormones

• • Plant hormones are produced in very low concentration, but a minute amount can greatly affect growth and development of a plant organ In general, hormones control plant growth and development by affecting the division, elongation, and differentiation of cells © 2011 Pearson Education, Inc.

Table 39.1

Auxin

• • • • The term

auxin

refers to any chemical that promotes elongation of coleoptiles Indoleacetic acid (IAA) is a common auxin in plants; in this lecture the term specifically to IAA

auxin

refers Auxin is produced in shoot tips and is transported down the stem Auxin transporter proteins move the hormone from the basal end of one cell into the apical end of the neighboring cell © 2011 Pearson Education, Inc.

Figure 39.7

RESULTS Epidermis Cortex Phloem Xylem Pith 100

m Cell 1 Cell 2 25

m Basal end of cell

Figure 39.7a

Epidermis Cortex Phloem Xylem Pith 100

m

Figure 39.7b

Cell 1 Cell 2 25

m Basal end of cell

• • •

The Role of Auxin in Cell Elongation

According to the acid growth hypothesis, auxin stimulates proton pumps in the plasma membrane The proton pumps lower the pH in the cell wall, activating

expansins

, enzymes that loosen the wall’s fabric With the cellulose loosened, the cell can elongate © 2011 Pearson Education, Inc.

Figure 39.8

Cross-linking polysaccharides Cell wall –loosening enzymes Expansin CELL WALL Cellulose microfibril H

H

H

H

H

H

H

H

H 2 O Plasma membrane Cell wall ATP H

Plasma membrane CYTOPLASM Nucleus Cytoplasm Vacuole

Figure 39.8a

Cross-linking polysaccharides Cell wall –loosening enzymes Expansin CELL WALL Cellulose microfibril H

H

H

H

H

H

H

H

ATP H

Plasma membrane CYTOPLASM

Figure 39.8b

H 2 O Plasma membrane Cell wall Nucleus Cytoplasm Vacuole

• Auxin also alters gene expression and stimulates a sustained growth response © 2011 Pearson Education, Inc.

• • • • •

Auxin’s Role in Plant Development

Polar transport of auxin plays a role in pattern formation of the developing plant Reduced auxin flow from the shoot of a branch stimulates growth in lower branches Auxin transport plays a role in phyllotaxy, the arrangement of leaves on the stem Polar transport of auxin from leaf margins directs leaf venation pattern The activity of the vascular cambium is under control of auxin transport © 2011 Pearson Education, Inc.

• •

Practical Uses for Auxins

The auxin indolbutyric acid (IBA) stimulates adventitious roots and is used in vegetative propagation of plants by cuttings An overdose of synthetic auxins can kill plants – For example 2,4-D is used as an herbicide on eudicots © 2011 Pearson Education, Inc.

Cytokinins

Cytokinins

are so named because they stimulate cytokinesis (cell division)

© 2011 Pearson Education, Inc.

• •

Control of Cell Division and Differentiation

Cytokinins are produced in actively growing tissues such as roots, embryos, and fruits Cytokinins work together with auxin to control cell division and differentiation © 2011 Pearson Education, Inc.

• •

Control of Apical Dominance

Cytokinins, auxin, and strigolactone interact in the control of apical dominance, a terminal bud’s ability to suppress development of axillary buds If the terminal bud is removed, plants become bushier © 2011 Pearson Education, Inc.

Figure 39.9

Lateral branches “Stump” after removal of apical bud (b) Apical bud removed Axillary buds (a) Apical bud intact (not shown in photo) (c) Auxin added to decapitated stem

Figure 39.9a

Axillary buds (a) Apical bud intact (not shown in photo)

Figure 39.9b

Lateral branches “Stump” after removal of apical bud (b) Apical bud removed

Figure 39.9c

(c) Auxin added to decapitated stem

Anti-Aging Effects

Cytokinins slow the aging of some plant organs by inhibiting protein breakdown, stimulating RNA and protein synthesis, and mobilizing nutrients from surrounding tissues © 2011 Pearson Education, Inc.

Gibberellins

Gibberellins

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

© 2011 Pearson Education, Inc.

• • •

Stem Elongation

Gibberellins are produced in young roots and leaves Gibberellins stimulate growth of leaves and stems In stems, they stimulate cell elongation and cell division © 2011 Pearson Education, Inc.

Figure 39.10

(a) Rosette form (left) and gibberellin-induced bolting (right) (b) Grapes from control vine (left) and gibberellin-treated vine (right)

Figure 39.10a

(a) Rosette form (left) and gibberellin-induced bolting (right)

• •

Fruit Growth

In many plants, both auxin and gibberellins must be present for fruit to develop Gibberellins are used in spraying of Thompson seedless grapes © 2011 Pearson Education, Inc.

Figure 39.10b

(b) Grapes from control vine (left) and gibberellin-treated vine (right)

Germination

After water is imbibed, release of gibberellins from the embryo signals seeds to germinate © 2011 Pearson Education, Inc.

Figure 39.11

Aleurone Endosperm 1 2 Water Scutellum (cotyledon) GA GA Radicle

-amylase 3 Sugar

Brassinosteroids

• • •

Brassinosteroids

are chemically similar to the sex hormones of animals They induce cell elongation and division in stem segments They slow leaf abscission and promote xylem differentiation © 2011 Pearson Education, Inc.

Abscisic Acid

• •

Abscisic acid (ABA)

slows growth Two of the many effects of ABA – Seed dormancy – Drought tolerance © 2011 Pearson Education, Inc.

• • •

Seed Dormancy

Seed dormancy ensures that the seed will germinate only in optimal conditions In some seeds, dormancy is broken when ABA is removed by heavy rain, light, or prolonged cold Precocious (early) germination can be caused by inactive or low levels of ABA © 2011 Pearson Education, Inc.

Figure 39.12

Red mangrove (Rhizophora mangle) seeds Coleoptile Maize mutant

Figure 39.12a

Red mangrove (Rhizophora mangle) seeds

Figure 39.12b

Coleoptile Maize mutant

• •

Drought Tolerance

ABA is the primary internal signal that enables plants to withstand drought ABA accumulation causes stomata to close rapidly © 2011 Pearson Education, Inc.

Strigolactones

• • • The hormones called

strigolactones

– Stimulate seed germination – Help establish mycorrhizal associations – Help control apical dominance Strigolactones are named for parasitic

Striga

plants

Striga

seeds germinate when host plants exude strigolactones through their roots © 2011 Pearson Education, Inc.

Ethylene

• • Plants produce

ethylene

in response to stresses such as drought, flooding, mechanical pressure, injury, and infection The effects of ethylene include response to mechanical stress, senescence, leaf abscission, and fruit ripening © 2011 Pearson Education, Inc.

• •

The Triple Response to Mechanical Stress

Ethylene induces the

triple response

, which allows a growing shoot to avoid obstacles The triple response consists of a slowing of stem elongation, a thickening of the stem, and horizontal growth © 2011 Pearson Education, Inc.

Figure 39.13

0.00

0.10

0.20

0.40

0.80

Ethylene concentration (parts per million)

• • Ethylene-insensitive mutants fail to undergo the triple response after exposure to ethylene Other mutants undergo the triple response in air but do not respond to inhibitors of ethylene synthesis © 2011 Pearson Education, Inc.

Figure 39.14

ein mutant ctr mutant (a) ein mutant (b) ctr mutant

Figure 39.14a

ein mutant (a) ein mutant

Figure 39.14b

ctr mutant (b) ctr mutant

• •

Senescence

Senescence

is the programmed death of cells or organs A burst of ethylene is associated with apoptosis, the programmed destruction of cells, organs, or whole plants © 2011 Pearson Education, Inc.

Leaf Abscission

A change in the balance of auxin and ethylene controls leaf abscission, the process that occurs in autumn when a leaf falls © 2011 Pearson Education, Inc.

Figure 39.15

0.5 mm Protective layer Abscission layer Stem Petiole

Figure 39.15a

0.5 mm Protective layer Abscission layer Stem Petiole

• • •

Fruit Ripening

A burst of ethylene production in a fruit triggers the ripening process Ethylene triggers ripening, and ripening triggers release of more ethylene Fruit producers can control ripening by picking green fruit and controlling ethylene levels © 2011 Pearson Education, Inc.

Systems Biology and Hormone Interactions

• • Interactions between hormones and signal transduction pathways make it hard to predict how genetic manipulation will affect a plant Systems biology seeks a comprehensive understanding that permits modeling of plant functions © 2011 Pearson Education, Inc.

Concept 39.3: Responses to light are critical for plant success

• • Light cues many key events in plant growth and development Effects of light on plant morphology are called

photomorphogenesis

© 2011 Pearson Education, Inc.

• • • Plants detect not only presence of light but also its direction, intensity, and wavelength (color) A graph called an

action spectrum

depicts relative response of a process to different wavelengths Action spectra are useful in studying any process that depends on light © 2011 Pearson Education, Inc.

Figure 39.16

1.0

436 nm 0.8

0.6

0.4

0.2

0 400 450 500 550 600 Wavelength (nm) (a) Phototropism action spectrum 650 700 Light Time

0 min Time

90 min (b) Coleoptiles before and after light exposures

Figure 39.16a

1.0

436 nm 0.8

0.6

0.4

0.2

0 400 450 500 550 600 Wavelength (nm) (a) Phototropism action spectrum 650 700

Figure 39.16b

Light Time

0 min Time

90 min (b) Coleoptiles before and after light exposures

Figure 39.16c

Time

0 min

Figure 39.16d

Time

90 min

• • Different plant responses can be mediated by the same or different photoreceptors There are two major classes of light receptors:

blue-light photoreceptors

and

phytochromes

© 2011 Pearson Education, Inc.

Blue-Light Photoreceptors

• Various blue-light photoreceptors control hypocotyl elongation, stomatal opening, and phototropism © 2011 Pearson Education, Inc.

Phytochromes as Photoreceptors

• • Phytochromes are pigments that regulate many of a plant’s responses to light throughout its life These responses include seed germination and shade avoidance © 2011 Pearson Education, Inc.

Phytochromes and Seed Germination

• • Many seeds remain dormant until light conditions change In the 1930s, scientists at the U.S. Department of Agriculture determined the action spectrum for light-induced germination of lettuce seeds © 2011 Pearson Education, Inc.

Figure 39.17

RESULTS Dark (control) Red Dark Red Far-red Dark Red Far-red Red Dark Red Far-red Red Far-red

Figure 39.17a

Dark (control)

Figure 39.17b

Red Dark

Figure 39.17c

Red Far-red Dark

Figure 39.17d

Red Far-red Red Dark

Figure 39.17e

Red Far-red Red Far-red

• • Red light increased germination, while far-red light inhibited germination The photoreceptor responsible for the opposing effects of red and far-red light is a phytochrome © 2011 Pearson Education, Inc.

Figure 39.18

Two identical subunits Chromophore Photoreceptor activity Kinase activity

Figure 39.UN01

Red light P r P fr Far-red light

• • • • • Phytochromes exist in two photoreversible states, with conversion of P r to P many developmental responses fr triggering Red light triggers the conversion of P r to P fr Far-red light triggers the conversion of P fr to P r The conversion to P fr conversion to P r is faster than the Sunlight increases the ratio of P fr triggers germination to P r , and © 2011 Pearson Education, Inc.

Figure 39.19

Synthesis P r Red light P fr Far-red light Slow conversion in darkness (some plants) Responses: seed germination, control of flowering, etc.

Enzymatic destruction

Phytochromes and Shade Avoidance

• • • • The phytochrome system also provides the plant with information about the quality of light Leaves in the canopy absorb red light Shaded plants receive more far-red than red light In the “shade avoidance” response, the phytochrome ratio shifts in favor of P tree is shaded r when a © 2011 Pearson Education, Inc.

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, even when kept under constant light or dark conditions © 2011 Pearson Education, Inc.

Figure 39.20

Noon Midnight

Figure 39.20a

Noon

Figure 39.20b

Midnight

• • •

Circadian rhythms

“clock” are cycles that are about 24 hours long and are governed by an internal Circadian rhythms can be entrained to exactly 24 hours by the day/night cycle The clock may depend on synthesis of a protein regulated through feedback control and may be common to all eukaryotes © 2011 Pearson Education, Inc.

The Effect of Light on the Biological Clock

• Phytochrome conversion marks sunrise and sunset, providing the biological clock with environmental cues © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Photoperiodism and Control of Flowering

• • • • Some processes, including flowering in many species, require a certain photoperiod Plants that flower when a light period is shorter than a critical length are called

short-day plants

Plants that flower when a light period is longer than a certain number of hours are called

long day plants

Flowering in

day-neutral plants

is controlled by plant maturity, not photoperiod © 2011 Pearson Education, Inc.

Critical Night Length

In the 1940s, researchers discovered that flowering and other responses to photoperiod are actually controlled by night length, not day length © 2011 Pearson Education, Inc.

• • Short-day plants are governed by whether the critical night length sets a minimum number of hours of darkness Long-day plants are governed by whether the critical night length sets a maximum number of hours of darkness © 2011 Pearson Education, Inc.

Figure 39.21

24 hours (a) Short day (long-night) plant Light Critical dark period Flash of light Darkness (b) Long-day (short-night) plant Flash of light

• • • Red light can interrupt the nighttime portion of the photoperiod A flash of red light followed by a flash of far-red light does not disrupt night length Action spectra and photoreversibility experiments show that phytochrome is the pigment that receives red light © 2011 Pearson Education, Inc.

Figure 39.22

24 hours R R FR R FR R R FR R FR Critical dark period Short-day (long-night) plant Long-day (short-night) plant

• • • Some plants flower after only a single exposure to the required photoperiod Other plants need several successive days of the required photoperiod Still others need an environmental stimulus in addition to the required photoperiod – For example,

vernalization

is a pretreatment with cold to induce flowering © 2011 Pearson Education, Inc.

A Flowering Hormone?

• • • Photoperiod is detected by leaves, which cue buds to develop as flowers The flowering signal is called

florigen

Florigen may be a macromolecule governed by the

FLOWERING LOCUS T (FT)

gene © 2011 Pearson Education, Inc.

Figure 39.23

24 hours 24 hours 24 hours Graft Short-day plant Long-day plant grafted to short-day plant Long-day plant

Concept 39.4: Plants respond to a wide variety of stimuli other than light

• Because of immobility, plants must adjust to a range of environmental circumstances through developmental and physiological mechanisms © 2011 Pearson Education, Inc.

Gravity

• • • Response to gravity is known as

gravitropism

Roots show positive gravitropism; shoots show negative gravitropism Plants may detect gravity by the settling of

statoliths

, dense cytoplasmic components © 2011 Pearson Education, Inc.

© 2011 Pearson Education, Inc.

Video: Gravitropism

Figure 39.24

Statoliths 20

m (a) Primary root of maize bending gravitropically (LMs) (b) Statoliths settling to the lowest sides of root cap cells (LMs)

Figure 39.24a

Figure 39.24b

Figure 39.24c

Statoliths 20

m

Figure 39.24d

Statoliths 20

m

• • Some mutants that lack statoliths are still capable of gravitropism Dense organelles, in addition to starch granules, may contribute to gravity detection © 2011 Pearson Education, Inc.

Mechanical Stimuli

• • The term

thigmomorphogenesis

refers to changes in form that result from mechanical disturbance Rubbing stems of young plants a couple of times daily results in plants that are shorter than controls © 2011 Pearson Education, Inc.

Figure 39.25

• • • •

Thigmotropism

is growth in response to touch It occurs in vines and other climbing plants Another example of a touch specialist is the sensitive plant

Mimosa pudica

, which folds its leaflets and collapses in response to touch Rapid leaf movements in response to mechanical stimulation are examples of transmission of electrical impulses called

action potentials

© 2011 Pearson Education, Inc.

© 2011 Pearson Education, Inc.

Video: Mimosa Leaf

Figure 39.26

(a) Unstimulated state Leaflets after stimulation Pulvinus (motor organ) (b) Stimulated state Side of pulvinus with flaccid cells Side of pulvinus with turgid cells Vein (c) Cross section of a leaflet pair in the stimulated state (LM)

Figure 39.26a

(a) Unstimulated state

Figure 39.26b

(b) Stimulated state

Figure 39.26c

Leaflets after stimulation Pulvinus (motor organ) (c) Cross section of a leaflet pair in the stimulated state (LM)

Figure 39.26d

Side of pulvinus with flaccid cells Side of pulvinus with turgid cells Vein (c) Cross section of a leaflet pair in the stimulated state (LM)

Environmental Stresses

• • • • Environmental stresses have a potentially adverse effect on survival, growth, and reproduction Stresses can be

abiotic

(nonliving) or

biotic

(living) Abiotic stresses include drought, flooding, salt stress, heat stress, and cold stress Biotic stresses include herbivores and pathogens © 2011 Pearson Education, Inc.

Drought

• • During drought, plants reduce transpiration by closing stomata, slowing leaf growth, and reducing exposed surface area Growth of shallow roots is inhibited, while deeper roots continue to grow © 2011 Pearson Education, Inc.

Flooding

• Enzymatic destruction of root cortex cells creates air tubes that help plants survive oxygen deprivation during flooding © 2011 Pearson Education, Inc.

Figure 39.27

Vascular cylinder Air tubes (a) Control root (aerated) 100

m Epidermis 100

m (b) Experimental root (nonaerated)

Figure 39.27a

Vascular cylinder 100

m (a) Control root (aerated) Epidermis

Figure 39.27b

Vascular cylinder Air tubes Epidermis (b) Experimental root (nonaerated) 100

m

Salt Stress

• • • Salt can lower the water potential of the soil solution and reduce water uptake Plants respond to salt stress by producing solutes tolerated at high concentrations This process keeps the water potential of cells more negative than that of the soil solution © 2011 Pearson Education, Inc.

Heat Stress

• • Excessive heat can denature a plant’s enzymes

Heat-shock proteins

help protect other proteins from heat stress © 2011 Pearson Education, Inc.

Cold Stress

• • • • Cold temperatures decrease membrane fluidity Altering lipid composition of membranes is a response to cold stress Freezing causes ice to form in a plant’s cell walls and intercellular spaces Many plants, as well as other organisms, have antifreeze proteins that prevent ice crystals from growing and damaging cells © 2011 Pearson Education, Inc.