Reception

Signals, whether internal or external, are first detected by receptors, proteins that undergo
conformational changes in response to a specific stimulus.

The receptor involved in greening in plants is called phytochrome, which consists of a light-
absorbing pigment attached to a specific protein.

Unlike many receptors, which are built into the plasma membrane, the phytochrome that functions
in greening occurs in the cytoplasm

Phytochrome functions in light detection in the greening process

Transduction
The greening response is triggered by extremely low levels of light.

Receptors such as phytochrome are sensitive to very weak environmental and chemical signals.

How is the information from these extremely weak signals amplified, and how is their reception
transduced into a specific response by the plant? The answer is second messengers -- small,
internally produced chemicals that transfer and amplify the signal from the receptor to proteins
that cause the specific response

In the greening response, for example, each activated phytochrome may give rise to hundreds of
molecules of a second messenger, each of which, in turn, may lead to the activation of hundreds of
molecules of a specific enzyme.

By such mechanisms, the second messenger of a signal transduction pathway leads to a rapid
amplification of the signal. In Chapter 11, we examined this role of second messengers in general
(Fig 11.12).

Here, let’s specifically consider the production of second messengers and their function in the
greening response (Fig 39.3)

Phytochrome is such a receptor.

Light causes phytochrome to undergo a conformational change, and the phytochrome then interacts
with a specific G-protein.
During activation, guanosine diphosphate (GDP) that is bound to the inactive G-protein is
displaced by guanosine triphosphate (GTP).

The G-protein, now in its active form, in turn activates other enzymes in the signal-transduction
pathway that leads to greening

The cyclic nucleotides are second messengers that include cyclic adenosine monophosphate
(cyclic AMP) and cyclic guanosine monophosphate (cyclic GMP).

Experiments indicate that cyclic GMP (cGMP) is involved in the greening process

Changes in cytosolic Ca+ also play an important role in phytochrome signal transduction. However,
a wide range of hormonal and environmental stimuli can cause brief increases in cytosolic Ca+.

In many cases, Ca2+ then binds directly to small proteins called calmodulins. The Ca+-
calmodulin complex then binds to and activates several enzymes, most notably certain types of
protein kinases.

Fig 39.3 that phytochrome activation during the greening mechanism results in both cGMP and
Ca+-calmodulin as second messengers

Response
Ultimately, a signal-transduction pathway leads to the regulation of one or more cellular
activities.

The 2 main mechanisms by which a signaling pathway can activate an enzyme are by stimulating
transcription of mRNA for the enzyme or by activating existing enzyme molecules (post-
translational modification)

Transcriptional Regulation. Transcription factors bind directly to specific regions of DNA and
control the transcription of specific genes (see Fig 19.8).

In the case of phytochrome-induced greening, several transcription factors are activated by

phosphorylation in response to the appropriate light conditions. The activation of some of these
transcription factors depends upon cyclic GMP, whereas the activation of others requires Ca+-
calmodulin.

Post-Translational Modification of Proteins. Although the synthesis of new proteins by

transcription and translation are important molecular events associated with greening, so are post-
translational modifications of existing proteins.
Most often these existing proteins are modified by phosphorylation, the addition of a P group onto
the protein. The diverse proteins called protein kinases catalyze this phosphorylation of target
proteins (see Fig 11.11).

By such mechanisms, many signal pathways ultimately regulate the synthesis of new proteins,
usually by turning specific genes on or off (Fig 39.3)

The Greening Proteins. What sorts of proteins are either newly transcibed or activated by
phosphorylation during the greening process? Many are enzymes that function in photosynthesis
directly; others are enzymes involved in supplying the chemical precursors necessary for
chlorophyll production; still others affect the levels of plant hormones that regulate growth

Elements required by plants in relatively large amounts are called macronutrients. There are nine
macronutrients in all, including the six major ingredients of organic compounds: carbon (C),
oxygen (O2), hydrogen (H2), nitrogen (N), sulfur (S), and phosphorus (P). The other three
macronutrients are potassium (K), calcium (Ca), and magnesium (Mg). (table 37.1)

Elements that plants need in very small amounts are called micronutrients. The eight
micronutrients are iron (Fe), chlorine (Cl), copper (Cu), manganese (Mn), zinc (Zn), molybdenum
(Mo), boron (Bo), and nickel (Ni). These elements function in plants mainly as cofactors of
enzymatic reactions

Form
Element vailable Major Functions
to Plants

Macronutrients

C CO2 Major component of plant’s organic compounds

O2 CO2 Major component of plant’s organic compounds

H2 H2O Major component of plant’s organic compounds

N NO3-, Component of nucleic acids, proteins, hormones, and coenzymes

NH4+

S SO42- Component of proteins, coenzymes

P H2PO42-, Component of nucleic acids, phospholipids, ATP, several

HPO42- coenzymes
K K+ Cofactor that functions in protein synthesis; major solute
functioning in water balance; operation of stomata

Ca Ca2+ Important in formation and stability of cell walls and in

maintenance of membrane structure and permeability; activates
some enzymes; regulates many responses of cells to stimuli

Mg Mg2+ Component of chlorophyll; activates many enzymes

Micronutrients

Cl Cl- Required for water-splitting step of photosynthesis; functions in

water balance

Fe Fe3+, Fe2+ Component of cytochromes; activates some enzymes

Bo H2BO3- Cofactor in chlorophyll synthesis; may be involved in

carbohydrate transport and nucleic acid synthesis

Mn Mn2+ Active in formation of amino acids; activates some enzymes;

required for water-splitting step of photosynthesis

Zn Zn2+ Active in formation of chlorophyll; activates some enzymes

Cu Cu+, Cu2+ Component of many redox and lignin-biosynthetic enzymes

Mo MoO42- Essential for nitrogen fixation; cofactor that functions in nitrate

reduction

Ni Ni2+ Cofactor for an enzyme functioning in nitrogen metabolism

Auxin (IAA, IBA, NAA, 2.4D)
Produce: Embryo of seed, meristems of apical buds, young leaves

Function: Stimulates stem elongation (low concentration only) root growth, cell differentiation,
and branching; regulates development of fruit; enhances apical dominance; functions in
phototropism and gravitropism.

Moving: Auxin seems to be transported directly through parenchyma tissue, from one cell-cell. It
moves only from shoot tip to base, not in the reverse direction.

This unidirectional transport of auxin is called polar transport . Polar auxin transport requires
energy.

Fig 39.6 illustrates how proton pumps in the plasma membrane, driven by ATP, couple metabolic
energy to auxin transport

The Role of Auxin in Cell Elongation. The apical meristem of a shoot is a major site of auxin
synthesis. As auxin from the shoot apex moves down to the region of cell elongation (see Chapter
35), the hormone stimulates growth of the cells, probably by binding to a receptor built into the
plasma membrane

According to a proposal called the acid growth hypothesis, proton pumps play a major role in the
growth response of cells to auxin. In a shoot’s region of elongation, auxin stimulates plasma
membrane proton pumps, an action that, within minutes, both increases the voltage across the
membrane (membrane potential) and lowers the pH in the cell wall (Fig 39.7)

Auxin also alters gene expression rapidly, causing cells in the region of elongation to produce new
proteins within minutes

Lateral and Adventitious Root Formation. Auxins are used commercially in the vegetative
propagation of plants by cuttings

Auxins as Herbicides. Synthetic auxins, such as 2,4-dinitrophenol (2,4-D), are widely used as
herbicides

Other Effects of Auxin. In addition to stimulating cell elongation for primary growth, auxin
affects secondary growth by inducing cell division in the vascular cambium and by influencing the
differentiation of secondary xylem
Cytokinins (BA, kinetin, 2iP, zeatin, TDZ)
Produce: Synthesized in roots and

Moving: Transported to other organs

Function: Affect root growth and differentiation; stimulate cell division and growth; stimulate
germination; delay senescence

Despite much effort, the enzyme that produces cytokinins has neither been purified from plants nor
has the gene that encodes for it been identified . Regardless of the source of cytokinins, plant cells
do have cytokinin receptors

Control of Cell Division and Differentiation. Cytokinins are produced in actively growing tissues,
particularly in roots, embryos, and fruits

Control of Apical Dominance. Cytokinins, auxin, and other factors interact in the control of apical
dominance, the ability of the terminal bud to suppress the development of axillary buds (Fig 39.8)

Anti-Aging Effects. Cytokinins retard the aging of some plant organs by inhibiting protein
breakdown, by stimulating RNA and protein synthesis, and by mobilizing nutrients from
surrounding tissues

Gibberellins (GA3)
Produce: Meristems of apical buds and roots, young levels, embryo

Moving: Transported to other organs

Function: Promote seed and bud germination, stem elongation, and leaf growth; stimulate
flowering and development of fruit; affect root growth and differentiation

In 1926, the Japanese plant pathologist E. Kurosawa discovered that a fungus of the genus
Gibberella caused this "foolish seedling disease" (Fig 39.9).

Stem Elongation. Roots and young leaves are major sites of gibberellin production. Gibberellins
stimulate growth in both the leaves and the stem, but they have little effect on root growth. In stems,
gibberellins stimulate cell elongation and cell division (Fig 39.10)

Fruit Growth. In many plants, both auxin and gibberellins must be present for fruit to set. The
most important commercial application of gibberellins is in the spraying of Thompson seedless
grapes (Fig 39.11). The hormone makes the individual grapes grow larger, a trait prized by the
consumer, and it makes the internodes of the grape bunch elongate, allowing more space for the
individual grapes.

Germination. The embryo of seeds is a rich source of gibberellins. After water is imbibed, the
release of gibberellins from the embryo signals the seeds to break dormancy and germinate

Gibberellins support the growth of cereal seedlings by stimulating the synthesis of digestive
enzymes such as α-amylase that mobilize stored nutrients (see Fig 38.13).

Abscisic Acid (ABA)

Produce: Leaves, stems, roots, green fruit

Moving: Transported to other organs

Function: Inhibits growth; closes stomata during water stress; counteracts breaking of dormancy

ABA generally slows down growth. Often ABA antagonizes the actions of the growth hormones,
and it is the ratio of ABA to one or more growth hormones that determines the final physiological
outcome

Seed Dormancy. Seed dormancy has great survival value because it ensures that the seed will
germinate only when there are optimal conditions of light, temperature, and moisture

The high levels of ABA in maturing seeds inhibit germination and induce the production of
special proteins that help the seeds withstand the extreme dehydration that accompanies maturation.
Many types of dormant seeds will germinate when ABA is removed or inactivated in some way.
The seeds of some desert plants break dormancy only when heavy rains wash ABA out of the seed

Abscisic acid induces dormancy in seeds. When its action is blocked--in this case, by a mutation
affecting an ABA-regulated transcription factor--precocious germination results (Fig 39.12)

Drought Stress. ABA is the primary internal signal that enables plants to withstand drought. When
a plant begins to wilt, ABA accumulates in leaves and causes stomata to close rapidly, reducing
transpiration and preventing further water loss.
Ethylene
Produce: Tissues of ripening fruits, nodes of stems, aging leves and flowers

Moving: Transported to other organs

Function: Promotes fruit ripening, opposes some auxin effects; promotes or inhibits growth and
development of roots, leaves, and flowers, depending on species

The Triple Response to Mechanical Stress: Using Mutants to Dissect a Signal-Transduction

Pathway. As the stem pushes against the obstacle, the mechanical stress in its delicate tip induces
the seedling to start producing ethylene. Ethylene, in turn, instigates the seedling to perform a
growth maneuver called the triple response that enables it to circumvent the obstacle.

The 3 parts of this response, which you can see in Fig 39.13, are a slowing of stem elongation, a
thickening of the stem (which makes it stronger), and a curvature that causes the stem to start
growing horizontally

Ethylene-insensitive (ein) mutants fail to undergo the triple response after exposure to ethylene
(Fig 39.14a). The phenotype of such ethylene-overproducing (eto) mutants can be restored to wild-
type by treating the seedlings with inhibitors of ethylene synthesis. Still other mutants, called
constitutive triple-response (ctr) mutants, undergo the triple response in air but do not respond to
inhibitors of ethylene synthesis (Fig 39.14b). The affected gene in ctr mutants turns out to code for
a protein kinase . Fig 39.15 summarizes the responses of ein , eto , and ctr mutants to ethylene and
ethylene synthesis inhibitors

Apoptosis: Programmed Cell Death. Consider the shedding of a leaf in autumn or the death of an
annual after flowering. The cells, organs and plants that are genetically programmed to die on a
particular schedule do not simply shut down their cellular machinery and await death. Rather, the
onset of programmed cell death, called apoptosis, is one of the busiest times in a cell’s life,
requiring new gene expression.

Leaf Abscission. The loss of leaves each autumn is an adaptation that keeps deciduous trees from
desiccating during winter when the roots cannot absorb water from the frozen ground

When an autumn leaf falls, the breaking point is an abscission layer located near the base of the
petiole. The small parenchyma cells of this layer have very thin walls, and there are no fiber cells
around the vascular tissue. The abscission layer is further weakened when enzymes hydrolyze
polysaccharides in the cell walls. Finally, the weight of the leaf, with the help of wind, causes a
separation within the abscission layer . Even before the leaf falls, a layer of cork forms preventing
pathogens from invading the plant (Fig 39.16).

A change in the balance of ethylene and auxin controls abscission

Fruit Ripening. Immature (unripe) fruits that are tart, hard, and green become edible at the time of
seed maturation. A burst of ethylene production in the fruit triggers this ripening

A chain reaction occurs during ripening: ethylene triggers ripening, and ripening, in turn,
triggers even more ethylene production -- one of the rare examples of positive feedback in
physiology (Fig 1.8). The result is a huge burst in ethylene production

Given the importance of ethylene in the post-harvest physiology of fruits, the genetic engineering
of ethylene signal transduction pathways has potentially important commercial applications