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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.

2. 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. Here, let’s specifically consider the production of second messengers
and their function in the greening response.

3. 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. 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. By such mechanisms, many signal
pathways ultimately regulate the synthesis of new proteins, usually by turning specific genes on
or off. 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.
4. 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.
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. 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.

5. 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.
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 .

6. 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" . 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 . 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 .

7. 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 . 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.

8. 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 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 . 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.
9 and 10.
Element Major Functions


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


S SO42- Component of proteins, coenzymes

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


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

Fe Fe3+, 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+, Component of many redox and lignin-biosynthetic enzymes