The Control of Flowering
• How do plants keep track of the seasons of the year and the time of day?
• Which environmental signals control flowering, and how are
those signals perceived?
• How are environmental signals transduced to bring about the
developmental changes associated with flowering?
The events occurring in the shoot apex that specifically commit the apical
meristem to produce flowers are collectively referred to as floral evocation
FLORAL MERISTEMS AND FLORAL ORGAN DEVELOPMENT
The Four Different Types of Floral Organs Are Initiated as Separate Whorls
Shoot Meristems in Arabidopsis
Change with Development
• As plants initiate reproductive development, the vegetative meristem is
transformed into an primary inflorescence meristem that produces floral
meristems on its flanks
• Cauline leaves (inflorescence leaves) develop into secondary inflorescence
meristems, and their activity repeats the pattern of development of the
primary inflorescence meristem
Longitudinal sections through a vegetative (A) and a reproductive (B) shoot apical region
of Arabidopsis
THE SHOOT APEX AND PHASE CHANGES
Shoot Apical Meristems Have Three Developmental Phases
During postembryonic development, the shoot apical meristem passes through three
more or less well-defined developmental stages in sequence:
1. The juvenile phase
2. The adult vegetative phase
3. The adult reproductive phase
The transition from one phase to another is called phase change.
Phase Changes Can Be Influenced by Nutrients, Gibberellins, and Other Chemical Signals
Combinatorial model of shoot development
According to this model, shoot development can be
described as a series of independently regulated,
overlapping programs (juvenile, adult, and
reproductive) that modulate the expression of a
common set of developmental processes
Combinatorial model of shoot development
Competence and Determination :Two Stages in
Floral Evocation
SIGNALS IN FLOWERING
• The developmental signals that bring about floral evocation include
endogenous factors, such as circadian rhythms, phase change, and hormones,
and external factors, such as day length (photoperiod) and temperature
(vernalization).
• In the case of photoperiodism, transmissible signals from the leaves,
collectively referred to as the floral stimulus, are translocated to the shoot
apical meristem
• The interactions of these endogenous and external factors enable plants to
synchronize their reproductive development with the environment
• The interactions of these endogenous and external factors enable plants to
synchronize their reproductive development with the environment
Three Types of Genes Regulate Floral Development
1. Floral organ identity genes directly control floral identity. The proteins encoded
by these genes are transcription factors that likely control the expression of
other genes whose products are involved in the formation and/or function of
floral organs.
2. Cadastral genes act as spatial regulators of the floral organ identity genes by
setting boundaries for their expression.
3. Meristem identity genes are necessary for the initial induction of the organ
identity genes. These genes are the positive regulators of floral organ identity.
Homeotic Mutations Led to the Identification of
Floral Organ Identity Genes
Homeotic genes encode transcription factors that determine the locations at which
specific structures develop
PART OF THE SIGNALING CASCADE IN FLOWERING
The ABC Model Explains the Determination of
Floral Organ Identity
More recently discovered SEP genes led to modification of ABC model
into ABCE model
The ABC Model Explains the Determination of
Floral Organ Identity
• Activity of type A alone specifies sepals.
• Activities of both A and B are required for the formation of petals.
• Activities of B and C form stamens.
• Activity of C alone specifies carpels.
FLORAL EVOCATION: INTERNAL AND EXTERNAL CUES
• The case in which flowering occurs strictly in response to internal
developmental factors and does not depend on any particular environmental
conditions is referred to as autonomous regulation.
• Some plants exhibit an absolute requirement for the proper environmental cues
in order to flower. This condition is termed an obligate or qualitative response to
an environmental cue.
• In other plant species, flowering is promoted by certain environmental cues but
will eventually occur in the absence of such cues. This is called a facultative or
quantitative response to an environmental cue.
CIRCADIAN RHYTHMS: THE CLOCK WITHIN
CIRCADIAN RHYTHMS: THE CLOCK WITHIN
Circadian Rhythms Exhibit Characteristic Features
1. Period, the time between comparable points in the repeating cycle. Typically the
period is measured as the time between consecutive maxima (peaks) or
minima (troughs).
2. Phase, any point in the cycle that is recognizable by its relationship to the rest of
the cycle. The most obvious phase points are the peak and trough positions.
3. Amplitude, usually considered to be the distance between peak and trough. The
amplitude of a biological rhythm can often vary while the period remains
unchanged.
PHOTOPERIODISM: MONITORING
DAY LENGTH
• Circadian clock enables organisms to determine the time of day at
which a particular molecular or biochemical event occurs
• Photoperiodism, or the ability of an organism to detect day length,
makes it possible for an event to occur at a particular time of year,
thus allowing for a seasonal response
Plants Can Be Classified by Their Photoperiodic
Responses
1. Short-day plants (SDPs) flower only in short days (qualitative
SDPs), or their flowering is accelerated by short days
(quantitative SDPs)
2. Long-day plants (LDPs) flower only in long days (qualitative
LDPs), or their flowering is accelerated by long days
(quantitative LDPs)
LDPs vs. SDPs
LDPs vs. SDPs
Critical day length vs. critical night length
Treating short- and long-day plants with different photoperiods clearly shows
that the critical variable is the length of the dark period
Critical day length vs. critical night length
Dual–day length plants
• Long-short-day plants (LSDPs) flower only after a sequence of
long days followed by short days. LSDPs, such as Bryophyllum,
Kalanchoe, and Cestrum nocturnum (night-blooming jasmine),
flower in the late summer and fall, when the days are shortening.
• Short-long-day plants (SLDPs) flower only after a sequence of
short days followed by long days. SLDPs, such as Trifolium repens
(white clover), Campanula medium (Canterbury bells), and
Echeveria harmsii (echeveria), flower in the early spring in
response to lengthening days.
Day-neutral plants (DNPs) are insensitive to day length
Photoperiodic Stimulus
• The Leaf Is the Site of Perception of the Photoperiodic
Stimulus
• In response to photoperiod the leaf transmits a signal that
regulates the transition to flowering at the shoot apex
• The photoperiod-regulated processes that occur in the
leaves resulting in the transmission of a floral stimulus to
the shoot apex are referred to collectively as
photoperiodic induction
• The Floral Stimulus Is Transported via the Phloem
Red vs. far red light in flowering response
Signaling pathways in flowering