Mechanism of Flowering

What Makes Plants Flower?

What causes the transition from vegetative to flowering
apical meristem in plants?

Vegetative meristem (indeterminate) ? Floral meristem (determinate)

Maize shoot apical meristems

(a) Dome of the vegetative

meristem (arrowhead) flanked
by leaf primordia.

(b) Inflorescence primordium

consists of a large terminal
meristem (large arrowhead).
 Flowering is controlled by several factors
1. internal  control shoot
apex growth and
development
– Geneticically
– biochemically
– Circadian rhythms
– autonomous
• Numbers of node, leaf,
physiological age
2. External factors - trigger
flowering genes
– light
– Temperature

Flowering time – integrated signaling  complex pathway

Multiple developmental pathways for
flowering in Arabidopsis:
1. photoperiodism,
The photoperiodic pathway is located in the
leaves
 FT protein
– In LDPs  FT protein is produced in the
phloem in response to CO protein
accumulation under long days 
translocated to the apical meristem.
– In SDPs  Hd3a protein  translocated to
the apical meristem.
2. the autonomous (leaf number)
3. vernalization (low temperature) pathways,
– The autonomous (leaf number) and
vernalization (low temperature) pathways
act in the apical meristem to negatively
regulate FLC, a negative regulator of SOC1.
4. the energy (sucrose) pathway, and
5. the gibberellin pathway.
– The sucrose and gibberellin pathways, also
localized to the meristem, promote SOC1
expression. (After Blázquez 2005.)
Once the plant has made the decision to flower, the apical meristem
becomes an inflorescence meristem (IM). An inflorescence is a branch
that bears flowers. Inflorescence meristems produce flowers in the axils of
their leaves rather than vegetative axillary meristems.

The structure of the inflorescence is determined by the phyllotaxy of the

IM and whether it is determinate or indeterminate.
A simplified model for floral evocation at the shoot apex

and/or
and/or
Vernalization Florigen?
and/or

?
 What Determines the Transition from the
Vegetative to the Flowering State?

During vegetative growth…

– apical meristems continuously
produce leaves, stems, and
axillary buds  indeterminate
growth
– If apical meristem becomes
inflorescence meristem, it
produces bracts and new
meristems in angle between bract bract
and stem
– New meristems may be
inflorescence meristems or
floral meristems, which
give rise to a flower
• inflorescence is an orderly
cluster of flowers

Dancing Girl Ginger (Globba

schomburgkii) inflorescence
• Floral meristem produces four
whorls or spirals of organs, with
very short internodes
– Sepals
– Petals
– Stamens
– Carpels

Floral whorls of passionflower

http://www.eeob.iastate.edu/classes/bio366/terminology/flower/images/whorls-Passiflora.jpg
 Genetics control on flowering

Flowering genes :
• Meristem identity  positive regulation on flower
identity
– starts cascade of further gene expression
• Cadastral  determine spatial patterns of floral
organs
• Flower organ identity  directly control flowering
Genetic control of flowering
Control of flowering  activate several genes known as integrator
flowering pathway, incl. LEAFY (LFY) and APETALA1 (AP1).
– LFY and AP1 encoding transcription factor
– LFY  upregulated in response to GA and CO activity.

Expression of LFY in shoot meristem  flower meristem

– Cadastral genes trigger expression of floral organ
identity (FOI) genes
• Homeotic genes
• Products control transcription of other genes to assign identity to
cells in floral meristem
• The ABC model of flower formation
– Identifies how floral organ identity genes direct
the formation of the four types of floral organs
Sepals
Petals
Stamens

A Carpels
B
C

C gene
activity
A+B B+C
gene gene
activity activity

A gene
activity
Organ Identity Genes in Arabidopsis Flowers

Mutant plants
produce
abnormal
“flowers”
 This combination of a,
b and c regulatory
functions give each
whorl an unique
identity

www.mun.ca/biology/desmid/brian/BIOL3530/DB_Ch07
 What Determines the Transition from the
Vegetative to the Flowering State?
Environmental cues trigger transition to the flowering state
– Annuals: complete life cycle in one year.
– Biennials: complete life cycle in two years.
– Perennials: live for several to many years.

Impatiens (left) are

annuals; foxglove
(right) is biennial.

www.suburbanlawnandgarden.com/images www.greenbeam.com/features/images
 What Determines the Transition from the
Vegetative to the Flowering State?

Photoperiodism is control of flowering and other

responses by length of day or night
– First studied in 1920s in mutant tobacco plants
• Plants would not flower until day length was shorter than 14
hours — critical day length

Wild type and gigantea (gi) mutant

Arabidopsis plants. Mutation delays
flowering, allowing for large growth.

http://3e.plantphys.net/images/ch24/wt2405a.jpg
 What Determines the Transition from the
Vegetative to the Flowering State?
Photoperiodism
 synchronizes flowering in the same species so
that cross-pollination and successful reproduction
is promoted
– Day-neutral plants are more common than
photoperiodic plants – they rely on other cues

Buckwheat, a day-
neutral plant

http://botit.botany.wisc.edu/images/130/Tropisms/photoperiodism
 What Determines the Transition from the
Vegetative to the Flowering State?

Photoperiodic plants actually

measure length of night, not
day
– Experiments with cocklebur, (an
SDP):
– Day length was varied in one
group, night length in the other.
The critical night length was 9
Cocklebur fruit
hours.

http://tncweeds.ucdavis.edu/photos
 Day Length and Flowering
– Key variable - night length
 Photoperiod (relative night length)
Plants are classified into three categories: short-day
(long-night), long-day (short-night), or day-neutral,
depending on their response to the relative duration
of darkness.
 What Determines the Transition from the
Vegetative to the Flowering State?
– Short-day plants (SDPs)
• flower when day is shorter than critical maximum
• Flower in late summer or fall
• Chrysanthemums, poinsettias, Maryland Mammoth tobacco

http://216.119.109.157/content/images www.colostate.edu/Depts/CoopExt/4DMG/images
 What Determines the Transition from the
Vegetative to the Flowering State?
– Long-day plants (LDPs)
• flower when the day is longer than a critical maximum
• Flower in midsummer.
• Spinach, clover

www.nationmaster.com/wikimir/images/upload.
wikimedia.org/wikipedia/commons/thumb/3/37 www.tiger-photographic.com
 What Determines the Transition from the
Vegetative to the Flowering State?

– Short-long-day plants must

experience short days
followed by long days to
flower
• e.g., white clover flowers in long
days just before midsummer

– Long-short-day plants must

experience long days followed
by short days White clover (Trifolium
• bloom only in fall repens), a short-long-day plant

http://www.nps.gov/wica/naturescience/images/White-Clover.jpg
3. Day-neutral plants form flowers regardless of day length. Examples
are tomato, corn, cucumber, and some strawberry cultivars.
Some plants do not fit into any category, but may respond to
combinations of day lengths. Petunias, for example, flower
regardless of day length, but flower earlier and more profusely with
long days.
 What Determines the Transition from the
Vegetative to the Flowering State?

Plants show many

circadian rhythms.
– Clover leaflets are folded
at night, and unfold during
the day
– Many flowers close at night
and open during the day
– Night flowering has
evolved in many plants that
are pollinated by bats and
moths
http://gis.esri.com/library/userconf/proc99/proceed/papers/pap530/p5302.jpg
 What Determines the Transition from the
Vegetative to the Flowering State?

Phytochromes and blue-light receptors

interact with plant’s biological clock
– Arabidopsis has at least three “clock
genes” that encode regulatory proteins
that interact to produce a circadian
oscillation (CO)
 What Determines the Transition from the
Vegetative to the Flowering State?

Timing mechanism is present in each leaf

– Determined by “blindfold” experiments
• If only one leaf on a plant is exposed to
conditions conducive to flowering, it will flower —
an inductive treatment
• A signal must be sent from leaf to site of
flowering.
 38.2 What Determines the Transition from the
Vegetative to the Flowering State?
Evidence that signal is a hormone:
– If a photoperiodically induced leaf is removed from
plant immediately, it does not flower.
– If it remains attached for several hours, plant will
flower.
 What Determines the Transition from the
Vegetative to the Flowering State?
– If several plants are grafted together, inducing
one leaf will cause all plants to flower
 What Determines the Transition from the
Vegetative to the Flowering State?

Hypothetical flowering hormone was

named florigen
– In 2005, experiments with Arabidopsis
showed that mRNA (which can be
transferred through plasmodesmata)
produced by FLOWERING LOCUS T
(FT) gene in the leaf, traveled to shoot
apex
– In Aug, 2007 was reported that FT
protein translated from the mRNA
activates a second (FD) protein to
stimulate stem cells to form flower buds
Arabidopsis thaliana, the
plant used in the research,
after flowering.

http://www.jyi.org/articleimages/426/img0.jpg
 What Determines the Transition from the
Vegetative to the Flowering State?
Vernalization – process of inducing flowering by low
temperatures
– In some plants, flowering is controlled by temperature
– Wheat and rye have two types of flowering behavior: annual
and biennial
• Spring wheat is an annual;
• Winter wheat is biennial; it is planted in fall and flowers in spring
 What Determines the Transition from the
Vegetative to the Flowering State?
– Winter wheat must be exposed to cold in its first year or it will not
flower the second year
– Vernalization inhibits expression of a gene whose protein product
represses other genes for flower development

A cabbage plant that had been grown in the greenhouse

for 5 years. Cabbage is a biennial and requires exposure
to prolonged winter cold in order to flower the second
spring after planting. The large cabbage has never been
exposed to winter and therefore cannot flower.

http://pbi-ibp.nrc-cnrc.gc.ca/en/bulletin/2004issue1/images/page1_01.jpg
 Molecular basis of vernalization-induced
flowering in cereals versus
Arabidopsis

1. In the temperate cereals

 VRN2 represses FT but VRN2 is not expressed in the short days of winter,
 VRN1 expression (after prolonged exposure to cold) promotes inflorescence initiation and represses VRN2, to allow long-day induction of FT to accelerate reproductive development.
2. In Arabidopsis
 FLC is expressed before winter and represses FT.
 Vernalization represses FLC,
 FT (and SOC1) promote flowering in spring. FLC expression is reset during meiosis to establish the vernalization requirement in the next generation
A comparison of the molecular pathways regulating flowering time in Arabidopsis and the temperate
cereals. Vernalization and long days promote flowering in Arabidopsis (left) and in the temperate cereals
(right). The day-length response is conserved (grey). CONSTANS (CO) senses long days and activates
FLOWERING LOCUS T (FT) expression. This requires PHOTOPERIOD1 (PPD1) in cereals. Vernalization
is a prerequisite for long-day induction of FT in both Arabidopsis and cereals, but the vernalization pathway
(blue) has evolved independently. In Arabidopsis, FLOWERING LOCUS C (FLC) blocks long-day induction
of FT but is repressed by vernalization. In the temperate cereals, VRN2 blocks long-day induction of FT
before winter, but VRN1 is induced by vernalization to repress VRN2 and allow long-day induction of FT. In
Arabidopsis, the vernalization and day-length response pathways intersect at FT, which can be described
as a floral integrator gene (green). In cereals, VRN2 is a floral integrator gene. In both Arabidopsis and in
the temperate cereals, activation of flowering causes expression of genes that promote inflorescence
meristem identity (red), such as APETALLA1 (AP1) in Arabidopsis. VRN1 acts as both a flowering time
gene in the vernalization response pathway (blue) and as a meristem identity gene during reproductive
development (red).