Transition to flowering: Unit 5 & Unit 6

CHAPTER-8
BY JYOTI KUMARI
References:
Notes are made from(SOURCES):
1. Taiz and Zeiger 6ED (content & Images credit)
2. Authentic google websites.
3. Review and research papers(NCBI )
 Vernalization
➢ Vernalization is also known as chilling treatment or cold treatment.
➢ Vernalization is the process whereby repression of flowering is alleviated(relieve or lessen) by a cold treatment
given to a hydrated seed (i.e., a seed that has imbibed water) or to a growing plant (dry seeds do not respond to the
cold treatment because vernalization is an active metabolic process).
➢ Without the cold treatment, plants that require vernalization show delayed flowering or remain vegetative, and
they are not competent to respond to floral signals such as inductive photoperiods.
➢ In many cases these plants grow as rosettes with no elongation of the stem.
➢ Plant have evolve the ability to alter their developmental programme in response to environment stimuli.
➢ A major switch in the developmental programme is the transition to flowering.
➢ Photoperiod and temperature play a important role in determining the correct time to flower.
➢ Different plant may require slight different Vernalization temperature.
➢ Photoperiodism and vernalization are two of the most important mechanisms underlying seasonal responses.
Photoperiodism is a response to the length of day; vernalization is the promotion of flowering.
 Vernalization(contd.)
➢ Promoting flowering by cold. Vernalization help to shorten the vegetative phase.
➢ The effective temperature range for vernalization is from just below freezing to about 10°C, with a broad optimum
usually between about 1 and 7°C
➢ Cold temperatures are required for the germination of certain seeds (stratification) and for flowering in certain
species (vernalization)
➢ Plants differ considerably in the age at which they become sensitive to vernalization.
➢ Competence of the meristem to undergo the floral transition.

Arabidopsis thaliana
(no Vernalization)
Arabidopsis thaliana
with Vernalization
 What is the need of Vernalization?
• Many plants do not come to flower before they experience a low temperature. These plants remain
vegetative during warm season, experience low temperature during winter, grow further and then bear
flowers and fruits.
Vernalized for 1
Non- Vernalized month at 5OC

❑ Without the cold treatment, plants that require vernalization show delayed flowering or
remain vegetative. In many cases these plants grow as rosettes with no elongation of the
stem.
❑ After vernalization , plant do not necessarily initiate floweing, but acquire the
competence to do that. It is reversible and can be lost as a result to devernalizing
condition, such as high temperature.
 Range and exposure
➢ The effective temperature range for vernalization is from just below freezing to about 10°C, with a
broad optimum usually between about 1 and 7°C.
➢ The effect of cold increases with the duration of the cold treatment until the response is saturated. The
response usually requires several weeks of exposure to low temperature, but the precise duration
varies widely with species and variety.
➢ Vernalization can be lost as a result of exposure to devernalizing conditions, such as high temperature
but the longer the exposure to low temperature, the more permanent the vernalization effect.
➢ A vernalization requirement is often linked to a requirement for a particular photoperiod.
 Site of cold perception Apical meristem

➢ Studies shows that the apical meristem is the site of cold perception during
vernalization and that vernalization causes the meristem to become competent to
flower.
➢ Vernalization appears to take place primarily in the shoot apical meristem.
➢ Even any actively growing region can act as a site of cold perception like- SAM, RAM,
Young leaves, seeds ,apical bud.
➢ Localized cooling causes flowering, when the stem apex is chilled, and this effect
appears to be largely independent of the temperature experienced by the rest of the
plant.
➢ In developmental terms, vernalization results in the acquisition of competence of the
meristem to undergo the floral transition
➢ Later there comes a hypothesis that the vernalin compound is responsible for
vernalization stimulus which produced in plant after vernalization process.
➢ Vernalin get activated at low temperature. Vernaline is hypothetical Plant growth
compound as its not been isolated yet.
➢ Vernalin formed in meristematic region.
 Vernalization :Case Study
▪ Vernalisation→ a process of shortening of juvenile or vegetative phase and Enhancing
flowering by a cold treatment.
▪ Vernalisation does not occur in dry seeds. The seeds must be germinated so that they contain
an active embryo. For this the seeds are moistened before exposing them to low temperature.
In a whole plant, an active meristem is required.
▪ An example of winter rye → When the seeds of this variety of rye were germinated at 1°C for
four weeks, the plants flowered eleven weeks after planting, but at the same time seeds
germinated at 18°C did not produce flowering shoot in the same duration
 Vernalin
▪ The stimulus received by the actively dividing cells of shoot or embryo tip travels to all parts of the plant
and prepare it to flower. The stimulus has been named as vernalin(hypothetical compound)
▪ Attempts to isolate and chemically identify vernalin have not succeeded as yet.
▪ However, Anton Lang have demonstrated that treatment with Gibberellic acid (GA), a plant hormone,
substitute for cold treatment in some species of plants.
 Mechanism of floral induction in vernalized plant
➢ FLOWERING LOCUS C (FLC): FLC is highly expressed in nonvernalized shoot apical regions. After
vernalization, this gene is epigenetically switched off for the remainder of the plant’s life cycle, permitting
flowering in response to long days to occur. FLC gene acts as a repressor of flowering has been identified.
➢ In Arabidopsis it has been shown that FLC works by directly
repressing the expression of the key floral signal FT in the
leaves as well as the transcription factors SOC1 and FD at the
shoot apical meristem.
➢ The epigenetic regulation of FLC involves stable changes in
chromatin structure resulting from chromatin remodeling.
➢ FLC is the flowering repressor that is responsible for the
vernalization requirement in Arabidopsis.
➢ FLC encodes a MADS box protein.
 Detailed mechanism of FLC silencing(additional)
• The epigenetic regulation of FLC involves stable changes in chromatin structure resulting from chromatin
remodeling .Vernalization causes the chromatin of the FLC gene to lose histone modifications characteristic
of euchromatin (transcriptionally active DNA) and to acquire modifications, such as the methylation of
specific lysine residues, characteristic of heterochromatin (transcriptionally inactive DNA). The cold-induced
conversion of FLC from euchromatin to heterochromatin effectively silences the gene.
 FLC EXPRESSION

Winter annual Winter annual Winter annual without cold, but

after 40 days cold without cold with an FLC mutation
FLC is highly expressed in nonvernalized shoot apical meristems
In cereals……
• In cereals, a gene encoding a different type of protein, a zinc finger–containing protein called
VRN2 (vernalization 2), acts as the flowering repressor that creates a vernalization
requirement.
 Mechanism of floral induction in vernalized plant
▪ This process have been identified in Arabidopsis thaliana. some flowering genes are involved in this process.
▪ There are stable changes in the pattern of gene expression in the meristem after cold treatment
▪ FLC gene( Flower locus C)- gene that acts as a repressor of flowering.
▪ FT gene (Flower locus T)- Gene in leaves, florigenic factor (flowering gene)
▪ SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1) – gene in shoot apical meristem (flowering gene)
▪ FD( flower locus D)- Gene in shoot apical meristem (flowering gene)
▪ VRN2 gene is a flowering repressor down-regulated by vernalization.(in winter cereals)
▪ VRN1 and VIN3 is activated by Vernalization and help in flowering.
▪ FRIGIDA(FRI) serves to prevent plants from flowering through the work of FLC.it activate FLC.
Mechanism of floral induction in vernalized plant
 Mechanism of floral induction in vernalized plant
No vernalization Vernalization
VRN1 and VIN3(inactive) VRN1 and VIN3
Active FLC Inactive FLC

Inhibition of FD, FT, SOC1 Activation of FD, FT, SOC1

flowering
No flowering
 The Identification of Florigen-FT
➢ Floral stimulus, referred to as florigen, which serves as the long-distance signal during flowering
➢ The leaf-derived photoperiodic floral stimulus is translocated via the phloem to the shoot apical meristem, where
it promotes floral evocation.
➢ Since the 1930s, there have been many unsuccessful attempts to isolate and characterize florigen.
➢ The Arabidopsis protein FLOWERING LOCUS T (FT) is florigen.
➢ Arabidopsis occurs when the CONSTANS gene is expressed during the light period. CO gene expression appears to
be highest in the companion cells of the phloem of leaves and stems. The downstream target gene of CO,
FLOWERING LOCUS T (FT), is also specifically expressed in the companion cells.
➢ Biochemically, FT is a small globular protein that is related to a family of regulatory proteins conserved
between budding yeast and vertebrates.
➢ Expression of the FT is induced in a range of species during their floral-inductive photoperiods. When the FT
gene is introduced into a range of plant species whose flowering is not influenced by photoperiod, it causes
photoperiod independent flowering.
➢ In addition, FT protein can move from the leaves to the apical meristem and thus exhibits all the properties
that would be expected of florigen.
 The Identification of Florigen-FT(contd.)
➢ FT protein moves via the phloem from the leaves to the meristem under inductive photoperiods. There are two
critical steps in this process: the export of FT from companion cells to the sieve tube elements, and the activation
of FT target genes at the shoot apex, which triggers flower development.
➢ The ER-localized protein, FT INTERACTING PROTEIN1 (FTIP1), is required for FT movement into the phloem
translocation stream, which takes it to the meristem (Figure). Once in the floral meristem, the FT protein enters
the nucleus and forms a complex with FLOWERING D (FD), a basic leucine zipper (bZIP) transcription factor that
is expressed in the meristem. The complex of FT and FD then activates floral identity genes such as APETALA1
(AP1).
➢ After being activated by the FT protein, FD triggers the expression of SOC1 and AP1. Both of these targets
activate LEAFY (LFY; a floral identity gene) and LFY directly activates the expression of AP1 and FD, forming
two positive feedback loops
➢ Because of the action of these positive feedback loops, floral initiation in Arabidopsis is irreversible
Multiple factors regulate flowering in Arabidopsis.
Red arrows indicate the direction of FT transport.
Multiple factors regulate flowering
in Arabidopsis. Red arrows indicate
the direction of FT transport.
Multiple factors regulate flowering
in Arabidopsis. Red arrows indicate
the direction of FT transport.
lfy and fd double mutants fail to form flowers,
highlighting the roles of LFY and FD as floral meristem
identity genes that serve as master regulators for the
initiation of floral development.
 Gibberellins and ethylene can induce flowering
• Among the naturally occurring growth hormones, gibberellins can have a strong influence on flowering.
Exogenous gibberellin can evoke flowering when applied either to rosette LDPs such as Arabidopsis.
• Gibberellin appears to promote flowering in Arabidopsis by activating expression of the LEAFY gene. The
activation of LFY by gibberellin is mediated by the transcription factor GA-MYB, which is negatively regulated by
DELLA proteins. In addition, GA-MYB levels are also modulated by a microRNA that promotes the degradation of
the GA-MYB transcript.
• Exogenously applied gibberellins can also evoke flowering in a few SDPs in noninductive conditions and in cold-
requiring plants that have not been vernalized.
• Gibberellin metabolism in the plant is strongly affected by day length. For example, in the LDP spinach (Spinacia
oleracea), the levels of gibberellins are relatively low in short days, and the plants maintain a rosette form. After
the plants are transferred to long days, the levels of all the gibberellins of the 13-hydroxylated pathway (GA53 →
GA44 →GA19 → GA20 →GA1) increase. However, the fivefold increase in the physiologically active gibberellin,
GA1, is what causes the marked stem elongation that accompanies flowering.
• Promotion of flowering in pineapple (Ananas comosus) by ethylene.

NOTE: Will be discussed in GA Chapter

Mutation: Loss of function of VRN2, whether by natural
mutations or deletions, resulted in spring lines, which do not
require vernalization to flower.
Practice Question
Practice Question
CSIR SEP 2022
 CSIR DEC 2023

A researcher, while studying vernalization in plants, has made the following statements. Choose the
INCORRECT one.
1. Vernalization causes stable change in the competency of the meristem to form an inflorescence in
some plant species.
2. Vernalization causes changes in gene expression that do not involve alteration in the DNA sequence.
3. Flowering is not altered under normal growth conditions in plants defective in vernalization.
4. Vernalization alters the expression of FLC gene.
CSIR DEC 2023

The mobile signal, florigen, that controls the flowering status of the plants in encoded by which
one of the following?
1. Flowering locus C (FLC)
2. Flowering locus D (FD)
3. Flowering locus T (FT)
4. CONSTANS (CO)
KEEP EXPLORING……………..
Educator Name: JYOTI KUMARI (CSIR UGC NET-Life science)
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