The spatial separation between the site of perception of the photoperiodic signal (LEAVES) and the site of flowering

(SHOOT APICAL MERISTEM) logically requires a transmissible signal that carries the information from the leaf to the shoot apex.

Hypothesis of a Floral Hormone

Suggested by Russian plant physiologist Mikhail Chailakhyan in 1936 FLORIGEN - hormone that stimulates flowering - term used to describe the hypothesized hormone-like molecules responsible for controlling and/or triggering flowering in plants

Xanthium strumarium (cocklebur)

Short Day Plant

Grafting - a horticultural technique whereby tissues from one plant are inserted into those of another so that the two sets of vascular tissues may join together.

Hypothesis of a Floral Hormone

In Arabidopsis thaliana, florigen is likely the Flowering Locus T (FT) protein.

In some short day cereals (such as rice) florigen is likely a protein called Hd3a.

Hypothesis of a Floral Hormone

Mechanism: 1. Initiation (photoperiod-regulated initiation) - In Arabidopsis thaliana, the signal is initiated by the production of messenger RNA (mRNA) coding a transcription factor called CONSTANS (CO). - CO protein promotes transcription of another gene called Flowering Locus T (FT).

Hypothesis of a Floral Hormone

Mechanism: 2. Translocation (signal translocation via the phloem) - The FT protein resulting from the short period of CO transcription factor activity is then transported via the phloem to the shoot apical meristem.

Hypothesis of a Floral Hormone

Mechanism: 3. Flowering (induction of flowering at the shoot apical meristem) - The FT protein interacts with FD protein to activate floral identity genes, thus inducing flowering. - Specifically, arrival of FT at the shoot apical meristem and formation of the FT/FD heterodimer is followed by the increased expression of at least one direct target gene, APETALA 1 (AP1), along with other targets, such as SOC1 and several SPL genes, which are targeted by a microRNA.

FT protein
-acts as a long range signal

FD protein
-protein mediating signals from the floral pathway integrator FT at the shoot apex

APETALA 1 gene
-gene that specifies flower meristem identity and is also required for normal development of sepals and petals

SOC1 gene
- triggers LFY gene expression

LFY gene
- gene that causes meristems to develop into flowers instead of shoots

FLC
- responsible for vernalization - negative regulator of SOC1

Low Temperature Requirement of Winter Annuals and Biennial Flowering Plants

Vernalization = the acquisition of a plant's ability to flower or germinate in the spring by exposure to the prolonged cold of winter

Winter Cereals
Winter Strains = will not normally flower during a single growing season; but must be planted in the fall in order to flower and produce a crop the following year Spring Strains = will flower and produce grain in the same year they are planted

Winter Cereals (Experiment)

Spring strain of Rye (Secale cereale var. Petkus) = Long day plant = under long days, flowers are initiated after 7 leaves have been produced = under short days, flowers will not appear until 22 leaves have been produced

Winter Cereals (Experiment)

Spring strain of Rye (Secale cereale var. Petkus) = germinated and grown at normal temperature (18C) is not a LD plant and flowers only after 22 leaves have been produced regardless of photoperiod = if subjected at low temperature (1C) for several weeks, it will flower early in response to long days just as spring strain does

Hyoscyamus niger (Black Henbane)

Annual strain = typical Long day plant = grows in a rosette habit under short day, but undergoing extensive stem elongation (BOLTING) and flowering under long days

Hyoscyamus niger (Black Henbane)

Biennial strain = it has a cold requirement that must be satisfied before it will bolt and flower as a long day plant = optimal temperature is 3-17C

 Some day neutral plants have a cold requirement if satisfied, flowering will proceed regardless of photoperiod. Example: Chrysanthemum Campanula medium

Gibberellin as Substitute for Vernalization

Gibberellins will normally substitute for cold requirement in vernalizable long day plants and for long days in other LD plants. If such plants are treated with gibberellins, they produce flowers without subjecting the plants to cold and photoperiodic treatments.

Gibberellin as Substitute for Vernalization

Example: Henbane is a rosette-leaved long day plant which requires cold treatment for flowering. - If such untreated plants are sprayed with gibberellins, the plants produce flowers.

Gibberellin as Substitute for Vernalization

Gibberellins do not usually substitute for short days in Short Day plants. Exception: Campanula