Specification of floral meristem identity at the Arabidopsis shoot apex

After floral induction the Arabidopsis shoot apical meristem becomes an indeterminate inflorescence meristem whose lateral primordia must be assigned floral, rather than shoot, fate. Floral integrators supply the input: the florigen FT, a PEBP-family protein made in leaves, reaches the apex and acts as a transcriptional coactivator of the bZIP factor FD to activate APETALA1 (AP1) and the MADS-box integrators SOC1 and FRUITFULL (FUL); SOC1 (as a heterodimer with AGL24) binds the LEAFY (LFY) promoter and activates it, and FUL promotes the reproductive transition partially redundantly with SOC1. LFY, a plant-specific DNA-binding transcription factor, directly induces AP1 in the floral anlagen, and AP1 in turn up-regulates LFY, so the two lock in floral fate through positive feedback. The SQUAMOSA-clade MADS factors AP1, CAULIFLOWER (CAL) and FUL act partially redundantly in this step (ap1 cal ful triple mutants make no flowers); AP1 also acts as a master repressor in the young flower, directly silencing the flowering-time/inflorescence genes SOC1, AGL24 and SVP so that the shoot program does not continue in floral meristems. The F-box protein UNUSUAL FLORAL ORGANS (UFO) binds LFY and DNA to form a LFY-UFO transcriptional complex that redirects LFY to region-specific targets in the young flower. TERMINAL FLOWER1 (TFL1), a PEBP paralog of FT, is expressed in the centre of the inflorescence meristem and acts as an FD-dependent corepressor that keeps LFY and AP1 out of the shoot apex, maintaining indeterminate inflorescence identity; AP1 and LFY reciprocally repress TFL1 in floral primordia. The A-class factor APETALA2 contributes to floral meristem establishment alongside AP1. Floral organ identity (the ABCE program activated later by LFY, UFO and AP1 together with SEP proteins), the leaf-side photoperiod pathway that produces FT, and the age, vernalization and gibberellin pathways converging on the integrators are separate modules.

MODULE:floral_meristem_identityDRAFTCONCRETEDevelopmental Processmodules/floral_meristem_identity.yaml
floral meristem identity specification (closest GO term)GO:0010228 floral meristem determinacyGO:0010582
PMID:10368173
Interactions among APETALA1, LEAFY, and TERMINAL FLOWER1 specify meristem fate.
Floral fate of lateral meristems is assigned by LFY, AP1 and CAL; LFY and AP1 activate each other and both antagonize TFL1, which bars them from the inflorescence meristem.
Therefore, the normally sharp phase transition between the production of leaves with associated shoots and formation of the flowers, which occurs upon floral induction, is promoted by positive feedback interactions between LFY and AP1, together with negative interactions of these two genes with TFL1.
PMID:10648231
Redundant regulation of meristem identity and plant architecture by FRUITFULL, APETALA1 and CAULIFLOWER.
FUL, AP1 and CAL act redundantly; without them LFY is not up-regulated and TFL1 is ectopic.
We demonstrate that this phenotype is caused both by the lack of LFY upregulation and by the ectopic expression of the TERMINAL FLOWER1 (TFL1) gene.
PMID:17428825
Specification of Arabidopsis floral meristem identity by repression of flowering time genes.
AP1 directly represses SVP, AGL24 and SOC1 in emerging floral meristems.
These findings indicate that once AP1 is activated during the floral transition, it acts partly as a master repressor in floral meristems by directly suppressing the expression of flowering time genes, thus preventing the continuation of the shoot developmental program.
file:ARATH/AP1/AP1-ai-review.yaml
AP1 gene review
The AP1 review proposes a new GO term "floral meristem identity specification" (parent GO:0010228); this module uses GO:0010228 as the closest existing process term pending that request.

Concrete Arabidopsis instance. Every annoton is grounded on the Swiss-Prot accession of a reviewed gene in genes/ARATH and its function term is taken from that review's core_functions. GO has no process term for floral meristem identity specification as distinct from determinacy; GO:0010228 (vegetative to reproductive phase transition of meristem) is used as the module concept, with GO:0010582 (floral meristem determinacy) for the floral-fate step, and the AP1 review carries the proposed new term. FUL is modelled twice because it has two separable roles in this module: an integrator role at the shoot apex (with SOC1) and a redundant floral-fate role with AP1/CAL in primordia; ap1 cal ful plants never flower. AGL24 and SVP (SOC1 partners and AP1 repression targets), the 14-3-3 proteins of the florigen activation complex, the SPL/miR156 age-pathway inputs and BOP1/BOP2 are referenced in prose only. FD is included as the DNA-binding partner of FT; FT production in leaves is covered by the photoperiodic_flowering_co_ft module. The ABCE floral organ identity program (AP3/PI/AG/SEP) downstream of LFY, UFO and AP1 is covered by floral_organ_identity_abce. No production GO-CAM in gocams/index.tsv contains these Arabidopsis gene products, so no gocam_associations are given. PANTHER family ids are not asserted; all participants are concrete gene products.

5Nodes
4Parts
0Variant Sets
0Variants
11Annotons
8Connections

Derived QC

Recommended-field compliance

100.0% recommended fields populated

All recommended fields populated.

Module deep research

✗ none found

No MODULE:floral_meristem_identity deep-research report alongside the module YAML.

Leaf nodes lacking representative members

✓ every leaf node grounds to a representative protein.

Template conformance

✓ every declared conforms_to bundle matches its template motif.

Function consistency and core coverage

11 gene-function check(s) reflected in core functions · 0 conflict(s) · 0 evidence or coverage gap(s) (11 comparison(s))

Retained noncore functions are reported separately; they do not require promotion to core functions. Support from representative genes does not establish conservation across the whole family. Missing reviews or identifiers leave a claim unverified.

AnnotonParticipantFunctionAssessment
ft_coactivator FT (At1g65480)
UniProtKB:Q9SXZ2
Gene
transcription coactivator activity
GO:0003713
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
fd_bzip_activator FD (At4g35900)
UniProtKB:Q84JK2
Gene
DNA-binding transcription activator activity, RNA polymerase II-specific
GO:0001228
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
soc1_integrator SOC1 (At2g45660)
UniProtKB:O64645
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
ful_integrator FUL (AGL8, At5g60910)
UniProtKB:Q38876
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
lfy_master_tf LFY (At5g61850)
UniProtKB:Q00958
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
ufo_lfy_cofactor UFO (At1g30950)
UniProtKB:Q39090
Gene
transcription coactivator activity
GO:0003713
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
ap1_floral_fate_tf AP1 (At1g69120)
UniProtKB:P35631
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
cal_redundant_tf CAL (AGL10, At1g26310)
UniProtKB:Q39081
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
ful_redundant_tf FUL (AGL8, At5g60910)
UniProtKB:Q38876
Gene
DNA-binding transcription factor activity, RNA polymerase II-specific
GO:0000981
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
ap2_floral_meristem_establishment AP2 (At4g36920)
UniProtKB:P47927
Gene
DNA-binding transcription factor activity
GO:0003700
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
tfl1_corepressor TFL1 (At5g03840)
UniProtKB:P93003
Gene
transcription corepressor activity
GO:0003714
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).

Feedback loops (advisory, prose-driven: a regulator counts as "feedback" when its own description says so)

✓ every negative regulator described as feedback has an upstream edge closing the loop.

  • ap1_floral_fate ⊣ integrator_input [closed]
    ap1_floral_fate is regulated by integrator_input, lfy_program: feedback loop closed
  • tfl1_inflorescence_maintenance ⊣ lfy_program [input]
    tfl1_inflorescence_maintenance is a negative regulator with no upstream edge (not described as feedback)

Gene-review completeness (10/10 grounded genes reviewed)

10 complete review(s) · 6 with deep research · 0 missing review · 4 reviewed but lacking deep research

Gene Review Complete Deep research
AP1 P35631 ✓ ✓ ✓
AP2 P47927 ✓ ✓ ✓
CAL Q39081 ✓ ✓ ✗
FD Q84JK2 ✓ ✓ ✗
FT Q9SXZ2 ✓ ✓ ✓
FUL Q38876 ✓ ✓ ✗
LFY Q00958 ✓ ✓ ✓
SOC1 O64645 ✓ ✓ ✓
TFL1 P93003 ✓ ✓ ✓
UFO Q39090 ✓ ✓ ✗

Details

Context
Arabidopsis thalianaNCBITaxon:3702
nucleusGO:0005634
Floral meristem identity specificationDevelopmental Processfloral_meristem_identity
floral meristem identity specification (closest GO term)GO:0010228 floral meristem determinacyGO:0010582
Context
Arabidopsis thalianaNCBITaxon:3702
nucleusGO:0005634

Connections

integrator_input -> lfy_program Positively Regulates
SOC1 (with AGL24) binds and activates the LFY promoter.
PMID:18466303
SOC1-AGL24 heterodimerization activates LFY.
Thus, we propose that heterodimerization of SOC1 and AGL24 is a key mechanism in activating LFY expression.
integrator_input -> ap1_floral_fate Positively Regulates
The FT-FD complex activates AP1 at the shoot apex, independently of LFY.
PMID:16099979
FD-FT activate AP1 transcription.
FD and FT are interdependent partners through protein interaction and act at the shoot apex to promote floral transition and to initiate floral development through transcriptional activation of a floral meristem identity gene, APETALA1 (AP1).
lfy_program -> ap1_floral_fate Positively Regulates
LFY directly induces early AP1 expression in floral anlagen.
PMID:10417387
LFY directly activates AP1.
Use of steroid-inducible activation of LFY demonstrated that early expression of AP1 is a result of transcriptional induction by LFY.
ap1_floral_fate -> lfy_program Positively Regulates
AP1 (with CAL and FUL) up-regulates LFY, closing the LFY-AP1 positive feedback loop that makes the switch to floral fate sharp.
PMID:10368173
AP1 positively regulates LFY.
We have found that AP1, in turn, can positively regulate LFY, because LFY is expressed prematurely in the converted floral meristems of plants constitutively expressing AP1.
ap1_floral_fate -> integrator_input Negatively Regulates
Once induced, AP1 directly represses the integrators SOC1 (and AGL24/SVP) in emerging floral meristems, preventing continuation of the shoot program in flowers.
PMID:17428825
AP1 directly represses SVP, AGL24 and SOC1.
We show here that the emerging floral meristems require AP1 to partly specify their floral identities by directly repressing a group of flowering time genes, including SHORT VEGETATIVE PHASE (SVP), AGAMOUS-LIKE 24 (AGL24) and SUPPRESSOR OF OVEREXPRESSION OF CO1 (SOC1).
AP1 down-regulates TFL1; AP1/CAL/FUL keep TFL1 out of floral primordia.
PMID:10368173
Constitutive AP1 down-regulates TFL1.
We show here that this negative regulation can be mutual because TFL1 expression is downregulated in plants constitutively expressing AP1.
TFL1 bars LFY expression from the inflorescence shoot meristem.
PMID:10368173
TFL1 excludes AP1 and LFY from the inflorescence meristem.
Shoot meristems maintain an identity distinct from that of flower meristems, in part through the action of genes such as TERMINAL FLOWER1 (TFL1), which bars AP1 and LFY expression from the influorescence shoot meristem.
TFL1 bars AP1 expression from the inflorescence shoot meristem.
PMID:10368173
TFL1 excludes AP1 and LFY from the inflorescence meristem.
Shoot meristems maintain an identity distinct from that of flower meristems, in part through the action of genes such as TERMINAL FLOWER1 (TFL1), which bars AP1 and LFY expression from the influorescence shoot meristem.
Part 1: floral integrator input at the shoot apex
Floral integrators activate meristem identity genesRegulatory Stepintegrator_input

At the shoot apex the FT-FD complex and the MADS-box integrators SOC1 and FUL convert flowering signals into transcriptional activation of LFY and AP1.

Annotons

FT: florigen, coactivator of FD
ft_coactivator
Participant: Gene Product: FT (At1g65480)
Gene Product:
FT (At1g65480)UniProtKB:Q9SXZ2

Function

transcription coactivator activityGO:0003713

Processes

positive regulation of flower developmentGO:0009911

Locations

nucleusGO:0005634

Mobile florigen that, once at the apex, partners with FD to activate floral identity genes such as AP1.

PMID:16099980
An FT-FD complex activates floral identity genes such as AP1.
A complex of FT and FD proteins in turn can activate floral identity genes such as APETALA1 (AP1).
FD: bZIP DNA-binding partner of FT
fd_bzip_activator
Participant: Gene Product: FD (At4g35900)
Gene Product:
FD (At4g35900)UniProtKB:Q84JK2

Function

DNA-binding transcription activator activity, RNA polymerase II-specificGO:0001228

Processes

positive regulation of flower developmentGO:0009911

Locations

nucleusGO:0005634

Apex-expressed bZIP factor that provides the DNA-binding moiety for FT (and TFL1) action.

PMID:16099979
FD and FT act together at the apex to activate AP1.
FD and FT are interdependent partners through protein interaction and act at the shoot apex to promote floral transition and to initiate floral development through transcriptional activation of a floral meristem identity gene, APETALA1 (AP1).
SOC1: MADS-box integrator, activator of LFY
soc1_integrator
Participant: Gene Product: SOC1 (At2g45660)
Gene Product:
SOC1 (At2g45660)UniProtKB:O64645

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

vegetative to reproductive phase transition of meristemGO:0010228

Locations

nucleusGO:0005634

Binds the LFY promoter (as a SOC1-AGL24 heterodimer that enters the nucleus) and activates LFY expression.

PMID:18466303
SOC1 binds the LFY promoter directly.
Here, we show that SOC1 directly binds to the distal and proximal region of the LFY promoter where critical cis-elements are located.
FUL: MADS-box integrator at the shoot apex
ful_integrator
Participant: Gene Product: FUL (AGL8, At5g60910)
Gene Product:
FUL (AGL8, At5g60910)UniProtKB:Q38876

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

vegetative to reproductive phase transition of meristemGO:0010228

Locations

nucleusGO:0005634

Rapidly induced at the apex by flowering signals; promotes the meristem identity transition partially redundantly with SOC1 (FUL-SVP and FUL-SOC1 heterodimers).

PMID:24465009
FUL promotes the reproductive transition redundantly with SOC1.
A model is proposed where the sequential formation of FUL-SVP and FUL-SOC1 heterodimers may mediate the vegetative and meristem identity transitions
Part 2: LEAFY activation and LFY-UFO complex
LEAFY floral meristem identity programRegulatory Steplfy_program

LFY, expressed throughout young floral primordia, directly activates AP1 at the stage when floral fate is assumed; UFO binds LFY and DNA to redirect LFY to additional cis-elements in specific regions of the young flower.

Annotons

LFY: floral meristem identity transcription factor
lfy_master_tf
Participant: Gene Product: LFY (At5g61850)
Gene Product:
LFY (At5g61850)UniProtKB:Q00958

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

floral meristem determinacyGO:0010582

Locations

nucleusGO:0005634

Directly induces early AP1 expression in floral anlagen; LFY activity levels set the threshold for flower initiation.

PMID:10417387
Early AP1 expression results from direct transcriptional induction by LFY.
Use of steroid-inducible activation of LFY demonstrated that early expression of AP1 is a result of transcriptional induction by LFY.
UFO: F-box transcriptional cofactor of LFY
ufo_lfy_cofactor
Participant: Gene Product: UFO (At1g30950)
Gene Product:
UFO (At1g30950)UniProtKB:Q39090

Function

transcription coactivator activityGO:0003713

Processes

regulation of DNA-templated transcriptionGO:0006355

Locations

nucleusGO:0005634

Forms a LFY-UFO-DNA complex that redirects LFY to new cis-elements; required for full floral meristem identity and for region-specific LFY targets such as AP3. Its SCF (ubiquitin ligase adaptor) association is largely dispensable for this role.

PMID:36732360
UFO forms an active transcriptional complex with LFY at new regulatory elements.
we establish that UFO instead acts by forming an active transcriptional complex with LFY at newly discovered regulatory elements.
PMID:7780306
ufo mutants show defects in floral meristem identity.
UFO is required for the proper identity of the floral meristem and acts in three different aspects of the process that distinguishes flowers from shoots.
Part 3: AP1/CAL/FUL floral fate and repression of the shoot program
AP1/CAL/FUL establish floral fateRegulatory Stepap1_floral_fate

The SQUAMOSA-clade MADS factors AP1, CAL and FUL act partially redundantly in floral primordia to up-regulate LFY and to exclude TFL1; AP1 directly represses SOC1, AGL24 and SVP so that floral meristems do not revert to shoot identity. AP2 contributes to establishing the floral meristem.

Annotons

AP1: floral meristem identity MADS factor and repressor of flowering-time genes
ap1_floral_fate_tf
Participant: Gene Product: AP1 (At1g69120)
Gene Product:
AP1 (At1g69120)UniProtKB:P35631

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

floral meristem determinacyGO:0010582

Locations

nucleusGO:0005634

Binds cis-regulatory regions of SVP, AGL24 and SOC1 and represses them in emerging floral meristems; up-regulates LFY.

PMID:17428825
AP1 binds and represses flowering-time genes in floral meristems.
By post-translational activation of an AP1-GR fusion protein and chromatin immunoprecipitation assays, we further demonstrate the repression of these flowering time genes by induced AP1 activity and in vivo AP1 binding to the cis-regulatory regions of these genes.
CAL: AP1 paralog, partially redundant floral meristem identity factor
cal_redundant_tf
Participant: Gene Product: CAL (AGL10, At1g26310)
Gene Product:
CAL (AGL10, At1g26310)UniProtKB:Q39081

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

floral meristem determinacyGO:0010582

Locations

nucleusGO:0005634

Expressed in young flower primordia; ap1 cal double mutants form reiterating inflorescence meristems in place of flowers.

PMID:7824951
CAL and AP1 are partially redundant in floral meristem formation.
Genetic studies demonstrate that two Arabidopsis genes, CAULIFLOWER and APETALA1, encode partially redundant activities involved in the formation of floral meristems, the first step in the development of flowers.
FUL: redundant floral-fate MADS factor with AP1/CAL
ful_redundant_tf
Participant: Gene Product: FUL (AGL8, At5g60910)
Gene Product:
FUL (AGL8, At5g60910)UniProtKB:Q38876

Function

DNA-binding transcription factor activity, RNA polymerase II-specificGO:0000981

Processes

positive regulation of flower developmentGO:0009911

Locations

nucleusGO:0005634

Contributes to LFY up-regulation and TFL1 exclusion together with AP1 and CAL; ap1 cal ful plants elaborate leafy shoots in place of flowers.

PMID:10648231
ful ap1 cal triple mutants do not flower.
We have found that mutations in the FRUITFULL (FUL) MADS-box gene, when combined with mutations in AP1 and CAL, lead to a dramatic non-flowering phenotype in which plants continuously elaborate leafy shoots in place of flowers.
AP2: AP2/ERF factor contributing to floral meristem establishment
ap2_floral_meristem_establishment
Participant: Gene Product: AP2 (At4g36920)
Gene Product:
AP2 (At4g36920)UniProtKB:P47927

Function

DNA-binding transcription factor activityGO:0003700

Processes

flower developmentGO:0009908

Locations

nucleusGO:0005634

Accessory contributor; better known for A-class organ identity and AG repression (organ identity module).

PMID:7919989
AP2 participates in establishment of the floral meristem.
APETALA2 (AP2) plays a central role in the establishment of the floral meristem, the specification of floral organ identity, and the regulation of floral homeotic gene expression in Arabidopsis.

Optional/accessory member of this module.

Part 4: maintenance of inflorescence meristem identity
TFL1 maintains inflorescence meristem identityRegulatory Steptfl1_inflorescence_maintenance

TFL1, expressed in the centre of the inflorescence meristem, represses FD-dependent targets and keeps LFY and AP1 out of the shoot apex, so the main meristem stays indeterminate.

Annotons

TFL1: FD-dependent transcriptional corepressor
tfl1_corepressor
Participant: Gene Product: TFL1 (At5g03840)
Gene Product:
TFL1 (At5g03840)UniProtKB:P93003

Function

transcription corepressor activityGO:0003714

Processes

maintenance of inflorescence meristem identityGO:0010077

Locations

nucleusGO:0005634

Antagonist of FT; interacts with FD and represses genes that FT activates, preventing terminal flower formation.

PMID:21890645
TFL1 represses FD-dependent transcription of FT-activated genes.
We conclude that TFL1 negatively modulates the FD-dependent transcription of target genes to fine-tune flowering time and the development of the inflorescence meristem.