# Integrating meristem initiation and floral fate control: toward de novo flower organogenesis in vitro

**Authors:** Niels R. Peeters, Jason Gardiner, Marcel Proveniers

PMC · DOI: 10.3389/fpls.2026.1739266 · Frontiers in Plant Science · 2026-02-16

## TL;DR

This paper explores how to directly induce flowers in plant tissue culture by combining meristem formation with floral development, potentially speeding up plant breeding.

## Contribution

The novel hypothesis is that co-expressing UFO and LFY could bypass vegetative stages and enable direct floral meristem formation in vitro.

## Key findings

- Floral pathway integrators and floral meristem identity genes accelerate flowering but fail to induce direct floral organogenesis.
- UFO, a cofactor of LFY, may be essential for triggering direct floral meristem formation in tissue culture.
- Spatiotemporal coordination of UFO and LFY expression could bypass intermediate developmental stages.

## Abstract

Plant tissue culture is a cornerstone of green biotechnology, facilitating key applications such as genetic transformation and clonal propagation. In tissue culture, meristems serve as the source of regenerative growth, enabling complete plants to develop from small explants or single cells. While tissue culture accelerates propagation, inducing flowering remains a critical bottleneck for breeding programs, as flowering typically requires an extended vegetative phase. The ability to induce direct flowering in vitro, effectively bypassing the vegetative phase, would decouple reproduction from endogenous developmental timing and environmental cues. We frame this approach as de novo flower organogenesis (DNFO), analogous to de novo shoot (DNSO) and root organogenesis (DNRO), to conceptualize strategies for directly inducing floral meristems. DNFO could accelerate breeding cycles, streamline the development of transgenic lines, and enhance experimental throughput. Meristems are typically induced via hormone supplementation or ectopic expression of developmental regulators (DRs). However, regenerated plants still undergo sequential fate transitions before floral induction. This transition is mainly controlled by floral pathway integrators (FPIs), such as FLOWERING LOCUS T (FT) and SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1 (SOC1), and floral meristem identity (FMI) genes, including LEAFY (LFY) and APETALA1 (AP1). In this review, we examine the potential to simultaneously induce meristems and floral fate to enable DNFO in tissue culture. Although constitutive or early expression of FPIs and FMIs accelerates flowering in planta and in vitro, it has so far not been sufficient to induce flowers directly from undifferentiated tissue. We propose that this limitation may reflect the absence of UNUSUAL FLORAL ORGANS (UFO), a cofactor of LFY that is required for the full activation of floral identity programs. We hypothesize that spatiotemporally coordinated co-expression of UFO and LFY could bypass intermediate developmental stages and trigger direct formation of floral meristems, hence flowers, in in vitro culture.

## Linked entities

- **Genes:** FT (PEBP (phosphatidylethanolamine-binding protein) family protein) [NCBI Gene 842859], LFY (floral meristem identity control protein LEAFY (LFY)) [NCBI Gene 836307], AP1 (K-box region and MADS-box transcription factor family protein) [NCBI Gene 843244], UFO (F-box family protein) [NCBI Gene 839982]

## Full-text entities

- **Genes:** TFL1 (PEBP (phosphatidylethanolamine-binding protein) family protein) [NCBI Gene 831683] {aka MED24.6, TERMINAL FLOWER 1, TFL-1}, AGL24 (AGAMOUS-like 24) [NCBI Gene 828556] {aka AGAMOUS-like 24, F22K18.260, F22K18_260}, CLV3 (CLAVATA3) [NCBI Gene 817267] {aka AtCLV3, CLAVATA3, F12K2.17, F12K2_17}, WUS [NCBI Gene 543793], STM (KNOX/ELK homeobox transcription factor) [NCBI Gene 842534] {aka BUM, BUM1, BUMBERSHOOT, BUMBERSHOOT 1, F24O1.38, F24O1_38}, APO1 (APO RNA-binding protein (DUF794)) [NCBI Gene 842789] {aka ACCUMULATION OF PHOTOSYSTEM ONE 1, F13O11.11, F13O11_11}, FT (PEBP (phosphatidylethanolamine-binding protein) family protein) [NCBI Gene 842859] {aka F5I14.3, F5I14_3, FLOWERING LOCUS T, REDUCED STEM BRANCHING 8, RSB8}, NPH4 (Transcriptional factor B3 family protein / auxin-responsive factor AUX/IAA-like protein) [NCBI Gene 832196] {aka ARF7, AUXIN RESPONSE FACTOR 7, AUXIN-REGULATED TRANSCRIPTIONAL ACTIVATOR 7, AUXIN-RESPONSIVE TRANSCRIPTIONAL ACTIVATOR 7, BIP, BIPOSTO}, WOX5 (WUSCHEL related homeobox 5) [NCBI Gene 820297] {aka WOX5B, WUSCHEL related homeobox 5, WUSCHEL related homeobox 5B}, ARF19 (auxin response factor 19) [NCBI Gene 838505] {aka ARF11, AUXIN RESPONSE FACTOR11, IAA22, T29M8.9, T29M8_9, auxin response factor 19}, AG (K-box region and MADS-box transcription factor family protein) [NCBI Gene 827631] {aka AGAMOUS, AGAMOUS PROTEIN, F13C5.130, F13C5_130}, LFY (floral meristem identity control protein LEAFY (LFY)) [NCBI Gene 836307] {aka LEAFY, LEAFY 3, LFY3, MAC9.13, MAC9_13}, TSF (PEBP (phosphatidylethanolamine-binding protein) family protein) [NCBI Gene 827785] {aka F9F13.20, F9F13_20, TWIN SISTER OF FT}, AP1 (K-box region and MADS-box transcription factor family protein) [NCBI Gene 843244] {aka AGAMOUS-like 7, AGL7, APETALA1, AtAP1, F4N2.9, F4N2_9}, AGL8 (AGAMOUS-like 8) [NCBI Gene 836212] {aka AGAMOUS-like 8, FRUITFULL, FUL, MSL3.30, MSL3_30}, SPL (sporocyteless (SPL)) [NCBI Gene 828841] {aka F27G19.11, NOZZLE, NZZ, SPOROCYTELESS}, KNU (C2H2 and C2HC zinc fingers superfamily protein) [NCBI Gene 831249] {aka KNUCKLES, MAC12.2, MAC12_2}, CUC2 (NAC (No Apical Meristem) domain transcriptional regulator superfamily protein) [NCBI Gene 835478] {aka ANAC098, ATCUC2, Arabidopsis NAC domain containing protein 98, CUP-SHAPED COTYLEDON 2, K19P17.12, K19P17_12}, SHR (GRAS family transcription factor) [NCBI Gene 829919] {aka F19F18.140, F19F18_140, SGR7, SHOOT GRAVITROPISM 7, SHORT ROOT}, MIR172a (ncRNA) [NCBI Gene 28718310] {aka MICRORNA 172, MIR172, microRNA172A, p_MI0000215}, REV (Homeobox-leucine zipper family protein / lipid-binding START domain-containing protein) [NCBI Gene 836190] {aka HOMEODOMAIN-LEUCINE ZIPPER PROTEIN, HOMEODOMAIN-LEUCINE ZIPPER PROTEIN IFL1, IFL, IFL1, INTERFASCICULAR FIBERLESS, INTERFASCICULAR FIBERLESS 1}, CLE40 (CLAVATA3/ESR-RELATED 40) [NCBI Gene 831139] {aka CLAVATA3/ESR-RELATED 40, T24H18.160, T24H18_160}, CUC1 (NAC (No Apical Meristem) domain transcriptional regulator superfamily protein) [NCBI Gene 820748] {aka ANAC054, ATNAC1, Arabidopsis NAC domain containing protein 54, CUP-SHAPED COTYLEDON1}, AP2 (Integrase-type DNA-binding superfamily protein) [NCBI Gene 829845] {aka AP22.49, AP22_49, APETALA 2, AtAP2, FL1, FLO2}, ACR4 (ACT domain repeat 4) [NCBI Gene 843236] {aka ACT domain repeat 4, F4N2.2, F4N2_2}, MYB115 (myb domain protein 115) [NCBI Gene 834034] {aka AtMYB115, MPO12.8, MPO12_8, myb domain protein 115}, BZIP27 (basic region/leucine zipper motif 27) [NCBI Gene 816286] {aka ATBZIP27, FD PARALOG, FDP, T17A5.15, T17A5_15, basic region/leucine zipper motif 27}, CAL (K-box region and MADS-box transcription factor family protein) [NCBI Gene 839172] {aka AGAMOUS-like 10, AGL10, CAL1, CAULIFLOWER, F28B23.25, F28B23_25}, LEC2 (AP2/B3-like transcriptional factor family protein) [NCBI Gene 839724] {aka AtLEC2, F3H9.5, F3H9_5, LEAFY COTYLEDON 2}, CLV1 (Leucine-rich receptor-like protein kinase family protein) [NCBI Gene 843915] {aka ATCLV1, CLAVATA 1, CLV1 RECEPTOR KINASE PRECURSOR, FAS3, FASCIATA 3, FLO5}, WUS (Homeodomain-like superfamily protein) [NCBI Gene 816305] {aka PGA6, T27K22.18, T27K22_18, WUS1, WUSCHEL, WUSCHEL 1}, MYB118 (myb domain protein 118) [NCBI Gene 822399] {aka AT3G27780, ATMYB118, PGA37, PLANT GROWTH ACTIVATOR 37, myb domain protein 118}, UFO (F-box family protein) [NCBI Gene 839982] {aka F17F8.16, UNUSUAL FLORAL ORGANS}, SCR (GRAS family transcription factor) [NCBI Gene 824589] {aka SCARECROW, SGR1, SHOOT GRAVITROPISM 1}, LEC1 (Histone superfamily protein) [NCBI Gene 838800] {aka AtLEC1, EMB 212, EMB212, EMBRYO DEFECTIVE 212, LEAFY COTYLEDON 1, NF-YB9}, AGL20 (AGAMOUS-like 20) [NCBI Gene 819174] {aka AGAMOUS-like 20, ATSOC1, F17K2.19, MADS-BOX PROTEIN AGL20, SOC1, SUPPRESSOR OF OVEREXPRESSION OF CO 1}, ATH1 (homeobox protein ATH1) [NCBI Gene 829435] {aka F26P21.100, F26P21_100, homeobox gene 1}, BBM (Integrase-type DNA-binding superfamily protein) [NCBI Gene 831609] {aka BABY BOOM, T10B6.90, T10B6_90}
- **Diseases:** SE (MESH:D013001), SAM (MESH:D010485), VPC (MESH:D000210), floral organ abnormalities (MESH:D009102)
- **Chemicals:** sugar (MESH:D000073893), FMIs (MESH:C000625557), CIM (-), Auxin (MESH:D007210), cytokinin (MESH:D003583)
- **Species:** Medicago sativa (alfalfa, species) [taxon 3879], Petunia (petunia, genus) [taxon 4101], Lotus (genus) [taxon 3867], Powellomyces sp. EA (species) [taxon 252690], Solanum lycopersicum (tomato, species) [taxon 4081], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Oryza sativa (Asian cultivated rice, species) [taxon 4530], Nicotiana benthamiana (species) [taxon 4100]

## Full text

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## Figures

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## References

140 references — full list in the complete paper: https://tomesphere.com/paper/PMC12950686/full.md

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Source: https://tomesphere.com/paper/PMC12950686