Flower Structure and Developmental Models

Understanding flower development involves recognizing diverse floral structures and the genetic models that explain their formation. Flowers can be classified based on the presence of their parts and their reproductive structures.

A complete flower is one that possesses all four main whorls of floral organs: sepals, petals, stamens, and carpels. The image below illustrates the typical anatomy of such a flower.

Complete Flower Anatomy Diagram

Figure 1: General anatomy of a complete flower, showing key reproductive and accessory organs.

The specification of these floral organs is largely governed by the ABCE model. In the classic ABCE model, distinct classes of homeotic genes interact combinatorially to determine the identity of the four floral organ types arranged in concentric whorls:

  • Sepals: A-function genes alone (plus E-function).
  • Petals: A-function and B-function genes together (plus E-function).
  • Stamens: B-function and C-function genes together (plus E-function).
  • Carpels: C-function genes alone (plus E-function).

The E-function genes (often SEPALLATA genes) are generally required for the proper development of all four whorls. The A-function and C-function genes are typically mutually antagonistic.

ABCE Model and Fading Borders Model

Figure 2: Comparison of the classic ABCE model (A) and the fading borders model (B) of floral organ identity. Modified from Chanderbali et al. (2016).[3]

While the classic ABCE model (Figure 2A) defines sharp boundaries, the "fading borders model" (Figure 2B)[3] suggests more gradual transitions in gene activity, potentially leading to morphologically intergrading organs like tepals, where gene class influence might be lower (indicated by lowercase letters like "aBC").

An incomplete flower is one that lacks one or more of the four main floral whorls. A prime example is found in the Poaceae family (grasses), such as rice. These flowers are highly specialized for wind pollination.

Rice (Grass) Flower Structure

Figure 3: Structure of a rice flower (floret of Poaceae), an example of an incomplete flower adapted for wind pollination.

Key features of a grass floret include:

  • Reduced Perianth: The sepals and petals are typically absent or highly reduced to small, scale-like structures called lodicules, which help in opening the flower.
  • Protective Bracts: The reproductive organs are enclosed by specialized bracts: the lemma and the palea. Additional sterile lemmas or glumes may be present at the base.
  • Wind Pollination Adaptations: Stamens often have long, pendulous filaments and large anthers to release copious amounts of pollen into the wind. Stigmas are typically large, feathery, and branched to effectively capture airborne pollen.

Unisexual flowers (also called imperfect flowers) possess either stamens (male reproductive parts) or carpels/pistils (female reproductive parts), but not both. Pumpkin (Cucurbita pepo) is a classic example of a plant with unisexual flowers, typically being monoecious (having separate male and female flowers on the same plant).

Pumpkin Male and Female Flower Structure

Figure 4: Structure of pumpkin (Cucurbita pepo) male and female flowers, illustrating unisexual flower characteristics.

Distinguishing features:

  • Male Flower: Contains only stamens, which produce pollen. The filaments of stamens in pumpkin are often fused. After releasing pollen, the male flower typically withers.
  • Female Flower: Contains only a pistil (composed of stigma, style, and ovary). A key identifying feature is the presence of a prominent inferior ovary at the base of the flower, which looks like a miniature pumpkin. This ovary, if pollinated, will develop into the fruit.

Pollination in pumpkin is usually carried out by insects, such as bees, which transfer pollen from male flowers to female flowers.

Plants are broadly classified into monocotyledons (monocots) and dicotyledons (dicots or eudicots). Their flower structures often exhibit distinct characteristics. The image below shows a typical dicot flower structure.

Dicotyledon Flower Structure

Figure 5: General structure of a dicotyledon flower.

Typical characteristics of dicot flowers include:

  • Floral Parts in 4s or 5s: The number of sepals, petals, and stamens is often in multiples of four or five.
  • Differentiated Perianth: There is usually a clear distinction between sepals (forming the calyx) and petals (forming the corolla).
  • Net-like Leaf Venation: While not a floral trait, dicots typically have leaves with reticulate (net-like) venation.
  • Taproot System: Dicots often possess a primary taproot.

In contrast, monocot flowers (like grasses or lilies) typically have floral parts in multiples of three, often undifferentiated perianth (tepals), parallel leaf venation, and a fibrous root system.

Epigenetic mechanisms, as discussed below, play a crucial role in establishing and maintaining the precise expression domains of genes like those in the ABCE model, thereby ensuring correct floral patterning across these diverse flower types.

Mechanisms of Epigenetic Regulation

Histone Acetylation in Floral Control

Histone acetylation, generally associated with gene activation, is a dynamic process catalyzed by histone acetyltransferases (HATs) and reversed by histone deacetylases (HDACs). The HAT GENERAL CONTROL NON DEREPRESSIBLE5 (GCN5) has emerged as a key regulator in Arabidopsis flower development.[1]

  • GCN5 and Floral Meristem (FM) Maintenance:
    • GCN5 catalyzes histone acetylation (e.g., H3K9Ac, H3K14Ac) at the chromatin of crucial FM maintenance genes WUSCHEL (WUS) and CLAVATA3 (CLV3), thereby activating their expression.
    • The GCN5ALTERATION/DEFICIENCY IN ACTIVATION 2 (ADA2b) histone acetyltransferase module interacts with the SWI/SNF ATPase SPLAYED (SYD) to form a GCN5-ADA2b-SYD ternary complex.
    • This ternary complex is recruited by cytokinin-responsive type-B Arabidopsis response regulators (ARRs) for WUS activation.
  • GCN5 and FM Determinacy:
    • During floral organogenesis, the transcription factor PERIANTHIA (PAN) recruits the GCN5-ADA2b module for AGAMOUS (AG) activation, which is essential for FM determinacy.
    • GCN5 also activates KNUCKLES (KNU), ensuring timely termination of FM activity.
    • Expression of SUPERMAN (SUP) and CRABS CLAW (CRC), also involved in FM determinacy, is promoted by GCN5.
  • GCN5 and Floral Organ Patterning:
    • GCN5 modulates the expression pattern of the B-class gene APETALA3 (AP3), contributing to the correct formation of reproductive floral organs.
  • Phenotypes: Mutants like gcn5-7 exhibit abnormal flower development, including reduced carpel numbers and varied stamen numbers, highlighting GCN5's indispensable role.[1]

Overall, GCN5 establishes a permissive chromatin environment critical for the timely and spatially accurate expression of genes governing floral meristem activity and organ development.[1]

DNA Methylation and Ethylene Signaling Crosstalk

DNA methylation is another key epigenetic mark involved in regulating gene expression. It typically occurs in CG, CHG, and CHH contexts (where H is A, T, or C). S-adenosylmethionine (SAM) is the primary methyl group donor for DNA methylation, and S-adenosylmethionine synthase (SAMS) is the enzyme responsible for its synthesis.[2]

Overexpression of AtSAMS (SAMOE) in Arabidopsis leads to unexpected floral phenotypes, revealing a complex interplay between SAM metabolism, DNA methylation, and ethylene signaling.[2]

  • SAMOE and DNA Demethylation:
    • Despite increased SAM levels (the methyl donor), SAMOE plants exhibit genome-wide DNA demethylation.
    • This is attributed to the reduced expression of SAM-dependent methyltransferase genes and components of the RNA-directed DNA methylation (RdDM) pathway (e.g., DRM2, DCLs, RDRs, AGO6), alongside increased expression of DNA demethylase genes (ROS1, DME).
  • SAMOE and Ethylene Biosynthesis:
    • SAMOE plants show increased ethylene content. SAM is also a precursor for ethylene.
    • This is linked to upregulated expression of ACC synthase genes (ACS), key enzymes in ethylene biosynthesis.
  • Impact on ABCE Model Genes:
    • DNA demethylation and increased ethylene in SAMOE plants lead to altered expression of ABCE floral homeotic genes:
      • A-function genes (AP1, AP2) and B-function genes (AP3, PI) are down-regulated.
      • C-function gene (AG) and E-function genes (SEP1, SEP2, SEP3) are up-regulated.
    • Methylation changes correlate with expression changes for A, C, and E class genes. However, B-class gene downregulation appears to be independent of direct methylation changes at their loci and is likely mediated by ethylene signaling.
  • Phenotypes: SAMOE plants display abnormal floral organs (e.g., secondary flowers, sepal-to-carpel transformations, altered petal/stamen numbers). Similar phenotypes are observed when wild-type plants are treated with the DNA methylation inhibitor 5-azacytidine (5'-Aza), or with ethephon (ethylene-releasing compound), highlighting the roles of both pathways.[2]

This research indicates a crosstalk mechanism where SAMS-mediated changes in SAM levels impact both DNA methylation landscapes and ethylene signaling, collectively altering ABCE gene expression and leading to floral developmental abnormalities.[2]

Key Epigenetic Regulators and Their Roles

The following table summarizes some of the key genes and protein complexes involved in the epigenetic regulation of flower development discussed in the referenced articles.

Gene/Protein Type / Function Key Role in Flower Epigenetics Primary Reference
GCN5 (GENERAL CONTROL NON DEREPRESSIBLE5) Histone Acetyltransferase (HAT) Catalyzes histone acetylation (H3K9Ac, H3K14Ac) to activate key floral meristem (FM) and organ identity genes (WUS, CLV3, AG, KNU, AP3). [1]
ADA2b (ALTERATION/DEFICIENCY IN ACTIVATION 2b) Adaptor protein, part of SAGA-like complex Forms a module with GCN5, crucial for GCN5's recruitment and activity in FM regulation and AG activation. [1]
SYD (SPLAYED) SWI/SNF Chromatin Remodeling ATPase Interacts with GCN5-ADA2b to form a ternary complex for WUS activation; antagonizes PRC2. [1]
ARRs (Type-B Arabidopsis Response Regulators) Transcription Factors (Cytokinin-responsive) Recruit the GCN5-ADA2b-SYD complex for WUS activation. [1]
PAN (PERIANTHIA) Transcription Factor Recruits the GCN5-ADA2b module for AGAMOUS activation. [1]
SAMS (AtSAMS1/2) S-adenosylmethionine Synthase Produces SAM. Its overexpression leads to DNA demethylation and increased ethylene, affecting ABCE gene expression. [2]
Various DNA Methyltransferases Enzymes (e.g., MET1, CMTs, DRMs) Establish and maintain DNA methylation. Their pathway components (e.g. DRM2) are downregulated in SAMOE plants. [2]
DNA Demethylases (ROS1, DME) Enzymes Actively remove DNA methylation. Upregulated in SAMOE plants, contributing to demethylation. [2]
PRC2 (Polycomb Repressive Complex 2) Histone Methyltransferase Complex Catalyzes H3K27me3 for gene silencing (e.g., involved in KNU-mediated repression of WUS). Antagonistic to SYD. [1]

Key References