In several eudicots, including members of the Asteraceae family, the CYCLOIDEA (CYC) genes, which belong to the TCP class of transcription factors, are key players for floral symmetry. The sunflower inflorescence is heterogamous (radiate capitulum) with sterile monosymmetric ray flowers located in the outermost whorl of the inflorescence and hermaphrodite polysymmetric disk flowers. In inflorescence of Heliantheae tribe, flower primordia development initiates from the marginal ray flowers while disk flowers develop later in an acropetal fashion in organized parastichies along a number found to be one of Fibonacci patterns. Mutants for inflorescence morphology can provide information on the role of CYC-like genes in radiate capitulum evolution. The tubular ray flower (turf) mutant of sunflower shows hermaphrodite ray flowers with a nearly polysymmetric tubular-like corolla. Here, we demonstrate that this mutation is caused by the insertion in the TCP motif of a sunflower CYC-like gene (HaCYC2c) of non-autonomous transposable element (TE), belonging to the CACTA superfamily of transposons. We named this element Transposable element of turf1 (Tetu1). The Tetu1 insertion changes the reading frame of turf-HaCYC2c for the encoded protein and leads to a premature stop codon. Although in Tetu1 a transposase gene is lacking, our results clearly suggest that it is an active TE. The excision of Tetu1 restores the wild type phenotype or generates stable mutants. Co-segregation and sequence analysis in progenies of F(2) and self-fertilized plants derived from reversion of turf to wild type clearly identify HaCYC2c as a key regulator of ray flowers symmetry. Also, HaCYC2c loss-of-function promotes the developmental switch from sterile to hermaphrodite flowers, revealing a novel and unexpected role for a CYC-like gene in the repression of female organs.

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The Helianthus annuus LEAFY COTYLEDON1-LIKE (HaL1L) gene encodes a heme-activated protein 3 subunit of the CCAAT box-binding factor. The phylogenetic analysis indicates that HaL1L is closely related to LEAFY COTYLEDON1 (LEC1)-type of Arabidopsis thaliana. In particular, the peptide results homologous to the LEC1-LIKE gene of A. thaliana, with which it shares a high amino acid sequence identity (56%). HaL1L transcripts are accumulated primarily at an early stage of sunflower embryogenesis. High levels of HaL1L messenger RNA (mRNA) have been detected in the developing embryo proper, suspensor, endosperm, integument, and integumentary tapetum cells, while in unfertilized ovules, HaL1L mRNA was present at rather low levels. In an attempt to examine the involvement of HaL1L on somatic embryogenesis, a somaclonal variant of H. annuus x H. tuberosus (EMB-2) that produces ectopic embryo- and shoot-like structures, arranged in clusters along leaf veins, was used. We found that the epiphyllous proliferation of ectopic embryos on EMB-2 leaves was associated to HaL1L mRNA accumulation. The detection of HaL1L transcripts was evident in somatic embryos at the heart- and early cotyledon-stage. On the contrary, no signal related to HaL1L transcript accumulation was observed in EMB-2 leaves characterized by the presence of shoot-like structures. Together, these results support the conclusion that the transcription of the HaL1L gene is maintained both in zygotic and in somatic embryogenesis. In addition, the ectopic accumulation of HaL1L mRNA in parenchymal cells around the vascular bundles of epiphyllous leaves opens the possibility that HaL1L could also be involved in switching somatic cell fate towards embryogenic competence.

The radiate sunflower inflorescence is composed by zygomorphic ray flowers and actinomorphic disk flowers. Studies performed on mutants identify HaCYC2c, a CYCLOIDEA (CYC)-like gene, as one of the key players controlling flower symmetry in sunflower. turf and tub mutants are characterized by a shift from zygomorphic to actinomorphic ray flowers, caused by insertion of transposable elements (TEs) in HaCYC2c gene. In dbl or Chry mutants, an insertion upstream the coding region of HaCYC2c causes the ectopic expression of the gene and the shift from actinomorphic to zygomorphic disk flowers. We focused on Chry2 mutant: a 1034 bp insertion placed 558 bp before the start codon of HaCYC2c was identified. The insertion is a truncated version of a CACTA TE. Unexpectedly, phenotypic and genetic co-segregation analysis in F2 and F3 progenies derived from the crosses Chry2 × turf and turf × Chry2 demonstrated that CACTA insertion is not always sufficient to alter the expression of HaCYC2c gene and generate Chry2 phenotype. F3 plants homozygous for the CACTA insertion displayed either HaCYC2c transcription pattern identical to wild-type plants or a normal heterogamous inflorescence. Stated these results, we conclude that a much more complex regulatory system stays behind the Chry2 phenotype.

The inflorescence of sunflower (Helianthus annuus L.) is heterogamous with zygomorphic ray flowers located in the outermost whorl of the head and actinomorphic disk flowers arrayed in arcs radiating from the center of the head. Two mutants with altered corolla symmetry have been described. The Chrysanthemoides (Chry) mutant is characterized by a shift from the polysymmetric corolla of disk flowers into a monosymmetric ray-like corolla. The tubular ray flower (turf) mutant is characterized by a shift from the zygomorphic corolla of ray flowers into a nearly actinomorphic tubular-like corolla. We performed a genetic analysis of turf, showing that a single nuclear recessive gene controls the trait. Furthermore, we characterized in detail the morphological floral features of Chry and turf, demonstrating that both mutations also affect the development of stamens and carpels. Most disk flowers found in the peripheral whorls of Chry heads showed drastic reduction in stamen length, as well as absence of ovules, and developed an unbranched style. By contrast, tubular-like ray flowers of turf achieved the ability to differentiate both fertile stamens and ovules. Homeotic transformations were also identified in the tubular-like ray flowers of turf, affecting both filaments and anthers that displayed petaloid-like traits. Our results point to a primary role for TURF and CHRY in the programming of the corolla symmetry and suggest a key interaction of both genes with floral organ identity genes.Résumé : L'inflorescence du tournesol (Helanthus annuus L.) est hétérogame, avec des fleurs à rayons asymétriques localisées sur le verticille le plus externe du capitule, et des fleurs tubuliformes symétriques, sur la surface du disque, arrangées selon des arcs s'irradiant du centre vers la périphérie du capitule. On a déjà décrit deux mutants dont la symétrie de la corolle est altérée. Le mutant Chrysanthémoïde (Chry) se caractérise par un déplacement de la corolle polysymétrique des fleurs du disque vers une corolle monosymétrique d'aspect ligulé. Le mutant à fleur tubuliflore (turf) se caractérise par un déplacement d'une corolle asymétrique, constituée de fleurs ligulées, vers une corolle de fleurs d'aspect tubulé presque symétrique. Les auteurs ont conduit une analyse génétique sur le turf, laquelle montre qu'un simple gène nucléique récessif contrôle ce caractère. De plus, ils ont caractérisé en détail les particularités de la morphologie florale du Chry et du turf, démontrant que les deux mutations affectent également le développement des éta-mines et des carpelles. La plupart des fleurs du disque, localisées dans les verticilles périphériques des capitules Chry, montrent une nette réduction de la longueur des étamines, ainsi qu'une absence d'ovule, et développent un style nonramifié. Par opposition, les fleurs des rayons d'aspect tubulé du turf parviennent à différencier des étamines et des carpelles fertiles. On a également identifié des transformations homéotiques dans les fleurs d'aspect tubulé des rayons du turf, qui affect...

Undoubted lines of evidence point out that members of CYCLOIDEA (CYC) 2 clade are essential players to control flower symmetry and, amusingly, also are determinants of capitula architecture (pseudanthium). In several species, CYC-like genes influence the androecium patterning, but to date, the function of these genes in the development of gynoecium organs is less clear. In this review, we first reported details about floral symmetry and an overview of genes and molecular mechanisms regulating the development of zygomorphism in different angiosperm lineages (e.g., basal and core eudicots and monocots). Then, we paid emphasis on the role of CYC-like genes in the development of heterogamous inflorescence of sunflower as well as other Asteraceae and some species within the Dipsacaceae family. Helianthus annuus is particularly attractive because it represents a useful model to study the role of CYC-like genes on shaping floral corolla as well as the differentiation of reproductive organs in different flowers of pseudanthia. A special attention was reserved to inflorescence morphology mutants of sunflower (i.e., Chrysanthemoids2 and tubular ray flower) because they provide useful information on the role of CYC-like genes in the radiate capitulum evolution. Finally, we discuss data from literature to suggest that CYC-like genes are also co-opted to regulate stamen and carpel differentiation likely throughout their interaction with the cell cycle and flower organ identity genes. The recruitment of reproductive organs in ray flowers also supports the phylogenetic origin of a radiate inflorescence of sunflower from a discoid capitulum and suggests that in sterile zygomorphic ray flower primordia the latent identity to differentiate both microsporangium and macrosporangium was conserved

Plant trichomes are outgrowths developed from an epidermal pavement cells of leaves and other organs. Trichomes (also called ‘hairs’) play well-recognized roles in defense against insect herbivores, forming a physical barrier that obstructs insect movement and mediating chemical defenses. In addition, trichomes can act as a mechanosensory switch, transducing mechanical stimuli (e.g., insect movement) into physiological signals, helping the plant to respond to insect attacks. Hairs can also modulate plant responses to abiotic stresses, such as water loss, an excess of light and temperature, and reflect light to protect plants against UV radiation. The structure of trichomes is species-specific and this trait is generally related to their function. These outgrowths are easily analyzed and their origin represents an outstanding subject to study epidermal cell fate and patterning in plant organs. In leaves, the developmental control of the trichomatous complement has highlighted a regulatory network based on four fundamental elements: (i) genes that activate and/or modify the normal cell cycle of epidermal pavement cells (i.e., endoreduplication cycles); (ii) transcription factors that create an activator/repressor complex with a central role in determining cell fate, initiation, and differentiation of an epidermal cell in trichomes; (iii) evidence that underlines the interplay of the aforesaid complex with different classes of phytohormones; (iv) epigenetic mechanisms involved in trichome development. Here, we reviewed the role of genes in the development of trichomes, as well as the interaction between genes and hormones. Furthermore, we reported basic studies about the regulation of the cell cycle and the complexity of trichomes. Finally, this review focused on the epigenetic factors involved in the initiation and development of hairs, mainly on leaves.

The indeterminate inflorescence of sunflower (Helianthus annuus) is heterogamous with zygomorphic ray flowers in the outer whorl of the head and actinomorphic disc flowers arrayed in arcs radiating from the centre of the head. The Chrysanthemoides (Chry) mutant is characterised by a change of the radially symmetric corolla of tubular disc flowers into a monosymmetric ligulate-like corolla. Zygomorphy is pronounced in the disc flowers placed on the peripheral whorls of inflorescences, while the monosymmetry is less marked toward the centre of the inflorescence. Although the Chry phenotype was one of first known morphological mutants in plants, studies on the genetic control of this trait are scarce and contradictory. Our results indicate that the Chry mutation is semidominant and exclude a maternal influence. Moreover, the data gathered in F 2 , BC 1 and F 3 progenies support a genetic model involving one major locus and an unknown number of modifiers. The improved knowledge on genetic control of the Chry mutation should enhance the introduction of this trait in crossbreeding programmes designed to produce new varieties of ornamental sunflower.