Search

The shoot concept of the flower suggests that flowers correspond to vegetative short-shoots except the fact that their lateral appendages are floral and not vegetative leaves. However, in view of the different properties of vegetative and flower meristems, this concept should be questioned. Differential meristem activity resulting in tubes, hypanthia and inferior ovaries, continuous meristem expansion providing space for stamen fascicles and additional structures and the process of (repeated) fractionation using a given space completely, are characteristics of flower meristems hardly explainable with the shoot concept. Linking instead flower development with recent findings in molecular biology and computational modeling widens the view to the fundamental relation between growth and form. Given that the same general principles characterize plant growth at all life stages, the loss of apical growth appears to play the major role in changing geometry, space availability and genetic regulation in flower meristems. The flower, thus, turns out to be the sporangia bearing tip of a shoot.

Pepper flowers are hermaphroditic; the plant's male sterility trait is characterized by its inability to produce pollen grains. In the ABC model of flower development, B-function genes play roles in petal and stamen development in the angiosperm. In this study, a B-class gene designated as PAP3 (GenBank accession no. HM104635) was isolated in pepper. The gene encoded 226 amino acids and shared high similarity with the MADS-box protein family, with a conservative MADS domain and semiconservative K domain. Furthermore, the expression of PAP3 was abundant only in petals and anthers but not in leaves. A functional study employing virus-induced gene silencing (VIGS) showed that knockdown of PAP3 led to the shriveling of pollen grains and male sterility; however, it did not affect petal development. These results suggest an essential role for PAP3 in the development of the pepper stamen and in contributing to the variation of floral traits.

During reproductive development, rice plants develop unique flower organs which determine the final grain yield. OsMADS1, one of SEPALLATA-like MADS-box genes, has been unraveled to play critical roles in rice floral organ identity specification and floral meristem determinacy. However, the molecular mechanisms underlying interactions of OsMADS1 with other floral homeotic genes in regulating flower development remains largely elusive. In this work, we studied the genetic interactions of OsMADS1 with B-, C-, and D-class genes along with physical interactions among their proteins. We show that the physical and genetic interactions between OsMADS1 and OsMADS3 are essential for floral meristem activity maintenance and organ identity specification; while OsMADS1 physically and genetically interacts with OsMADS58 in regulating floral meristem determinacy and suppressing spikelet meristem reversion. We provided important genetic evidence to support the neofunctionalization of two rice C-class genes (OsMADS3 and OsMADS58) during flower development. Gene expression profiling and quantitative RT-PCR analyses further revealed that OsMADS1 affects the expression of many genes involved in floral identity and hormone signaling, and chromatin immunoprecipitation (ChIP)–PCR assay further demonstrated that OsMADS17 is a direct target gene of OsMADS1. Taken together, these results reveal that OsMADS1 has diversified regulatory functions in specifying rice floral organ and meristem identity, probably through its genetic and physical interactions with different floral homeotic regulators.

Auxin and cytokinin regulate different critical processes involved in plant growth and environmental feedbacks. These plant hormones act either synergistically or antagonistically to control the organisation, formation and maintenance of meristem. Meristem cells can be divided to generate new tissues and organs at the locations of plant postembryonic development. The aboveground plant organs are created by the shoot apical meristem (SAM). It has been proposed that the phytohormone, cytokinin, plays a positive role in the shoot meristem function, promotes cell expansion and promotes an increasing size of the meristem in Arabidopsis, whereas it has the reverse effects in the root apical meristem (RAM). Over the last few decades, it has been believed that the apically derived auxin suppresses the shoot branching by inactivating the axillary buds. However, it has recently become clear that the mechanism of action of auxinis indirect and multifaceted. In higher plants, the regulatory mechanisms of the SAM formation and organ separation are mostly unknown. This study reviews the effects and functions of cytokinin and auxin at the shoot apical meristem. This study also highlights the merger of the transcription factor activity with the actions of cytokinin/auxin and their complex interactions with the shoot meristem in rice.

Flower development is one of the major developmental processes that governs seed setting in angiosperms. However, little is known about the molecular mechanisms underlying flower development in legumes. Employing RNA-seq for various stages of flower development and few vegetative tissues in chickpea, we identified differentially expressed genes in flower tissues/stages in comparison to vegetative tissues, which are related to various biological processes and molecular functions during flower development. Here, we provide details of experimental methods, RNA-seq data (available at Gene Expression Omnibus database under GSE42679) and analysis pipeline published by Singh and colleagues in the Plant Biotechnology Journal (Singh et al., 2013), along with additional analysis for discovery of genes involved in shoot apical meristem (SAM) development. Our data provide a resource for exploring the complex molecular mechanisms underlying SAM and flower development and identification of gene targets for functional and applied genomics in legumes.

Plant apical meristems (AMs) grow continuously by delicately balancing cells leaving at the periphery to form lateral organs with slowly dividing central domain cells that replenish reservoirs of pluripotent cells. This balance can be modified by signals originating from within and outside the meristem, and their integration results in a gradual maturation process that often culminates with the meristem differentiating into a flower. Accompanying this 'meristem maturation' are changes in spacing and size of lateral organs and in rates at which lateral meristems are released from apical dominance. Modulation of distinct meristem maturation parameters through environmental and genetic changes underlies the remarkable diversity of shoot architectures. Here, we discuss recent studies relating the dynamics of meristem maturation with organization of floral branching systems--inflorescences--in the nightshades. From this context, we suggest general principles on how factors coordinating meristem maturation impact shoot organization more broadly.

A key feature of the development of a higher plant is the continuous formation of new organs from the meristems. Originally patterned during embryogenesis, the meristems must activate cell division de novo at the time of germination, in order to initiate post-embryonic development. In a mutagenesis screen aimed at finding new players in early seedling cell division control, we identified ELONGATA3 (ELO3) as a key regulator of meristem cell cycle activation in Arabidopsis. Our results show that plants carrying a hypomorphic allele of ELO3 fail to activate cell division in the meristems following germination, which leads to seedling growth arrest and lethality. Further analyses suggest that this is due to a failure in DNA replication, followed by cell cycle arrest, in the meristematic tissue. Interestingly, the meristem cell cycle arrest in elo3 mutants, but not the later leaf developmental defects that have been linked to the loss of ELO3 activities, can be relieved by the addition of metabolic sugars in the growth medium. This finding points to a new role by which carbohydrate availability promotes meristem growth. Furthermore, growth arrested elo3 mutants suffer a partial loss of shoot meristem identity, which provides further evidence that cell cycle activities can influence the control of tissue identity.

A long-standing question in plants and animals is how spatial patterns are maintained within stem cell niches despite ongoing cell divisions. Here we address how, during shoot meristem formation in Arabidopsis thaliana, the three apical cell layers acquire stem cell identity. Using a sensitized mutant screen, we identified miR394 as a mobile signal produced by the surface cell layer (the protoderm) that confers stem cell competence to the distal meristem by repressing the F box protein LEAF CURLING RESPONSIVENESS. This repression is required to potentiate signaling from underneath the stem cells by the transcription factor WUSCHEL, maintaining stem cell pluripotency. The interaction of two opposing signaling centers provides a mechanistic framework of how stem cells are localized at the tip of the meristem. Although the constituent cells change, the surface layer provides a stable point of reference in the self-organizing meristem.

Alstroemeria sp. cv. Green Coral has numerous bracts instead of flowers, and its cyme structures are repeated eternally. Observations of the development and morphology of inflorescence in cv. Green Coral revealed that transition from inflorescence to floral meristem was restricted. We isolated and characterized floral meristem identity genes LEAFY-like (AlsLFY) and SQUAMOSA-like (AlsSQa and AlsSQb) genes from Alstroemeria ligtu. In situ hybridization results indicated that AlsSQa and AlsSQb were expressed in the dome-shaped floral meristems and all floral organ primordia in A. ligtu. Transcripts of AlsLFY accumulated early in the dome-shaped floral meristems; the signals were restricted later to the outer region of the floral meristem. These results indicate that AlsLFY, AlsSQa, and AlsSQb function as floral meristem identity genes. Expression profiles of AlsLFY, AlsSQa, AlsSQb, and other MADS-box genes were compared between A. ligtu and cv. Green Coral. AlsLFY, AlsDEFa, and AlsAGL6 transcripts were not detected at the shoot apices of cv. Green Coral but were detected in A. ligtu. The early induction and accumulation of AlsLFY transcripts in the inflorescence meristem of A. ligtu prior to development of the floral meristem suggest that downregulation of AlsLFY is likely to restrict the inflorescence-to-floral meristem transition in cv. Green Coral.

VFL and VvTFL1 genes expression patterns and the effects of sucrose on the expression of VFL and VvTFL1 genes in different organs of the “Xiangfei” grapevine (Vitis vinifera L.) were investigated. VvTFL1 gene expression was detected in the meristem of the apical bud and lateral bud, but was not detected during inflorescence differentiation and flower organ development. After sucrose treatment, VvTFL1 gene expression increased in the apical bud, but decreased in the lateral bud. These results suggested that the VvTFL1 gene might be mainly involved in the apical growth process of shoots, and exogenous sucrose had an effect on the VvTFL1 gene by increasing shoot apical meristem initiation of apical buds. The VFL gene was expressed primarily during inflorescence differentiation and early flower organ development, but it gradually reduced in later flower development. After sucrose treatment, VFL gene expression increased in the inflorescence and small or middle flower, but a little change was seen in the large flower. These results suggested that the VFL gene plays important roles in the initiation of inflorescence meristems and the morphological formation of flower organs. Exogenous sucrose had an effect on VFL gene expression at the early stage of flower development.

Floral homeotic genes encode transcription factors and act as master regulators of flower development. The homeotic protein complex is expressed in a specific whorl of the floral primordium and determines floral organ identity by the combinatorial action. Homeotic proteins continue to be expressed until late in flower development to coordinate growth and organogenesis. Recent genomic studies have shown that homeotic proteins bind thousands of target sites in the genome and regulate the expression of transcription factors, chromatin components and various proteins involved in hormone biosynthesis and signaling and other physiological activities. Further, homeotic proteins program chromatin to direct the developmental coordination of stem cell maintenance and differentiation in shaping floral organs.

The shoot apical meristem of angiosperm plants generates leaf, stem and floral structures throughout the plant's lifetime. To do this, the plant must maintain a population of stem cells within the meristem while at the same time carefully controlling the placement and establishment of new leaf primordia. As there is little cell rearrangement in plants, underlying patterning mechanisms must exert careful control of cell division rates and orientations to achieve the correct final form. It has been twenty years since the first genes controlling meristem development were molecularly cloned. In the intervening decades, our understanding of the inner workings directing meristem development has increased enormously. This review summarizes our current knowledge of how the meristem functions as a persistent organ generating center. The story that emerges is one in which transcription factor activity combines with the action of the classic plant growth regulators auxin and cytokinin and with the action of more recently discovered small peptides to control proliferation and cell fate in the shoot apical meristem.

20 years after establishment of the ABC model many of the molecular mechanisms underlying development of the angiosperm flower are relatively well understood. Central players in the gene regulatory network controlling flower development are SQUA-like, DEF/GLO-like, AG-like and AGL6/SEP1-like MIKC-type MADS-domain transcription factors. These provide class A, class B, class C and the more recently defined class E floral homeotic functions, respectively. There is evidence that the floral homeotic proteins recognize the DNA of target genes in an organ-specific way as multimeric protein complexes, thus constituting 'floral quartets'. In contrast to the detailed insights into flower development, how the flower originated during evolution has remained enigmatic. However, while orthologues of all classes of floral homeotic genes appear to be absent from all non-seed plants, DEF/GLO-like, AG-like, and AGL6-like genes have been found in diverse extant gymnosperms, the closest relatives of the angiosperms. While SQUA-like and SEP1-like MADS-box genes appear to be absent from extant gymnosperms, reconstruction of MADS-box gene phylogeny surprisingly suggests that the most recent common ancestor of gymnosperms and angiosperms possessed representatives of both genes, but that these have been lost in the lineage that led to extant gymnosperms. Expression studies and genetic complementation experiments indicate that both angiosperm and gymnosperm AG-like and DEF/GLO-like genes have conserved functions in the specification of reproductive organs and in distinguishing male from female organs, respectively. Based on these findings novel models about the molecular basis of flower origin, involving changes in the expression patterns of DEF/GLO-like or AGL6/SEP1/SQUA-like genes in reproductive structures, were developed. While in angiosperms SEP1-like proteins play an important role in floral quartet formation, preliminary evidence suggests that gymnosperm DEF/GLO-like and AG-like proteins alone can already form floral quartet-like complexes, further corroborating the view that the formation of floral quartet-like complexes predated flower origin during evolution.

During the floral transition the shoot apical meristem changes its identity from a vegetative to an inflorescence state. This change in identity can be promoted by external signals, such as inductive photoperiod conditions or vernalization, and is accompanied by changes in expression of key developmental genes. The change in meristem identity is usually not reversible, even if the inductive signal occurs only transiently. This implies that at least some of the key genes must possess an intrinsic memory of the newly acquired expression state that ensures irreversibility of the process. In this review, we discuss different molecular scenarios that may underlie a molecular memory of gene expression.

EMBRYONIC FLOWER (EMF) genes are required to maintain vegetative development via repression of flower homeotic genes in Arabidopsis. Removal of EMF gene function caused plants to flower upon germination, producing abnormal and sterile flowers. The pleiotropic effect of emf1 mutation suggests its requirement for gene programs involved in diverse developmental processes. Transgenic plants harboring EMF1 promoter::glucuronidase (GUS) reporter gene were generated to investigate the temporal and spatial expression pattern of EMF1. These plants displayed differential GUS activity in vegetative and flower tissues, consistent with the role of EMF1 in regulating multiple gene programs. EMF1::GUS expression pattern in emf mutants suggests organ-specific auto-regulation. Sense- and antisense (as) EMF1 cDNA were expressed under the control of stage- and tissue-specific promoters in transgenic plants. Characterization of these transgenic plants showed that EMF1 activity is required in meristematic as well as differentiating tissues to rescue emf mutant phenotype. Temporal removal or reduction of EMF1 activity in the embryo or shoot apex of wild-type seedlings was sufficient to cause early flowering and terminal flower formation in adult plants. Such reproductive cell memory is reflected in the flower MADS-box gene activity expressed prior to flowering in these early flowering plants. However, temporal removal of EMF1 activity in flower meristem did not affect flower development. Our results are consistent with EMF1's primary role in repressing flowering in order to allow for vegetative growth.

Through multifaceted genome-scale research involving phylogenomics, targeted gene surveys, and gene expression analyses in diverse basal lineages of angiosperms, our studies provide insights into the most recent common ancestor of all extant flowering plants. MADS-box gene duplications have played an important role in the origin and diversification of angiosperms. Furthermore, early angiosperms possessed a diverse tool kit of floral genes and exhibited developmental 'flexibility', with broader patterns of expression of key floral organ identity genes than are found in eudicots. In particular, homologs of B-function MADS-box genes are more broadly expressed across the floral meristem in basal lineages. These results prompted formulation of the 'fading borders' model, which states that the gradual transitions in floral organ morphology observed in some basal angiosperms (e.g. Amborella) result from a gradient in the level of expression of floral organ identity genes across the developing floral meristem.

We report a recessive mutation of rice, aberrant panicle organization 1 (apo1), which severely affects inflorescence architecture, floral organ identity, and leaf production rate. In the wild-type inflorescence, the main-axis meristem aborts after forming 10–12 primary branch primordia. However, in apo1, the main-axis meristem was converted to a spikelet meristem after producing a small number of branch primordia. In addition, the branch meristems in apo1 became spikelet meristems earlier than in wild type. Therefore, in the inflorescence, the apo1 mutation caused the precocious conversion of the meristem identity. In the apo1 flower, lodicules were increased at the expense of stamens, and carpels were formed indeterminately by the loss of meristem determinacy. Vegetative development is also affected in the apo1. Leaves were formed rapidly throughout the vegetative phase, indicating that APO1 is also involved in temporal regulation of leaf production. These phenotypes suggest that the APO1 plays an important role in the temporal regulation of both vegetative and reproductive development.

The ABC model of flower organ identity is widely recognized as providing a framework for understanding the specification of flower organs in diverse plant species [1]. Recent studies in Arabidopsis thaliana have shown that three closely related MADS-box genes, SEPALLATA1 (SEP1), SEP2 and SEP3, are required to specify petals, stamens, and carpels because these organs are converted into sepals in sep1 sep2 sep3 triple mutants [3, 4]. Additional studies indicate that the SEP proteins form multimeric complexes with the products of the B and C organ identity genes. Here, we characterize the SEP4 gene, which shares extensive sequence similarity to and an overlapping expression pattern with the other SEP genes. Although sep4 single mutants display a phenotype similar to that of wild-type plants, we find that floral organs are converted into leaf-like organs in sep1 sep2 sep3 sep4 quadruple mutants, indicating the involvement of all four SEP genes in the development of sepals. We also find that SEP4 contributes to the development of petals, stamens, and carpels in addition to sepals and that it plays an important role in meristem identity. These and other data demonstrate that the SEP genes play central roles in flower meristem identity and organ identity.

The transition from vegetative to reproductive development establishes new growth patterns required for flowering. This switch is controlled by environmental and/or intrinsic developmental cues that converge at the shoot apical meristem (SAM) [1–4]. During this developmental transition, floral inductive signals cause the vegetative meristem to undergo morphological changes that are essential for flowering [5, 6]. Arabidopsis plants containing null mutations in two paralogous BEL1 -like ( BELL ) homeobox genes, PENNYWISE ( PNY ) and POUND-FOOLISH ( PNF ), disrupt the transition from vegetative to reproductive development. These double mutants are completely unable to flower even though the SAM displays morphological and molecular changes that are consistent with having received floral inductive signals. These studies establish a link between the competence to receive floral inductive signals and restructuring of the SAM during floral evocation.

Histone acetyltransferases, which are able to acetylate histone and non-histone proteins, play important roles in gene regulation. Many histone acetyltransferases are related to yeast Gcn5, a component of two transcription regulatory complexes SAGA and ADA. In this work, by characterizing a mutation in the Arabidopsis GCN5 gene (AtGCN5) we studied the regulatory function of this gene in controlling floral meristem activity. We show that in addition to pleiotropic effects on plant development, this mutation also leads to the production of terminal flowers. The flowers show homeotic transformations of petals into stamens and sepals into filamentous structures and produce ectopic carpels. The phenotypes correlate to an expansion of the expression domains within floral meristems of the key regulatory genes WUSCHEL (WUS) and AGAMOUS (AG). These results suggest that AtGCN5 is required to regulate the floral meristem activity through the WUS/AG pathway. This study brings new elements on the elucidation of specific developmental pathways regulated by AtGCN5 and on the control mechanism of meristem regulatory gene expression.

Two CONSTANS-LIKE (COL) genes, PdCO1 and PdCO2, were isolated from eastern cottonwood (Populus deltoides Bartr. ex. Marsh. var. deltoides). PdCO1 is unique among all the characterized CONSTANS-LIKE genes in the plant kingdom in that the deduced protein sequence contains a putative signal peptide with a potential transmembrane helix at the N-terminal end. Current bioinformatics tools suggest that PdCO1 protein may be secreted. Immuno-blot analysis indicated that although the PdCO1 was abundant in leaf, it was not detected in the subtended bud. Temporal mRNA and protein expression of PdCO1 and PdCO2 was determined in the leaf and bud at the ninth node of shoots, where transition from vegetative to floral phase has been observed to occur. The expression of both genes in the leaf increased dramatically from March 30 to April 21 and decreased on May 9. The increased expression coincided with morphological changes occurring in the axillary meristem of the subtended bud and with increased abundance of PdCO1 protein in the leaf. These changes sequentially included (1) the cessation of leaf-primordium formation, (2) the transition of the axillary shoot apical meristem to an inflorescence meristem, (3) and the subsequent formation of bracts and floral meristems in the axils of bracts flanking the inflorescence meristem. The temporal expression of PdCO1 and PdCO2 prior to flower bud development suggests that these genes might be involved in floral induction.

Considerable progress has been made in identifying genes that are involved in the evolution of plant morphologies. Elements of the ABC model of flower development are conserved throughout angiosperms, and homologous MADS-box genes function in gymnosperm reproduction. Candidate gene and mapping analyses of floral symmetry, sex determination, inflorescence architecture, and compound leaves provide intriguing glimpses into the evolution of morphological adaptations.

The development of the floral organs is specified by the combinations of three classes of gene for organ identity in the 'ABC' model. Recently, molecular genetic studies have shown this model is applicable to grass plants as well as most eudicots. Transcription factor complexes of ABC and homologous proteins form the molecular basis of the ABC model.

Over the past decade, the ABC model of flower development has been widely promulgated. However, correct flower-organ development requires not only the ABC genes but also the SEPALLATA genes. When the SEPALLATA genes are expressed together with the ABC genes, both vegetative and cauline leaves are converted to floral organs. Most of the ABC genes and all three SEPALLATA genes encode MADS transcription factors, which bind to DNA as dimers. Here, amendments to the ABC model are considered that incorporate both the SEPALLATA genes and the ability of MADS proteins to form higher-order complexes.

The homeotic gene AGAMOUS (AG) has dual roles in specifying organ fate and limiting stem cell proliferation in Arabidopsis flowers. We show that the floral identity protein LEAFY (LFY), a transcription factor expressed throughout the flower, cooperates with the homeodomain protein WUSCHEL (WUS) to activate AG in the center of flowers. WUS was previously identified because of its role in maintaining stem cell populations in both shoot and floral meristems. The unsuspected additional role of WUS in regulating floral homeotic gene expression supports the hypothesis that floral patterning uses a general meristem patterning system that was present before flowers evolved. We also show that AG represses WUS at later stages of floral development, thus creating a negative feedback loop that is required for the determinate growth of floral meristems.

Floral meristems and shoot apical meristems (SAMs) are homologous, self-maintaining stem cell systems. Unlike SAMs, floral meristems are determinate, and stem cell maintenance is abolished once all floral organs are initiated. To investigate the underlying regulatory mechanisms, we analyzed the interactions between WUSCHEL (WUS), which specifies stem cell identity, and AGAMOUS (AG), which is required for floral determinacy. Our results show that repression of WUS by AG is essential for terminating the floral meristem and that WUS can induce AG expression in developing flowers. Together, this suggests that floral determinacy depends on a negative autoregulatory mechanism involving WUS and AG, which terminates stem cell maintenance.

Shoot apical meristems are self-renewing stem cell populations that generate all of the above-ground organs (i.e. stems, leaves and flowers) of higher plants. Recent studies have identified new molecular components required for proper shoot meristem activity, and they have revealed that complex, intercellular communication pathways play important roles in coordinating meristem function.

The ABC model of flower development represents a milestone in explaining how the fate of emerging floral organ primordia is specified. This model states that organ identity is specified by different combinations of the activities of the A, B and C class homeotic genes. In spite of the remarkable simplicity of this model, the complex regulatory interactions that establish the initial pattern of A, B and C gene activity have yet to be fully explained. It has been shown that the LEAFY gene functions early to promote flower meristem identity, and that it is subsequently required for the normal expression of the ABC genes. Recently, LEAFY has been identified as an immediate upstream regulator of the floral homeotic genes, thus opening up an avenue to examine the transcriptional interactions that underlie floral patterning.

The shoot meristem gives rise to the aerial parts of higher plants by continuously initiating new organs. The basis of this activity is its ability to maintain a pool of pluripotent stem cells, which are the ultimate source of all tissues of the shoot. In Arabidopsis plants mutant for the WUSCHEL (WUS) gene, the stem cells are misspecified and appear to undergo differentiation. Here, we show that WUS encodes a novel homeodomain protein which presumably acts as a transcriptional regulator. The pattern of WUS expression suggests that stem cells in the shoot meristem are specified by an underlying cell group which is established in the 16-cell embryo and becomes localized to its prospective domain of function by asymmetric cell divisions.

The Arabidopsis flower is composed of four concentric whorls of organs: the sepals, petals, stamens and carpels. The development of this pattern depends on two general processes: that of establishing a florally determined meristem, and that of establishing particular organ identities. Mutational analyses have been instrumental in defining classes of genes that participate in each of these processes. The floral meristem identity genes are required to specify a florally determined meristem, and also appear to be required in part to activate the floral homeotic genes. The floral homeotic genes function in discrete regions of the florally determined meristem to specify the different organ types. The molecular analysis of these genes is beginning to shed light on the details of their regulation and function. The challenge that remains is to understand how the action of these genes in the dividing meristematic cell population can be coordinated and maintained to result in the differentiation of the complex array of cell and tissue types that make up the Arabidopsis flower.

All aerial parts of a plant are generated by groups of dividing cells called shoot meristems. These structures determine the arrangement and number of the different organs they produce, and, as a consequence the architecture of the plant largely depends on meristem activity. Extensive morphological studies have led to a model which postulates that shoot meristems are organised into functionally and morphologically distinct zones. According to this model, the centre of the meristem is responsible for cell production and meristem maintenance, whereas the periphery will initiate the organ primordia. Recently, genetic analysis allowed the identification of mutants affected in different functions of the meristem and some of the genes involved have now been identified. Meristem function also depends on the strict coordination of morphogenesis at the cellular level, as illustrated by the highly ordered patterns of cell division, division plane alignment and cell expansion observed in many species. Over the past few years, our understanding of the molecular mechanisms that control cell proliferation and growth has progressed significantly. Therefore, we can now start to unravel the networks that link these basic cellular processes to the regulators that control plant development at the level of the whole meristem.

The shoot apical meristem is responsible for above-ground organ initiation in higher plants, accomplishing continuous organogenesis by maintaining a pool of undifferentiated cells and directing descendant cells toward organ formation. Normally, proliferation and differentiation are balanced, so that the structure and size of the shoot meristem is maintained. However, Arabidopsis plants homozygous for mutations at the CLAVATA1 (CLV1) locus accumulate excess undifferentiated cells. We describe the molecular cloning and expression pattern of the CLV1 gene. It encodes a putative receptor kinase, suggesting a role in signal transduction. The extracellular domain is composed of 21 tandem leucine-rich repeats that resemble leucine-rich repeats found in animal hormone receptors. We provide evidence that CLV1 expression in the inflorescence is specifically associated with meristematic activity.

Flowers and shoots are derived from specialized groups of stem cells termed meristems. Recent studies in Arabidopsis have identified factors that contribute to meristem structure and identity, such as CLAVATA1, CLAVATA3, and SHOOTMERISTEMLESS, which act in both shoot and flower meristems, as well as LEAFY and APETALA1 which specifically determine a floral fate.

In higher plants, organ formation occurs throughout life. This remarkable process occurs at a collection of stem cells termed the shoot meristem. The shoot meristem originates during embryogenesis and is later responsible for generating the above-ground portion of the plant. The shoot meristem can be thought of as having two zones, a central zone containing meristematic cells in an undifferentiated state, and a surrounding peripheral zone where cells enter a specific developmental pathway toward a differentiated state. Recent advances have revealed several genes that specifically regulate meristem development in Arabidopsis . The function of these genes and their genetic interactions are described.

The physiological genetics of flowering in the pea (Pisum sativumL.) is examined starting with events in the leaves and concluding with events in the inflorescence, flower and floral organ meristems. WT peas have a late flowering, quantitative long day habit. The photoperiod response results from production of a flower inhibitor under short days. The phytochrome system controls inhibitor production via two mechanisms—a red (R) activated switch and a far-red (FR) mechanism involving extended exposure. The phyB deficient mutant,lv, is early flowering under long or short days. Thus thelvmutation may prevent R activation of the inhibitor synthesis pathway. The long term FR mechanism probably involves phyA and we suggest a phyA defective pea would be unable to down regulate inhibitor synthesis and would thus flower late in long or short days. Thelwmutant shows delayed flower initiation and enhanced response to photoperiod, traits indicative of an enlarged response to phyB. The phytochrome chromophore deficient mutant,pcd1, shows a small flowering delay under all photoperiods. Three genesSn, DneandPpdact in the inhibitor synthesis pathway and two genesEandHrcontrol ontogenetic expression of the system. Evidence of an additional gene which may regulate inhibitor synthesis has recently been obtained following identification of a dominant day neutral mutant, A23. The primary action of the graft-transmissible inhibitor may be to direct assimilate flow. Thegimutation blocks synthesis of the floral stimulus. TheLfalleles appear to determine apical reception of the flowering signal. GeneDetdetermines identity of the primary (I1) inflorescence meristem andVeg2identity of the lateral (I2) inflorescence meristem. It is not yet clear whetherVeg1determines I1or I2identity. Mutantsveg1andveg2-1never flower. Five mutantspim, A324,uni, M175 andlefloshow floral proliferation and aberrant flowers; these genes may determine floral (F) meristem identity. In addition, five mutants with floral homeotic changes are described. In two cases stamens are replaced by pistils and in two cases stamens are replaced by petals. In thekmutant, the change is within the petal whorl itself and the two wing petals express keel-like traits.

Background: Homeotic genes controlling the identity of flower organs have been characterized in several plant species. To determine whether cells expressing these genes are specified to follow particular developmental fates, we have studied the pattern of cell lineages in developing flowers of Antirrhinum . Each flower has four whorls of organs, and progenitor cells of these can be marked at particular stages of development using a temperature-sensitive transposon. This allows the cell lineages in the flower to be followed, as well as giving information about rates of cell division. Results We show here that, prior to the emergence of organ primordia, cells in the floral meristem have not been allocated organ identities. After this time, lineage restrictions arise between whorls, correlating with the onset of expression of genes that control organ identity. A further lineage restriction appears slightly later on, between the dorsal and ventral surfaces of the petal. Our results further suggest that the rates of cell division fluctuate during key stages of meristem development, perhaps as a consequence of meristem-identity gene expression. Conclusion The patterns of lineage restriction and organ-identity gene expression in early floral meristems are consistent with some cells being allocated specific identities at about this stage of development. Plant cells cannot move relative to each other, so lineage restrictions in plants may reflect particular orientations and/or rates of growth at boundary regions.

Homeotic mutants, that is, mutants with a normal organ in a place where an organ of another type is typically found, were first recognized in plants. The earliest descriptions of mutants in which petals replace stamens, giving double flowers, go back to ancient Greece and Rome. Similar accounts can be found in the botanical literature of China more than a thousand years ago, and in the books of the herbalists of Renaissance Europe (Meyerowitz et al., 1989). The use of such mutants (and similar but noninherited developmental abnormalities) to understand developmental processes in plants is more recent, dating from Linnaeus in the mid-eighteenth century (see Cullen and Stevens, 1990), and from Goethe (1790), who derived the ideas of organ homology and homological comparison from plants showing what was then called abnormal metamorphosis. Our term homeosis dates from Bateson’s work (1894) on organismal variation, in which he expanded Masters’treatment (1889) of abnormal metamorphosis in plants to animals and introduced the term homoeosis as a replacement for the older term. Goethe (1790) used homeotic variation in plants as the basis for a specific model explaining the developmental origin of different organ types in flowers. In his view, the four types of floral organs (sepals, petals, stamens, and carpels) are all modified leaves. As sap rises through developing flowers it is progressively refined, thereby inducing different organ types in different positions.

In recent years, isolation of several genes affecting flower development in Antirrhinum majus made this species a major model system to study this important developmental process. Genes like SQUAMOSA and FLORICAULA are involved in determination of the floral meristem. Their mutation results in the development of bract-forming shoots at positions where normally flowers would develop. The phenotypes obtained upon mutation of the genes found to affect floral organogenesis fall into three major categories. In each category, always the floral organs in two adjacent whorls become homeotically transformed. Based on this observation a simple genetic model has been proposed to explain the establishment of floral organ identity in the four concentric whorls of the flower. The model hypothesizes the independent induction of two developmental pathways specifying floral organ identity after the formation of sepals as the basic type of organ following induction of a floral meristem. One of these pathways is under the control of the PLENA gene, the other is controlled by the DEFICIENS and GLOBOSA genes. These genes, as well as SQUAMOSA, encode transcription factors sharing a conserved DNA binding domain: the MADS-box. In vitro DNA-binding studies complemented with molecular genetic analysis of the respective mutants show that the DEF and GLO proteins may act together in the form of a heterodimer in the regulation of their target genes as well as in autoregulation. The possible interactions between other MADS-box proteins and their role in flower development is under current investigation.

Over the last several years, a number of mutations that disrupt flower development have been described in both Arabidopsis thaliana and Antirrhinum majus. These mutations define genes that affect flower induction, meristem identity, and floral organ identity. Many of these regulatory genes have been cloned and found to contain a conserved protein-coding domain present in transcription factors from yeast and mammals. Transcripts from these genes are found in regions of the flower that exhibit defects in the corresponding mutants. By examining the spatial expression patterns of these genes in various mutant backgrounds, the molecular interactions that control flower development are beginning to be elucidated.

More than 200 years ago, Goethe proposed that each of the distinct flower organs represents a modified leaf [1]. Support for this hypothesis has come from genetic studies, which have identified genes required for flower organ identity. These genes have been incorporated into the widely accepted ABC model of flower organ identity, a model that appears generally applicable to distantly related eudicots as well as monocot plants. Strikingly, triple mutants lacking the ABC activities produce leaves in place of flower organs, and this finding demonstrates that these genes are required for floral organ identity [2]. However, the ABC genes are not sufficient for floral organ identity since ectopic expression of these genes failed to convert vegetative leaves into flower organs. This finding suggests that one or more additional factors are required [3, 4]. We have recently shown that SEPALLATA (SEP) represents a new class of floral organ identity genes since the loss of SEP activity results in all flower organs developing as sepals [5]. Here we show that the combined action of the SEP genes, together with the A and B genes, is sufficient to convert leaves into petals.

The first step in flower development is the generation of a floral meristem by the inflorescence meristem. We have analyzed how this process is affected by mutant alleles of the Arabidopsis gene LEAFY. We show that LEAFY interacts with another floral control gene, APETALA1, to promote the transition from inflorescence to floral meristem. We have cloned the LEAFY gene, and, consistent with the mutant phenotype, we find that LEAFY RNA is expressed strongly in young flower primordia. LEAFY expression precedes expression of the homeotic genes AGAMOUS and APETALA3, which specify organ identity within the flower. Furthermore, we demonstrate that LEAFY is the Arabidopsis homolog of the FLORICAULA gene, which controls floral meristem identity in the distantly related species Antirrhinum majus.

We characterized the distribution of AGAMOUS (AG) RNA during early flower development in Arabidopsis. Mutations in this homeotic gene cause the transformation of stamens to petals in floral whorl 3 and of carpels to another ag flower in floral whorl 4. We found that AG RNA is present in the stamen and carpel primordia but is undetectable in sepal and petal primordia throughout early wild-type flower development, consistent with the mutant phenotype. We also analyzed the distribution of AG RNA in apetela2 (ap2) mutant flowers. AP2 is a floral homeotic gene that is necessary for the normal development of sepals and petals in floral whorls 1 and 2. In ap2 mutant flowers, AG RNA is present in the organ primordia of all floral whorls. These observations show that the expression patterns of the Arabidopsis floral homeotic genes are in part established by regulatory interactions between these genes.

The homeotic genes that determine floral organ identity in plants turn out to be regulated by trans-acting factors related to the Polycomb-group proteins that have long been known as regulators of homeotic gene expression in Drosophila.