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PubMed1992

The tomato 66.3-kD polyphenoloxidase gene: molecular identification and developmental expression.

A gene coding for a polypeptide abundant in tomato floral meristems was isolated and shown to represent a tomato 66.3-kD polyphenoloxidase. Analysis of cDNA clones and a corresponding intronless genomic clone indicated that the plastid-bound 587-residue-long polypeptide, designated P2, contains two conserved copper-binding domains, similar to those found in fungal and mammalian tyrosinases. P2 transcripts and polypeptides are accumulated in the arrested floral primordia of the anantha mutant inflorescences and are equally abundant in primordia of wild-type flowers; the gene continues to be expressed at high levels in developing floral organs. In young expanding leaves, P2 protein is concentrated in palisade cells and in epidermal trichomes. Expression patterns of P2 in plant meristems permit molecular distinction between floral and vegetative primordia, and, in a companion study, comparison with dUTPase suggests that the two genes mark two alternative complementary developmental programs in the floral and vegetative meristems of the tomato plants.

Base Sequence↗

PubMed1992

LEAFY Interacts with Floral Homeotic Genes to Regulate Arabidopsis Floral Development.

In the leafy mutant of Arabidopsis, most of the lateral meristems that are fated to develop as flowers in a wild-type plant develop as inflorescence branches, whereas a few develop as abnormal flowers consisting of whorls of sepals and carpels. We have isolated several new alleles of leafy and constructed a series of double mutants with leafy and other homeotic mutants affecting floral development to determine how these genes interact to specify the developmental fate of lateral meristems. We found that leafy is completely epistatic to pistillata and interacts additively with agamous in early floral whorls, whereas in later whorls leafy is epistatic to agamous. Double mutants with leafy and either apetala1 or apetala2 showed a complete loss of the whorled phyllotaxy, shortened internodes, and suppression of axillary buds typical of flowers. Our results suggest that the products of LEAFY, APETALA1, and APETALA2 together control the differentiation of lateral meristems as flowers rather than as inflorescence branches.

Journal Article↗

PubMed1994

Root nodulation of Sesbania rostrata.

The tropical legume Sesbania rostrata can be nodulated by Azorhizobium caulinodans on both its stem and its root system. Here we investigate in detail the process of root nodulation and show that nodules develop exclusively at the base of secondary roots. Intercellular infection leads to the formation of infection pockets, which then give rise to infection threads. Concomitantly with infection, cortical cells of the secondary roots dedifferentiate, forming a meristem which has an "open-basket" configuration and which surrounds the initial infection site. Bacteria are released from the tips of infection threads into plant cells via "infection droplets," each containing several bacteria. Initially, nodule differentiation is comparable to that of indeterminate nodules, with the youngest meristematic cells being located at the periphery and the nitrogen-fixing cells being located at the nodule center. Because of the peculiar form of the meristem, Sesbania root nodules develop uniformly around a central axis. Nitrogen fixation is detected as early as 3 days following inoculation, while the nodule meristem is still active. Two weeks after inoculation, meristematic activity ceases, and nodules then show the typical histology of determinate nodules. Thus, root nodule organogenesis in S. rostrata appears to be intermediate between indeterminate and determinate types.

Cell Differentiation↗

PubMed2003

Quiescent center formation in maize roots is associated with an auxin-regulated oxidizing environment.

Embedded within the meristem of all Angiosperm roots is a population of slowly dividing cells designated the quiescent center (QC). In maize roots the QC can constitute upwards of 800-1200 cells, most of which spend an extended period of time (180-200 hours) in the G(1) phase of the cell cycle. How the QC forms and is maintained is not known. Here we report that cells of the QC are characterized by their highly oxidized status. Glutathione and ascorbic acid occur predominately in the oxidized forms in the QC. This is contrasted with the status of these redox intermediates in adjacent, rapidly dividing cells in the root meristem, in which the reduced forms of these two species are favored. Using a redox sensitive fluorescent dye we were able to visualize an overall oxidizing environment in the QC, and we also made comparisons with the adjacent, rapidly dividing cells in the root meristem. Altering the distribution of auxin and the location of the auxin maximum in the root tip activates the QC, and cells leave G(1) and enter mitosis. Commencement of relatively more rapid cell division in the QC is preceded by changes in the overall redox status of the QC, which becomes less oxidizing. We discuss how the position of the auxin maximum may influence the redox status of the QC and thereby modulate the cell cycle.

Ascorbic Acid↗

PubMed2004

CUPULIFORMIS establishes lateral organ boundaries in Antirrhinum.

Cupuliformis mutants are defective in shoot apical meristem formation, but cup plants overcome this early barrier to development to reach maturity. CUP encodes a NAC-domain transcription factor, homologous to the Petunia NAM and Arabidopsis CUC proteins. The phenotype of cup mutants differs from those of nam and cuc1 cuc2 in that dramatic organ fusion is observed throughout development. In addition to cotyledon and floral organ fusions, severe lateral organ fusion is found in leaves and inflorescences, and the apical meristem becomes highly fasciated. These features reveal a role for CUP in the establishment of all above ground organ boundaries. Consistent with this function, CUP is expressed at the boundaries of all lateral organs and meristems. It is not currently known how NAC-domain genes act to establish organ boundaries. Here, we show that CUP directly interacts with a TCP-domain transcription factor. Members of the TCP-domain family have previously been shown to regulate organ outgrowth. Our results suggest a model for the establishment of organ boundaries based on the localised expression of NAC-domain and TCP-domain factors.

Antirrhinum↗

PubMed2004

HUA ENHANCER3 reveals a role for a cyclin-dependent protein kinase in the specification of floral organ identity in Arabidopsis.

In plants, organs are generated post-embryonically from highly organized structures known as meristems. Cell division in the meristem is closely integrated with cell fate specification and organ formation. The presence of multiple cyclin-dependent kinases (CDKs) and their partner cyclins in plants and other multicellular organisms probably reflects the complexity of cell cycle regulation within developmental contexts. The Arabidopsis genome encodes at least eight CDKs and 30 cyclins. However, no mutants in any CDKs have been reported, and the function of the great majority of these genes in plant development is unknown. We show that HUA ENHANCER3 (HEN3), which encodes CDKE, a homolog of mammalian CDK8, is required for the specification of stamen and carpel identities and for the proper termination of stem cells in the floral meristem. Therefore, CDK8 plays a role in cell differentiation in a multicellular organism.

Amino Acid Sequence↗

PubMed2005

Transcriptional program controlled by the floral homeotic gene AGAMOUS during early organogenesis.

Floral organs, whose identity is determined by specific combinations of homeotic genes, originate from a group of undifferentiated cells called the floral meristem. In Arabidopsis, the homeotic gene AGAMOUS (AG) terminates meristem activity and promotes development of stamens and carpels. To understand the program of gene expression activated by AG, we followed genome-wide expression during early stamen and carpel development. The AG target genes included most genes for which mutant screens revealed a function downstream of AG. Novel targets were validated by in situ hybridisation and binding to AG in vitro and in vivo. Transcription factors formed a large fraction of AG targets, suggesting that during early organogenesis, much of the genetic program is concerned with elaborating gene expression patterns. The results also suggest that AG and other homeotic proteins with which it interacts (SEPALLATA3, APETALA3, PISTILLATA) are coordinately regulated in a positive-feedback loop to maintain their own expression, and that AG activates biosynthesis of gibberellin, which has been proposed to promote the shift from meristem identity to differentiation.

AGAMOUS Protein, Arabidopsis↗

PubMed1991

The effect of a heterochronic mutation, Teopod2, on the cell lineage of the maize shoot.

Teopod2 (Tp2) is a semi-dominant mutation of maize that prolongs the expression of characteristics normally confined to the juvenile phase of development. Two of the many dramatic morphological effects of this mutation are an increase in the number of vegetative nodes, and a reduction in the overall size of the shoot. To determine the cellular basis of these phenotypes, the technique of clonal analysis was used to compare the cell division patterns of wild-type and Tp2 plants. Our results indicate that Tp2 increases the number of vegetative nodes produced by the apicalmost cells in the meristem but does not affect the cell lineage of the basal, juvenile, part of the shoot. This result demonstrates that Tp2 does not act uniquely in a 'juvenile' domain of the meristem, but instead causes cells that are normally destined to produce adult structures to express juvenile traits inappropriately. Clonal analysis also demonstrates that Tp2 does not affect the size of the meristem prior to germination, nor does it affect the cell lineage of the basic structural unit of the stem, the phytomer. Thus the effects of this mutation on the size of the shoot are the result of changes in cell fate late in development.

Cloning, Molecular↗

PubMed1996

Mutations in the PERIANTHIA gene of Arabidopsis specifically alter floral organ number and initiation pattern.

An open question in developmental biology is how groups of dividing cells can generate specific numbers of segments or organs. We describe the phenotypic effects of mutations in PERIANTHIA, a gene specifically required for floral organ patterning in Arabidopsis thaliana. Most wild-type Arabidopsis flowers have 4 sepals, 4 petals, 6 stamens, and 2 carpels. Flowers of perianthia mutant plants most commonly show a pentamerous pattern of 5 sepals, 5 petals 5 stamens, and 2 carpels. This pattern is characteristic of flowers in a number of plant families, but not in the family Brassicaceae, which includes Arabidopsis. Unlike previously described mutations affecting floral organ number, perianthia does not appear to affect apical or floral meristem sizes, nor is any other aspect of vegetative or floral development severely affected. Floral organs in perianthia arise in a regular, stereotypical pattern similar to that in distantly related species with pentamerous flowers. Genetic analysis shows that PERIANTHIA acts downstream of the floral meristem identity genes and independently of the floral meristem size and floral organ identity genes in establishing floral organ initiation patterns. Thus PERIANTHIA acts in a previously unidentified process required for organ patterning in Arabidopsis flowers.

Arabidopsis↗

PubMed2001

The PINOID protein kinase regulates organ development in Arabidopsis by enhancing polar auxin transport.

Arabidopsis pinoid mutants show a strong phenotypic resemblance to the pin-formed mutant that is disrupted in polar auxin transport. The PINOID gene was recently cloned and found to encode a protein-serine/threonine kinase. Here we show that the PINOID gene is inducible by auxin and that the protein kinase is present in the primordia of cotyledons, leaves and floral organs and in vascular tissue in developing organs or proximal to meristems. Overexpression of PINOID under the control of the constitutive CaMV 35S promoter (35S::PID) resulted in phenotypes also observed in mutants with altered sensitivity to or transport of auxin. A remarkable characteristic of high expressing 35S::PID seedlings was a frequent collapse of the primary root meristem. This event triggered lateral root formation, a process that was initially inhibited in these seedlings. Both meristem organisation and growth of the primary root were rescued when seedlings were grown in the presence of polar auxin transport inhibitors, such as naphthylphtalamic acid (NPA). Moreover, ectopic expression of PINOID cDNA under control of the epidermis-specific LTP1 promoter provided further evidence for the NPA-sensitive action of PINOID. The results presented here indicate that PINOID functions as a positive regulator of polar auxin transport. We propose that PINOID is involved in the fine-tuning of polar auxin transport during organ formation in response to local auxin concentrations.

Arabidopsis↗

PubMed2002

Roles of PIN-FORMED1 and MONOPTEROS in pattern formation of the apical region of the Arabidopsis embryo.

In dicotyledonous plants, the apical region of the embryo shifts from radial to bilateral symmetry as the two cotyledon primordia develop on opposite sides of the shoot meristem. To further elucidate the mechanisms regulating this patterning process, we analyzed functions of two Arabidopsis genes, PIN-FORMED1 (PIN1) and MONOPTEROS (MP), encoding a putative auxin efflux carrier and a transcription factor thought to mediate auxin signaling, respectively. The corresponding mutants show similar defects in apical patterning, including cotyledon fusion and dissymmetric organ positioning. Both mutations perturb the spatial expression patterns of CUP-SHAPED COTYLEDON1 (CUC1) and CUC2, which are redundantly required for cotyledon separation and meristem formation. During early embryogenesis, both CUC genes are affected differently: the area of CUC1 expression is expanded while that of CUC2 expression is reduced. In addition, genetic analysis indicates that PIN1 and MP are required for the activity of CUC2 while CUC1 activity is only slightly affected by both mutations. These results suggest a differential regulation of the CUC genes by PIN1 and MP. Furthermore, genetic analysis suggests that SHOOT MERISTEMLESS (STM), another regulator for cotyledon separation and meristem formation, promotes CUC1 activity in parallel with PIN1. Our results suggest a model where PIN1 and MP regulate apical patterning partially through the control of CUC gene expression.

Arabidopsis↗

PubMed2002

ASYMMETRIC LEAVES1 reveals knox gene redundancy in Arabidopsis.

The shoot apical meristem comprises undifferentiated stem cells and their derivatives, which include founder cells for lateral organs such as leaves. Meristem maintenance and lateral organ specification are regulated in part by negative interactions between the myb domain transcription factor ASYMMETRIC LEAVES1, which is expressed in lateral organ primordia, and homeobox transcription factors which are expressed in the shoot apical meristem (knox genes). The knox gene SHOOT MERISTEMLESS (STM) negatively regulates ASYMMETRIC LEAVES1 (AS1) which, in turn, negatively regulates other knox genes including KNAT1 and KNAT2, and positively regulates the novel gene LATERAL ORGAN BOUNDARIES (LOB). Genetic interactions with a second gene, ASYMMETRIC LEAVES2 (AS2), indicate it acts at the same position in this hierarchy as AS1. We have used a second-site suppressor screen to isolate mutations in KNAT1 and we show that KNAT1 is partially redundant with STM in regulating stem cell function. Mutations in KNAT2 show no such interaction. We discuss the regulation and evolution of redundancy among knox genes.

Amino Acid Sequence↗

PubMed1998

Cell fate in plants. Lessons from the Arabidopsis root.

Classical studies in plant development have indicated that the fate of plant cells is fixed late, after cell division has ceased. Earlier commitment events are therefore considered reversible. To gain a mechanisatic understanding of the processes involved in specification and fixation of cell fate in plants, we are using the Arabidopsis root epidermis as a model system. The Arabidopsis root epidermis is composed of two cell types whose pattern of differentiation is directed by positional cues during development. Examination of mutations has identified genes involved in the establishment of cell fate specification in this tissue. TRANSPARENT TESTA GLABRA (TTG) and GLABRA2 (GL2) are positive regulators of non-hair fate and are active during the early differentiation of the epidermis in the meristem. GL2 encodes a homeobox protein which is expressed in non-hair cells in the meristem and is positively regulated by TTG. Mutations in genes involved in the regulation of ethylene biosynthesis and signal transduction indicate that ethylene is a positive regulator of hair cell fate. Treatment of ttg and gl2 plants with modulators of ethylene biosynthesis indicate that ethylene acts down stream of TTG and GL2 during the fate specification process. The relationship between meristem organisation and the mechanism underpinning the establishment of cell fate in other systems is also discussed.

Arabidopsis↗

PubMed2000

Evolutionary loss of sepals and/or petals in detarioid legume taxa Aphanocalyx, Brachystegia, and Monopetalanthus (Leguminosae: Caesalpinioideae).

Floral development using scanning electron microscopy is compared in several taxa of the Brachystegia subtribal group of caesalpinioid tribe Detariae. This group is characterized by missing sepals and/or petals. In Aphanocalyx djumaensis, Monopetalanthus durandii, and two Brachystegia species, one sepal is initiated in median abaxial position. In the first two, one or two additional sepal rudiments may initiate late. Brachystegia species have all five sepals, which remain scalelike. In Aphanocalyx and Monopetalanthus, one petal initiates adaxially and medianly (a position atypical for the first initiated petal in the family); additional petal rudiments may form in lateral sites. In Brachystegia, five petals are initiated unidirectionally on a meristem ring, but all are suppressed after initiation. In all taxa, ten stamens are initiated on a ring meristem: unidirectionally in Monopetalanthus, bidirectionally in Brachystegia, vs. in erratic order in Aphanocalyx. Carpel and petal initiation are concurrent. Different organ whorls overlap in time in Monopetalanthus and Brachystegia. In all, the floral apex characteristically is elongate radially and narrow tangentially after bracteole initiation. Two ontogenetic features, the meristem ring and the radially elongate post-bracteole floral apex, appear to be possible synapomorphies for the Brachystegia group.

Journal Article↗

PubMed2000

LEAFY and the evolution of rosette flowering in violet cress (Jonopsidium acaule, Brassicaceae).

Arabidopsis and most other Brassicaceae produce an elongated inflorescence of mainly ebracteate flowers. However, the early-flowering species violet cress (Jonopsidium acaule) and a handful of other species produce flowers singly in the axils of rosette leaves. In Arabidopsis the gene LEAFY (LFY) is implicated in both the determination of flower meristem identity and in the suppression of leaves (bracts) that would otherwise subtend the flowers. In this study we examined the role of LFY homologs in the evolution of rosette flowering in violet cress. We cloned two LFY homologs, vcLFY1 and vcLFY2, from violet cress. Their exon sequences show ∼90% nucleotide similarity with Arabidopsis LFY and 99% similarity to each other. We used in situ hybridization to study vcLFY expression in violet cress. The patterns were very similar to LFY in Arabidopsis except for stronger expression in the shoot apical meristem outside of the region of flower meristem initiation. It is possible that the relatively diffuse expression of vcLFY contributes to the lack of bract suppression in violet cress. Additionally, the earliest flowers produced by violet cress express vcLFY, suggesting that accelerated flowering in violet cress could also result from changes in the regulation of vcLFY.

Journal Article↗

PubMed2000

Cortical development in roots of the aquatic plant Pontederia cordata (Pontederiaceae).

Adventitious roots of marsh-grown Pontederia cordata were examined to determine cortical development and structure. The innermost layer of the ground meristem forms the endodermis and aerenchymatous cortex. The outermost layer of the early ground meristem undergoes a precise pattern of oblique and periclinal cell divisions to produce a single or double layer of prohypodermis with an anchor cell for each radial file of aerenchyma cells. At maturity, endodermal cell walls are modified only by narrow Casparian bands. The central regions of the ground meristem become proaerenchyma and exhibit asymmetric cell division and expansion. They produce an aerenchymatous zone with barrel-shaped large cells and irregularly shaped small cells traversing the aerenchyma horizontally along radii; some crystalliferous cells with raphides are present in the aerenchyma. The walls of the hypodermis are modified early by polyphenols. The outermost layer of the hypodermis later matures into an exodermis with Casparian bands that are impermeable to berberine, an apoplastic tracer dye. The nonexodermal layer(s) of the hypodermis has suberin-modified walls. Radial files of aerenchyma are usually connected by narrow protuberances near their midpoints, the aerenchyma lacunae having been produced by expansion of cells along walls lining intercellular spaces. We are terming this type of aerenchyma development, which is neither schizogenous nor lysigenous, "differential expansion."

Journal Article↗

PubMed2000

Floral ontogeny, pattern formation, and evolution in Hibbertia and Adrastaea (Dilleniaceae).

Floral development was compared with scanning electron microscopy in 12 Australian species of Hibbertia representing most of its morphological variation, and in the related Adrastaea (Dilleniaceae). Calyx and corolla arise in quincuncial helices in radially symmetrical species, while the petals initiate unidirectionally from one side in zygomorphic species. Stamen number (3-200+) proliferates by centrifugal addition of individual primordia or by innovations of common primordia and ring meristems. Common primordia arise in single-stamen positions alternately with petals, and each produces one to several stamens centrifugally that remain attached to a shared base and form a stamen fascicle. A ring meristem in Adrastaea initiates a whorl of five stamens, alternate with the first stamens but outside their whorl. In radially symmetrical species of Hibbertia, a first ring of stamens is supplemented centrifugally by additional stamens on a meristem ring. The first stamens in zygomorphic species of Hibbertia initiate as a terminal ridge on the floral apex, with subsequent stamens added centrifugally on one side and two carpels initiated on the opposite side. The carpels arise as a simultaneous ring in radially symmetrical flowers, or as a simultaneous pair in zygomorphic species. Staminodial presence is viewed as of minor significance. Four pollinator syndromes are proposed for Hibbertia, related to differing floral architecture.

Journal Article↗

PubMed2002

[Variability of the number of chloroplasts in a population of stoma guard cells in the sugar beet (Beta vulgaris L.)].

Diploid sugar beet plants demonstrate a broad variability of the number of chloroplasts in stoma guard cells, which is related to myxoploidy of cell populations in leaf apical meristems (epigenomic variability). In addition to random organelle segregation between daughter cells, this variability is affected by factors disrupting the mitotic cycle: (1) plant treatment with a mitotic poison, such as colchicine; (2) duration of the life cycle of a plant; the variability in second-year plants is greater than in first-year ones; (3) the mode of plant reproduction; the variability in inbred plants is greater than in the initial population. Treatment of germinating seeds with a diluted colchicine solution increases the number of organelles in cells in the myxoploid generation (generation C0) and the variance of the distributions in the first vegetation year. The variability in the organelle number in stoma cells correlates with that in maternal meristem cells. It is concluded that the epidermal cell monolayer (including stoma guard cells) keeps record of the epigenomic and epiplastome variability in meristem cells. The variability of the number of chloroplasts in stoma guard cells is approximated by binomials with negative powers.

Beta vulgaris↗