Specification of chimeric flowering shoots in wild-type Arabidopsis

Within wild-type Arabidopsis populations, a subset of the plants were found to have a single chimeric shoot on their primary shoot axes. The chimeric shoots were located below the lowest primary-axis flower; and they exhibited features of both flowers and paraclades (lateral flowering shoots). Morphological analyses of chimeric shoots indicated that they developed from single primordia. In each chimeric shoot, the side furthest from the apical meristem was specified as 'flower'—while the side closest to the meristem was specified as 'paraclade'—suggesting that a stimulus from outside the apical meristem can directly induce primordia to develop as flowers. It is concluded that the development of the teratological chimeric shoots resulted from the overlap of the vegetative and floral specification processes within single primordia.     

Auxin Depletion from the Leaf Axil Conditions Competence for Axillary Meristem Formation in Arabidopsis and Tomato

The enormous variation in architecture of flowering plants is based to a large extent on their ability to form new axes of growth throughout their life span. Secondary growth is initiated from groups of pluripotent cells, called meristems, which are established in the axils of leaves. Such meristems form lateral organs and develop into a side shoot or a flower, depending on the developmental status of the plant and environmental conditions. The phytohormone auxin is well known to play an important role in inhibiting the outgrowth of axillary buds, a phenomenon known as apical dominance. However, the role of auxin in the process of axillary meristem formation is largely unknown. In this study, we show in the model species Arabidopsis thaliana and tomato (Solanum lycopersicum) that auxin is depleted from leaf axils during vegetative development. Disruption of polar auxin transport compromises auxin depletion from the leaf axil and axillary meristem initiation. Ectopic auxin biosynthesis in leaf axils interferes with axillary meristem formation, whereas repression of auxin signaling in polar auxin transport mutants can largely rescue their branching defects. These results strongly suggest that depletion of auxin from leaf axils is a prerequisite for axillary meristem formation during vegetative development.     

Revisiting phase transition during flowering in Arabidopsis

Single-phase transition during flowering has been suggested by Hempel and Feldman (1994) [Planta 192: 276]. When early flowering ecotypes of Arabidopsis were microscopically observed, a long day signal simultaneously induced the acropetal (bottom to top) production of flower primordia and the basipetal (top to bottom) differentiation of paraclades (axillary flowering shoots) from the axils of pre-existing leaf primordia. However, this model could not account for the production of an extra number of secondary shoots in the TERMINAL FLOWER 1 overexpressor line or AGL20 overexpressor line in Columbia background with a functional allele of FRIGIDA. We report here that Columbia with a functional allele of FRIGIDA under long days and Columbia under short days show an inflorescence-producing phase between the vegetative and the flower-producing phases, supporting two-step phase transition during flowering. In addition, a late-flowering mutant, fwa shows an inflorescence phase but fca, fy and fve follow a single-phase transition, suggesting flowering time mutations have different effects on phase transition during flowering.     

Identification of a MADS-box gene, FLOWERING LOCUS M, that represses flowering

The timing of flowering is important for the reproductive success of plants. Here we describe the identification and characterization of a new MADS-box gene, FLOWERING LOCUS M (FLM), which is involved in the transition from vegetative to reproductive development. FLM is similar in amino-acid sequence to FLC, another MADS-box gene involved in flowering-time control. flm mutants are early flowering in both inductive and non-inductive photoperiods, and flowering time is sensitive to FLM dosage. FLM overexpression produces late-flowering plants. Thus FLM acts as an inhibitor of flowering. FLM is expressed in areas of cell division such as root and shoot apical regions and leaf primordia.     

Growth correlations in shoot apices ofBrassica campestris L. during transition to flowering

Growth correlations in the shoot apical meristem during transition to flowering were studied in a quantitative long day plant,Brassica campestris L. cv. Ceres, requiring only one long day for floral initiation. During photo-inductive exposure of the plants, an overall increase in cell number could be observed at the shoot apex concomitant with promotion of leaf initiation. Release from apical dominance and decline in relative growth rate of leaf primordia are reported as early effects of photo-induction. With the onset of floral differentiation, production of new leaf primordia had stopped altogether. Maximum increase in RNA concentration could be noticed in axillary meristems following photoperiodic treatment, whereas in vegetative plants the highest RNA concentration was found in leaf primordia. The significance of these changes occurring during transition to flowering is discussed.     

Flowering and apical meristem growth dynamics

The shoot apical meristem generates stem, leaves, and lateralshoot meristems during the entire shoot ontogeny. Vegetativeleaves are generated by the meristem in the vegetative developmentalphase, while in the reproductive phase either bracts subtendinglateral flower primordia (or paraclades), or perianth and strictlyreproductive organs are formed. Meristem growth is fully characterizedby the principal growth rates, directions, volumetric, and arealgrowth rates. Growth modelling or sequential in vivo methodsof meristem observation complemented by growth quantificationallow the above growth variables to be estimated. Indirectly,growth is assessed by cell division rates and other cell cycleparameters. Temporal and spatial changes of growth and geometrytake place at the meristem during the transition from the vegetativeto the reproductive phase. During the vegetative phase, meristemgrowth is generally indeterminate. In the reproductive phaseit is almost always determinate, but the extent of determinacydepends on the inflorescence architecture. In the vegetativephase the central meristem zone is the slowest growing region.The transition from the vegetative to the reproductive phaseis accompanied by an increase in mitotic activity in this zone.The more determinate is the meristem growth, the stronger isthis mitotic activation. However, regardless of the extent ofthe activation, in angiosperms the tunica/corpus structure ofthe meristem is preserved and therefore the mitotic activityof germ line cells remains relatively low. In the case of thethoroughly studied model angiosperm plant Arabidopsis thaliana,it is important to recognize that the flower primordium developsin the axil of a rudimentary bract. Another important featureof growth of the inflorescence shoot apical meristem is theheterogeneity of the peripheral zone. Finally, the role of mechanicalfactors in growth and functioning of the meristem needs furtherinvestigation. Key words: Flower primordium, geometry, growth, inflorescence, shoot apical meristem, transition from vegetative to reproductive phase Received 4 October 2007; Revised 5 November 2007 Accepted 6 November 2007     

Apical Growth and Modification of the Development of Primordia During Re-flowering of Reverted Plants of Impatiens balsamina L

Impatiens balsamina L. was induced to flower by exposure to5 short days and then made to revert to vegetative growth byreturn to long days. After 9 long days reverted plants wereinduced to re-flower by returning them to short days. Petalinitiation began immediately and seven primordia already presentdeveloped into petals instead of into predominantly leaf-likeorgans. However, the arrangement of primordia at the shoot apex,their rate of initiation and size at initiation remained unchangedfrom the reverted apex, as did apical growth rate and the lengthof stem frusta at initiation. The more rapid flowering of thereverted plants than of plants when first induced, and the lackof change in apical growth pattern, imply that the revertedapices remain partially evoked, and that the apical growth patternand phyllotaxis typical of the flower, and already present inthe reverted plants, facilitate the transition to flower formation. Impatiens balsamina, flower reversion, partial evocation, shoot meristem, determination, leaf development     

Appearance and disappearance of proteins in the shoot apical meristem of Sinapis alba in transition to flowering

Vegetative plants of Sinapis alba L. grown under short days were induced to flower by exposure to one long day or continuous long days. Irrespective of the number of long days, the first flower primordia were initiated by the shoot apical meristem 60 h after the start of the inductive treatment. An indirect histoimmunofluorescence technique was used to search in the apical meristem for three antigenic proteins which had been previously detected by immunodiffusion tests in the whole apical bud (Pierard et al. (1977) Physiol. Plant. 41, 254–258). One protein called protein A, present in the vegetative meristem, increased in concentration during the first 48 h following the start of the inductive treatment. It stayed constant up to 96 h and disappeared completely at a later time. Two other proteins called B and C, absent in the vegetative meristem, appeared in the meristem of induced plants between 30 and 36 h after the start of the inductive treatment and progressively accumulated at later times up to 240 h. These proteins appeared 8 h before the irreversible commitment of the meristem to produce flower primordia (point of no return) was reached and 24 h before start of flower production. These observations support an interpretation of floral evocation as consisting, at least partially, of an early and qualitative change in gene expression.Abbreviations AVB anti-vegetative-bud antiserum - ARB antireproductive-bud antiserum - IgG immunoglobulins G - TRITC tetramethylrhodamine isothiocyanate - GAR IgG goat antirabbit IgG - S₀ IgG non-immune rabbit IgG     

Competence to respond to floral inductive signals requires the homeobox genes PENNYWISE and POUND-FOOLISH

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). During this developmental transition, floral inductive signals cause the vegetative meristem to undergo morphological changes that are essential for flowering. 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.     

The inhibition and stimulation of DNA synthesis in shoot apices ofChenopodium rubrum L. during photoperiodic induction of flowering

Three short-day inductive cycles bring about inhibition followed by transitional enhancement of growth, not only in roots and leaves but also in different zones of shoot apical meristem, as shown by measurement of DNA synthesis using³H-thymidine autoradiography. The first inductive cycle resulted in marked inhibition of the cells of the central zone (CZ), rib meristem (RM), and peripheral zone (PZ). Subsequent enhancement of DNA synthesis occurs in RM during the second inductive cycle, but in CZ only in the third cycle. The growth activation in PZ is counteracted by decrease in apical dominance which results in further inhibition of leaf primordia and increases in bud primordia. In plants induced only by one cycle, which later reverse the vegetative pattern of growth and differentiation, increased DNA synthesis in RM and CZ was not observed. The significance of inhibitory and stimulatory processes in particular zones of the shoot apex is discussed considering flower morphogenesis.     

Cytophotometric Study of the 2C Nuclear DNA Content in the Apical Bud of Sinapis alba during Floral Transition

Feulgen cytophotometry was used to detect possible changes inthe 2C DNA content in the various parts of the apical bud ofSinapis alba during floral evocation and flower development.This study showed that there was no significant difference inthe 2C DNA content between the vegetative, evoked or reproductivemeristems. In vegetative plants, the 2C DNA content was lowerin the leaf primordia than in the meristem. This content inthe leaf exhibited a further decrease during the floral transition.In the flower primordia, the 2C value never exceeded the typicalvalue of the meristem. In the flower at anthesis, the DNA contentwas lower in the pistil and stamen than in the meristem. Apical bud, floral transition, 2C DNA content, cytophotometry, Sinapis alba L.     

Reversion of flowering

Reversion from floral to vegetative growth is under environmental control in many plant species. However the factors regulating floral reversion, and the events at the shoot apex that take place when it occurs, have received less attention than those associated with the transition to flowering. Reversions may be categorized as flower reversion, in which the flower meristem resumes leaf production, or inflorescence reversions, in which the meristem ceases to initiate bracts with flowers in their axils and begins instead to make leaves with vegetative branches in their axils. Related to these two types of reversion, but distinct from them, are examples of partial flowering, when non-floral meristems grow out so that the plant begins to grow vegetatively again. Anomalous or proliferous flowers may form as a result of unfavourable growth conditions or viral infection, but these do not necessarily involve flower reversions.     

A tobacco calcium/calmodulin-binding protein kinase functions as a negative regulator of flowering

A tobacco calcium/calmodulin-binding protein kinase (NtCBK1) was isolated and identified. The predicted NtCBK1 protein has 599 amino acids, an N-terminal kinase domain, and shares high homology with other calmodulin (CaM)-related kinases. Whereas NtCBK1 phosphorylates itself and substrates such as histone IIIS and syntide-2 in the absence of CaM, its kinase activity can be stimulated by tobacco CaMs. However, unlike another tobacco protein kinase designated NtCBK2, NtCBK1 was not differentially regulated by the different CaM isoforms tested. The CaM-binding domain of NtCBK1 was located between amino acids 436 and 455, and this domain was shown to be necessary for CaM modulation of kinase activity. RNA in situ hybridization showed that NtCBK1 was highly regulated in the transition to flowering. Whereas NtCBK1 mRNA was accumulated in the shoot apical meristem during vegetative growth, its expression was dramatically decreased in the shoot apical meristem after floral determination, and in young flower primordia. The expression of NtCBK1 was up-regulated to high levels in floral organ primordia. Fluctuations in NtCBK1 expression were verified by analysis of tobacco plants expressing green fluorescent protein under the control of the NtCBK1 promoter, suggesting a role of NtCBK1 in the transition to flowering. This conclusion was confirmed by overexpressing NtCBK1 in transgenic tobacco plants, where maintenance of high levels of NtCBK1 in the shoot apical meristem delayed the switch to flowering and extended the vegetative phase of growth. Further work indicated that overexpression of NtCBK1 in transgenic tobacco did not affect the expression of NFL, a tobacco homologue of the LFY gene that controls meristem initiation and floral structure in tobacco. In addition, the promotion of tobacco flowering time by DNA demethylation cannot be blocked by the overexpression of NtCBK1.     

Regulation of extent of vegetative development of the maize shoot meristem

In maize plants ( Zea mays L.), the extent of vegetative development in the shoot is precisely regulated such that the apical meristem produces a predictable number of leaves before converting to tassel development. In previous experiments using shoot apex culture, we showed that the developmental program that limits vegetative development in maize is not intrinsic to the shoot apical meristem. Rather, the meristem receives information from elsewhere in the plant and responds by either continuing leaf initiation or becoming determined for determinate growth and forming an inflorescence, the tassel. Here we examine leaf primordia as potential sources for that information using shoot apex culture. Our results show that the presence of the four to six youngest leaf primordia on the shoot apex is sufficient to provide such information. The ability to reset shoot development by meristem culture also allows us to examine the basis for expression of a specific phenotype at a particular developmental stage. We found that the mutation hcf106 , which is typically expressed only during seedling stages, is not re-expressed when the shoot morphogically has regained a juvenile phase.     

The rate of cell division in the shoot apical meristem during photoperiodic induction and transition to flowering

Cell division contributing to longitudinal growth of the shoot apex was investigated inChenopodium rubrum in segments marked by the axils of leaf primordia. Plants treated with two short days (16h of darkness and 8h of light) were compared with two non-induced controls (cultivated in continuous light or treated by alternations of 8 h of darkness and 4 h of light for two days). During the short-day treatments the rate of cell division contributing to the longitudinal growth decreases in all segments of the shoot apex irrespective of whether the darkness was given in inductive or non-inductive photoperiods. The rate of cell division contributing to longitudinal growth increases in the upper internodes of the shoot apex after the termination of the photoperiodic treatment and transfer of the plants to continuous light. However, cell division remains inhibited in the lowest segment of the shoot apex. This inhibition in the differentiating parts of the shoot apical meristem is a direct consequence of photoperiodic induction. It is supposed that this inhibition is related to evocation similarly as the well-known phenomenon of stimulation of cell division in the apical dome.     

Growth and Development of the Banana Plant: II. The Transition from the Vegetative to the Floral Shoot in Musa acuminata cv. Gros Michel

The changes that occur in the shoot apex of the banana, as itpasses from the vegetative to the flowering stage, are described.The crucial events occur well before floral primordia are evident,and they require a redistribution of activity in the variousgrowing regions. The vegetative shoot apex is in a central depressionin the rhizome; there is virtually no internodal growth in theaxis, the most active growth is in the leaf bases; vegetativebuds do not form in the leaf axils but only appear adventitiouslyfar from the tip of the shoot. With the onset of flowering thisis changed; growth in the axis itself, previously suppressed,occurs and flower buds arise as primordia in the axils of subtendingbracts. The bracts do not show the market growth in their baseswhich is so characteristic of leaves. Thus, the shoot apex risesto the level of the rhizome and then above it; as it does so,its tip changes in shape from a broad flattened some to a pointedcone. At the transitional stage, more activity occurs in thecells of the mantle, or tunica, which now consists of 3 to 4layers over the central dome. Below, in the central or mothercell zone of the corpus, which was quiescent in the vegetativeshoot, the cells spring into greater activity, becoming moreprotoplasmic and stain more deeply. Directly below this regionin the rib meristem, cells show transverse divisions. Bractprimordia occur high on the flanks of the apex, and, thoughthey originate in the manner of leaves, their subsequent growthis different. Flower primordia occur even in the axils of bractsclose to the shoot tip. Thus, the problem now is to designatethe source, nature, and mode of action of the stimuli whichinitiate and control this quite different distribution of growthin the floral, as contrasted with the vegetative, shoot. Thesignificance of the previously more quiescent central, or mothercell zone, of the apex as the source of such stimuli, is stressed.Thus, flowering first requires that the limiting controls whichapply to the vegetative shoot be released, and, secondly, thatthe apex of the shoot, rather than the leaf base, becomes themain centre of growth and development.     

A complex case of simple leaves: indeterminate leaves co-express ARP and KNOX1 genes

The mutually exclusive relationship between ARP and KNOX1 genes in the shoot apical meristem and leaf primordia in simple leaved plants such as Arabidopsis has been well characterized. Overlapping expression domains of these genes in leaf primordia have been described for many compound leaved plants such as Solanum lycopersicum and Cardamine hirsuta and are regarded as a characteristic of compound leaved plants. Here, we present several datasets illustrating the co-expression of ARP and KNOX1 genes in the shoot apical meristem, leaf primordia, and developing leaves in plants with simple leaves and simple primordia. Streptocarpus plants produce unequal cotyledons due to the continued activity of a basal meristem and produce foliar leaves termed “phyllomorphs” from the groove meristem in the acaulescent species Streptocarpus rexii and leaves from a shoot apical meristem in the caulescent Streptocarpus glandulosissimus. We demonstrate that the simple leaves in both species possess a greatly extended basal meristematic activity that persists over most of the leaf’s growth. The area of basal meristem activity coincides with the co-expression domain of ARP and KNOX1 genes. We suggest that the co-expression of ARP and KNOX1 genes is not exclusive to compound leaved plants but is associated with foci of meristematic activity in leaves.     

植物花发育的分子机理研究进展

花的发育分为开花决定、花的发端和花器官的发育三个阶段。植物开花由多条途径诱导,包括光周期和光质诱导、春化作用、自主途径、赤霉素诱导、碳水化合物诱导等;植物体本身也存在着开花抑制途径。各种开花诱导途径能激活花分生组织特性基因,使茎端分生组织转变为花分生组织。花器官的发育由器官特性基因决定,这些基因的精确表达需要花分生组织特性基因的激活和多个正、负调节因子的调控;另有一类基因控制着花发育的对称性。花发育机理的研究具有重要的理论意义和广泛的应用前景。