Regulation of node number in day-neutral Nicotiana tabacum: a factor in plant size

Employing genotypes of day-neutral tobacco that exhibit a wide range in the number of nodes produced, it has been established that node number, and plant size, in tobacco is regulated, in large part, by two endogenous signals and one developmental state, competence. All genotypes have the same level of a root signal that maintains a vegetative pattern during early growth. The number of nodes produced before the formation of the terminal flower, as well as plant size, is a function of the strength of the floral stimulus from the leaves and the competence of the terminal meristem to respond to the floral stimulus by initiating the terminal flower.     

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.     

A yeast mitotic activator sensitises the shoot apical meristem to become floral in day-neutral tobacco

True day-neutral (DN) plants flower regardless of day-length and yet they flower at characteristic stages. DN Nicotiana tabacum cv. Samsun, makes about forty nodes before flowering. The question still persists whether flowering starts because leaves become physiologically able to export sufficient floral stimulus or the shoot apical meristem (SAM) acquires developmental competence to interpret its arrival. This question was addressed using tobacco expressing the Schizosaccharomyces pombe cell cycle gene, Spcdc25, as a tool. Spcdc25 expression induces early flowering and we tested a hypothesis that this phenotype arises because of premature floral competence of the SAM. Scions of vegetative Spcdc25 plants were grafted onto stocks of vegetative WT together with converse grafts and flowering onset followed (as the time since sowing and number of leaves formed till flowering). Spcdc25 plants flowered significantly earlier with fewer leaves, and, unlike WT, also formed flowers from axillary buds. Scions from vegetative Spcdc25 plants also flowered precociously when grafted to vegetative WT stocks. However, in a WT scion to Spcdc25 stock, the plants flowered at the same time as WT. SAMs from young vegetative Spcdc25 plants were elongated (increase in SAM convexity determined by tracing a circumference of SAM sections) with a pronounced meristem surface cell layers compared with WT. Presumably, Spcdc25 SAMs were competent for flowering earlier than WT and responded to florigenic signal produced even in young vegetative WT plants. Precocious reproductive competence in Spcdc25 SAMs comprised a pronounced mantle, a trait of prefloral SAMs. Hence, we propose that true DN plants export florigenic signal since early developmental stages but the SAM has to acquire competence to respond to the floral stimulus.     

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.     

Development of Meristems of Weigela florida Variety 'Bristol Ruby' Following Reproductive Induction in Long Days

Weigela florida variety ‘Bristol Ruby’ has longday requirements for its growth and, in general, for its flowering.Vegetative development, floral initiation and floral organogenesisare described using scanning electron microscopy during photoperiodictreatment in long days, under controlled conditions. Flowering of axillary buds of cuttings has been studied. Theapex of Weigela at the vegetative phase is characterized bya very small hollow meristem. After 9 long days, the meristemenlarges and, after 12 long days, early axillary buds are initiatedin the axils of the leaves, which become bracts. When the numberof long days was increased, flowers were initiated in the budson the induced branches; first at the proximal part of the branchwhere development afterwards slowed down, then on the medianparts of the branch where development was accelerated. Two bracteoles are differentiated soon after floral initiation;first initiation of the calyx required 18 long days. Petals,stamens and ovary were rapidly initiated after that. Weigelaflowers are clustered; the inflorescence ceased growth by abortionof the terminal meristem or by formation of a terminal flower.In axillary buds of the fifth node the formation of the clusterwas completed about 20 days after the beginning of floral induction. Weigela florida ‘Bristol Ruby’, scanning electron microscopic analysis, vegetative meristem, floral development stages, long days induction     

Morpho-histogenic studies on tendrils of Passiflora

The ontogenetic development of the tendril and its associatedorgans is investigated in 17 species of Passiflora. The shootapex shows a single tunica layer though the second layer simulatestunica. The cytohistological zonation is not a constant feature.In P. caerulea Linn., it is distinct at leaf initiation butin P. pruinosa Mast., P. vespertilio Linn., and P. watsonianaMast., it is indistinct. The main axillary bud differentiatesfrom the peripheral meristem of the shoot apex. The differentiationof this bud into floral and tendril menstems occurs at a nodeimmediately below the shoot apex in P. minima Blanco. and Pracemosa Brot. In other species this differentiation generallyoccurs at the lower nodes. The floral meristem is initiatedas an accessory bud from this bud, thus forming a bud complex.The residuum of the bud complex develops as a tendril. The thirdaccessory bud which does not originate from this bud complex,develops into a vegetative branch. The fundamental nature ofthe vascular relationship between the flower, tendril, accessorybud, subtending leaf, and the axis is similar in most of theinvestigated species.     

EMF Genes Interact with Late-Flowering Genes to Regulate Arabidopsis Shoot Development

To investigate the genetic mechanisms regulating the transitionfrom vegetative to reproductive phase in Arabidopsis, doublemutants between two embryonic flower (emf) and 12 differentlate-flowering mutants were constructed and analyzed. Doublemutants in all combinations displayed the emf phenotypes withoutforming rosettes during early development; however, clear variationsbetween different double mutants were observed during late development,fwa significantly enhanced the vegetative property of both emfmutants by producing a high number of sessile leaves withoutany further reproductive growth in emf1 fwa double mutants.It also produced numerous leaf-like flower structures similarto those in leafy ap1 double mutant in emf1 fwa double mutants.Nine late-flowering mutants, ft, fca, ld, fd, fpa, fe, fy, fha,and fve, caused different degrees of increase in the numberof sessile leaves, the size of inflorescence, and the numberof flowers only in weak emf1 and emf2 mutant alleles background.Two late-flowering mutants, co and gi, however, had no effecton either emf1 and emf2 mutant alleles in double mutants. Ourresults suggest that FWA function in distinct pathways fromboth EMF genes to regulate flower competence by activating geneswhich specify floral meristem identity. CO and GI negativelyregulate both EMF genes, whereas the other nine late-floweringgenes may interact with EMF genes directly or indirectly toregulate shoot maturation in Arabidopsis. ¹ To whom correspondence should be addressed.     

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.     

The shoot apical meristem of Sinapis alba L. expands its central symplasmic field during the floral transition

The shoot apical meristem (SAM) is functionally subdivided into zones with distinct tasks. During vegetative growth the peripheral zone of the meristem gives rise to leaf primordia that develop into dorsiventral leaves under the influence of signals from the central zone. During the floral transition the function of the SAM is altered and its peripheral zone starts to form floral structures in a specific pattern. This requires alterations in the signal networks that coordinate the activities of the peripheral and central zone of the SAM. These signal networks are partly housed in the symplasmic space of the SAM. Dye-coupling experiments demonstrate that in the superficial layer of the Sinapis alba meristem this space is radially subdivided. The cells of the central zone are coupled into a symplasmic field, which is shielded from the peripheral zone by the positional closing of plasmodesmata. In the vegetative meristems, most of these central symplasmic fields have a triangular geometry and are relatively small in size. Plants that are induced to flower by exposure to a single long day alter the geometry as well as the size of their central symplasmic field. After two subsequent days under short photoperiod the central symplasmic fields exhibit a circular form. Simultaneously, their size strongly increases both in an absolute sense and relative to the enlarging meristem. The geometric change in the fields is hypothesized to be due to recruitment of extra initial cells, required to support the increase in phyllotactic complexity. The proportional increase in field size is interpreted as an adjustment in the balance between the central and peripheral zone of the SAM, accompanying the shift from leaf production to flower formation.     

Regulatory networks that function to specify flower meristems require the function of homeobox genes PENNYWISE and POUND-FOOLISH in Arabidopsis

Flowering is a major developmental phase change that transforms the fate of the shoot apical meristem (SAM) from a leaf-bearing vegetative meristem to that of a flower-producing inflorescence meristem. In Arabidopsis, floral meristems are specified on the periphery of the inflorescence meristem by the combined activities of the FLOWERING LOCUS T (FT)–FD complex and the flower meristem identity gene, LEAFY ( LFY ). Two redundant functioning homeobox genes, PENNYWISE ( PNY ) and POUND-FOOLISH ( PNF ), which are expressed in the vegetative and inflorescence SAM, regulate patterning events during reproductive development, including floral specification. To determine the role of PNY and PNF in the floral specification network, we characterized the genetic relationship of these homeobox genes with LFY and FT . Results from this study demonstrate that LFY functions downstream of PNY and PNF. Ectopic expression of LFY promotes flower formation in pny pnf plants, while the flower specification activity of ectopic FT is severely attenuated. Genetic analysis shows that when mutations in pny and pnf genes are combined with lfy , a synergistic phenotype is displayed that significantly reduces floral specification and alters inflorescence patterning events. In conclusion, results from this study support a model in which PNY and PNF promote LFY expression during reproductive development. At the same time, the flower formation activity of FT is dependent upon the function of PNY and PNF.     

Changes in Apical Growth and Phyllotaxis on Flowering and Reversion in Impatiens balsamina L.

When plants of Impatiens balsamina L were subjected to 5 shortdays and then re-placed in long days, they began to form a terminalflower and then reverted to vegetative growth at this terminalshoot apex The onset of flowering was accompanied by an increasein the rate of initiation of primordia, an increase in the growthrate of the apex, a change in primordium arrangement from spiralto whorled or pseudo-whorled, a lack of internodes, and a reductionm the size at initiation of the primordia and also of the stemfrusta which give rise to nodal and internodal tissues On reversion,parts intermediate between petals and leaves were formed, followedby leaves, although in reverted apices the size at initiationand the arrangement of primordia remained the same as in thefloweing apex The apical growth rate and the rate of primordiuminitiation were less in the reverted apices than in floral apicesbut remained higher than in the original vegetative apex Sincethe changes in apical growth which occur on the transition toflowering are not reversed on reversion, the development oforgans as leaves or petals is not directly related to the growthrate of the apex, or the arrangement, rate of initiation orsize at initiation of primordia Impatiens balsamina L, flower reversion, evocation, phyllotaxis, shoot meristem     

Development of the floral shoot and pre‐anthesis cleistogamy in Hydrobryopsis sessilis (Podostemaceae)

The reproductive biology of Hydrobryopsis sessilis (Podostemaceae, subfamily Podostemoideae), a reduced, threatened, aquatic angiosperm endemic to the Western Ghats of India, was examined. This is the first report on the transition from the vegetative to the reproductive phase in this plant, describing floral ontogeny, pollination and the breeding system. The cytohistological zonation of the apical meristem of the reproductive thallus is identical to that of the apical meristem of the vegetative thallus. The floral shoots do not replace vegetative shoots (i.e. the vegetative shoots never bear flowers), but form at new sites at the tip of the flattened plant body. Each floral shoot meristem is tiny, deep‐seated and concave and arises endogenously following lysigeny. The floral shoot meristem gives rise to four to six bracts in a distichous manner. The development of spathe, stamens and carpels is described. The ab initio dorsiventrality of the carpels and the occurrence of endothelium in the ovules are reported. The mature stigmas and anthers lie close to each other. The pollen germinates within undehisced anthers and the pollen tubes enter the stigmas in the unopened floral bud, leading to pre‐anthesis, complete, constitutional cleistogamy under water. The seed set is 63.2%. A significant finding is the penetration of several pollen tubes into the filaments of stamens in 16% of the flower buds, indicating a trend towards cryptic self‐fertilization. The Indian Podostemoideae appear to show a shift from xenogamy or geitonogamy or autogamy in a chasmogamous flower to complete autogamy in a cleistogamous flower. The floral modifications leading to cleistogamy in H. sessilis have been identified. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 222–236.     

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.     

The Role of Leaves in the Perception of Vernalizing Temperatures in Sugar Beet

Annual and biennial sugar beet varieties require long days toinduce flowering but the biennial genotypes additionally requirevernalization. Previous research has suggested that the inabilityof non-vernalized biennial plants to flower can be explainedby a lack of competence of the leaves to respond to long days.In this study defoliation experiments were used to investigatewhich leaves could perceive long daylengths and, in particular,whether leaves initiated from a non-vernalized shoot apicalmeristem could perceive vernalizing temperatures and producea floral stimulus in long days. Annual and vernalized biennialplants flowered if young leaves (i.e. those formed during orafter vernalization) were kept on the plants, but they did notflower if only older expanded leaves (including those expandedprior to vernalization) were present. No evidence was obtainedto indicate that the older leaves contained inhibitors of floweringand it seems most likely that there is a decline in responsivenessto daylength with increasing leaf age. Exposure to vernalizingtemperatures accelerated flowering of the annual and was essentialfor flowering of the biennial. The presence of a single leafinitiated, but not expanded, prior to the transfer of biennialplants to vernalizing temperatures was sufficient to induceflowering. This indicates that expanding leaves do not needto be initiated from a vernalized apical meristem to becomecompetent to produce a floral stimulus in long days. Key words: Beta vulgaris L., sugar beet, vernalization, flowering     

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。     

Developmental morphology of branching flowers in Nymphaea prolifera

Nymphaea and Nuphar (Nymphaeaceae) share an extra-axillary mode of floral inception in the shoot apical meristem (SAM). Some leaf sites along the ontogenetic spiral are occupied by floral primordia lacking a subtending bract. This pattern of flower initiation in leaf sites is repeated inside branching flowers of Nymphaea prolifera (Central and South America). Instead of fertile flowers this species usually produces sterile tuberiferous flowers that act as vegetative propagules. N. prolifera changes the meristem identity from reproductive to vegetative or vice versa repeatedly. Each branching flower first produces some perianth-like leaves, then it switches back to the vegetative meristem identity of the SAM with the formation of foliage leaves and another set of branching flowers. This process is repeated up to three times giving rise to more than 100 vegetative propagules. The developmental morphology of the branching flowers of N. prolifera is described using both microtome sections and scanning electron microscopy.     

Photoperiod and Temperature Regulation of Floral Initiation and Anthesis in Soya Bean

Floral development includes initiation of floral primordia andsubsequent anthesis as discrete events, even though in manyinvestigations only anthesis is considered. For ‘Ransom’soya bean [Glycine max (L.) Merrill] grown at day/night temperaturesof 18/14, 22/18, 26/22, 30/26, and 34/30 °C and exposedto photoperiods of 10, 12, 14, 15, and 16 h, time of anthesisranged from less than 21 days after exposure at the shorterphotoperiods and warmer temperatures to more than 60 days atlonger photoperiods and cooler temperatures. For all temperatureregimes, however, floral primordia were initiated under shorterphotopenods within 3 to 5 days after exposure and after notmore than 7 to 10 days exposure to longer photoperiods. Onceinitiation had begun, time required for differentiation of individualfloral primordia and the duration of leaf initiation at shootapices increased with increasing length of photoperiod. Whileproduction of nodes ceased abruptly under photoperiods of 10and 12 h, new nodes continued to be formed concurrently withinitiation of axillary floral primordia under photoperiods of14, 15 and 16 h. The vegetative condition at the main stem shootapex was prolonged under the three longer photoperiods and issuggestive of the existence of an intermediate apex under theseconditions. The results indicate that initiation and anthesisare controlled independently rather than collectively by photoperiod,and that floral initiation consists of two independent steps—onefor the first-initiated flower in an axil of a main stem leafand a second for transformation of the terminal shoot apex fromthe vegetative to reproductive condition. Apical meristem, intermediate apex, floral initiation, anthesis, photoinduction, Glycine max(L.) Merrill, soya bean, photoperiod, temperature     

Expression of CENTRORADIALIS (CEN) and CEN-like genes in tobacco reveals a conserved mechanism controlling phase change in diverse species.

Plant species exhibit two primary forms of flowering architecture, namely, indeterminate and determinate. Antirrhinum is an indeterminate species in which shoots grow indefinitely and only generate flowers from their periphery. Tobacco is a determinate species in which shoot meristems terminate by converting to a flower. We show that tobacco is responsive to the CENTRORADIALIS (CEN) gene, which is required for indeterminate growth of the shoot meristem in Antirrhinum. Tobacco plants overexpressing CEN have an extended vegetative phase, delaying the switch to flowering. Therefore, CEN defines a conserved system controlling shoot meristem identity and plant architecture in diverse species. To understand the underlying basis for differences between determinate and indeterminate architectures, we isolated CEN-like genes from tobacco (CET genes). In tobacco, the CET genes most similar to CEN are not expressed in the main shoot meristem; their expression is restricted to vegetative axillary meristems. As vegetative meristems develop into flowering shoots, CET genes are downregulated as floral meristem identity genes are upregulated. Our results suggest a general model for tobacco, Antirrhinum, and Arabidopsis, whereby the complementary expression patterns of CEN-like genes and floral meristem identity genes underlie different plant architectures.     

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     

LEAFY同源基因研究进展

LEAFY(LFY)同源基因存在于所有的陆生植物中,在植物花发育早期表达,并在花发育过程中抑制茎端分生组织的营养生长,调控花分生组织和花器官的形成,使转LFY基因植株提前开花,LFY同源基因与其上下游基因共同调控花发育过程.LFY同源基因的蛋白质结构在不同物种间保守性很高,但它们的表达部位差异很大.该文总结了近年来国内外已经克隆到的LFY同源基因的表达、功能及其在果树、花卉、粮食作物上的应用,以期为植物花发育的深入研究提供参考.