A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005     

Structural cost of shoot modules and its implications on plant potential fitness in a Mediterranean perennial shrub, Retama sphaerocarpa (L.) Boiss

This paper introduces a methodology to analyse the structural costs on plant potential fitness, empirically exemplified in the hierarchical shoot system of a Mediterranean perennial plant, Retama sphaerocarpa (L.) Boiss. During growing season every year (March-August), the terminal shoot (which is the basic unit of growth) develops inflorescences, flowers and fruits, as well as new shoots (first-, second- and third-order branching shoots) which have the potential to "behave" as terminal shoots in the following year. Different morphological and demographical aspects of the modules within the terminal shoot were measured in 100 terminal shoots selected from different plants of a natural population of R. sphaerocarpa. Complementary samples of 100 shoots of different branching orders were collected to obtain biomass estimations of the terminal shoots. We propose a simple procedure to estimate structural cost (biomass investment) on plant potential fitness (flowering buds) as a methodology for interpreting and comparing the consequences on fitness of different plant growth patterns. The results of this study exemplify how differential allocation patterns among plant structural modules, depending on their position within the shoot system, can be quantified to estimate their influence upon plant potential fitness.     

Endorhythmic development in Choisya ternata Kunth (Rutaceae), with a further elucidation of lammas shoots, as well as sylleptic and proleptic shoots

Most apical resting buds of Choisya tenata include inflorescence buds in the axils of their lower consecutive paired scales. These inflorescences develop as apical buds which burst in spring. The whole of the lateral inflorescence system on a shoot originating from an apical bud may be viewed as a single, proliferous inflorescence. After the spring flush there are usually two other flushes of the same shoot within the same season, each of which may be accompanied by the development of lateral inflorescences as in the spring flush. Each further flush produces an apical 'lammas shoot'. As an apical lammas shoot elongates, lateral lammas shoots may also develop from upper, previously resting, axillary buds on the underlying stem segment of the preceding flush. Lateral inflorescences on apical lammas shoots arise from axillary buds preformed within the briefly-dormant apical buds terminating the preceding flush. These inflorescences, as well as the spring ones, represent proleptic shoots. The production of resting apical buds between two intra-season flushes of a shoot may be fugacous, without the differentiation of perfect bud-scales, and with curtailmenl ol internode elongation. As no environmental influence seems to be responsible for intra-season rhythmicity in development, this is said to be endorhythmic. The interrelations of proleptic to sylleptic shoots are discussed.     

AXR1 acts after lateral bud formation to inhibit lateral bud growth in Arabidopsis

The AXR1 gene of Arabidopsis is required for many auxin responses. The highly branched shoot phenotype of mature axr1 mutant plants has been taken as genetic evidence for a role of auxin in the control of shoot branching. We compared the development of lateral shoots in wild-type Columbia and axr1-12 plants. In the wild type, the pattern of lateral shoot development depends on the developmental stage of the plant. During prolonged vegetative growth, axillary shoots arise and develop in a basal-apical sequence. After floral transition, axillary shoots arise rapidly along the primary shoot axis and grow out to form lateral inflorescences in an apical-basal sequence. For both patterns, the axr1 mutation does not affect the timing of axillary meristem formation; however, subsequent lateral shoot development proceeds more rapidly in axr1 plants. The outgrowth of lateral inflorescences from excised cauline nodes of wild-type plants is inhibited by apical auxin. axr1-12 nodes are resistant to this inhibition. These results provide evidence for common control of axillary growth in both patterns, and suggest a role for auxin during the late stages of axillary shoot development following the formation of the axillary bud and several axillary leaf primordia.     

Flowering Gradient Along the Stem Axis in an Orchid Hybrid Aranda deborah

Decapitation stimulated the production of axillary shoots inAranda deborah. The nature of these axillary shoots, whethervegetative or reproductive, was shown to be correlated withtheir positions along the stem axis. The higher ones near theapex developed into inflorescences while the lower ones awayfrom the apex developed into vegetative shoots. These resultsindicated that a flowering gradient exists along the stem axisin this orchid hybrid.     

QUANTITATIVE LAWS IN REGENERATION : III. THE QUANTITATIVE BASIS OF POLARITY IN REGENERATION.

It is well known that a long defoliated piece of stem of Bryophyllum calycinum forms shoots only at the apical or the two apical nodes, while when such a stem is cut into as many pieces as there are nodes each node produces shoots. It is shown in this paper that the dry weight of shoots produced in the apical nodes of a long piece of stem is approximately equal to the dry weight of shoots the same stem would have produced if it had been cut into as many pieces as it possesses nodes. Hence all the material which can be used for the growth of shoots goes into the most apical part of the stem and this accounts for the polar character of regeneration in this case. It seems that the mass of basal roots produced by a piece of defoliated stem also increases with the mass of the stem.     

Growth Dynamics and Morphology of Annual Shoots, According to their Architectural Position, in Young Cedrus atlantica (Endl.) Manetti ex Carriere (Pinaceae)

The influence of shoot architectural position on growth andbranching pattern of young Cedrus atlantica (Endl.) Manettiex Carrière trees were studied. Extension growth andtype of axillary products (lateral bud, sylleptic short or longshoots) of annual shoots of increasing branching order (mainstem, branches and branchlets) were recorded weekly during the1993 growing season. Annual final shoot length, duration ofextension, and maximum extension rate decreased with increasingbranching order. Sylleptic axillary shoots occurred only onannual shoots of the main stem and branches and were producedwhen extension rate was at its highest. Differences in growthrate and final length of annual shoots, according to their architecturalposition, were related to differences in the total number anddiversity of types of sylleptic axillary shoots produced. Itis suggested that types and numbers of sylleptic axillary shootsproduced are linked with threshold values for both final lengthand extension rate of the parent shoot. Copyright 1999 Annalsof Botany Company Atlas cedar, extension growth, sylleptic branching, tree architecture, morphology.     

Flower removal increases rhizome mass in natural populations of Alstroemeria aurea (Alstroemeriaceae)

Plant architecture and phenotypic plasticity under natural conditions remain little known for many rhizomatous species. This study evaluates, in situ, the plastic responses of Alstroemeria aurea plants from three Patagonian populations to flower or flowering-shoot removal. The size and architecture of treated and untreated plants were assessed. Nutrient contents (N, P and K) were evaluated for rhizomes and roots developed in two successive years. Those plants that were deprived of their inflorescences developed, on average, a heavier rhizome than both control plants and plants from which flowering shoots had been removed. Neither of the two treatments applied altered the number of metamers or the branching pattern of the rhizomes. The contents of N, P and K were higher in rhizomes than in roots. In summer, nutrients were more concentrated in inflorescences and the new rhizome segment than in the rhizome segment developed in the previous year. The idea that fruiting failure in A. aurea promotes resource re-assignment from aerial shoots to rhizomes without altering the architecture of plants is supported. The development of the underground portion of aerial shoots in late summer-autumn allows A. aurea plants to take full advantage of short growth periods, but would impose a limit to plasticity.     

Morphogenesis in Selaginella. III. Meristem determination and cell differentiation.

Selaginella willdenovii Baker is a prostrate vascular cryptogam with a dorsiventral stem. At each major branching of the stem tip a dorsal and a ventral angle meristem are formed. The ventral meristem becomes determined as a root and the dorsal meristem as a shoot. Indoleacetic acid (IAA) is transported basipetally in the stem and has been found to be the regulatory agent for meristem determination both in vitro and in vivo.Growth measurements of intact plants indicated that the sequence of development for each stem unit is frond expansion, internodal elongation, ventral meristem growth as a root, and dorsal meristem growth as a shoot. The principal experimental findings of this study are as follows. Triiodobenzoic acid (TIBA), an inhibitor of auxin transport alters the normal pattern of development in intact plants, causing ventral meristems to develop as shoots and dorsal meristems to develop precociously. Dorsal meristems grown in sterile culture on an auxin-free medium develop as shoots, but in the presence of IAA develop as roots. Meristems transferred after excision from auxin-free to plus-auxin medium on successive days showed an increasing tendency to develop as shoots, with more than 50% doing so by day 5. The mitotic index is low at the time of excision of the meristem, rises to a peak on day 5 and then declines.     

Observations on the reproductive ecology of Thalassia testudinum (Hydrocharitaceae) II. Leaf width as a secondary sex character

Short-shoots of Thalassia testudinum Banks ex König were collected from 5 sites in Florida. A total of 284, 625-cm² quadrat samples, containing 6182 shhort-shoots was analyzed for leaf width, inflorescence number and sex (the latter if determinable). Although leaf widths and reproductive densities differed at the 5 sites, leaf width was consistently greater when reproductive structures were present, and when the number of inflorescences increased. The mean number of inflorescences per short-shoot was significantly higher for shoots bearing male inflorescences compared to female shoots. Female inflorescences were normally solitary; male short-shoots usually had 2 or 3 inflorescences. Sex ratios were male-biased for 4 of the 5 sites. Comparisons of leaf widths between the sexes indicated that leaf width constituted a secondary sex character for this species. Female short-shoots tend to have narrower leaves than male short-shoots. This relationship should also be considered when evaluating the significance of morphogeographic and stress-related variation in leaf width for this species.     

Growth patterns and endogenous indole-3-acetic acid concentrations in current-year coppice shoots and seedlings of two Betula species

Apical dominance, internode elongation, radial growth and xylem cell size in coppice and apical shoots of Betula pubescens B. Pendula were determined and related to endogenous indole-3-acetic acid (IAA) levels, measured by gas chromatography-selected ion monitoring in the apical bud and at three positions along the stem. The effects of defoliation and debudding on morphological and anatomical characters and endogenous IAA levels were also investigated. The coppice shoots displayed superior stem elongation and increased branching during the initial phase of growth, after which their growth pattern was similar to that of the seedlings; however, their radial growth was greater throughout the experiment. Both plant types produced smaller-sized xylem cells at the top of the shoot than at the bottom with coppice shoots tending to form larger tracheids and smaller vessels than the seedlings. There was no consistent difference in IAA concentration between the coppice shoots and the seedlings. Defoliation and debudding reduced the IAA level in the stem within 36 h and it was still low after 25 days. Although the extent of the IAA decrease was similar in both coppice shoots and seedlings, the treatments affected the morphological and anatomical characters differently in the two plant types. The results suggest that the observed differences between seedlings and coppice shoots were not mediated through a drastic change in IAA level.     

Genotypic differences in branching pattern and fruiting habit in common walnut (Juglans regia L.)

Architectural analysis of 840 Slovenian walnut (Juglans regia L.) genotypes was performed to determine the most typical and frequent morphological types and to evaluate their vegetative and generative potential. Four branching and fruiting patterns (I-IV) were detected. A 3-year-old fruiting branch, consisting of a 3-year-old shoot plus corresponding 2-year-old and 1-year-old shoots, was used as a structural unit for quantitative analysis. In the intermediate fruit-bearing types with mesotonic and acrotonic branching pattern (types II and III), the total lengths of 3-, 2- and 1-year-old shoots were 385 and 380 cm, respectively, compared with 275 and 253 cm in the terminal and lateral-fruiting types (types I and IV). In type I, 1-year-old shoots had significantly fewer nodes than in other types. In addition, they had a thinner basal diameter than types III and IV, and their angles were the most erect (39 degrees ). Only 0.4 out of 3.6 1-year-old shoots were flowering with one mixed bud with 1.9 female flowers. In type IV, 2-year-old shoots had significantly more nodes and a larger basal diameter than other types. One-year-old shoots in type IV are thicker than those in other types. Ratios between the number of flowering and the total number of 1-year-old shoots were 0.7 in type IV, 0.6 in type III, 0.5 in type II and 0.1 in type I. On 1-year-old shoots in type IV, 1.7 mixed buds with a mean of three female inflorescences per bud were counted. Consequently, the generative potential is highest in type IV and lowest in type I. In types II and III, growth and the ability to bear fruits are more balanced.     

Redesign of the castorbean plant body plan for optimal combine harvesting

The seed oil of castorbean (Ricinus communis, Euphorbiaceae) is highly appreciated in several sectors of the chemical industry. A tropical perennial, with sympodial branching, this plant presents, at the same time, both vegetative shoots and inflorescences, as well as racemes with ripe seeds. This architecture is an obstacle to harvesting the plants using the standard combine harvester which requires, as with any other seed crop, that the plants be dry. One agronomic solution is to spray the plants with desiccants. However, in this paper a genetic solution is suggested, that is, an annual plant. This model is discussed within the present knowledge of plant monocarpic senescence. A mutation that changes the quantity and/or the quality of cambial activity could reduce the secondary growth in castorbean, thus favouring the plants death. Moreover, to stop iterative growth which is at the base of the perennial form of castorbean, a non‐branching model is also proposed. The occurrence of this phenotype in some crops (maize and sunflower) and mutants (tomato, barley, etc.) is examined. Considering apical dominance as the primary mechanism at the base of a non‐branching form, a mutation capable of inducing permanent dormancy of axillary buds is discussed. Both mutation breeding and genetic engineering are suggested as major tools to obtain an annual plant of castorbean.     

SPIRAL DEVELOPMENTAL PATTERNS IN THE STEM AND INFLORESCENCE OF LILIUM TIGRINUM

Extensive correlations in spirality were observed among vegetative and floral organs in Lilium tigrinum Ker. Organs involved were vegetative leaves, bracts, and bracteoles. These correlations varied in their degree of constancy depending upon the organs involved. The mature inflorescences of L. tigrinum appeared to fit the common definition of a raceme. In 67.3% of the flowers at node 3 on the raceme, the bract-bracteole spirals reversed the spiral of vegetative leaves on the stem. These reversals resembled those observed on essentially cymose inflorescences of certain members of the Caryophyllaceae. Cymose branching was found to be an invariable feature of the inflorescence of L. tigrinum when secondary flowers appear. The apparently indeterminate tips of inflorescence main axes were interpreted as exhibiting stages in progression from a basically determinate (cymose) inflorescence. It was concluded that the ancestors of L. tigrinum had well-developed cymose branching patterns in the inflorescence. Reversal of stem spirals by the bract-bracteole spirals at the apices of many inflorescences was considered to be the result of complete utilization of the inflorescence meristem. Explanations for those reversals were provided by the field theory and by the theory of the first available space.     

THEORY OF REGENERATION BASED ON MASS ACTION

1. The writer''s older experiment, proving that equal masses of isolated sister leaves of Bryophyllum regenerate under equal conditions and in equal time equal masses (in dry weight) of shoots and roots, is confirmed. It is shown that in the dark this regeneration is reduced to a small fraction of that observed in light. 2. The writer''s former observation is confirmed, that when a piece of stem inhibits or diminishes the regeneration in a leaf, the dry weight of the stem increases by as much or more than the weight by which the regeneration in the leaf is diminished. It is shown that this is also true when the axillary bud in the stem is removed or when the regeneration occurs in the dark. 3. These facts show that the regeneration of an isolated leaf of Bryophyllum is determined by the mass of material available or formed in the leaf during the experiment and that such a growth does not occur in a leaf connected with a normal plant for the reason that in the latter case the material available or formed in the leaf flows into the stem where it is consumed for normal growth. 4. It is shown that the sap sent out by a leaf in the descending current of a stem is capable of increasing also the rate of growth of shoots in the basal parts of the leaf when the sap has an opportunity to reach the anlagen for such shoots. 5. The fact that a defoliated piece of stem forms normally no shoots in its basal part therefore demands an explanation of the polar character of regeneration which lays no or less emphasis on the chemical difference between ascending and descending sap than does Sachs'' theory of specific root- or shoot-forming substances (though such substances may in reality exist), but which uses as a basis the general mass relation as expressed in the first three statements of this summary. 6. It is suggested that the polar character of the regeneration in a stem of Bryophyllum is primarily due to the fact that the descending sap reaches normally only the root-forming tissues at the base of the stem, while the ascending sap reaches normally only the shoot-forming anlagen at the apex of the stem. 7. This suggestion is supported by the fact that when the anlagen for shoots and roots are close together as they are in the notch of a leaf, the sap of the leaf causes the growth of both roots and shoots from the same notch and the influence of the sap of the leaf on this growth increases for both roots and shoots in proportion with the mass of the leaf.     

A quantification of shoot shedding in pedunculate oak (Quercus robur L.)

Almost 37 000 fragments (twigs, peduncles, inflorescences) were shed from a single mature oak tree over one year. Shedding showed strong seasonality with abrupt peaks in June and October, yet only weak correlations to weather parameters. The majority of shed twigs were short shoots, consisting of one or few extension units, which were not older than two years. In many twigs, shedding was preceded by the outgrowth of the terminal bud into either a male inflorescence or a defective extension unit. The large majority of shed twigs belong to a category of summer annual and biannual deciduous short shoots of an exploitative nature, only a minority are explorative long shoots. Implications for the architecture of the tree crown are that twigs of higher branching order are mostly affected by shedding, as they consist of short modules which exhibit a high proportion of flower buds.     

Vegetative Proliferation and Secondary Proliferated Inflorescences Development in Grass

We report the vegetative proliferation and new phenomenon of “secondary proliferated inflorescences” in the grass Ischaemum barbatum Retz, as determined by anatomical analysis of prepared sections of inflorescences. Leaves and shoots could be developed from the original spikelets of inflorescences and plantlets developed when these shoots were transplanted to moist soil. “Secondary proliferated inflorescences” is the first name here because some inflorescences that developed inadequacy are grown from the spikelet on the mother inflorescence. Our investigation showed that this form of vegetative proliferation and secondary proliferated inflorescences development of I. barbatum has arisen following late autumn fires of the previous year. It is suggested that the sudden onset of a fire could lead to a hormone imbalance or a chemical induction, which results in ephemeral vegetative proliferation even secondary proliferated inflorescences development in wild populations.(Author for correspondence. Tel: +86 (0)20 3725 2993; E-mail: magh@scib.ac.cn)     

The shoot apex of Podostemaceae: De novo structure or reduction of the conventional type?

The paper reviews and discusses various interpretations of the shoot apex of Podostemaceae with special reference to subfamily Podostemoideae. Main questions concern (1) the proposed absence of a shoot apical meristem (SAM) in apical “meristemless” shoot tips of Podostemoideae and, as the consequence, the endogenous inception of leaf-borne leaves and branches and (2) the predicted stem bifurcation below a “terminal” dithecous (double-sheathed) leaf positioned instead of a shoot apex, as it is reported for subfamily Podostemoideae. Does the “meristemless” shoot apex represent a true evolutionary novelty? Does the view of stem bifurcation represent a new ramification pattern with the consequence that the “classical root–shoot model” of angiosperms is not valid for Podostemaceae? Both interpretations do not conform to previous studies that are complemented here by new data on the SAM of Zeylanidium olivaceum and Thelethylax minutiflora (Podostemoideae). Although a SAM is difficult to observe in the vegetative shoots of many Podostemoideae, it becomes well visible when the shoot passes into the flowering stage approaching the conspicuous shoot apex of floriferous shoots. The arguments of the absence of a SAM in vegetative shoots are not convincing and the endogenous origin of “leaf-borne leaves” appears questionable. Consequently, the “meristemless” shoot apex cannot be considered as a structure having evolved de novo. In the less advanced subfamilies Tristichoideae and Weddellinoideae, the leaf primordia develop only from a few apical cells of the outer shoot layer. This allows the conclusion that the surface layer of the apex in these subfamilies corresponds to the horizontally spread single-layered apical meristem of subfamily Podostemoideae. Similarly, the view of shoot bifurcation does not conform to the diachsial–sympodial branching pattern occurring in the cymose inflorescences of many Podostemoideae. This fact contradicts the presence of a terminal leaf.