Flower Development in Silene: Morphology and Sequence of Initiation of Primordia

The initiation and development of the flower of Silene coeli-rosawas followed by examining apices by scanning electron microscopy.The sepals, stamens and carpets are initiated in a spiral sequence,the direction of the spiral king the opposite of the acropetalhelix of unequal axillary buds at the nodes below the flower.The petals are initiated almost simultaneously and at the sametime as the first few stamens. The change in phyllotaxis fromopposite and decussate in the vegetative shoot to spiral inthe flower occurs with the displacement of the first two sepalsaway from the mid-line of the apex and towards the axillarybud at the node below the flower. The sizes of the sepals andstamens are a function of their age since initiation but thepetals grow more slowly. The Silene flower can be interpretedas a shoot bearing primordia with associated axillary primordia,some of the latter being precocious in their development. Silent weli-rosa, flower initiation, flower development, phyllotaxis, primordia     

Regulation of Branching in Decussate Species with Unequal Lateral Buds

In the decussate plants Alternanthera philoxeroides and Hygrophilasp. the opposite axillary bud primordia are of unequal sizefrom the time of their inception; the larger or + buds lie alongone helix and the smaller or – buds along another (helicoidalsystem). In decapitated plants of Alternanthera both buds grewout, but unequally; if the node was vertically split growthof the two shoots was more equal, and if the + buds were excisedgrowth of the – shoots approximately equalled that ofcontrol + shoots. In decapitated shoots of Hygrophila grownin sterile culture only one bud, the + or larger one, grew outat each of the upper nodes. In excised cultured nodes, also,only the + bud grew out; but if the nodes were split longitudinallyboth buds grew out, initially rather unequally. These experimentssupport the view that the regulation of branching in these specieshas two components, apical dominance and the dominance of thelarger (+) bud over the smaller (–) bud at the same node.The restriction of growth potentiality imposed on the –bud is not permanent but can be modified. Further correlativeeffects on bud outgrowth include those of the subtending leavesand of buds at other nodes.     

Branch geometry in Cornus kousa (Cornaceae): computer simulations

Computer simulations similar to actual trees were constructed using simple branching rules. Branch orientation with respect to the direction of gravity was a fundamental consideration. In Cornus kousa BUERG. ex HANCE, several types of branches develop from winter buds, varying from orthotropic shoots to plagiotropic ones. Based on actual observations and measurements of branching structures with a wide range of orientations, we made a flexible geometrical model consisting of five forking branches that varied in outgrowth depending on the direction of the shoot with respect to gravity. Repetition of the branching by computer generated a realistic tree pattern, which was close to the shape of a young C. kousa tree. Reproductive shoots seem to be under a branching rule that was a modification of vegetative branching, although the reproductive branch size was considerably smaller than the vegetative one, and reproductive branching was bifurcated instead of five-forked. We conclude that all branchings in orthotropic and plagiotropic shoots in the vegetative phase and shoots in the reproductive phase are formed under the same branching rule, but each has different parameter values.     

Maintenance mechanisms of the pipe model relationship and Leonardo da Vinci’s rule in the branching architecture of <Emphasis Type="Italic">Acer rufinerve</Emphasis> trees

The pipe model relationship (constancy of branch cross-sectional area/leaf area) and Leonardo da Vinci’s rule (equality of total cross-sectional area of the daughter branches and cross-sectional area of their mother branch) are empirical rules of tree branching. Effects of branch manipulation on the pipe model relationships were examined using five Acer rufinerve trees. Half the branches in each tree were untreated (control branches, CBs), and, for the others (manipulated branches, MBs), either light intensity or leaf area (both relating to photosynthetic source activity), or shoot elongation (source + sink activities), was reduced, and responses of the pipe model relationships were followed for 2 years. The pipe model relationship in MBs changed by suppression of source activity, but not by simultaneous suppression of source + sink activities. The manipulations also affected CBs in the year of manipulation and both branches in the next year. The branch diameter growth was most affected by light, followed by shoot elongation and leaf area, in that order. Because of the decussate phyllotaxis of A. rufinerve, one branching node can potentially have one main and two lateral branches. Analysis of 295 branching nodes from 13 untreated trees revealed that the da Vinci’s rule held in branching nodes having one shed branch but not in the nodes without branch shedding, indicating the necessity of natural shedding of branches for da Vinci’s rule to hold. These analyses highlight the importance of the source–sink balance and branch shedding in maintenance of these empirical rules. This article was contributed at the invitation of the Editorial Committee.     

Shoot development ofAucuba japonica I. Morphological study

The shoot development ofAucuba japonica was studied morphologically. The shoot shows dichasial branching in connection with the formation of a terminal inflorescence and shows a decussate phyllotaxis even in the reproductive phase. The sequence of initiation of successive foliar appendages is very precise, hence the foliage leaf, scale leaf and bract can be compared with each other even at their stages of initiation. In the stage of proximal foliage leaf formation the shoot apex is flat, while in the stage of formation of distal foliage leaves, bud scales and proximal bracts, it becomes concave. In the stage of formation of distal bracts the apex becomes domed. Plastochron durations are relatively long in the vegetative phase in comparison with other plants, and the duration from initiation of the first pair of appendages to that of the second is about one and a half months. Both male and female inflorescences exhibit basically a thyrsoid type of monotelic synflorescence.     

Development of Axillary and Leaf-opposed Buds in Rattan Palms

Axillary vegetative buds are present in Calamus, Ceratolobus,and Plectocomiopsis. Two species of Daemonorops Sect. Piptospathaalso have axillary vegetative buds. All species of Daemonoropshave only displaced adnate axillary inflorescence buds. A singlebud is initiated in the axil of the first or second leaf primordiumin a way similar to that for axillary inflorescence buds. Themeristem is displaced during development on to the internodeabove and sometimes on to the base of the leaf above. Leaf-opposedvegetative buds occur in five species of Daemonorops Sect. Cymbospathaand in one species of Daemonorops Sect. Piptospatha. This typeof bud is initiated 180° away from the axil of the firstor second leaf primordium. It is not a displaced axillary bud,but does become adnate to the internode above like the axillarybuds. One or more leaves, transitional between juvenile andadult, on a shoot often subtend both types of buds. Myrialepishas leaf-opposed vegetative buds, but their development wasnot observed. Korthalsia has buds that are displaced about 130°from the leaf axil and are intermediate between the axillaryand the leaf-opposed condition. Other forms of vegetative budsare described: multiple buds in Plectocomia, aerial forkingin Korthalsia, and suckering from inflorescences and from aerialstems in Calamus. bud development, rattan palms, palm taxonomy, branching     

Bud Content and its Relation to Shoot Size and Structure in Nothofagus pumilio(Poepp. et Endl.) Krasser (Nothofagaceae)

Buds of shoots from the trunk, main branches, secondary branchesand short branches of 10–21 year-old Nothofagus pumiliotrees were dissected and their contents recorded. The numberof differentiated nodes in buds was compared with the numberof nodes of sibling shoots developed at equivalent positionsduring the following growing season. Axillary buds generallyhad four cataphylls, irrespective of bud position in the tree,whereas terminal buds had up to two cataphylls. There were morenodes in terminal buds, and the most distal axillary buds, oftrunk shoots than in more proximal buds of trunk shoots, andin all buds of shoots at all other positions. The highest numberof nodes in the embryonic shoot of a bud varied between 15 and20. All shoots had proximal lateral buds containing an embryonicshoot with seven nodes, four with cataphylls and three withgreen leaf primordia. The largest trunk, and main branch, shootswere made up of a preformed portion and a neoformed portion;all other shoots were entirely preformed. In N. pumilio, theacropetally-increasing size of the sibling shoots derived froma particular parent shoot resulted from differences in: (1)the number of differentiated organs in the buds; (2) the probabilityof differentiation of additional organs during sibling shootextension; (3) sibling shoot length; (4) sibling shoot diameter;and (5) the death of the apex and the most distal leaves ofeach sibling shoot. Copyright 2000 Annals of Botany Company Axis differentiation, branching, bud structure, leaf primordia, neoformation, Nothofagus pumilio, preformation, size gradient     

AXILLARY AND DICHOTOMOUS BRANCHING IN THE PALM CHAMAEDOREA

In both Chamaedorea seifrizii Burret and C. cataractarum Martius each adult foliage leaf subtends one axillary bud. The proximal buds in C. seifrizii are always vegetative, producing branches (= new shoots or suckers); and the distal buds on a shoot are always reproductive, producing inflorescences. The prophyll and first few scale leaves of a vegetative branch lack buds. Transitional leaves subtend vegetative buds and adult leaves subtend reproductive buds. Both types of buds are first initiated in the axil of the second or third leaf primordia from the apex, P2 or P3. Later development of both types of bud tends to be more on the adaxial surface of the subtending leaf base than on the shoot axis. Axillary buds of C. cataractarum are similarly initiated in the axil of P2 or P3 and also have an insertion that is more foliar than cauline. However, all buds develop as inflorescences. Vegetative branches arise irregularly by a division of the apex within an enclosing leaf (= P1). A typical inflorescence bud is initiated in the axil of the enclosing leaf when it is in the position of P2 and when each new branch has initiated its own P1. No scale leaves are produced by either branch and the morphological relationship among branches and the enclosing leaf varies. Often the branches are unequal and the enclosing leaf is fasciated. The vegetative branching in C. cataractarum is considered to be developmentally a true dichotomy and is compared with other examples of dichotomous (= terminal) branching in the Angiospermae.     

Bud Structure in Relation to Shoot Morphology and Position on the Vegetative Annual Shoots of Juglans regia L. (Juglandaceae)

The length and basal diameter of all lateral and terminal budsof vegetative annual shoots of 7-year-oldJuglans regia treeswere measured. All buds were dissected and numbers of cataphylls,embryonic leaves and leaf primordia were recorded. Each axillarybud was ranked according to the position of its associated leaffrom the apex to the base of its parent shoot. Bud size andcontent were analysed in relation to bud position and were comparedwith the size and number of leaves of shoots in equivalent positionswhich extended during the following growing season. Length andbasal diameter of axillary buds varied according to their positionon the parent shoot. Terminal buds contained more embryonicleaves than any axillary bud. The number of leaves was smallerfor apical and basal axillary buds than for buds in intermediatepositions on the parent shoot only. All new extended shootswere entirely preformed in the buds that gave rise to them.Lateral shoots were formed in the median part of the parentshoot. These lateral shoots derived from buds which were largerthan both apical and basal ones. Copyright 2001 Annals of BotanyCompany Juglans regia L., Persian walnut tree, branching pattern, preformation, bud content, shoot morphology     

Effect of Severity of Defoliation on the Viability of Reproductive and Vegetative Axillary Buds of Trifolium repens L.

This glasshouse experiment was performed to assess the effectsof a range of constant defoliation regimes applied to cuttingsof a single large-leaved genotype ofTrifolium repens L. on theviability of its axillary buds. Plants were established to comprisea single main stolon (axillary branches were removed) and defoliationtreatments were applied by removing the older (basal) leavesuntil leaf complements of 1·0, 1·5, 2·0,2·5, 3·0 or all leaves (control) remained. Basalleaves were subsequently removed as necessary to maintain thetarget leaf complements. Only severe defoliation (leaf complements of 1·0 and1·5) induced a loss of viability in axillary buds. Lossof viability was greatest in reproductive buds present withinthe apical bud when the treatments were first imposed. Althoughthe most severe treatment (leaf complement 1·0) resultedin death of half the plants, in plants surviving that treatment,death of vegetative axillary buds was restricted to 21% of thevegetative buds at the three youngest node positions withinthe apical bud at the time of treatment application. No othertreatment induced any loss of viability of vegetative buds.There was no loss of viability of axillary buds at nodes formedafter the treatments were imposed. The frequency of initiationof inflorescences at nodes formed after treatments were imposeddecreased as defoliation severity increased. Severe defoliation resulted in marked changes in plant morphologyindicative of a sharp decrease in availability of intraplantresources. It was concluded that under severe defoliation: (1)the potential for vegetative growth (as represented by viablevegetative axillary buds) was maintained at the expense of reproductivegrowth; and (2) that the loss of viability of axillary budswas associated with the sudden changes in physiological processesinduced by defoliation as there was no loss of viability inbuds formed after plants had adjusted their phenotype to oneof smaller size. Trifolium repens L.; white clover; defoliation; axillary buds; viability; inflorescences     

Grafting Experiments on Correlative Effects between Lateral Buds

Shoots of Hygrophila sp., which are decussate and have budsof unequal size at a node, were grown in liquid culture. Inexcised nodes it is known that the larger (+) bud inhibits thesmaller (–) bud in the axil of the opposite leaf, andonly one shoot grows out; in nodes split longitudinally bothbuds grow out. When nodes were split and grafted together again(+/– grafts), in general only one bud grew out; if aluminiumfoil was introduced at the nodal region both buds grew out.Thus the inhibitory effect of a + on a – bud is laterallytransmissible across a graft union. In +/– grafts of half-nodesdiffering in age by two plastochrones, a higher proportion yieldedtwo shoots, suggesting that the age differential had some importance.This view is supported by observations on sectioned material.Grafts having two + or two – buds (+/+ grafts) were madebetween half-nodes differing in age by two plastochrones; inthe majority both buds grew out. Thus a + bud inhibits a –bud but usually not another + bud; in either case a considerabledifference in stage of development of the half-nodes may affectthe results. It is concluded that bud dominance resembles apicaldominance, and is probably mediated by hormonal means.     

FLORAL INDUCTION DOES NOT ALTER THE GROWTH PARAMETERS OF FUCHSIA

Floral induction by night interruption of Fuchsia hybrida cv. Lord Byron, a quantitative long-day plant with decussate phyllotaxis and an indeterminate flowering habit, altered neither the rate of leaf initiation nor the rate of leaf expansion; nor did flower initiation and development change the vegetative growth of the plants. This was diagnosed using plastochron duration and plastochron ratio measurements before, during, and after a 10-day induction period. A comparison between indeterminate and determinate flowering is made using these two parameters.     

Growth Context and Fate of Axillary Meristems of Young Peach Trees. Influence of Parent Shoot Growth Characteristics and of Emergence Date

The relationship between several growth components of a shootand the fates of the axillary meristems (developing in the axilsof the leaves) borne by that shoot were studied, on first-ordershoots of young peach trees. A comprehensive picture of thoserelationships was obtained by a discriminant analysis. Shootgrowth at meristem emergence date was characterized by internodelength, leaf-production rate and leaf-unfolding duration. Allpossible fates of axillary meristems at the end of the growingseason (i.e. blind nodes, single vegetative or flower bud, budassociations, sylleptic or proleptic shoots) were considered.Shoot-elongation rate determined meristem fates quantitatively.The number of buds produced by a meristem increased when theshoot-elongation rate increased. Qualitatively, the fate of axillary meristems was related tothe balance between shoot-growth components. If the subtendingleaf unfolded slowly, sylleptic or proleptic shoots were morelikely to develop than bud associations, for high shoot-elongationrates; and flower buds were more frequent than vegetative buds,for low shoot-elongation rates. Compared to flower buds, blindnodes appeared for similar shoot-elongation rates but longerinternodes and lower leaf-production rates. The emergence dateslightly modified the relation between shoot growth and axillary-meristemfates, but the main features held true throughout the growingseason. The relationships between shoot growth and meristem fates mayresult from competitive interactions between the growing subtendingleaf and the developing axillary meristem. Growing conditionsmight also influence both shoot growth and meristem fates byfavouring either cell enlargement or cell division.Copyright1995, 1999 Academic Press Peach tree, Prunus persica (L.) Batsch, axillary meristem, meristem fate, branching, flowering, shoot growth, discriminant analysis, exploratory analysis     

The diversity and ontogenetic changes of phyllotaxis in wild-types and two leaf form mutants of Antirrhinum majus L.

We have analysed the phyllotactic patterns of the main shoot in vegetative and generative phases of growth in wild type and mutant plants of Antirrhinum majus L. Wild types 'Sippe50' and 'W l08' were compared to mutants grminifolia and phanlastica . The normal vegetative phyllotaxis of the wild type plants is decussate, but the inflorescence phyllotaxis is spiral and of the Fibonacci type. The phyllotaxis patterns of the mutants differ strongly from that of the wild type. Besides decussate phyllotaxis, whorls of three or four elements as well as spiral patterns in vegetative phase were observed. The vegetative phyllotaxis in mutants is ontogenetically unstable with frequent transitions between patterns, including the reversion of chirality of spiral phyllotaxis. The number of transitions per plant was larger in graminifolia than in phantastica . The inflorescence phyllotaxis was more stable and occasional non-typical phyllotaxis patterns finally transformed to a Fibonacci pattern. The results suggest a possible role of genetic factors in determining the regularity of spatial arrangement of organs.     

Unusual branch development in the palm, Chrysalidocarpus

Vegetative branch buds of C. lutescens are non-axillary and occur within an abaxial, tubular extension of the leaf sheath, either at the base of a shoot or aerially, a position unusual for palms. Buds are initiated on the abaxial surface of a leaf during its first plastochron (the youngest leaf primordium). The foliar origin of the vegetative bud appears to be unique for angiosperms. In contrast, inflorescence buds are axillary and are initiated as an adaxial ridge on the base of a leaf during its third plastochron (the third primordium from the apex). Aerial branches and basal suckers are developmentally identical and changes in their phyllotaxis are described. As far as can be established by comparative morphology, other species of Chrysalidocarpus have the same type of branch development as in C. lutescens. The development of branches is related to the morphogenetic characteristics of arborescent monocotyledons.     

PHOTOPERIOD INDUCED CHANGE IN PHYLLOTAXIS IN XANTHIUM

Photoperiodic floral induction in Xanthium, achieved by subjecting the plants to two long nights, is accompanied by a transient change of the phyllotaxis from the (2, 3) contact parastichy pattern of vegetative plants, to a (3, 5) pattern during the transition. To specify the phyllotaxis, two parameters were estimated from transverse sections of apical buds of control and treated plants: the divergence angle, α, and the plastochron ratio, a. The plastochron ratio decreased progressively during transition from the vegetative to the reproductive state of growth, from a = 1.48 initially to a = 1.15 six days after the beginning of induction. The divergence angle was not altered during the transition. This change in phyllotaxis is interpreted as a change in the relative positioning of leaf primordia on the transitional apex. This transient change appears to be identical with the previously described long-term change of the phyllotaxis of Xanthium brought about by treatment of plants with gibberellic acid.     

Preformation of Node Number in Vegetative and Reproductive Proleptic Shoot Modules of Persea (Lauraceae)

The numbers of nodes on single flush terminal and axillary shootmodules were determined in a range of Persea species and cultivars.They were compared with node numbers in apical and axillarybuds to investigate whether preformation or neoformation ofnodes occurred. Mean number of nodes on terminal shoots was14 for vegetative shoot modules and 21 for reproductive shootmodules, and was similar across species, cultivars, rootstocks,locations and climates. In the cultivar 'Hass', numbers of nodeson axillary shoot modules were variable, and lower than thosefor primary shoot modules forming the dominant growth axis ofannual growth modules. There was a mean of 12 nodes for vegetativeproleptic shoot modules, 15 for reproductive proleptic shootmodules and six for sylleptic shoot modules, which were invariablyvegetative. All nodes were preformed within both apical andaxillary proleptic buds. This was not the case in syllepticbuds, which burst contemporaneously with extension of the parentaxis. The majority (63%) of reproductive buds formed indeterminatecompound inflorescences. They carried six basal bud scales,six axillary inflorescences and their subtending bracts, andup to nine true leaves.Copyright 1994, 1999 Academic Press Persea Clus., avocado, Persea americana Mill., bud morphology, shoot growth, preformation, prolepsis     

Anisophylly of Lateral Shoots

The inequality shown by anisophyllous leaves in plants withdecussate phyllotaxis can be measured as the difference in sizebetween the two leaves at a node, expressed as a percentageof their mean size. This is referred to as the degree of anisophylly. In Acer anisophylly of two types with different causation occurs.

1. 1. Primary anisophylly which is shown only by first year shootsdeveloped from axillary buds and which is due to the inequalityof the leaf primordia. Primary anisophylly is thus determinedby events occurring during the formation of the bud and is irreversible.
2. 2. Secondary amsophylly which is shown chiefly by the second(and subsequent) year's extension growth of lateral shoots frombuds which are isophyllous and which is due to the positionin which the shoot develops. This type of anisophylly is reversibleif the position of the developing shoot be changed.

Effects of secondary anisophylly may be superimposed on thoseof primary anisophylly. The recognition of these two causally distinct types of anisophyllyoccurring in one plant does much to resolve the apparently conflictinginformation available relating to lateral anisophylly.     

Shoot initiation in the leaflet axils of compound leaves from micropropagated shoots of juvenile and mature common ash (Fraxinus excelsior L.)

Micropropagation of a mature ash tree has been achieved forthe first time. The main obstacle encountered was contaminationof the initial explants with microorganisms. However, once apparentlysterile shoots had been obtained, shoot proliferation was achievedmost effectively by culturing nodes on Driver and Kuniyuki walnutmedium containing 22.2 µM benzyladenine. After severalsubcultures, a species of Bacillus appeared with the matureculture line, but it did not affect shoot or root developmentadversely. With successive subculturing, shoots of the matureash clone became progressively easier to root. Pinnately compoundleaf explants from micropropagated shoots of two seedlings andthe mature tree, placed on to Murashige and Skoog-based culturemedia supplemented with 4.4µM-phenyl-N-1,2,3-thiadiazol-5-ylurea,produced shoot buds. Most buds developed from the rachis atthe points of attachment of the leaflets. When transferred toDriver and Kuniyuki walnut medium, buds from one of the seedlinglines and from the mature tree, elongated into shoots, and weresubsequently transferred to media for micropropagation and rooting.Shoot initiation is discussed in relation to possible occurrenceof vestigial meristems in the axils of leaflets and the partialshoot theory of leaf structure. Key words: Forestry, leaf structure, partial shoot theory, thidiazuron, tree