Pattern of Shoot Growth in Zizyphus mauritiana and Z. oenoplia

Shoot polymorphism and patterns of sylleptic branching and shoottip abscission in two Zizyphus species are reported. Z. mauritiana(ber) produced three types of proleptic shoots: vigorous, normaland spur-type, by reiteration or following pruning;Z. oenopliahad only one type of shoot. Both species revealed up to thirdorder sylleptic branching on the vigorous proleptic shoots witha characteristic pattern and rhythm. Bud dormancy breaking chemicalsgiven as pre-pruning foliar sprays in ber, did not alter thepattern of sylleptic branching. Zizyphus oenoplia produced morefirst and second order sylleptic branches compared to Z. mauritiana.All main axes and 97% of first order sylleptic branches ofZ.mauritiana abscised their apical buds during summer, but inZ. oenoplia all main axes and 54% of first order sylleptic branchesremained active. Shoot tip abscission was almost complete onthe higher order sylleptic branches of both species. Activefirst order branches in Z. oenoplia were confined to the tophalf of the shoot. Both species had a few dormant apical budsduring the summer on their sylleptic branches, emerging mostlyfrom the middle portion of the shoot. Some of the most vigorousfirst order branches of Z. oenoplia, which had dormant apicalbuds during summer, showed a change in the frequency of syllepticbranching when they resumed growth following monsoon showers.These characteristic growth and branching patterns may haveadaptive value for canopy development under arid and semi-aridconditions. Copyright 1999 Annals of Botany Company Ber, branching pattern, shoot polymorphism, shoot tip abscission, sylleptic branching, Zizyphus mauritiana, Zizyphus oenoplia.     

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     

Architectural analysis of 1-year-old apple seedlings according to main shoot growth and sylleptic branching characteristics

Main shoot and sylleptic shoot growth characteristics were measured during and after the first year of growth of 255 Telamon x Braeburn apple seedlings. Although mean main shoot growth characteristics between branched and non-branched trees were significantly different, many non-branched trees expressed similar main shoot growth to branched trees. The variables describing length, number and position of the sylleptic shoots were used to classify branched trees into architecturally different groups. A continuum from trees with few and short shoots to trees with many long shoots is observed. The release of axillary buds from apical dominance is not under complete control by the apical meristem. Genetic seedling difference at the level of roots presumably plays an important role in sylleptic branching. Genetic variation in terms of number, position, and subsequent elongation of sylleptic shoots is clearly observed.     

Geographical Transition of Sylleptic/Proleptic Branching in Three Cinnamomum Species with Different Bud Types

Three native Japanese Cinnamomum(Lauraceae) species exhibiteither sylleptic and proleptic, or proleptic branching dependingon species-specific types of bud-structure (hypsophyllary andscaled-buds) and geographical distribution. Major branchingshifted from sylleptic at the axils of cataphylls and hypsophyllsin the southernmost C. doederleinii(latitudinal distributionrange 24–28° N) to both sylleptic at cataphyllary-and proleptic at hypsophyllary-axils in mid-latitudinal C. daphnoides(27–33°N), and to proleptic at the axils of foliage-leaves in the northernmostspecies C. japonicum(25–37° N). In all three Cinnamomumspecies, the sylleptic and proleptic branches expand aroundthe boundary between two successive yearly shoot-modules. Thepredominant, fertile axils having branches shifted from cataphylls(n year) and hypsophylls (n-1 year) in C. doederleinii and C.daphnoides, to foliage-leaves (n-1) in C. japonicum. The budsof proleptic branches at the axils of hypsophylls and foliage-leavesinevitably experience winter dormancy, hence the shoots areequipped with protective cataphylls instead of a hypopodium.Shoot length and leaf number gradually increased from the mostdistal cataphyllary sylleptic (n) shoots towards the most basalproleptic shoots at the axils of foliage-leaves (n-1). Prolepticshoots at the axils of foliage-leaves perform similarly to mainshoots in all three Cinnamomum species studied. The relativenumber of leaves borne on proleptic shoots gradually increasedfrom C. doederleinii, to C. daphnoides, and C. japonicum. Underthe seasonal, short growing periods of temperate latitudes,the increased shoot performance and bud protection of prolepticshoots are made possible through the earlier onset of bud formation(prolepsis) at the axils of the previous year's hypsophyllsand/or foliage-leaves in trees with rhythmic growth. Copyright2001 Annals of Botany Company Syllepsis, prolepsis, bud type, cataphyll, hypsophyll, shoot-growth cycle, evergreen broad-leaved tree, Cinnamomum doederleinii, Cinnamomum daphnoides, Cinnamomum japonicum     

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     

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     

Stand Density Effects on Branching in an Annual Legume (Senna obtusifolia)

Effects of proximity to nearest neighbours on shoot morphologyand branch growth were examined for Senna obtusifolia (L.) I.& B., an annual legume species that displays wide variationin branching. Periodic surveys described location and type ofgrowth at all nodes on plants in regularly spaced monocultures.Stands with interplant distances of 15-50 cm (51-5 plants m^-2)formed closed canopies with similar amounts of leaf area andbiomass. Number of lower primary branches, their degree of curvature,and location of branch apices relative to the main stem respondedto interplant distance relatively early in shoot growth, beforecanopy closure. The final effects of ten-fold differences inplanting density on the formation of these ascending lower brancheswere only two-fold differences in number of vertical shoot axeswithin the stand. An additional study examined response of lowerbranches to an adjacent gap in the stand. Initial number anddirection of branch growth were not affected by location ofthe gap. The first effect of crowding by neighbours was on number ofbranches initiated by developing shoots. Differences in branchingsubsequently increased through differentiaI survival and directionof branch growth. Such alterations principally affected lateralexpansion of individual shoots, with little effect on radialsymmetry or vertical distribution of leaves.Copyright 1994,1999 Academic Press Branch morphology, canopy, competition, light quality, Senna obtusifolia, shoot growth, sicklepod     

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     

桃树修剪分枝模式的模拟

在不同修剪手法下,对栽培桃树(Prunuspersica(L.)Batsch)不同母枝上的分枝模式进行了比较研究.从分枝模式来看:修剪后的母枝基本由3个不同的区域组成,基部是不萌发的潜伏芽形成的未分枝区域;中部是延迟分枝和多次分枝组成的分枝区域(主要的枝条类型有短枝、长枝和多次枝);顶部是被剪除的部分.我们通过隐式半马尔可夫模型来模拟这一分枝模式,主要是定量描述1次枝和多次枝在母枝上的数量及其分布状况.在上述模型中,未分枝区、延迟分枝区和多次分枝区称为瞬时态,被剪除的部分称为吸收态.模拟的结果与观察的结果进行对比后发现,两者具有很好的一致性.这说明隐式半马尔可夫模型是模拟植物分枝过程的一种有效方法,尽管隐式半马尔可夫链模型只是一个描述性的模型,但仍能对其所描述的生物现象进行解释,在预测修剪手法对母枝分枝模式影响方面比传统的方法具有明显的优势.本研究结果是建立三维虚拟桃树树冠分枝结构的基础.     

Preformed and neoformed extension of shoots and sylleptic branching in relation to shoot length in Tsuga canadensis

Summary Shoot systems developed over 3 successive years were investigated on 55 understorey Tsuga canadensis (L.) Carr. trees. Paired comparisons of preformed-leaf content of terminal buds and numbers of leaves produced on new shoots showed that neoformed leaves were produced in large numbers. Parent-shoot character was not useful in predicting numbers of preformed leaves, was better related to total leaves produced, but left the majority of the variation unexplained. This reflected the capacity of any terminal bud to produce a shoot with more or less neoformation, depending on conditions for growth. All shoots over 6 cm long produced sylleptic shoots that bore from two to many leaves and were arranged in a mesitonic pattern along the parent. Some of the longer sylleptic shoots produced lateral buds or second-order sylleptic shoots. Monopodial second-year extensions of sylleptic-shoot axes followed an acrotonic pattern, as did proleptic shoots from the few lateral buds borne on the parent shoots. Such lateral buds were more frequent on shorter parent shoots: they typically occurred near the proximal and distal ends. Duration of shoot extension was positively correlated with shoot length: terminal buds became evident as shoot extension neared cessation.     

Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees (Olea europaea L.)

The development of new shoots plays a central role in the complex interactions determining vegetative and reproductive growth in woody plants. To explore this role we evaluated the new shoots in the olive tree, Olea europaea L., and the effect of fruiting on new shoot growth and subsequent flowering. Five-year-old branches served as canopy subunits in order to obtain a global, whole-tree view of new shoot number, size and morphological origin. The non-bearing trees had many more shoots than the fruit-bearing trees, and a greater number of longer shoots. In both bearing conditions, however, the majority of shoots were less than 4 cm long, with shoots of progressively longer lengths present in successively decreasing frequencies. Six major shoot types were defined on the basis of apical or lateral bud origin and of parent shoot age. On fruit-bearing trees, the new shoots originated predominantly from the shoot apex, while on non-fruiting trees, they formed mainly from axillary buds, but in both cases, they tended to develop on younger parent shoots. The previous bearing condition of the tree was the main determinant for subsequent inflorescence development, which was independent of both shoot type and length. Thus, reproductive behavior strongly affected both the amount and type of new branching, but subsequent flowering level was more influenced by previous bearing than by the potential flowering sites on new shoots.     

桃树修剪分枝模式的模拟

在不同修剪手法下,对栽培桃树(Prunuspersica(L.)Batsch)不同母枝上的分枝模式进行了比较研究。从分枝模式来看修剪后的母枝基本由3个不同的区域组成,基部是不萌发的潜伏芽形成的未分枝区域;中部是延迟分枝和多次分枝组成的分枝区域(主要的枝条类型有短枝、长枝和多次枝);顶部是被剪除的部分。我们通过隐式半马尔可夫模型来模拟这一分枝模式,主要是定量描述1次枝和多次枝在母枝上的数量及其分布状况。在上述模型中,未分枝区、延迟分枝区和多次分枝区称为瞬时态,被剪除的部分称为吸收态。模拟的结果与观察的结果进行对比后发现,两者具有很好的一致性。这说明隐式半马尔可夫模型是模拟植物分枝过程的一种有效方法,尽管隐式半马尔可夫链模型只是一个描述性的模型,但仍能对其所描述的生物现象进行解释,在预测修剪手法对母枝分枝模式影响方面比传统的方法具有明显的优势。本研究结果是建立三维虚拟桃树树冠分枝结构的基础。     

Branching responses in Silphium integrifolium (Asteraceae) following mechanical or gall damage to apical meristems and neighbor removal

Branching in plants increases plant access to light and provides pathways for regrowth following damage or loss of the apical meristem. We conducted two experiments in an eastern Kansas tallgrass prairie to determine how apical meristem loss (by clipping), apical meristem damage (by insect galling), and increased light availability affected growth, reproduction, and branching in Silphium integrifolium (Asteraceae). The first experiment compared clipping with galling. Clipping increased axillary shoot numbers, while galling increased axillary shoot lengths, reflecting different allocation responses among damage types and inhibition of branching by galls. However, total capitulum production was less in all gall/clip treatments than in intact shoots. The second experiment compared clipping with mowing the surrounding vegetation to increase light availability. Mowing increased total leaf, total capitulum, and axillary shoot length and axillary capitulum production in clipped and unclipped plants and in large vs. small shoots. The presence of the neighboring canopy, not of an intact apical meristem, was therefore the stronger limitation on leaf and capitulum production. These experiments suggest that damage and light competition affected both branching frequency and the partitioning of resources among shoots, branches, and leaves. Because Silphium's growth form is widespread, similar responses may occur in other grassland forbs.     

Differences in Architecture and Shoot Growth during Stagnant and Extension Growth Phases of Acanthopanax sciadophylloides(Araliaceae)

The shoot growth of a deciduous tree, Acanthopanax sciadophylloidesFranch. et Savat. shows inter-annual intermittent repetitionof two distinctive phases, a stagnant growth phase (S-phase)and vigorous extension-growth (E-phase). To help understandthe differentiation mechanism, shoot development was studiedover time in both shoot phases. S-phase and E-phase shoots weredistinguished from each other by morphological traits: S-phaseshoots are characterized by higher allocation to leaves anda shorter period of stem growth, while E-phase shoots show continuousstem extension over the growing season. Specific leaf area didnot differ between the two phases. This shoot differentiationwas similar to the morphological differentiation of shoots betweenlong vs. short shoots found in some temperate trees. Leavesof both phases were well-dispersed through adjustment of petiolelength and leaf-blade size to reduce mutual shading within ashoot. Stem-wood density of current-year shoots was lower inE-phase compared with S-phase shoots. Leaves produced earlyin the season affected the growth phase of the following year.These results suggest that annual shoot differentiation of A.sciadophylloides was determined during the previous season andreflects leaf productivity in a given habitat during that growingseason. Copyright 2001 Annals of Botany Company Acanthopanax sciadophylloides, Araliaceae, biomass allocation, intermittent shoot growth, leaf display, shoot architecture, shoot differentiation     

Application of architectural analysis and AMAPmod methodology to study dwarfing phenomenon: the branch structure of 'Royal Gala' apple grafted on dwarfing and non-dwarfing rootstock/interstock combinations

Architectural analysis was applied to study branch development of 'Royal Gala' apple trees grafted with dwarfing and non-dwarfing rootstock/interstock combinations, which had been chosen to produce trees with a wide range of vigour. Using AMAPmod methodology, the structure of 3-year-old branches was described at four levels of representation: branch; annual shoot; growth unit; and node. Three types of growth units were distinguished: extension growth unit (vegetative unit with internode extension); vegetative spur with minimal internode extension; and fruiting spur or bourse. The aim of the analysis was to describe exactly how the rootstock/interstock combinations affected the structure building process. The number of extension growth units, vegetative spurs and fruiting spurs per annual shoot changed over the years, but this was not affected by rootstock/interstock combination. Compared with MM.106 rootstock, M.9 rootstock reduced the number of nodes per extension growth unit. In most cases, rootstock/interstock combination had no effect on the linear relationship between extension growth unit length and node number (R(2) = 0.88). Average internode length depended on unit node number, with internodes being shorter for units with fewer nodes. Thus the difference in apple branch size induced by the rootstock/interstock combinations was mainly due to a reduction in the length and number of neoformed nodes produced on extension growth units. As percentage budbreak of axillary buds on extension growth units was not affected by rootstock/interstock combination, differences in numbers of axillary annual shoots per branch were entirely due to differences in the total numbers of nodes extended during the previous year.     

Correlative Inhibition in the Shoot of Agropyron repens ( L.) Beauv

Correlative inhibition was investigated in plants of Agropyronrepens at two temperatures. Reciprocal inhibition ocrurred betweenthe main shoot apex and the outgrowing axillary shoots, withthe balance of inhibition varying with temperature. Apical dominancewas stronger at 10 °C than at 20 °C , but even at 10°C release of apical dominance by decapitation had onlyminor effects on the timing of outgrowth, growth pattern andrate of dry weight aocumulation of the axillary shoots. Dominanceof the main shoot apex by the axillary shoots was stronger at20 °C than at 10 °C. Removal of axillary buds preventeddecline in size and activity of the main shoot apex ard resultedin increased rates of primordium initiation, leaf emergenceand dry weight accumulation in the main shoot. It is suggestedthat a system of reciprocal dominance provides a mechanism formaintaining the characteristic habit of the grass plant andlimits growth in height of vegetative shoots. Agropyron repens (L.) Beauv, couch grass, correlative inhibition, apical dominance, shoot, apex     

In Vitro Shoot Regeneration from Callus, Leaf Axils and Petioles of Sugar Beet (Beta vulgaris L.)

Procedures are described for producing axillary shoots fromseedling apices and adventitious shoots from petioles and leaf-derivedcallus of sugar beet cultivars. The rate of adventitious shootregeneration from petioles was influenced by temperature, BAPconcentration of the medium, and the time in culture of theseedling apices from which the petioles were excised. Petiolesectioning confirmed that adventitious shoots originated inthe sub-epidermal parenchyma. Two distinct types of callus wereproduced from leaf explants, but only white friable callus wascapable of shoot development. This callus developed from browntissue and was composed of thin-walled cells with dense cytoplasmand prominent nuclei. Green compact callus with thick-walledlignified cells developed from green tissue, but did not produceshoots. Successful seed sterilization and shoot regenerationfrom petiole explants and callus was cultivar-dependent. Adventitiousshoots were rooted and successfully transplanted to pottingcompost under glasshouse conditions. Key words: Adventitious shoots, axillary shoots, callus, sugar beet (Beta vulgaris L.)     

The Effects of Temperature and Other Variables on the Contribution of Lateral Branches to Yield in Phaseolus vulgaris L.

At low temperatures (15/15 °C day/night) in controlled environmentsthe growth of lateral branches at the cotyledonary node ofPhaseolusvulgarts L is suppressed The suppression can be overcome byraising the temperature of the buds by approximately 6 °Cwith small electric heaters In order to test the practical significanceof the induction of branching for yields of the commercial crop,seedlings were raised in contrasted regimes and then transplantedto the field The effects of pretreatment regime on final yieldwere small, changes in the yield from axillary branches tendedto be balanced by compensatory changes in the yield from themain stem In another field experiment, synthetic growth substanceswere applied in order to suppress or enhance branching Changesin the amount of yield carned on branches were again offsetby compensatory changes in the yield from the main stem Compensatoryeffects between branches and main stem were also found in avariety trial However, in an experiment on a single cultivarand various levels of N fertilizer, compensatory effects werenot found, here, branch and main stem yields were positively,rather than negatively, correlated These results are discussedin relation to the intrinsic factors that govern yield in Pvulgaris Phaseolus vulgaris L, dwarf bean, axillary branches, correlative inhibition, temperature, growth substances, plant density, yield     

Sylleptic Shoot Formation in Young Apple Trees Exposed to Various Soil Temperature and Air Humidity Regimes in Three Successive Periods of the Growing Season

Under otherwise controlled conditions, the effect of two soiltemperatures (12 and 22 °C) and two air humidities (50 and90%) applied during three successive periods of 6 weeks (startingat the beginning of the growing season) on sylleptic shoot productionof young apple trees was evaluated during the first year ofgrowth. The treatment effects were mainly reflected on syllepticgrowth; the growth of the main shoot was much less influenced.In Period I sylleptic growth tended to be reduced at 12 °C.In Period II it responded greatly to the soil temperature inPeriod I resulting in a much stronger sylleptic growth at 12than at 22 °C. However, the actual soil temperature in PeriodII was not of any importance. In Period III growth activitywas low and there were no treatment effects. High humidity favouredsylleptic growth in Period I at the soil temperature of 22 °Cbut not at 12 °C, but in Period II the reverse was found.Humidity did not affect syllepsis in Period III. The varioustreatments greatly affected the distribution of sylleptic shootsalong the main shoot. The ability to grow out into syllepticshoots was restricted to buds in a certain stage of development.The results are discussed in terms of a negative feed back mechanismbetween shoot and root growth which controls sylleptic growth. Malus pumilaMill; soil temperature; air humidity; apple; syllepsis; branching