Winter peridermal conductance of apple trees: lammas shoots and spring shoots compared

Lammas shoots are flushes formed by some woody species later in the growing season. Having less time to develop, tissue formation is suggested to be incomplete leading to a higher peridermal water loss during consecutive months. In this study, we analysed morphological and anatomical parameters, peridermal conductance to water vapour and the level of native embolism in mid-winter and late-winter of lammas shoots and normal spring shoots of the apple varieties Malus domestica ‘Gala’ and ‘Nicoter’. Lammas shoots showed a significantly higher shoot cross-sectional area due to larger pith and corticular parenchyma areas. In contrast, phloem was significantly thicker in spring shoots. No pronounced differences were observed in xylem and collenchyma thickness or mean hydraulic conduit diameter. The phellem of spring shoots was composed of more suberinised cells compared to lammas shoots, which led to a significantly higher peridermal conductance in the latter. The amount of native embolism in mid-winter did not differ between shoot types, but in late-winter lammas shoots were more embolised than spring shoots. Data show that the restricted vegetation period of lammas shoots affects their development and, in consequence, their transpiration shield. This may also pose a risk for winter desiccation.     

Flowering Habit and Vegetative Behaviour in Tea (Camellia sinensis L) Seed Trees in North-East India

Apical growth of a tea shoot occurs by a succession of flushesseparated by short periods of rest. This paper describes theexternal morphology of flowering, fruiting, and abscission ofleaves of the tea plant in north-east India in relation to thephasic activity of shoot apices. All shoots on a tree make leafy growth when a new cycle of growthbegins in the spring, but terminal buds apparently become dormantas the season advances. Apparently dormant terminal buds shedbud scales, leaving on the stem a considerable number of scars,representing leafless cataphyllary flushes. These cataphyllaryflushes are produced at the same time as the leafy flushes onother shoots. A flower is formed only in the axil of a bud scale. Flowerswhich appear to develop in leaf axils are in fact inserted inthe axils of bud scales of the axillary buds. A distal leafy flush is without flowers. Flowers appear in itsleaf axils only when the terminal bud starts growth for thenext higher flush. A distal floriferous cataphyllary flush appearsas a terminal cluster of flowers. Thus, there is an acropetalsuccession of flowers, flush by flush on a caulome, determinedby the phasic activity of the apical bud. The main crop of flowers exposes anthers from the end of thethird flush (late September to early October) until the endof the winter period of growth (late January to early February).In some plants a second, minor crop of flowers appears in thespring between the end of the first and beginning of the secondflushes. In spite of considerable time lag between anthesis,the fruits produced by these two crops of flowers mature anddehisce at the same time during October to November. Abscission of leaves is also dependent upon the phasic activityof the apical buds. Only the top two flushes of a shoot possessleaves. Resumption of apical growth for a third flush, leafyor cataphyllary, causes the abscission of leaves on the lowermostof the three flushes. Two cataphyllary flushes therefore resultin the loss of all leaves on a shoot.     

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     

Periods of organogenesis in mono- and bicyclic annual shoots of Juglans regia L. (Juglandaceae)

The organogenetic cycle of shoots on main branches of 4-year-old Juglans regia trees was studied. Mono- and bicyclic floriferous and vegetative annual shoots were analysed. Five parent annual shoot types were sampled between October 1992 and August 1993. Organogenesis of summer growth units was monitored between 16 Jun. and 3 Aug. 1993. Variations over time in the number of nodes, cataphylls and embryonic green leaves of terminal buds were studied. The number of nodes of parent shoot buds was compared with the number of nodes of shoots derived from parent shoot buds. The spring growth units of mono- and bicyclic shoots consist exclusively of preformed leaves which were differentiated, respectively, during the spring flush of growth (mid-April until mid-May) or the summer flush of growth (mid-June until early August) in the previous growing season. Thus, winter buds may consist of flower and leaf primordia differentiated in two different periods during annual shoot extension. The summer growth units of bicyclic shoots consist of preformed leaves that were differentiated in spring buds during the spring flush of growth in the current growing season. Bud morphology is compared between spring and summer shoots.     

Execution of the auxin replacement apical dominance experiment in temperate woody species

The classic Thimann-Skoog or auxin replacement apical dominance test of exogenous auxin repression of lateral bud outgrowth was successfully executed in both seedlings and older trees of white ash, green ash, and red oak under the following conditions: (1) decapitation of a twig apex and auxin replacement were carried out during spring flush, (2) the decapitation was in the previous season's overwintered wood, and (3) the point of decapitation was below the upper large irrepressible lateral buds but above the lower repressible lateral buds. Although it has been suggested that neither auxin, the terminal bud, nor apical dominance have control over the outgrowth of the irrepressible buds during spring flush, there is evidence in the present study that indicates that such control over the repressible buds exists. In seedlings, second-order branching, which resulted from decapitation of elongating current shoots, was also inhibited by exogenous auxin in the three species. Hence, the auxin replacement experiments did work on year-old proleptic buds (of branches of older trees) that would have entered the bud bank and also on current buds of seedlings. Cytokinin treatments were ineffectual in promoting bud growth.     

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.     

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.     

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     

Differences in proleptic and epicormic shoot structures in relation to water deficit and growth rate in almond trees (Prunus dulcis)

Background and Aims

Shoot characteristics differ depending on the meristem tissue that they originate from and environmental conditions during their development. This study focused on the effects of plant water status on axillary meristem fate and flowering patterns along proleptic and epicormic shoots, as well as on shoot growth rates on ‘Nonpareil’ almond trees (Prunus dulcis). The aims were (1) to characterize the structural differences between proleptic and epicormic shoots, (2) to determine whether water deficits modify shoot structures differently depending on shoot type, and (3) to determine whether shoot structures are related to shoot growth rates.

Methods

A hidden semi-Markov model of the axillary meristem fate and number of flower buds per node was built for two shoot types growing on trees exposed to three plant water status treatments. The models segmented observed shoots into successive homogeneous zones, which were compared between treatments. Shoot growth rates were calculated from shoot extension measurements made during the growing season.

Key Results

Proleptic shoots had seven successive homogeneous zones while epicormic shoots had five zones. Shoot structures were associated with changes in growth rate over the season. Water deficit (1) affected the occurrence and lengths of the first zones of proleptic shoots, but only the occurrence of the third zone was reduced in epicormic shoots; (2) had a minor effect on zone flowering patterns and did not modify shoot or zone composition of axillary meristem fates; and (3) reduced growth rates, although patterns over the season were similar among treatments.

Conclusions

Two meristem types, with different latency durations, produced shoots with different growth rates and distinct structures. Differences between shoot type structure responses to water deficit appeared to reflect their ontogenetic characteristics and/or resource availability for their development. Tree water deficit appeared to stimulate a more rapid progression through ontogenetic states.     

Flowering and fruiting phenology in the apomictic blackberry, Rubus nessensis (Rosaceae)

Five stands of an apomictic blackberry, Rubus nessensis , were studied. Biennial shoots flower and fruit in their second year on lateral branches developed from axillary buds. Most premature death of reproductive units occurs by withering of entire laterals, presumably due to damage by frost during the preceeding autumn or winter. Health status of the floricane, assessed as percentage live axillary buds in the spring, was positively correlated with the reproduction parameters. Shoot height as well, is positively correlated with number of flower buds and flowers produced on each lateral but, however, negatively correlated with number of berries, probably because tall shoots tend to suffer more damage by frost which seriously afflicts the vascular tissues of the laterals.
The position of the axillary buds on the shoot affects survival and degree of fertility, those situated medially greatly surpassing apical and basal ones.     

Morphological studies on the genusClematis Linn

While the axes of branched axillary inflorescences seen in most species ofClematis show fundamentally the same features in nodal vasculature as the vegetative stems, those of simple axillary inflorescences, having only a pair of opposite sterile bracts at specific positions, exhibit an exclusive feature in nodal vasculature because they have entirely lost lateral branches or buds in the bract axils. Noticeably, at the nodal level of the axis of the simple inflorescence ofC. japonica andC. Williamsii, trace bundles to the missing lateral branches are formed and extend unaltered beyond the node. Thus, in these two species stelar bundles in the inflorescence axis generally increase in number upwardly from six to eight or more through the node with sterile bracts. On the basis of both anatomical and morphological data, a probable evolutionary trend in floral shoots with simple axillary inflorescences is proposed. The type of floral shoot ofC. tosaensis bearing simple, scale-subtended and basally fascicled inflorescences is the most primitive and constitutes an initial phase of evolution, and progressive changes in a kind of inflorescence-subtending leaves as well as in the shape and position of the sterile bracts resulting in the type of floral shoot ofC. japonica and further in that ofC. obvallata. A floral shoot ofC. Williamsii follows another line of evolution from an ancestral type similar to that ofC. tosaensis.     

Thorn and hook ontogeny in Artabotrys hexapetalus (Annonaceae)

Artabotrys hexapetalus is widely planted in the tropics and is known as "climbing ylang-ylang," an ornamental liana or woody climber. New natural sprouts, or water shoots, and those induced by the damage of Hurricane Andrew (24 August 1992) were collected and fixed in formalin/acidic acid/alcohol. Seeds from these plants were planted and grown in a greenhouse where seedling morphology was observed and young material collected and fixed. The development of lateral plagiotropic and orthotropic shoots was studied using both epi-illumination light microscopy and scanning electron microscopy. A series of buds develops in the axils of leaves on the orthotropic shoot. At the lateral margins of the axillary shelf, plagiotropic shoots form that will develop into either vegetative shoots, or thorns, or sympodial shoots that bear hooks and flowers. In between the two marginal buds, a series of median vertical buds develop that either remain dormant or grow out as renewal orthotropic shoots. Previous work that suggested that the plagiotropic shoot buds were displaced out of the median vertical series of supernumerary buds is not supported. The sympodial development of plagiotropic branches as inflorescence hooks is documented.     

Development of shoot inHydrangea macrophylla II. sequence and timing

The development of axillary buds, terminal buds, and the shoots extended from them was studied inHydrangea macrophylla. The upper and lower parts in a nonflower-bearing shoot are discernible; the preformed part of a shoot develops into the lower part and the neoformed part into the upper part (Zhou and Hare, 1988). These two part are formed by the different degrees of internode elongation at early and late phases during a growth season, respectively. Leaf pairs in the neoformed part of the shoot are initiated successively with a plastochron of 5–20 days after the bud burst in spring. The upper axillary buds are initiated at approximately the same intervals as those of leaf pairs, but 10–30 days later than their subtending leaves. Changes in numbers of leaf pairs and in lengths of successive axillary buds show a pattern similar to the changes in internode lengths of the shoot at the mature stage. The uppermost axillary buds of the flower-bearing shoot often begin extending into new lateral shoots when the flowering phase has ended. The secondary buds in terminal and lower axillary buds are initiated and developed in succession during the late phase of the growth season. Internode elongation seems to be important in determining the degrees of development of the axillary buds. Pattern of shoot elongation is suggested to be relatively primitive. Significances of apical dominance and environmental conditions to shoot development are discussed.     

Correlation of Endogenous Gibberellic Acid with Initiation of Mango Shoot Growth

Stems of mango (Mangifera indica L.) rest in a nongrowing, dormant state for much of the year. Ephemeral flushes of vegetative or reproductive shoot growth are periodically evoked in apical or lateral buds of these resting stems. The initiation of shoot growth is postulated to be primarily regulated by a critical ratio of root-produced cytokinins, which accumulate in buds and by leaf-produced auxin, which decreases in synthesis and transport over time. Exogenously applied gibberellic acid (GA3) delays initiation of bud break but does not determine whether the resulting flush of growth is vegetative or reproductive. We tested the hypothesis that endogenous GA3, which influences release of these resting buds, may decrease in stem tips or leaves with increasing age of mango stems. GA3 and several other GAs in stem tip buds and leaves were identified and quantified in stems of different ages. The major endogenous GAs found in apical buds and leaves of vegetative mango stems were early 13-hydroxylation pathway gibberellins: GA1, epi-GA1, GA3, GA19, GA20, and GA29, as identified by gas chromatography-mass spectrometry (GC-MS). A novel but unidentified GA-like compound was also present. The most abundant GAs in apical stem buds were GA3 and GA19. Contrary to the hypothesis, the concentration of GA3 increased within buds with increasing age of the stems. The concentrations of other GAs in buds were variable. The concentration of GA3 did not change significantly with age in leaves, whereas that of most of the other GAs declined. GA1 levels were greatest in leaves of elongating shoots. These results are consistent with the concept that rapid shoot growth is associated with synthesis of GAs leading to GA1. The role of GA3 in delaying bud break in mango is not known, but it is proposed that it may enhance or maintain the synthesis or activity of endogenous auxin. It, thereby, maintains a high auxin/cytokinin ratio similar to responses to GA3 that maintain apical dominance in other plant species.     

A morphological study of the development of the second inflorescences in strawberry (Fragaria×ananassa Duch.)

To clarify the timing of the differentiation of the first and second inflorescences in strawberry (Fragaria × ananassa Duch.), morphological changes on shoot apices during short day and low night temperature treatments were observed by scanning electron microscopy (SEM) and optical microscopy. Axillary buds just below the first inflorescence (axillary bud 1) became visible when sepal primordia of the primary flower were differentiated. By this time, other axillary buds had already developed. Axillary bud 1 developed four leaf primordia, and then a differentiated inflorescence at its summit. The phase transition of shoot apices from the vegetative to the reproductive phase may therefore trigger the differentiation of axillary bud 1 which is destined to develop into extension crowns.     

Correlation of Endogenous Gibberellic Acid with Initiation of Mango Shoot Growth

Stems of mango (Mangifera indica L.) rest in a nongrowing, dormant state for much of the year. Ephemeral flushes of vegetative or reproductive shoot growth are periodically evoked in apical or lateral buds of these resting stems. The initiation of shoot growth is postulated to be primarily regulated by a critical ratio of root-produced cytokinins, which accumulate in buds and by leaf-produced auxin, which decreases in synthesis and transport over time. Exogenously applied gibberellic acid (GA3) delays initiation of bud break but does not determine whether the resulting flush of growth is vegetative or reproductive. We tested the hypothesis that endogenous GA3, which influences release of these resting buds, may decrease in stem tips or leaves with increasing age of mango stems. GA3 and several other GAs in stem tip buds and leaves were identified and quantified in stems of different ages. The major endogenous GAs found in apical buds and leaves of vegetative mango stems were early 13-hydroxylation pathway gibberellins: GA1, epi-GA1, GA3, GA19, GA20, and GA29, as identified by gas chromatography-mass spectrometry (GC-MS). A novel but unidentified GA-like compound was also present. The most abundant GAs in apical stem buds were GA3 and GA19. Contrary to the hypothesis, the concentration of GA3 increased within buds with increasing age of the stems. The concentrations of other GAs in buds were variable. The concentration of GA3 did not change significantly with age in leaves, whereas that of most of the other GAs declined. GA1 levels were greatest in leaves of elongating shoots. These results are consistent with the concept that rapid shoot growth is associated with synthesis of GAs leading to GA1. The role of GA3 in delaying bud break in mango is not known, but it is proposed that it may enhance or maintain the synthesis or activity of endogenous auxin. It, thereby, maintains a high auxin/cytokinin ratio similar to responses to GA3 that maintain apical dominance in other plant species.     

CONTROL OF SHOOT-RHIZOME DIMORPHISM IN THE WOODY MONOCOTYLEDON,CORDYLINE (AGAVACEAE)

In Cordyline terminalis negatively geotropic leafy shoots and positively geotropic rhizomes develop from single axillary buds on either shoots or rhizomes. All axillary buds have similar morphogenetic potential when released from apical dominance. Experiments in which the orientation of the apex is changed, organs removed, or growth regulators applied indicate that after a rhizome is initiated, it is maintained as a rhizome by auxin originating in the leafy shoot. When auxin levels are lowered by changes in the orientation of the axis or shoot removal, the rhizome apex becomes a shoot apex, which appears to be the stable state of the actively growing apex. Benzyl adenine when applied exogenously to the apex or lateral buds has the same effect as lowering the auxin level. Gibberellic acid has no effect on the apex or lateral buds. High levels of exogenous naphthaleneacetic acid cause bud release and development of rhizomes from previously inhibited axillary buds of the shoot. However, it was not possible to convert a shoot apex into a rhizome apex by auxin treatment. It is suggested that the release of buds on the lower side of horizontal branches and of buds directly above a stem girdle is caused by high auxin levels on the lower side or distal to the girdle. The experimental results are discussed in relation to naturally occurring shoot-rhizome dimorphism.     

The growth pattern of shoots in Whytockia (Gesneriaceae) with phylogenetic implications

The morphogenesis of shoots in Whytockia W. W. Smith was investigated in order to reveal its growth pattern. The shoot in Whytockia has lost apical growth, which is contrary to the present knowledge about its growth pattern. Its stem is in fact a lateral branch system formed by sprouting of lateral buds in axils of small leaves substituting for the thoroughly restrained phyllogens. The unbranched stem of the genus is due to the restrained state of axillary buds in axils of large leaves. This so-called simple stem is secondary in phylogeny rather than relict in Epithemateae. According to the revealed growth pattern of the shoot in Whytockia, the present paper discusses the phylogenetic relationships between Whytockia and Loxonia, Monophyllaea and Rhynchoglossum in Epi-themateae.     

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     

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