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     

Bud and growth-unit structure in seedlings and saplings of Nothofagus alpina (Nothofagaceae)

In temperate trees, axis length growth generally results from the differentiation of organs at the end of a growing season and the extension of such "preformed organs" in the next growing season. Neoformation, i.e., the simultaneous differentiation and extension of organs, has been studied for only a few species. Here we evaluated bud composition and growth unit (GU) size for seedlings and saplings of Nothofagus alpina, a valuable South American forest tree. Trunk GUs of seedlings and saplings included preformed and neoformed organs, whereas main-branch GUs of saplings were entirely preformed. The size of a GU was more closely related to the number of preformed green leaves than to the number of cataphylls of its preceding bud. Proximal buds of a trunk GU had more cataphylls and less green-leaf primordia than distal buds. Individual leaf area increased from proximal to distal positions on trunk GUs. For trunk and main-branch GUs, the length/width ratio was maximum for leaves in intermediate positions. The development of large neoformed leaves at the end of the growing season could increase the photosynthetic capacity of this species in late summer, when the activity of preformed organs is likely to be decreasing.     

Periods of organogenesis in shoots of Nothofagus dombeyi (Mirb.) oersted (Nothofagaceae)

The organogenetic cycle of main-branch shoots of Nothofagus dombeyi (Nothofagaceae) was studied. Twelve samples of 52-59 parent shoots were collected from a roadside population between September 1999 and October 2000. Variations over time in the number of nodes of terminal and axillary buds, and the length, diameter and number of leaves of shoots derived from these buds (sibling shoots) were analysed. The number of nodes of buds developed by parent shoots was compared with the number of nodes of buds developed, I year later, by sibling shoots. The length, diameter and number of leaves of sibling shoots increased from October 1999 to February 2000 in those shoots with a terminal bud. However, extension of most sibling shoots, including the first five most distal leaf primordia, ceased before February due to abscission of the shoot apex. Axillary buds located most distally on a shoot had more nodes than both terminal buds and more proximal axillary buds. The longest shoots included a preformed part and a neoformed part. The organogenetic event which initiated the neoformed organs continued until early autumn, giving rise to the following year's preformation. The absence of cataphylls in terminal buds could indicate a low intensity of shoot rest. The naked terminal bud of Nothofagus spp. could be interpreted as a structure less specialized than the scaled bud found in genera of Fagaceae and Betulaceae.     

Relative extents of preformation and neoformation in tree shoots: Analysis by a deconvolution method

BACKGROUND AND AIMS: Neoformation is the process by which organs not preformed in a bud are developed on a growing shoot, generally after preformation extension. The study of neoformation in trees has been hindered due to methodological reasons. The present report is aimed at assessing the relative importance of preformation and neoformation in the development of shoots of woody species. METHODS: A deconvolution method was applied to estimate the distribution of the number of neoformed organs for eight data sets corresponding to four Nothofagus species and a Juglans hybrid. KEY RESULTS: The number of preformed organs was higher and less variable than the number of neoformed organs. Neoformation contributed more than preformation to explain full-size differences between shoots developed in different positions within the architecture of each tree species. CONCLUSIONS: Differences between the distributions of the numbers of preformed and neoformed organs may be explained by alluding to the duration of differentiation and extension for each of these groups of organs. The deconvolution of distributions is a useful tool for the analysis of neoformation and shoot structure in trees.     

HETEROPHYLLY AND NEOFORMATION OF LEAVES IN SUGAR MAPLE (ACER SACCHARUM)

Variation in leaf form and timing of leaf initiation were investigated in vigorous leader shoots of open-grown saplings and larger forest trees of sugar maple (Acer saccharum Marsh.). Winter buds of leader shoots usually contained 6 or 8 leaf primordia and embryonic leaves, whereas 12 to 18 leaves typically expanded along the shoots each year. Preformed (early) leaves differ in form from neoformed (late) leaves. As in some other Acer species, the first-formed late leaves have large angles of secondary lobe divergence and deeply indented sinuses. This pattern of heterophylly contributes to the multilayered nature of open-grown saplings and leader shoots of forest trees of sugar maple.     

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.     

Preformation and neoformation in shoots of Nothofagus antarctica (G. Forster) Oerst. (Nothofagaceae) shrubs from northern Patagonia

The size (length and diameter) and number of leaf primordia of winter buds of Nothofagus antarctica (G. Forster) Oerst. shrubs were compared with the size and number of leaves of shoots derived from buds in equivalent positions. Buds developed in two successive years were compared in terms of size and number of leaf primordia. Bud size and the number of leaf primordia per bud were greater for distal than for proximally positioned buds. Shoots that developed in the five positions closest to the distal end of their parent shoots had significantly more leaves than more proximally positioned shoots of the same parent shoots. The positive relationship between the size of a shoot and that of its parent shoot was stronger for proximal than for distal positions on the parent shoots. For each bud position on the parent shoots there were differences in the number of leaf primordia per bud between consecutive years. The correlations between the number of leaf primordia per bud and bud size, bud position and parent shoot size varied between years. Only shoots produced close to the distal end of a parent shoot developed neoformed leaves; more proximal sibling shoots consisted entirely of preformed leaves. Leaf neoformation, a process usually linked with high shoot vigour in woody plants, seems to be widespread among the relatively small shoots developed in N. antarctica shrubs, which may relate to the species' opportunistic response to disturbance.     

Partial shoot reiteration in Wollemia nobilis (Araucariaceae) does not arise from 'axillary meristems'

Background and Aims

Conifers are characterized by the paucity of axillary buds which in dicotyledonous trees usually occur at every node. To compensate, conifers also produce ‘axillary meristems’, which may be stimulated to late development. In juvenile material of Wollemia nobilis (Araucariaceae: Massart''s model) first-order (plagiotropic) branches lack both axillary buds and, seemingly, axillary meristems. This contrasts with orthotropic (trunk) axes, which produce branches, either within the terminal bud or as reiterated orthotropic axes originating from axillary meristems. However, plagiotropic axes do produce branches if they are decapitated. This study investigated how this can occur if axillary meristems are not the source.

Methods

The terminal buds of a series of plagiotropic branches on juvenile trees were decapitated in order to generate axillary shoots. Shoots were culled at about weekly intervals to obtain stages in lateral shoot development. Serial sections were cut with a sliding microtome from the distal end of each sample and scanned sequentially for evidence of axillary meristems and early bud development.

Key Results

Anatomical search produced no clear evidence of pre-existing axillary meristems but did reveal stages of bud initiation. Buds were initiated in a group of small starch-rich cortical cells. Further development involved de-differentiation of these small cells and the development of contrasting outer and inner regions. The outer part becomes meristematic and organizes the apex of the new branch. The inner part develops a callus-like tissue of vacuolated cells within which vascular cambia are developed. This kind of insertion of a branch on the parent axis seems not to have been described before.

Conclusions

Axillary meristems in Wollemia characterize the leaf axils of trunk axes so that the origin of reiterated shoots is clear. Plagiotropic axes seemingly lack axillary meristems but still produce axillary branches by distinctive developmental processes. These observations demonstrate limited understanding of branch initiation in trees generally.     

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.     

Winter bud content according to position in 3-year-old branching systems of 'Granny Smith' apple

An investigation was made of the number of preformed organs in winter buds of 3-year-old reiterated complexes of the 'Granny Smith' cultivar. Winter bud content was studied with respect to bud position: terminal buds were compared on both long shoots and spurs according to branching order and shoot age, while axillary buds were compared between three zones (distal, median and proximal) along 1-year-old annual shoots in order 1. The percentage of winter buds that differentiated into inflorescences was determined and the flowers in each bud were counted for each bud category. The other organ categories considered were scales and leaf primordia. The results confirmed that a certain number of organs must be initiated before floral differentiation occurred. The minimum limit was estimated at about 15 organs on average, including scales. Total number of lateral organs formed was shown to vary with both bud position and meristem age, increasing from newly formed meristems to 1- and 2-year-old meristems on different shoot types. These differences in bud organogenesis depending on bud position, were consistent with the morphogenetic gradients observed in apple tree architecture. Axillary buds did not contain more than 15 organs on average and this low organogenetic activity of the meristems was related to a low number of flowers per bud. In contrast, the other bud categories contained more than 15 differentiated organs on average and a trade-off was observed between leaf and flower primordia. The ratio between the number of leaf and flower primordia per bud varied with shoot type. When the terminal buds on long shoots and spurs were compared, those on long shoots showed more flowers and a higher ratio of leaf to flower primordia.     

Comparison of the bud morphology and shoot growth dynamics of four species of Mediterranean subshrub growing along an altitudinal gradient

Bud morphology, bud activity, and shoot growth dynamics were studied over 13 months in four species of Mediterranean subshrubs growing along an altitudinal gradient in north-east Spain. The species selected were representative of the main functional types of subshrubs that grow naturally along altitudinal gradients on Mediterranean mountains. All species bore naked buds as their renewal structures, but the degree of protection of the buds varied between species, increasing with altitude. The period of morphogenesis and the level of shoot pre-formation also varied along the gradient. The species growing at low altitude had partially neoformed shoots, longer shoot growth periods, and started morphogenesis earlier than those growing at high altitude. These differences may indicate different morphological and phenological adaptations of Mediterranean subshrubs in response to increased winter cold along the gradient. At low altitudes, brachyblast-bearing species with partially neoformed shoots may be favoured because of the plastic shoot growth these structures confer. However, as the climate becomes colder and more predictable, brachyblast-bearing species may be replaced by species without brachyblasts, with tightly protected buds and more constrained patterns of shoot development.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 527–539.     

Bud structure and resprouting in coppiced stools of Salix viminalis L., S. eriocephala Michx., and S. amygdaloides Anders

Fast-growing willows are cultivated as coppice in short rotation biomass plantations. The production and sustainability of the system is based on the ability of trees to resprout after repeated harvesting. The large variation in coppicing ability is due to plant genotypic differences in structure and physiology as well as environmental factors. Morphological and structural prerequisites for resprouting were compared in two shrubby willows with high coppicing ability, S. viminalis and S. eriocephala, and one tree-formed species, S. amygdaloides, with low coppicing ability. The initiation and development of buds and the resprouting pattern of coppiced stools were compared. All buds were axillary in origin and showed the same principal structure consisting of one main shoot primordium and two lateral primordia. In S. viminalis and S. eriocephala the lateral buds contained several leaf primordia and sprouted shortly after the main bud. In S. amygdaloides further development of lateral buds was inhibited after formation of two budscales, and leaf primordia were not formed until the buds were forced to sprout. The number of sprouts developing after coppicing were correlated to the structure and number of buds and their position on the stools. Self-thinning rate was high and many shoots originating from lateral buds died. Most buds were located above ground on the remaining basal portions of harvested stems. No adventitious buds were found on the stools. Significantly different bud differentiation pattern and frequent sylleptic sprouting resulted in lower coppice response in S. amygdaloides compared to S. viminalis and S. eriocephala.     

Development of shoot inHydrangea macrophylla I. Terminal and axillary buds

The structure of shoots, in particular of winter buds, ofHydrangea macrophylla was examined. The non-flower-bearing shoot is usually composed of a lower and an upper part, between which a boundary is discernible by means of a distinctly short internode. This internode is the lowermost of the upper part, and it is usually shorter than the internodes immediately above and below, although the internodes tend to shorten successively from the proximal to the distal part of the shoot. Variations exist in the following characters among the terminal bud, the axillary bud on the lower part of the shoot and the axillary bud on the upper part: (1) length of bud; (2) character of the outermost pair of leaf primordia; (3) degree of development of secondary buds in the winter bud; and (4) the number of leaf primordia. Usually, the terminal bud contains several pairs of foliage leaf primordia with a primordial inflorescence at the terminal of the bud, but the axiallary bud contains only the primordia of foliage leaves in addition to a pair of bud scales.     

Secondary buds in Scots pine trees infested with <Emphasis Type="Italic">Gremmeniella abietina</Emphasis>

In winter 2000–2001, there was a serious outbreak of Gremmeniella abietina Morelet in southeastern Norway. During the outbreak, we noted that injured Scots pine trees (Pinus sylvestris L.) developed secondary buds in response to the fungus attack, and we decided to study the relationship between injury, appearance of secondary buds and recovery of the trees thereafter. For this purpose, 143 trees from 10 to 50 years of age were chosen and grouped into crown density classes. Injury was assessed in detail, and buds were counted before bud burst in the spring of 2002. In addition, a subset of 15 trees was followed through the summer of 2002 to assess recovery. All injured trees developed secondary buds, with a clear overweight of dormant winter buds in proportion to interfoliar buds. Healthy control trees did not develop secondary buds at all. The secondary buds appeared predominantly on the injured parts of the tree; interfoliar buds in particular developed just beneath the damaged tissue. Most of the secondary buds died during the winter of 2001–2002, mainly because the fungus continued to spread after the first outbreak. Many of the remaining buds developed shoots with abnormal growth during the summer. Secondary buds may help trees to recover from Gremmeniella attacks, but this strategy may fail when the fungus continues to grow and injure the newly formed buds and shoots.     

Plantlet regeneration from fascicular buds of seedling shoot apices of <Emphasis Type="Italic">Pinus roxburghii</Emphasis> Sarg

Shoot apices of Pinus roxburghii Sarg were cultured on Murashige and Skoog’s medium (MS) supplemented with cytokinins [6-benzyladenine (BA), kinetin and N-benzyl-9-(2-tetrahydropyranyl) adenine (BPA)] alone and in combination with auxin, α-napthaleneacetic acid (NAA). Of the three cytokinins tested at varying concentrations, medium supplemented with 10 μM BA was found optimal in respect of explant responsiveness (97.22 %) and average number of buds induced per explant (7.42). The concentration of cytokinins in the induction medium had a profound effect on rate of elongation of induced buds on MS basal medium containing 0.5 % activated charcoal. Further, shoots induced on lower concentrations of BA increased up to 2.4 times in length in 4 weeks. Decapitation of the explant enhanced the rate of axillary bud elongation. Proliferating shoot cultures were established by sub-culturing the axillary shoots on MS supplemented with 10 μM BA. Shoots 2–3 cm in length were suitable for culturing as more buds were induced on them compared to longer or shorter shoots. Root primordia were induced on 70.83 % shoots when transferred to 1/2 MS medium supplemented with 5.0 μM NAA. Elongation of root primordia (60 %) was achieved in liquid 1/2 MS basal medium. The plantlets were successfully transferred to soil after hardening; the time period from initiation of shoot buds to transplantation being 20–22 weeks.     

Further studies on branch buds in mosses; “Pseudoparaphyllia” and “scaly leaves”

The development of so-called foliose pseudoparaphyllia in the species of theClimacium-type branch development was studied with a paraffin sectioning method and SEM. Scaly leaves (or scale-like leaves) and “foliose” pseudoparaphyllia proved to originate as leaves by segmentation of an apical cell of a branch initial in the very first stage of development into a branch bud. Branch buds of two species among the six species examined develop linear-lanceolate appendages which serve to protect the buds as well as scaly leaves. These appendages originate in the peripheral part of the epidermal layer of buds, and therefore they can be homologous to trichomes previously reported for species with branch primordia. Emendation of two terms are proposed; every organ originating from buds as leaves that are effective for bud protection should be called scaly leaves, while those which homologous to trichomes should be called pseudoparaphyllia. Both terms are used here in a narrower sense than before: There might befilamentous orfoliose scaly leaves, andfilamentous orfoliose (linear-to broad-lanceolate) pseudoparaphyllia. A scheme is given to show the pattern of branch development and the manner in which branch buds (or primordia) are protected.     

Anatomical inferences on aerial bud protection of three Eugenia shrub species from the Cerrado

• Location and degree of protection of aerial buds are important functional traits in disturbance- or stress-prone environments since aerial buds ensure the development of new organs under favourable growing conditions. This study was carried out in a Brazilian Cerrado area under regeneration after long-term Pinus cultivation, where the trees were clear-cut in 2012 and the remaining material was burned in 2014.
• After the fire treatment, several species resprouted from belowground organs and their aboveground organs were directly exposed to full sunlight. We collected 15 terminal branches with fully expanded leaves from three individuals of each of three Eugenia species to investigate if those with well-developed belowground organs invest in bark for aboveground bud protection. The samples were analysed using light and electron microscopy.
• In addition to terminal and axillary buds, all species presented accessory buds, and the number varied according to the node analysed. None of the aerial buds were protected by bark, but all were well protected by cataphylls and densely pubescent leaf primordia. There were also inter- and intra-petiolar colleters that released a mucilaginous protein exudate. The distance between the shoot apical meristem and the outer surface was longer in the terminal bud than in axillary buds. The bud leaf primordia covering the shoot apical meristem had a thick cuticle, unicellular non-glandular trichomes that accumulate phenolic and lipophilic compounds, and secretory cavities.
• Our study shows that all three Eugenia species studied here had highly protected aerial buds allocated from belowground organs. These morphological traits may improve the chances of the species' persistence in areas subjected to frost events, low relative humidity, high irradiance and harmful UV levels.

     

Suppression of growth and death of meristematic tissues in <Emphasis Type="Italic">Abies sachalinensis</Emphasis> under strong shading: comparisons between the terminal bud,the terminally lateral bud and the stem cambium

The suppression of apical growth and radial trunk growth in trees under shade is a key factor in the competition mechanism among individuals in natural and artificial forests. However, the timing of apical and radial growth suppression after shading and the physiological processes involved have not been evaluated precisely. Twenty-one Abies sachalinensis seedlings of 5-years-old were shaded artificially under a relative light intensity of 5% for 70 days from August 1, and the histological changes of the terminal bud and terminally lateral bud of terminal leader and the cambial zone of the trunk base were analyzed periodically. In shade-grown trees, cell death of the leaf primordia in a terminal bud of terminal leader was observed in one of the three samples after 56 and 70 days of shading, whereas the leaf primordia in a terminal bud of terminal leader in all open-grown trees survived until the end of the experiment. In addition, the leaf primordia of the terminally lateral buds of terminal leader retained their cell nuclei until the end of the experiment. No histological changes were observed in the cambial cells after shading, but the shade-grown trees had less cambial activity than the open-grown trees through the experiment. Strong shading appeared to inhibit the formation and survival of cells in the terminal bud of terminal leader rather than the terminally lateral buds of terminal leader and the cambium. The suppression of the terminal bud growth and elongation of the surviving lateral buds would result in an umbrella-shaped crown under shade.     

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