Ontogeny of the proventitious epicormic buds in Quercus petraea.

In the present work, we described the fate of proventitious epicormic buds on the trunks of 40-year-old Quercus petraea trees and in parallel the vascular trace they produced in the wood. Our results show that small and large individual epicormic buds can survive as buds for 40 years and that both are composed of a terminal meristem and scales. Meristematic areas are detected in the scale axils of small buds; in addition to these meristems the large buds also have secondary bud primordia. The small buds are connected to the pith of the main stem by a unique trace, whereas the large buds are connected by one or multiple traces. A single trace might imply that the whole bud is still alive and multiple traces might indicate that the terminal meristem has died. In the latter case, each trace is connected to a secondary bud of the large bud. The buds found in a cluster are composed of a terminal meristem and scales with axillary meristems in the scale axils. A cluster is connected to the pith of a stem either by a unique trace when it seems to be the result of partial abscission of an epicormic shoot or multiple traces when it might have originated from an epicormic bud in which the terminal meristem has died. Whatever the type of the bud, the vascular trace in the bark is composed of a cambium, secondary xylem and parenchyma cells and the trace present in the wood had parenchyma cells with vestiges of secondary xylem. Each year, the vascular trace should be produced in the bark by the cambium of the tree but not by the bud itself. On 40-year-old Q. petraea, we observed a proliferation of epicormic buds and in parallel a multiplication of the number of vascular traces in the trunk, but the knots caused by the traces of epicormic buds in the wood, either as individuals or in clusters, are minor since their colours are only slightly darker than those of woody rays and they are less than 2 mm in diameter. The knots will appear when epicormic buds develop into shoots. Received: 30 March 1999 / Accepted: 09 June 1999     

Ontogeny of proventitious epicormic buds in Quercus petraea. I. In the 5 years following initiation

 The persistence of large epicormic shoots is one of the main factors that reduces timber quality and value in Quercus petraea. The early phases of epicormic shoot formation, i.e. the initiation of the epicormic buds, their survival and their proliferation over the years, are not clearly understood. In the present work, we studied the initiation of the axillary buds giving rise to epicormic buds and shoots, and followed their behaviour during the first 5 years using both scanning electron microscopy and light microscopy. Two types of proventitious epicormic buds have been identified. The first type has small axillary buds associated with the rings of bud-scale scars which are found at the base and tip of each growth unit. These buds are made of a terminal meristem surrounded only by scales; no leaf primordium is detected. During the second and third years of epicormic life, meristematic areas appear in the scale axil. Progressively, the meristematic areas organize into secondary bud primordia composed solely of the terminal meristem surrounded by scales. The second type of epicormic bud has secondary buds produced by a large axillary bud when this large bud either developed into a shoot or partially abscised. The epicormic potential in Q. petraea is characterized by a balance between the epicormic buds in apparent rest, enclosing meristematic areas and secondary bud primordia, and their mortality over the years. Received: 22 January 1998 / Accepted: 8 May 1998     

Bud banks of perennial savanna grasses in Botswana

Three semi‐arid savanna grasses in Botswana (Stipagrostis uniplumis, Eragrostis lehmanniana, and Aristida stipitata) were sampled to quantify their belowground bud banks during the dormant season and to estimate their relative allocation to vegetative and sexual reproduction. Bud banks of these African perennial caespitose grasses were also compared with four perennial caespitose grasses of semi‐arid North American grasslands. The three African grasses each maintained approximately two buds per tiller and showed a high percentage (88–99%) of tillers producing seed. Only E. lehmanniana produced new aerial tillers from axillary buds at elevated nodes on the stem as well as from the belowground bud bank. Compared with species of North American grasslands, these African grasses produced fewer belowground buds but showed a much higher percentage of tillers producing seed. These patterns indicate relatively greater belowground meristem limitation, lower allocation to vegetative reproduction (tillering) and higher allocation to seed reproduction in these African grasses, although studies of more species are needed to assess the generality of this pattern. The management of savannas in ways that favour the maintenance of a reserve population of belowground buds may increase the ability of grasses to respond to pulses of resource availability, increase their compensatory growth capacity following grazing or drought, and decrease the invasibility of these plant communities by exotic species, whereas maintaining allocation to sexual reproduction may be important for conserving genetic variation and enhancing their capacity to adapt to environmental change.     

NODE COUNTING IN AXILLARY BUDS OF NICOTIANA TABACUM CV. WISCONSIN 38, A DAY-NEUTRAL PLANT

A mature, quiescent, primary axillary bud on the main axis of a flowering Nicotiana tabacum cv. Wisconsin 38 plant, when released from apical dominance and before forming its terminal flower, produced a number of nodes which was dependent upon its position on the main axis. Each bud produced about one more node than the next bud above it. The total number of nodes produced by an axillary bud was about 6 to 8 greater than the number of nodes present above this bud on the main axis. At anthesis of the terminal flower on the main axis, mature, quiescent, primary axillary buds had initiated 7 to 9 leaf primordia while secondary axillary buds, sometimes present in addition to the primary ones, had initiated 4 to 5 leaf primordia. When permitted to grow out independently, primary and secondary axillary buds located at the same node on the main axis produced the same number of nodes before forming their terminal flowers. In contrast, immature primary axillary buds which had produced only 5 leaf primordia and which were released from apical dominance prior to the formation of flowers on the main axis produced only as many nodes as would be produced above them on the main axis by the terminal meristem, i.e., “extra” nodes were not produced. Therefore, it is the physiological status of the plant and not the number of nodes on the bud at the time of release from apical dominance that influenced the node-counting process of a bud. When two axillary buds were permitted to develop on the same main axis, each produced the same number of nodes as single axillary buds developing at these nodes. Thus, the counting process in an axillary bud of tobacco is independent of other buds. Axillary buds on main axes of plants that had been placed horizontally produced the same number of nodes as identically-positioned axillary buds on vertical plants, indicating that gravity does not play a major role in the counting, by an axillary bud, of the nodes on the main axis.     

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.     

THE DEVELOPMENTAL ANATOMY OF LONG-BRANCH TERMINAL BUDS OF PINUS BANKSIANA

A study of the composition of long-branch terminal buds (LBTB) of Pinus banksiana Lamb. and the yearly periodicity associated with their formation, development, and elongation was undertaken. Each LBTB has lateral bud zones and zones of cataphylls lacking axillary buds. When present, staminate cone primordia differentiate from the lowest lateral buds in the lowest lateral bud zone of the LBTB. Ovulate cone primordia and lateral long-branch buds can differentiate from the upper lateral buds in any lateral bud zone. When both types of buds are present, lateral long-branch buds are uppermost. Dwarf-branch buds occur in all lateral bud zones. During spring LBTB internodes elongate, new cataphylls are initiated, dwarf branches elongate, needles form and elongate, pollen forms and is released, and ovulate cones are pollinated. During summer buds form in the axils of the newly formed cataphylls. By early fall the new LBTB are in overwintering condition and the four types of lateral buds are discernable. The cytohistological zonation of the LBTB shoot apex is similar to that of more than 20 other conifer species. Cells in shoot apices of pine are usually arranged in distinct zones: apical initials, subapical initials, central meristem, and peripheral meristem. Periclinal divisions occur in the surface cells of the apex; therefore no tunica is present. At any given time, shoot apex volume and shape vary among LBTB in various positions on a tree. In any one LBTB on a tree, shoot apex shape changes from a low dome during spring to a high dome during summer to an intermediate shape through fall and winter.     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

THE OCCURRENCE OF ADVENTITIOUS AND PREVENTITIOUS BUDS WITHIN THE BARK OF SOME TEMPERATE AND TROPICAL TREES

Endogenous adventitious buds develop in situ from dedifferentiated parenchyma cells of the trunk-bark in the temperate trees Tilia platyphyllos, Acer pseudoplatanus, and Fraxinus excelsior, as well as in the tropical cauliflorous trees Artocarpus integrifolia, Swartzia schomburgkii, and Couroupita guianensis. On aerial roots of Clusia rosea endogenous adventitious buds originate within the proliferated phelloderm beneath lenticels. In Salix alba, Fraxinus excelsior, and Terminalia arjuna, exogenous dormant buds are overgrown during secondary growth and engulfed within the bark tissue, so that they give the impression of apparent endogeny. In the leaf axils of young shoots of Araucaria angustifolia, superficial tissue layers divide and form axillary protrusions, which soon become parenchymatic and partly suberized. A few cells at their bases stay meristematic, however, and develop as rudimentary endogenous bud primordia, which persist in the bark for many years.     

CYTOKININ- AND GIBBERELLIC ACID-INDUCED EFFECTS ON THE STRUCTURE AND METABOLISM OF SHOOT APICAL MERISTEMS IN OPUNTIA POLYACANTHA (CACTACEAE)

The dormant axillary buds of Opuntia polyacantha can be activated by either cytokinins or gibberellic acid. Under the influence of benzylaminopurine (BAP), the axillary bud meristem increases greatly in size and becomes mitotically active. The primordia produced by the meristem develop as normal photosynthetic leaves. Gibberellic acid (GA) also causes the meristem to become mitotically active, but the meristem does not increase in size. The primordia produced under the influence of GA develop as normal cactus spines. Leaf-producing meristems and spine-producing meristems have the same zonation, despite the differences in size. The meristems are composed of a uniseriate tunica, a central mother cell zone, peripheral zone, and a pith rib meristem. The mitotic activity of each of the zones in the leaf-producing meristem differs significantly from the mitotic activity of the corresponding zones in the spine-producing meristem.     

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.     

DEVELOPMENT AND PHYLLOTAXIS OF THE VEGETATIVE AXILLARY BUD OF MICHELIA FUSCATA

Tucker, Shirley C. (Northwestern U., Evanston, III.) Development and phyllotaxis of the vegetative axillary bud of Michelia fuscata . Amer. Jour. Bot. 50(7): 661–668. Illus. 1963.—The vegetative axillary buds of Michelia fuscala are dorsiventrally symmetrical with 2 ranks of alternately produced leaves. The direction of the ontogenetic spiral in each of these buds is related both to the symmetry of the supporting branch and to the position of the bud along the branch. On a radially symmetrical branch, all the axillary buds are alike—all clockwise, for example. But in a dorsiventrally organized branch the symmetry alternates from clockwise in 1 axillary bud to counterclockwise in the next bud along the axis. Leaf initiation and ontogeny of the axillary apical meristem conform with those of the terminal vegetative bud. The axillary bud arises as a shell zone in the second leaf axil from the terminal meristem. During this process the axillary apex develops a zonate appearance. The acropetally developing procambial supply of the axillary bud consists wholly of leaf traces. At the nodal level the bud traces diverge from the same gap as the median bundle trace of the subtending leaf. Only the basal 1–2 axillary buds which form immediately after the flowers elongate each year, while the majority remains dormant with 3 leaves or fewer.     

Leaf buds, a factor in host selection by Battus philenor butterflies

ABSTRACT.

• 1 Field and laboratory experiments identified a character intrinsic to Aristolochia reticulata Nutt. host plants, the terminal leaf bud, that is involved in host-selection behaviour by female pipevine swallowtail butterflies (Battus philenor L.) searching for oviposition sites.
• 2 In the field, the frequency with which females landed on non-host buds declined seasonally as the proportion of host foliage that consisted of buds decreased. Female butterflies did not land on non-host species in proportion to their abundance; rather, females landed on those non-host species whose buds resembled those of A.reticulata.
• 3 A.reticulata plants whose terminal leaf bud was concealed by plastic tape were less susceptible to oviposition in the field than were control plants.
• 4 Female butterflies released in a large, outdoor enclosure were conditioned to search for leaf buds only when exposed to a host species bearing a prominent terminal leaf bud.
• 5 The significance of conditioning of leaf-bud searching behaviour is discussed with respect to discrimination between hosts and non-hosts, between host species, and among plants within a host species.

     

LIGNOTUBER ONTOGENY IN THE CORK-OAK (QUERCUS SUBER; FAGACEAE) I. LATE EMBRYO

Changes at the cotyledonary node of the cork-oak (Quercus suber L.) were examined during the embryo maturation phase using light microscopy and scanning electron microscopy techniques. During the maturation phase the embryo axis elongates by diffuse growth, the apical meristem forms the first leaf primordia, and the radicle meristem remains inactive. The primary axis of the embryo bears, axillary to the cotyledons, in the range of five to seven pairs of lateral buds at differing stages of development. Two or three pairs of these buds are visible, occurring on the upper unfused portion of the embryonic axis, while the remaining buds are hidden by the fused cotyledonary tissues. Lateral buds develop from clusters of cells in the peripheral meristem forming a shell zone delimiting the bud meristem. Lateral buds do not undergo much development until germination begins. The results are discussed with reference to the possible role of the cotyledonary node as the source of the lignotuber in the cork-oak.     

Effect of Assimilate Supply on Development and Growth Potential of Axillary Buds in Roses

The effect of assimilate supply on axillary bud developmentand subsequent shoot growth was investigated in roses. Differencesin assimilate supply were imposed by differential defoliation.Fresh and dry mass of axillary buds increased with increasedassimilate supply. The growth potential of buds was studiedeither by pruning the parent shoot above the bud, by graftingthe bud or by culturing the bud in vitro. Time until bud breakwas not clearly affected by assimilate supply during bud development,Increase in assimilate supply slightly increased the numberof leaves and leaf primordia in the bud; the number of leavespreceding the flower on the shoot grown from the axillary budsubstantially increased. No difference was found in the numberof leaves preceding the flower on shoots grown from buds attachedto the parent shoot and those from buds grafted on a cutting,indicating that at the moment of release from inhibition thebud meristem became determined to produce a specific numberof leaves and to develop into a flower. Assimilate supply duringaxillary bud development increased the number of pith cells,but the final size of the pith in the subsequent shoot was largelydetermined by cell enlargement, which was dependent on assimilatesupply during shoot growth. Shoot growth after release frominhibition was affected by assimilate supply during axillarybud development only when buds sprouted attached to the parentshoot, indicating that shoot growth is, to a major extent, dependenton the assimilate supply available while growth is taking place.Copyright1994, 1999 Academic Press Assimilate supply, axillary bud, cell number, cell size, defoliation, development, growth potential, meristem programming, pith, Rosa hybrida, rose, shoot growth     

Vegetative regeneration of biennial Oenothera species after disturbance: Field observations and experiment

It has been recently shown that some annual and biennial species of man-made habitats cope with severe disturbance by resprouting (vegetative regeneration) from their bud bank and do not only rely on regeneration from seeds. Nevertheless, information on the ecology of this phenomenon is rare. In a field study, we answered the question how frequent is resprouting from root buds in populations of the ruderal biennial herb Oenothera biennis, and how it is affected by habitat conditions. In an experiment, we tested the hypothesis that higher severity of injury and later life-cycle phase of the injured plants suppress resprouting from both axillary and root buds in O. biennis and also in its closely related congeners O. fallax and O. glazioviana.In 25 out of 29 studied ruderal populations of O. biennis severely injured individuals were found; however, only half of these populations included injured individuals that resprouted from roots. Among these populations, the number of root-sprouting individuals varied highly (from 3% to 67% of injured individuals). The largest populations and the highest percentage of root-resprouting individuals were found in urban habitats with sandy/gravelly substrate, a low vegetation cover, and a high frequency of disturbance.In the experiment with three Oenothera congeners, removal of aboveground biomass with all axillary buds largely led to the death of plants of all three species. When a portion of the basal axillary buds remained intact, individuals of O. biennis mostly failed to regenerate, whereas individuals of O. fallax and O. glazioviana survived and formed seeds. A higher severity of injury suppressed resprouting in Oenothera congeners in this experiment. However, the relationship between life-cycle phase and the ability to resprout remains unclear in Oenothera species.This study showed that resprouting after severe injury is an important feature of Oenothera individuals occurring in man-made habitats and may represent an alternative strategy to regeneration from the seed bank under disturbance conditions.     

Axillary bud proliferation and organogenesis of <Emphasis Type="Italic">Euphorbia pulchurrima</Emphasis> winter rose

Summary Protocols for both axillary bud proliferation and shoot organogenesis of Euphorbia pulchurrima Winter Rose^TM were developed using terminal buds and leaf tissues. Greenhouse-grown terminal buds were placed on Murashige-Skoog (MS) basal medium supplemented with various concentrations of either benzlyaminopurine (BA) or thidiazuron (TDZ). Explants produced the greatest number of axillary buds on media containing between 2.2 and 8.8 μM BA. The number of explants that produced axillary buds increased with increasing BA concentration. TDZ at concentrations between 2.3 and 23.0 μM caused hyperhydricity of shoots and were not effective in promoting shoot proliferation. The most calluses and shoots were produced from leaf midvein sections from in vitro grown plants placed on the medium containing 8.8–13.3 μM BA and 17.1 μM indole-3-acetic acid (IAA) for 1 mo. before transferring to the medium containing only BA. Adventitious buds were produced only from red-pigmented callus, and explants that produced callus continued to produce adventitious shoots in the presence of IAA. Five-mo.-old shoots derived from shoot culture or organogenesis rooted readily in artificial soil with or without treatment with indolebutyric acid, and were acclimatized in the greenhouse.     

Shoot development and bud mite infestation in hazelnut (Corylus avellana)*

The interaction of environmental and genetic variation in hazelnut (Corylus avellana) shoot development and the behaviour, survival, and colonisation of eriophyid bud mites (Phytoptus avellanae and Cecidophyopsis vermiformis) were studied. The distribution of galled buds on shoots indicated that mites colonised only those buds formed during the mite migration period. The point of entry is probably the growing shoot tip. Once within this structure, as the shoot develops the mites have access to a succession of newly-formed, bud primordia that are unprotected by bud scales. The relative accessibility of the apical meristem and bud primordia may affect host susceptibility.     

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     

The induction of sun and shade leaves of the European beech (Fagus sylvatica L.): anatomical studies

Summary Primordia from buds of sun and shade twigs of European beech (Fagus sylvatica L.) were collected six times a year for anatomical investigations. Differentiation into sun-leaf and shade-leaf primordia was first observed in early August. Sun-leaf primordia had five, and shade-leaf primordia four layers of mesophyll meristem cells. With potted graft unions of beeches possible structural changes of leaf primordia were investigated. Trees adapted to shade develop sun-leaf primordia when put into full daylight, provided the transfer happened before July. Trees adapted to full daylight developed leaf primordia which remained structurally sun-leaf primordia when the plant was kept under shade conditions. Shadeleaf branches of young beech trees cut in February in order to expose the shade buds to full daylight developed either shade leaves or intermediate shade/sun leaves. These experiments show that the subtending leaf may provide the developing axillary bud with photoassimilates, but its character, whether sun or shade leaf, has no influence on the character of the developing leaf primordia.