Cell and tissue dynamics of olive endocarp sclerification vary according to water availability

Endocarp developmental timing in drupe‐type fruits, involving tissue expansion and sclerification processes, is increasingly used as marker for biological studies and crop management. In spite of its wide application, however, little is known regarding how these morphogenetic processes unfold or the factors that modify it. This study evaluates endocarp expansion and sclerification of olive (Olea europaea) fruits, used as an example of drupe‐type fruits, from trees growing under different water regimes: full irrigated, deficit irrigated (moderate reduction of water availability) and rainfed (severe reduction of water availability). Fruits were sampled weekly until pit hardening, and fruit and endocarp areas were evaluated in histological preparations. An image analysis process was tested and adjusted to quantify sclerified area and distribution within the endocarp. Individual stone cells differentiated independently but distribution and timing indicated the overall coordination of endocarp tissue sclerification. Increase in sclerified area was initially gradual, accelerated abruptly the week prior to the end of endocarp expansion and then continued at an intermediate rate. These results suggest that the end of the expansion period is driven by sclerification and the morphogenetic signals involved act first on sclerification rather than endocarp size. Intensification of sclerification and the end of expansive growth occurred first with lowest water supply. Moderate and severe reductions in water availability proportionately decreased endocarp expansion and prolonged the sclerification, delaying the date of physically perceived hardening but not affecting the final degree of endocarp sclerification.     

Axillary bud flowering after apical decapitation in Pharbitis in relation to photoinduction

The flowering response of axillary buds of seedlings of Pharbitis nil Choisy, cv. Violet, was examined in relation to the timing of apical bud removal (plumule including the first leaf or second leaf) before or after a flower-inductive 16-h dark period. When the apical bud was removed well before the dark period, flower buds formed on the axillary shoots that subsequently developed, but when removed just before, or after, the dark period, different results were observed depending on the timing of the apical bud removal and plant age. In the case of 8-day-old seedlings, fewer flower buds formed on the axillary shoots developing from the cotyledonary node when plumules were removed 20 to 0 h before the dark period. When the apical bud was removed after the dark period, no flower buds formed. Using 14-day-old seedlings a similar reduction of flowering response was observed on the axillary shoots developing from the first leaf node when the apical bud was removed just after the dark period. To further elucidate the relationship between apical dominance and flowering, kinetin or IAA was applied to axillary buds or the cut site where the apical bud was located. Both chemicals influenced flowering, probably by modulating apical dominance which normally forces axillary buds to be dormant.     

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.     

Plant regeneration from seedling explants of green bean (Phaseolus vulgaris L) via organogenesis

Green bean (Phaseolus vulgaris L.) plants were regenerated from 3-day old seedling explants via organogenesis. The explants contained a cotyledon and a small portion (2–3 mm) of embryonic axis split in half. Explants were cultured on a defined medium containing glutamine as the sole nitrogen source. A ring of meristematic tissue was produced at the base of the axillary bud located at the cotyledonary node. The meristematic tissue was produced only if the axillary bud was present together with the cotyledon in the explant. Buds and shoots developed from the meristematic ring. Selected shoots produced roots when excised from the cluster of buds and transferred to root induction medium. Rooted shoots (plantlets) grew well and produced viable seeds when grown in the greenhouse. Histological studies revealed the origin of buds from the peripheral layers of the meristematic ring.Production of buds and shoots was a continuous process, so that new shoots could be removed from the explant for plantlet production every 10–14 days. With the cultivar Dark Red Kidney, an average of 49 buds and 8 shoots were regenerated per explant by 30 days after culture initiation. Sixty-seven percent of the shoots produced roots, and 90–95% of the plantlets survived greenhouse acclimatization to produce healthy plants.     

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     

Molecular role of cytokinin in bud activation and outgrowth in apple branching based on transcriptomic analysis

Key message

Axillary bud activation and outgrowth were dependent on local cytokinin, and that bud activation preceded the activation of cell cycle and cell growth genes in apple branching.

Abstract

Cytokinin is often applied to apple trees to produce more shoot branches in apple seedlings. The molecular response of apple to the application of cytokinin, and the relationship between bud activation and cell cycle in apple branching, however, are poorly understood. In this study, RNA sequencing was used to characterize differential expression genes in axillary buds of 1-year grafted “Fuji” apple at 4 and 96 h after cytokinin application. And comparative gene expression analyses were performed in buds of decapitated shoots and buds of the treatment of biosynthetic inhibitor of cytokinin (Lovastatin) on decapitated shoots. Results indicated that decapitation and cytokinin increased ZR content in buds and internodes at 4–8 h, and induced bud elongation at 96 h after treatment, relative to buds in shoots receiving the Lovastatin treatment. RNA-seq analysis indicated that differential expression genes in auxin and cytokinin signal transduction were significantly enriched at 4 h, and DNA replication was enriched at 96 h. Cytokinin-responsive type-A response regulator, auxin polar transport, and axillary meristem-related genes were up-regulated at 4 h in the cytokinin and decapitation treatments, while qRT-PCR analysis showed that cell cycle and cell growth genes were up-regulated after 8 h. Collectively, the data indicated that bud activation and outgrowth might be dependent on local cytokinin synthesis in axillary buds or stems, and that bud activation preceded the activation of cell cycle genes during the outgrowth of ABs in apple shoots.

     

Influence of Bud Position and Plant Ontogeny on the Morphology of Branch Shoots in Pea (Pisum sativum L. cv. Alaska)

The morphology of axillary shoots of pea plants (Pisum sativumL. cv. Alaska) was analysed as a function of the position ofthe bud on the plant axis and the stage of plant developmentwhen the buds began to grow. Buds from the three most basalnodes were stimulated to develop by decapitating the main shootwhen buds were still growing (4 d plants), shortly after budsbecame dormant (7 d plants) or after the initiation of floweringon the main shoot (post-flowering plants, about 21 d after sowing).Branch shoots were scored for node of floral initiation (NFI),shoot length, and node of multiple leaflets (NML), a measureof leaf complexity. Shoots that developed spontaneously fromupper nodes (nodes 5-9) on intact post-flowering plants werescored for NFI. NFI for basal buds on 4 and 7 d plants variedas a function of nodal position and ranged from 5 to 6·7nodes. NFI on these plants was not influenced by bud size orwhether a bud was growing or dormant when the plant was decapitated.NFI for shoots derived from basal buds on decapitated post-floweringplants and upper nodes on intact post-flowering plants was about4. Reduced NFI on post-flowering plants may be due to depletionof a cotyledon-derived floral inhibitor. Basal axillary shootson 4 d plants were about 20% longer than those on 7 d plantsand about five times longer than those on post-flowering plants.These differences may be due to depletion of gibberellic acidsfrom the cotyledons. NFI and NML for the main shoot and forbasal axillary shoots were similar under some experimental conditionsbut different under other conditions, so it is likely that eachdevelopmental transition is regulated independently.Copyright1995, 1999 Academic Press Apical dominance, bud development, garden pea, initiation of flowering, Pisum sativum L., shoot morphology     

DIFFERENTIATION OF LATERAL SHOOTS AS THORNS IN ULEX EUROPAEUS

Ulex europaeus is a much-branched shrub with small, narrow, spine-tipped leaves and axillary thorn shoots. The origin and development of axillary shoots was studied as a basis for understanding the changes that occur in the axillary shoot apex as it differentiates into a thorn. Axillary bud primordia are derived from detached portions of the apical meristem of the primary shoot. Bud primordia in the axils of juvenile leaves on seedlings develop as leafy shoots while those in the axils of adult leaves become thorns. A variable degree of vegetative development prior to thorn differentiation is exhibited among these secondary thorn shoots even on the same axis. Commonly the meristems of secondary axillary shoots initiate 3–9 bracteal leaves with tertiary axillary buds before differentiating as thorns. In other cases the meristems develop a greater number of leaves and tertiary buds as thorn differentiation is delayed. The initial stages in the differentiation of secondary shoot meristems as thorns are detected between plastochrons 10–20, depending on vigor of the parent shoot. A study of successive lateral buds on a shoot shows an abrupt conversion from vegetative development to thorn differentiation. The conversion involves the termination of meristematic activity of the apex and cessation of leaf initiation. Within the apex a vertical elongation of cells of the rib meristem initials and their immediate derivatives commences the attenuation of the apex which results in the pointed thorn. All cells of the apex elongate parallel to the axis and proceed to sclerify basipetally. Back of the apex some cortical cells in which cell division has persisted longer differentiate as chlorenchyma. Although no new leaves are initiated during the extension of the apex, provascular strands are present in the thorn tip. Fibrovascular bundles and bundles of cortical fibers not associated with vascular tissue differentiate in the thorn tip and are correlated in position with successive incipient leaves in the expected phyllotactic sequence, the more developed bundles being related to the first incipient leaves. Some secondary shoots displayed variable atypical patterns of meristem differentiation such as abrupt conversion of the apex resulting in sclerification with limited cell elongation and small, inhibited leaves. These observations raise questions concerning the nature of thorn induction and the commitment of meristems to thorns.     

The effect of the gaseous state on bud induction and shoot multiplication in vitro in eastern white cedar

The effect of vessel type and the gaseous phase on the morphogenic response of Thuja occidentalis L. explants in vitro was studied. Explants were cultured in container types that varied in their degree of gas exchange. Traps for ethylene and CO₂ were employed. During shoot bud induction from embryonic explants, the number and elongation of shoot buds improved significantly when gastight, serum-capped flasks were used compared to the foam bung-capped flasks or the regularly used Petri dishes. Elimination of the two gases from the headspace of the flasks either singly or together reduced shoot bud induction and especially elongation of shoots. A similar response was seen during axillary bud development from cultured shoots. Ethylene and CO₂ accumulation promoted development and elongation of axillary shoots. An increase in the zeatin concentration in the medium produced a greater number of axillary shoots and higher levels of ethylene in the culture vessels. Removal of CO₂ caused gradual death of the shoots, while removal of ethylene alone reduced axillary shoot lengths significantly. Inclusion of aminoethoxyvinylglycine in the medium combined with ethylene traps produced an effect similar to the use of ethylene traps alone.     

Effect of phytoregulators and physiological characteristics of the explants on micropropagation of Maytenus ilicifolia

Micropropagated shoots of Maytenus ilicifolia Mart. were obtained from axillary buds cultured in Murashige & Skoog medium supplemented with 13.3 M 6-benzyladenine (BA). Addition of 1.1 M 1-indole-3-acetic acid (IAA) to the medium increased shoot elongation. The number of shoots formed was influenced by BA concentration, degree of juvenility of the explant, and by bud explant position on the stem. Cultures of buds taken from stem parts located close to the shoot tip yielded more callus than shoots, whereas axillary buds at distant positions from the apical bud yielded more shoots.Abbreviations BA 6-benzyladenine - IAA indole-3-acetic-acid     

Development and Growth Potential of Axillary Buds in Roses as Affected by Bud Age

The effect of axillary bud age on the development and potentialfor growth of the bud into a shoot was studied in roses. Ageof the buds occupying a similar position on the plant variedfrom 'subtending leaf just unfolded' up to 1 year later. Withincreasing age of the axillary bud its dry mass, dry-matterpercentage and number of leaves, including leaf primordia, increased.The apical meristem of the axillary bud remained vegetativeas long as subjected to apical dominance, even for 1 year. The potential for growth of buds was studied either by pruningthe parent shoot above the bud, by grafting the bud or by culturingthe bud in vitro. When the correlative inhibition (i.e. dominationof the apical region over the axillary buds) was released, additionalleaves and eventually a flower formed. The number of additionalleaves decreased with increasing bud age and became more orless constant for axillary buds of shoots beyond the harvestablestage, while the total number of leaves preceding the flowerincreased. An increase in bud age was reflected in a greaternumber of scales, including transitional leaves, and in a greaternumber of non-elongated internodes of the subsequent shoot.Time until bud break slightly decreased with increasing budage; it was long, relatively, for 1 year old buds, when theysprouted attached to the parent shoot. Shoot length, mass andleaf area were not clearly affected by the age of the bud thatdeveloped into the shoot. With increasing bud age the numberof pith cells in the subsequent shoot increased, indicatinga greater potential diameter of the shoot. However, final diameterwas dependent on the assimilate supply after bud break. Axillarybuds obviously need a certain developmental stage to be ableto break. When released from correlative inhibition at an earlierstage, increased leaf initiation occurs before bud break.Copyright1994, 1999 Academic Press Age, axillary bud, cell number, cell size, pith, shoot growth, Rosa hybrida, rose     

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     

Nauclea diderrichii: rooting of stem cuttings,clonal variation in shoot dominance,and branch plagiotropism

Summary Nauclea diderrichii (De Wild, and Th. Dur.) Merill (Rubiaceae), an indigenous hardwood of West Africa, is increasingly being grown commercially. This study investigates the potential for vegetative propagation and clonal selection, and raises some fundamental questions about the physiology of apical dominance and of plagiotropism. Rooting ability was high, with up to 100% rooting in 2–4 weeks, when different Indole-3-butyric acid (IBA) concentrations and leaf areas were tested. Auxin applications greatly increased the numbers of roots per cutting. The decapitation of unbranched plants revealed clonal variation in apical dominance and also in the establishment of outright dominance by the two shoots formed from the outgrowth of the axillary buds of the opposite leaves at the top node. Regression analysis of the Dominance Ratio (length of dominant: length of the sub-dominant shoot at the time of achieving dominance) against overall lateral bud activity (r = 0.82), showed that when the two top shoots co-dominate they provide a more powerful source of Correlative Inhibition than when one of the top shoots dominates the other. The imposition of plagiotropism in the axillary bud occurred over a period of a few days as the terminal and axillary buds emerged from the stipule. Growth of accessory buds on intact plants and debranched cuttings was orthotropic. These results are discussed with regard to the role of the leaf in root formation and the understanding of dominance relationships, branching and crown development in trees.     

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.     

Micropropagation of chinese redbud (<Emphasis Type="Italic">Cercis yunnanensis</Emphasis>) through axillary bud breaking and induction of adventitious shoots from leaf pieces

Summary Factors affecting in vitro shoot production and regeneration of Cercis yunnanensis Hu et Cheng were investigated by comparing various growth regulators and explant types. For optimum shoot production from axillary buds, Murashige and Skoog (MS) media containing 6-benzyladenine, either alone or in combination with a low concentration of thidiazuron, resulted in the greatest number of shoots formed per explant (>3). Explants (2 mm long) containing one axillary bud placed in directcontact with the medium yielded the most shoots per bud (1.6) when grown on growth regulator-free medium. Root formation on 70–80% of shoot explants was accomplished using either indole-3-butyric acid or α-naphthaleneacetic acid in the medium, with significantly more roots formed on explants possessing and apical bud than those without the bud. Direct shoot organogenesis from leaf explants occurred on MS medium containing 10–30 μM thidiazuron, with up to 42% of leaf explants producing shoots.     

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     

Occurrence of spontaneous shoot regeneration on leaf stipules in relation to hyperflowering response in micropropagated strawberry plantlets

Summary Stipular bud (SB) formation occurred spontaneously on “Gorella” strawberry leaf stipules during proliferation phase on Boxus medium. These buds gave rise to normal shoots on the inner median zone between the stipule tips. Some stipular buds also have been observed on other parts of adaxial surface of the stipule. Stipular bud formation took place directly on the stipule without an intermediate callus. The first stage consisted of subspherical protrusions of small cells which progressively differentiated into shoots that were able to proliferate and to root. Scanning electron microscopy was used to examine the developmental stages of this adventitious organogenesis and to describe morphologic abnormalities, e.g., multiapex formation and fasciation. In vitro cloning of plantlets was made from axillary and SBs produced by the same tufts. The greater flowering abundance of stipular plants resulted in a drastic reduction of the commercial production and the fruit caliber when compared to axillary plants. However, the general phenotype of the mother cultivar (Gorella) was not affected in either case.     

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     

棉花花芽分化及部分内源激素变化规律的研究

棉花（Ｇｏｓｓｙｐｉｕｍ ｈｉｒｓｕｔｕｍ）的腋芽原基,有的将来发育成叶枝;有的将来发育成果枝。这２种不同命运的腋芽,在其刚分化的初期就表现出了不同的解剖学特征。将来发育为叶枝的腋芽,其生长锥呈圆锥形或扁圆球形,体积较小,原套层数为１－２层;而将来发育为果枝的腋芽,其生长锥为圆柱形,顶端表面平坦,体积较大,原套层数为２－３层。从子叶展平后到肉眼可见花芽（现蕾）,连续测茎尖的内源ＡＢＡ及ＩＡＡ的含量     

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