Article

Bud Structure in Relation to Shoot Morphology and Position on the Vegetative Annual Shoots of Juglans regia L. (Juglandaceae)

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Abstract

The length and basal diameter of all lateral and terminal buds of vegetative annual shoots of 7-year-oldJuglans regia trees were measured. All buds were dissected and numbers of cataphylls, embryonic leaves and leaf primordia were recorded. Each axillary bud was ranked according to the position of its associated leaf from the apex to the base of its parent shoot. Bud size and content were analysed in relation to bud position and were compared with the size and number of leaves of shoots in equivalent positions which extended during the following growing season. Length and basal diameter of axillary buds varied according to their position on the parent shoot. Terminal buds contained more embryonic leaves than any axillary bud. The number of leaves was smaller for apical and basal axillary buds than for buds in intermediate positions on the parent shoot only. All new extended shoots were entirely preformed in the buds that gave rise to them. Lateral shoots were formed in the median part of the parent shoot. These lateral shoots derived from buds which were larger than both apical and basal ones. Copyright 2001 Annals of Botany Company

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... In one or more year old wood, past events such as the presence of leaves and buds are possible to identify from external morphological marks (scars) (Barthélémy and Caraglio 2007). Sabatier and Barthélémy (2001b) observed that in young walnut trees the type of axillary meristems is related to the node rank. In vegetative shoots, branching probability is positively associated with the size of the buds and is higher in middle positions of the parent shoot (node rank 8-10 out of 23). ...
... In vegetative shoots, branching probability is positively associated with the size of the buds and is higher in middle positions of the parent shoot (node rank 8-10 out of 23). The probability of latency and bud mortality is greater in axillary buds located in more distal and basal positions on the parent shoot (Sabatier and Barthélémy 2001b). Moreover, in walnut saplings, branching pattern and shoot length can be influenced by changes in local light environment (Taugourdeau and Sabatier 2010;Taugourdeau et al. 2011). ...
... Moreover, in walnut saplings, branching pattern and shoot length can be influenced by changes in local light environment (Taugourdeau and Sabatier 2010;Taugourdeau et al. 2011). In monocyclic shoots of adult trees, however, branching probability is higher in distal positions of the parent shoot (Sabatier and Barthélémy 2001b). In addition, architectural traits and pattern of adult walnut trees have been described according to different fruiting habits (Solar and Štampar 2003;Solar et al. 2004;Kelc et al. 2007Kelc et al. , 2010. ...
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Key message In adult walnut trees, poor light environments increase the number of catkins on a shoot, reducing the probability of bearing fruits. Growth units become shorter with the aging of the tree, tending to reach a stable length. Abstract Tree architecture results from the functioning of populations of meristems across consecutive periods of extension or growing cycles. The study of growth units (GUs) and the type of their axillary meristem lead to a better understanding of tree growth and allow the prediction of tree development under different environmental conditions. To study the effect of position inside the canopy on branch development and the relationship between parent wood and new shoots, 348 GUs from 14 five-to-six-year-old limbs (7 from upper/exposed and 7 from lower/shaded zones within the canopy) were analyzed. Each node was classified according to its axillary meristem type as: 0 (bud, dormant bud, bud scar and dead bud), 1 (fruit and fruit peduncle scar), 2 (male inflorescence or “catkin”, catkin scar and dead catkin) and 3 (branch and branch scar). Data were analyzed through a multinomial logistic model to summarize the distribution of axillary meristem type as a function of explanatory variables. Length of GUs and the number of catkins were modeled through a generalized linear mixed model, whereas the probability of finding fruit on apical nodes was analyzed through generalized additive models. Results show that axillary meristem type is affected by zone, node rank and wood age. On GUs of wood age two or younger, apical buds from upper/exposed zones showed higher probabilities of producing fruits compared with apical buds from lower/shaded zones. The latter was not observed for older wood. In upper/exposed zones, the slope of the relationship between the length of GUs and the length of parent wood increased with wood age. The number of catkins was higher in the lower/shaded than in the upper/exposed zone. The results of this study improve our understanding about walnut tree growth as affected by factors like ontogenetic changes and different light conditions within the tree canopy and might be useful to address tree managements such as training and pruning systems.
... Le Noyer est un objet d'étude intéressant car sa production de fruits et de bois à forte valeur économique a suscité de nombreuses recherches fondamentales et appliquées : l'amélioration et la sélection génétique des Noyers à fruit (Germain 1997) et à bois (Fady et Lefèvre 1995), la morphogenèse (Mauget 1984), le contrôle de la rhizogenèse et la culture in vitro d'embryons (Jay-Allemand et al. 1991), la compréhension du fonctionnement physiologique (Le Dizès et al. 1997) et biochimique (Le Roux et al. 1999) de l'arbre, l'analyse de la couleur du bois (Janin et al. 1993). Parmi ces travaux, la confrontation des connaissances acquises sur la variabilité architecturale et morphologique du Noyer d'une part (Sabatier et Barthélémy 2001 ;Sabatier et al. 2003), et celles acquises sur la physiologie de sa croissance d'autre part (Lacointe et al. 1993 ;Lacointe et al. 2004) nous offre la possibilité de mieux comprendre le développement de la couronne. ...
... Une tige peut également être constituée d'une partie préformée suivie d'une partie néoformée (figure II-2). De nombreux auteurs ont montré que le nombre d'organes préformés et néoformés dans une pousse dépend de l'espèce, du stade de différentiation et de la position de la pousse dans la couronne (Critchfield 1960 ;Kozlowski 1971 ;Remphrey et Powell 1984 ;de Reffye et al. 1991 ;Costes 1993 ;Puntieri et al. 2000 ;Sabatier et Barthélémy 2001 ;Puntieri et al. 2002). Chez certaines espèces, le passage de la préformation à la néoformation peut être observé par des marqueurs morphologiques tels que la variabilité de la forme des feuilles et présence d'un entre-noeud court (chez le Peuplier noir) ou bien les caractères de la floraison et de la ramification (chez l'Abricotier). ...
... Une diminution de la vigueur accompagne souvent une augmentation de l'ordre de ramification des axes (Costes et al. 1995 ;Barthélémy et al. 1997 ;Nicolini 1997). Barthélémy, 2001). Légende : = : arrêt de croissance inter-annuel, -: arrêt de croissance intra-annuel, ~: ralentissement de croissance intra-annuel, ○ : floraison. ...
Article
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Modelling tree architectural development according to environmental factors is an emerging field of research thanks to the development of adequate computational methods. However, modelling virtual plants is greatly constrained by the difficulty in collecting the architectural and geometric data of woody architecture and foliage. Moreover, partial understanding of functional processes that drive shoot growth limits the analysis and the modelling of the processes involved. Cross-fertilization of insights into architectural and morphological variability of Walnut tree on the one hand, and into growth physiology on the other hand, enables to better understand crown development. Thanks to 3-dimensional measurements of topology and geometry made on 7 to 9 years-old Walnut trees, we have analysed the impact on light interception of variables required for virtual plant modelling that are tedious to measure and that render extrapolation methods necessary. Secondly, we quantified the part of variance of growth variables which can be explained by radiation intercepted by comparison with other factors such as topological position. Our results point out the limitations of using allometric relationships and statistical laws for the reconstruction of 3D canopies. Leaf clumping inside the crown, including the clumping of leaflets at leaf scale, strongly determines the radiative properties of reconstructed crowns. Thanks to an approach based on correlations and statistics, an effect of light intercepted by mother shoots on the growth of daughter shoots was established. Strong correlations between shoot vigour (quantified by an index) and the length of the pathway between the collar and shoots on the one hand and shoot growth on the other hand suggest an effect of hydraulic architecture.
... Le Noyer est un objet d'étude intéressant car sa production de fruits et de bois à forte valeur économique a suscité de nombreuses recherches fondamentales et appliquées : l'amélioration et la sélection génétique des Noyers à fruit (Germain 1997) et à bois (Fady et Lefèvre 1995), la morphogenèse (Mauget 1984), le contrôle de la rhizogenèse et la culture in vitro d'embryons (Jay-Allemand et al. 1991), la compréhension du fonctionnement physiologique (Le Dizès et al. 1997) et biochimique (Le Roux et al. 1999) de l'arbre, l'analyse de la couleur du bois (Janin et al. 1993). Parmi ces travaux, la confrontation des connaissances acquises sur la variabilité architecturale et morphologique du Noyer d'une part (Sabatier et Barthélémy 2001 ;Sabatier et al. 2003), et celles acquises sur la physiologie de sa croissance d'autre part (Lacointe et al. 1993 ;Lacointe et al. 2004) nous offre la possibilité de mieux comprendre le développement de la couronne. ...
... Une tige peut également être constituée d'une partie préformée suivie d'une partie néoformée (figure II-2). De nombreux auteurs ont montré que le nombre d'organes préformés et néoformés dans une pousse dépend de l'espèce, du stade de différentiation et de la position de la pousse dans la couronne (Critchfield 1960 ;Kozlowski 1971 ;Remphrey et Powell 1984 ;de Reffye et al. 1991 ;Costes 1993 ;Puntieri et al. 2000 ;Sabatier et Barthélémy 2001 ;Puntieri et al. 2002). Chez certaines espèces, le passage de la préformation à la néoformation peut être observé par des marqueurs morphologiques tels que la variabilité de la forme des feuilles et présence d'un entre-noeud court (chez le Peuplier noir) ou bien les caractères de la floraison et de la ramification (chez l'Abricotier). ...
... Une diminution de la vigueur accompagne souvent une augmentation de l'ordre de ramification des axes (Costes et al. 1995 ;Barthélémy et al. 1997 ;Nicolini 1997). Barthélémy, 2001). Légende : = : arrêt de croissance inter-annuel, -: arrêt de croissance intra-annuel, ~: ralentissement de croissance intra-annuel, ○ : floraison. ...
... Gli studi architetturali sulla crescita degli alberi mostrano che la morfologia e il modello di crescita di un germoglio sono correlati alla posizione del germoglio e all'età della pianta. Più precisamente, le caratteristiche morfologiche dei germogli e dei rami possono variare in base a particolari andamenti noti come gradienti morfogenetici, legati alla posizione architettonica del germoglio e alla fase di sviluppo dell'albero (Sabatier e Barthélémy, 2001). ...
... I nuovi germogli laterali sono interamente preformati nelle gemme che li genereranno nella parte mediana del ramo genitore. Questi germogli laterali derivano da gemme più grandi di quelle distali e basali (Sabatier e Barthélémy, 2001). Sebbene la comprensione e la previsione della crescita e dello sviluppo delle piante possano aiutare nelle operazioni di formazione degli alberi di noce, è stata dedicata poca attenzione alla relazione tra la struttura delle gemme e la morfologia dei germogli risultanti in base alla loro posizione architettonica su rami di un anno (Sabatier e Barthélémy, 2001). ...
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To ascertain differences in bud topology related to the propagation technique, a study was carried on 1-yr-old cv Chandler walnut trees, micropropagated or grafted on seedlings (J. regia), in two orchards located in Emilia Romagna (Italy). Bud complexes were classified, and their number, position along the axis and length of shoots were also measured. The results evidenced that on average, micropropagated and grafted trees had a similar bud distribution along the shoot. Moreover, the internode length varied along the shoots according to the different fate of the buds. The bud topology was described also in 8 yrs-old micropropagated Chandler trees. Samples of 4 branches of different age were selected and all the buds were studied. The fate of each bud complex was evaluated according to the position along the axis. Terminal buds were investigated also in 3 micro-propagated plants and 4 apical shoots of 2-yrs-old trees. The female flower buds were more frequent in the lateral positions of the medium-terminal portion of long shoots, while the male flowers were more abundant in short shoots, only the terminal bud being female. The flowers in secondary buds were male. The material is available under Public License Creative Commons: Attribution 4.0 International (CC BY 4.0): http://amsacta.unibo.it/6228/
... Cevizde generatif ya da vejetatif dallar, ilkbahar başında birincil büyümelerini gerçekleştirirler. Özellikle genç ağaçlarda, belli bir durgunluk döneminin ardından ilkbahar ve yaz süresince ikincil bir büyüme, hatta üçüncül bir büyüme meydana gelebilir [24,26]. Cevizde dişi çiçekler ilkbaharda meydana gelen birincil büyüme sonucunda oluşurlar [24]. ...
... Özellikle genç ağaçlarda, belli bir durgunluk döneminin ardından ilkbahar ve yaz süresince ikincil bir büyüme, hatta üçüncül bir büyüme meydana gelebilir [24,26]. Cevizde dişi çiçekler ilkbaharda meydana gelen birincil büyüme sonucunda oluşurlar [24]. Generatif dallarda meyveler çeşitlerin dallanma karakteristikleri (dalların sıklığı, terminal ya da lateral pozisyonlu çiçek tomurcuklarının sayısı gibi), meyve verim ve kalitesi üzerinde büyük etkiye sahiptir [28]. ...
Article
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Tree architecture has a strong connection to tree training, fruit quantity and quality. In this study, the branch types in walnut are presented in three categories according to ((a) branching dynamic (sylleptic, proleptic, and epicormic), (b) branching mode (monopodial and sympodial), and (c) final length (short, medium and long). Also, some scientific papers on the issues of relationships between branch type and fruit quality, dominant branching layout on the parent branch (mesotony, basitony, acrotony and hypotony) and branch extinction are reviewed.
... Ces deux processus peuvent avoir lieu en même temps, constituant ainsi la partie préformée où les ébauches foliaires se développent à l'intérieur du bourgeon avant son débourrement, ou décalés dans le temps correspondant à une portion de tige non présente à l'état d'ébauches dans le bourgeon qualifié de partie néoformée (Rivals, 1966 ;Caraglio et al., 1997). Le rameau peut être entièrement préformé (Sabatier et al., 2001 ;Sabatier et al., 2003), ou entièrement néoformé, phénomène décrit chez différentes espèces tropicales (Borchert, 1969) ou encore constitué d'une portion préformée suivie d'une partie néoformée, phénomène cité par différents auteurs chez différentes espèces (Costes, 1993 ;Souza et al., 2002 ;Gordon et al., 2006 ;Guédon et al., 2006). Le nombre de pièces foliaires préformées ou néoformées dépend de l'espèce, du stade de développement de la plante et de la position du rameau étudié (Davidson et al., 1984 ;Remphrey et al., 1984 ;Costes, 2003 ;Sabatier et al., 2003 ;Costes et al., 2006). ...
... Les pousses printanières et automnales sont alors issues d'une préformation hivernale et estivale puis d'une néoformation printanière et automnale. Ces résultats corroborent les résultats obtenus chez le noyer (Juglans regia L.) qui montre une préformation hivernale donnant une partie de la pousse printanière et une préformation printanière, celle de la future pousse estivale (Sabatier et al., 1995 ;Sabatier et al., 2001). ...
Article
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Olive tree buds (Olea europaea L.) issued of the cultivar ‘Chemlali’, were sampled and described on two periods of olive growth season (on winter rest and summer stop) and on different types of shoots (short, medium and long shoot). The aim of this study was to characterize the structural and anatomical aspects of olive buds during the stop of growth and to analyze the relationship between the bud content during the rest and the structure of the produced shoot on olive tree on the end of the two growth flushes. The anatomical study of buds showed the classic zonation of buds: central zone, lateral zone and medullar zone. The bud section showed a number of nodes exactly half of that were present due to the decussate phyllotaxis of the olive tree. All buds for all forms deducted before the spring bud break or before the autumnal flush, showed between 3 to 4 pairs of leaf primordia with an average of 3.25 pairs for winter bud and 3.80 for summer bud. In fact, they developed between 6 to 7 pairs of preformed leaves on the resulting shoot. To analyze the relationship between the bud content during the rest and the structure of the subsequently produced shoot, it appears that some shoots may be entirely preformed and another may consist of a preformed portion followed by a neoformed portion. These types of shoots contained preformed metameres before spring and autumnal growth (winter or summer preformation) and neoformed metameres formed and simultaneously expanded without bud formation after the preformed part (spring and autumnal neoformation).
... Studies considering the periods of the year in which organogenesis takes place are rare, possibly due to the destructive techniques used for organogenesis assessment. Morphological categorization of individuals, axes and shoots of a species allows repeated sampling of homogeneous shoots, and thus is a precondition for the evaluation of organogenesis periods (Sabatier et al., 2001a;Puntieri et al., 2002a). ...
... In J. regia, the lateral buds of monocyclic¯oriferous shoots possess round and undivided cataphylls without lea¯et primordia (Fig. 5A), while the terminal bud of bicyclic shoots has cataphylls with distal incisions (Fig. 5B) This difference in form of bud protective leaves is in accordance with previous results on form variation between terminal and lateral buds of vegetative monocyclic shoots of J. regia (Sabatier and Barthe Âle Âmy, 2001a). In this species, winter buds, which give rise to spring growth units (Fig. 5A), and spring buds, from which summer growth units derive (Fig. 5B), differ in their composition. ...
Article
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Shoot extension during a growth season has been investigated in three morphological types of floriferous annual shoots (monocyclic with or without a small axis with immediate development and bicyclic) on four types of parental branches within 4-year-old Juglans regia L. 'Lara' growing in the south-west of France. The morphological type of parental branches has an influence on the time of inception of monocyclic shoot extension. No correlation has been found between the parental shoot length and monocyclic shoot length, while a positive correlation has been found between parental shoot length and bicyclic shoot length. Bicyclic annual shoots expand during two successive flushes: in spring and summer, respectively. The intra-annual resting phase results in the presence of a series of cataphylls in the median part of the shoots. Spring shoots and summer shoots may be distinguished by variations in form and in number of foliar organs. Annual shoots exhibiting an acrotonic axis with immediate development or a summer shoot generally possess greater extension periods and lengths of spring shoots than the unbranched ones.
... All lateral axes borne by the same Gu remain equal during the simulation, despite biological knowledge about vigor gradients [26] or asymmetric allocation depending of axis local environment [20]. ...
... In order to control tree death, a life cost function may be added [31]. The lack of lateral axes vigor gradient may be solved by introducing refinements in SAM base demand computing according to SAM position along the bearer Gu [26]. ...
Conference Paper
Context: Architectural studies highlight recurrent morphogenetical gradients that are observed on some tree species. These morphogenetical gradients are linked to morphological trends among successive shoots throughout plant structure and ontogeny. This study aims at testing a potential origin of these gradients as the complex result of some core plant functions. It will be achieved through a minimalist mathematical modelling approach.
... Bud size also depends on the position of buds along the parent shoot (Buck-Sorlin and Bell 2000). For instance, it is well established that apical buds exert a strong dominance over the other buds in the shoot, i.e. they display higher vigour and have greater probabilities to produce shoots than the rest of buds (Sabatier and Barthélémy 2001; Puntieri et al. 2002). However, no previous study has assessed the relationship between bud size and bud fate on the following years, how such relationship depends on bud position within the shoot, and the inter-annual variability of the relationship between bud size and fate in broadleaved tree species. ...
... Therefore, we regard bud size as a proxy of bud vitality. Previous studies (Gill 1971; Kozlowski et al. 1973; Sabatier and Barthélémy 2001; Montserrat-Martí et al. 2009) emphasize the strong influence of bud size and their position along the parent shoot on the shoot length formed by that bud. Consequently , it can be hypothesised that larger buds will have a greater impact on the expansion of the tree canopy than smaller ones if bud size determines the shoot length produced in the following year. ...
Article
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Understanding the relationships between bud size and position and bud fate through time is crucial for identifying and subsequently modeling the mechanisms underlying tree architecture. However, there is a lack of information on how bud size drives crown architectural patterns in coexisting tree species. We studied bud demography in two coexisting Mediterranean oak species with contrasting leaf habit (Quercus ilex, evergreen; Q. faginea, deciduous). The main objective was to analyse the effect of bud size on the fate of buds with different positions along the shoot (apical, leaf axillary and scale-cataphyll axillary buds). The number, length and position of all buds and stems were recorded in marked branches during four years. Study species presented different strategies in bud production and lifespan. The evergreen species showed greater mortality rate than the deciduous one, which produced larger buds. Bud size and position were highly related since apical buds where longer than axillary ones and bud length declined basipetally along the stem. Apical buds had also higher chances of bursting than axillary ones. Within positions, longer buds presented a higher probability of bursting than shorter ones, although no absolute size threshold was found below which bud bursting was impaired. In Q. ilex, four-year-old buds were still viable and able to burst, whereas in Q. faginea practically all buds burst in their first year or died soon after. Such different bud longevities may indicate contrasting strategies in primary growth between both species. Q. ilex is able to accumulate viable buds for several ages, whereas Q. faginea seems to rely on the production of large current-year buds with high bursting probability under favourable environmental conditions.
... Lauri et al. (2001) found out that at least two factors are involved in fruit weight variability: the length of preformed subtending axes of the bearing annual shoot and the number of fruits developed from the inflorescence. Sabatier and Barthélémy (2001b) showed that the formation of new shoots depends on the number and dimensions of the buds and their position along the parent shoot. The authors mentioned above were not concerned neither with the relationship between the quantitative traits (length and diameter) of the parent shoot nor with the number, dimensions and type of new shoots. ...
... As reported by Lauri et al. (2001), the quality and quantity of walnut fruits are also related to the branching and fruiting behaviour of the tree. Additionally, the largeness and homogeneity of the fruits varies from year to year as a consequence of different exogenous factors which may also be involved in the determination of the final diameter and length of branches (Sabatier and Barthélémy 2001b). So, in the research, we were trying to define the impact of a certain genotype and the age of the tree on the morphology and activity of annual shoots. ...
Article
The basal diameter of the annual shoot (1YD) affects vegetative growth and fruiting of the walnut trees. In order to determine interdependency between the 1YD and the older parent wood, 64 walnut genotypes belonging to four different branching and fruiting habits (morphotypes M-I, M-II, M-III and M-IV) were investigated. Year-to-year stability of 1YD was tested with the architectural analysis of a 3-year-old fruiting branch and its constituents (a 3-year-old bearer + corresponding 2-year-old + annual shoots) during 3 successive years. Based on Pearson's correlation coefficients and the multiple regression analysis of 12 quantitative traits, 12 models (four morphotype in 3 successive years) of 1YD were formed. They were compared with the standard model which was calculated on the basis of 1-year measurements of 1Y with no respect to the branching and fruiting type and comprises three quantitative traits, i.e. basal diameter of a 2-year-old parent shoot (2YD), the length of 2Y shoot (2YL), and the length of annual shoot (1YL). In a single year, the 1YD was influenced by two–five parameters. Five out of 12 models agreed with the standard model: in the lateral fruiting genotypes (M-IV), 1YD was always under the influence of the 2Y diameter, and the 1Y length. In addition, the number of nodes of the 2Y parent shoot had an important influence on 1Y diameter. In the terminal bearers (M-I), the impact of 2YD on the 1YD slightly increased with the tree age, and some other parameters, like 1Ynumber and 1Ynodes, which became to be important for 1YD. In the intermediate genotypes with mezotonic ramification (M-II), the number of vegetative buds per 1Y and angles of 1Y had significant effects on 1YD. In the intermediate bearers with acrotonic ramification (M-III), one to four other parameters were included into the model each year beside the 1Y number. Since the traits of a 2-year-old parent shoot have a great influence on the 1YD, the information from the year N can be used for the prediction of the annual shoot development in the year N+1. Such a prediction is more reliable in M-I and M-IV than in M-II and M-III. When we deal with the intermediate fruiting cultivars, 1Y number has to be considered in prediction of 1Y diameter beside 2YD and 1YL.
... bei seinen Untersuchungen an den fertigen Endknospen von Aesculus Hippocastanum fest, dass diese 7-8 Paare Schuppenblätter enthalten und die Anlagen von 3-4 Paaren Laubblättern des diesjährigen Zweiges. Daran anschliessend fand er oft 1-2 Paare Schuppenblattanlagen der Knospe des nächsten Jahres. Foster bezeichnet dieses Phänomen als Preformation.Sabatier (2001) stellte die Preformation bei Juglans regia, Studer-Ehrensberger (2015) fand sie bei Quercus robur. In den Endknospen können bereits die ersten Schuppenanlagen für die nächste Endknospe vorhanden sein. Dieses Prinzip erinnert an die Matrjoschka Puppen, bei denen eine Puppe die nächste Puppe enthält. Das Gleiche trifft in abgewandelter Fo ...
Preprint
Homology, Type, basic organs, ontogeny, scale leaf, leaf, bud, flower, ontogeny, Angiosperms, methodology. Review. Die Arbeit untersucht die erkenntnistheoretischen und methodischen Grundlagen der vergleichenden Morphologie. Die Begriffe Homologie und Analogie spielen eine zentrale Rolle. Teil der vergleichenden Morphologie ist die Organogenese. Die wichtigsten Konzepte sind das Homologie‐konzept, das Grundorgankonzept, das Typuskonzept. Die Arbeit enthält ein Review der verschiedenen Grundorgankonzepte und ein Review der Homologisierungen der Blüte mit dem Spross.
... La position des entités sur la structure est associée à des différences morphologiques qui affectent leur probabilité de débourrement, en fonction des espèces. On observe chez le noyer (Junglans regia) que les bourgeons axillaires dans la zone médiane de la branche ont des dimensions supérieures à celles des bourgeons apicaux et des bourgeons axillaires à la base (Sabatier et Barthélémy, 2001). Dans ce contexte, l'effet positif de la position latérale et du diamètre du bourgeon sur sa probabilité de débourrement explique la mésotonie de la ramification. ...
Thesis
A tree crown is the result of the repeated appearance of vegetative (metamers, axes) and reproductive elements, and of their disappearance. The appearance and disappearance of these elements are conditioned by architectural and environmental factors, and possible human manipulation. In particular, pruning of fruit trees stimulates bud burst and vegetative growth. Mango is a tropical fruit tree of economic, cultural and nutritional interest, whose cultivation meets major agronomic problems, such as generally low and irregular yield from one year to the next one, and phenological asynchronisms leading to the presence of stages susceptible to pests and diseases over long periods in orchards. These constraints stem in part from the strong interactions between vegetative growth and reproduction that have been evidenced in this species. Thus, the importance and dynamics of vegetative growth prior to flowering largely determine the success of the latter. The functional structural plant model V-Mango that simulates mango yield and quality build-up is based on this knowledge. It aims at the model-assisted design of more sustainable orchard management practices, based in particular on pruning optimization. This practice has recently been integrated into V-Mango. However, the simulations show an overestimation of vegetative growth and production, linked to an overproduction of new axes and to the non-consideration of axis mortality. We hypothesized that light, as a source of energy and a signal, has an essential role in the processes of vegetative bud burst, leaf fall and axes mortality in the mango tree, and that integrating these effects into V-Mango can lead to a significant improvement of model outputs. To evaluate this hypothesis, we quantified the variability of the light environment within a mature mango tree crown, and investigated functional relationships between light quantity and quality. We characterized the effects of architecture, light and pruning on axes appearance and disappearance. The annual dynamics of leaf fall and the factors affecting it were analyzed at the axis level. Finally, all these results were integrated into V-Mango after coupling it to a light interception model. Our results showed an important heterogeneity of light environment within the mango tree crown, as well as strong relationships between light quality and quantity. Light environment, axis morphology and topology, and pruning had marked effects on vegetative bud burst and axis mortality. Leaf fall dynamics was irregular during the year and depended mainly on architectural factors and little on light. The implementation in V-Mango of the effects of architecture, light and pruning on vegetative bud burst, after its coupling with a light interception model, allowed to simulate realistically the quantity and dynamics of the appearance of new axes following pruning, and to improve their spatialization in the crown. The simulations, performed on the 3D mock-up of a digitized adult mango tree, were validated at the axis scale and at the whole tree scale with independent data collected through original methodologies of analysis of hemispherical photographs and aerial images taken by drone. The significant improvement of the V-Mango model brought by this research makes it possible to consider using it in the short term to improve pruning practices on the mango tree and thus design more sustainable orchard management practices.
... Axillary buds have a major relevance in the annual growth of tree species with a predominantly sympodial construction (e.g. Macdonald et al. 1984;Barthélémy et al. 1999;Sabatier & Barthélémy 2001). This study showed clear qualitative differences between the two types of buds for most of the species analysed, especially due to the scaly nature of the proximal leaves of the axillary buds, i.e. the prophylls. ...
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The development of scaly buds (= cataphylls) has been traditionally associated with seasonally cold climates, although only few species from the southern hemisphere were investigated in this regard. The present work focuses on apical and axillary buds of seven tree species native to the South-American Temperate Rainforests (STR). Due to differences in the lineages from which these species derived, high levels of inter-specific variation in bud structure were expected. Apical and axillary buds were dissected under stereomicroscope, and the sizes of their parent shoots were evaluated. Cataphylls and leaf primordia were counted, and the presence of colleters and/or trichomes registered. Both intra- and inter-specific variations in bud structure were found. The apical buds were scaly in two out of seven species, and naked in the other species. Axillary buds were scaly in all but one species. In general terms, larger shoots developed buds with more organs. The presence of colleters (in four species) was not restricted to those buds lacking an outer cover of cataphylls. Further studies should focus on the relevance at a broader scale of colleters and trichomes as protective structures in tree buds. Keywords: Tree buds; South-American Temperate Rainforest; naked buds; scaly buds; preformation
... rhoifolia, Pl. strobilacea and C. ovata) represent three of the four estimated independent character-state changes. Consistently, Juglans regia, part of the fourth group with true cataphylls, is known to have entirely preformed shoots (Sabatier and Barthélémy, 2001). It should be noted, however, that, as in Pt. rhoifolia, saplings of J. regia have a terminal bud structure that differs from that of mature trees. ...
Article
Background and Aims Dormant resting buds are frequently regarded as static units, with protective cataphylls on the outside and embryonic foliage leaves on the inside. The developmental consequences of cataphyll insertion into the dynamic, cyclical, annually repeating sequence of leaf forms that a resting bud gives rise to have rarely been interrogated. To examine the connection between dormant structure and growing-season development, we compare the complete seasonal heteroblastic sequence of leaf forms of six species of temperate Juglandaceae with distinctly different vegetative resting bud structures. These include buds with cataphylls; buds without cataphylls; and buds with caducous cataphylls that are lost before the onset of winter. Methods In a common garden setting over a seven-month growing season, the dimensions of 2249 individual vegetative metamers were tracked from first exposure to abscission along the shoots of saplings and mature trees. The timing of metamer initiation within terminal buds was investigated using micro-CT scanning. Character state transitions of resting bud types were estimated using a phylogenetic tree of Juglandaceae. Key Results The presence of cataphylls within a heteroblastic sequence is associated with a single cohort of foliage leaves that flush and abscise synchronously. This growing pattern is highly determinate, with next year’s terminal-bud cataphylls already initiated before spring leaf out. In contrast, in sequences without cataphylls, shorter-lived foliage leaves appear and abscise in a staggered fashion. Despite these differences in leaf demography, all examined heteroblastic sequences produce a series of small, caducous leaf forms that precede terminal bud set. Conclusions The ubiquity of caducous leaf forms in Juglandoideae may point to the importance of shoot tip protection far beyond the dormant season. In addition, the presence or absence of cataphylls in resting buds is indicative of distinct shoot ontogenetic patterns, and functional strategies, in summer.
... To our knowledge, the first documented studies on the morphology of vegetative and floral buds of Juglans regia L. were conducted by De Candolle (1862) as cited by Nast (1935). Several researchers have subsequently studied the anatomy of walnut buds, especially staminate ones (Luza and Polito, 1988;Polito and Pinney, 1997;Polito, 1998;Germain et al., 1999;Sabatier and Barthélémy, 2001;Sabatier et al., 2003;Li et al., 2011;Gao et al., 2012Gao et al., , 2014. The temporal sequencing of anatomical and morphological changes in walnut buds varies according to the cultivar and the climatic conditions (Zhang et al., 1995;Polito and Pinney, 1997). ...
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Temperate deciduous fruit trees survive winter temperatures by entering a dormant phase in their aerial meristematic organs. Release from bud dormancy occurs after chill requirements (CR) have been satisfied, whereas bud burst/flowering follows heat requirement (HR) fulfillment. The physiological basis behind these metrics remains elusive. In this study, we are presenting the first multidisciplinary dormancy progression analysis in northern Patagonia, linking (1) forcing/field phenology, (2) bud anatomical development, and (3) soluble sugar (sucrose, glucose, and fructose) dynamics in Juglans regia L. CR and HR were determined for ‘Chandler’ and ‘Franquette,’ two walnut cultivars with markedly different CR, in artificial chill/forced heat trials (three seasons) and in-field chill/forced heat tests (five seasons) using excised twigs either with or without apical buds (non-decapitated and decapitated). The soluble sugar dynamics of ‘Chandler’ (high-performance liquid chromatography) and the anatomical changes of the buds (light microscopy) of the two cultivars were analyzed during endo-ecodormancy progression in one and two seasons, respectively. The CR defined by artificial chill tests proved to be an overestimation compared to the field determinations. Moreover, HR was the main driver in the phenology dynamics, as expected for a high-chill region. ‘Chandler’ showed an average of 10.3 field chill portions (CP) and 2,163 Growing Degree Hours (GDH°C) less than ‘Franquette’ for dormancy release and bud burst, respectively. These results were consistent with the transition of the shoot apex from the vegetative to the reproductive phase and the soluble sugar profile. The decrease in sucrose between 15 and 30 days after CR fulfillment could be a reliable biological marker for endodormancy release in walnut, while the increase in fructose and glucose is likely an osmolyte and cellulosic carbon source in pre-sprouting. In addition, we discuss the effect of paradormancy thanks to our apical bud experiment (with or without). Our results improve the current understanding of endo-ecodormancy progression in walnut and provide insightful results for walnut production (i.e., cultivation practices such as pruning) as well as for further application in dormancy modeling, to infer the ideotypes that should be bred for future climate conditions.
... Foster referred to this phenomenon as preformation. Sabatier and Barthélémy (2001) identified preformation in Juglans regia, while Studer and Schilperoord (2015) identified the phenomenon in Quercus robur. The terminal bud may contain the first scale primordia for the next terminal bud. ...
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The role of the vegetative bud in the structural development of the flower has received little research attention to date. The bud is an independent organ composed of mature scale leaves enveloping young, immature cauline leaves. There are some obvious parallels between vegetative buds and reproductive, or flower buds. It is generally accepted that the leaf serves as the morphological and genetic basis for the various flower organs. The term ‘leaf’ normally refers to a traditional foliage leaf. Study of the vegetative bud reveals that the leaf assumes two basic forms, namely those of the scale leaf (cataphyll) and the cauline leaf (trophophyll). This paper posits that the vegetative bud is the structural basis for the flower. The scale leaf serves as the basis for the first three categories of organs in the flower: the sepals, petals and stamens. The foliage leaf, on the other hand, serves as the basis for the carpels. The prevailing model used to elucidate the structure of the angiosperm is based on annual flowering plants. This paper proposes the use of a perennial, woody plant as an alternative, complementary model. This model draws from the primary organ concept proposed by Julius Sachs and Karl Goebel, which differentiates between the primary vegetative and reproductive organs. The two sporophylls – the stamen (microsporophyll) and the carpel (macrosporophyll) – are produced by the interactions between these two classes of primary organ. In this context, the stamen can be described as a catasporophyll, while the carpel can be described as a trophosporophyll. The morphological significance of the vegetative bud in defining the basic structure of the plant has been habitually overlooked, a fact which is reflected in the lack of genetic models focused on the development of the vegetative bud and the differentiation between the scale leaves and the cauline leaves.
... In the middle of the 20th century, many works were published, the authors of which investigated the properties of dormant buds (Kozhevnikov, 1950;Baldini and Mosse, 1956;Lyashenko, 1958Lyashenko, , 1964etc.). Today, the role of dormant buds in the formation of the overall body structure of woody plants has been determined (Serebryakov, 1955;Mazurenko and Khokhryakov, 1977;Mazurenko, 1986;Sabatier and Barthélémy, 2001;etc.). In describing the general architecture of the tree and shrub crown, the axes of regeneration are often differentiated in terms of their position and nature of occurrence; some of these axes are associated with the realization of dormant buds (Millet et al., 1998;Khutornoi et al., 2001;Nikolini et al., 2001;Getmanets, 2015;Kostina et al., 2015;etc.). ...
Article
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The role of dormant buds in the formation and transformation of herbaceous life forms of plants was determined by the example of a number of species of the genus Nepeta. Different variants of shoot formation and mechanisms reorganization of biomorphs with the participation of dormant buds are described. It has been established that regular and irregular development of dormant buds may lead to the formation of shortened, remote-nodal, and elongated residue, which are the basis for the construction of new shoot structures (shoot systems and rooting partial formations) typical for long-rhizome biomorphs.
... The early growth cessation leads to short shoot formation whereas growth continuation leads to medium and long shoots. The number of preformed organs within a winter bud as well as the final number of expanded phytomers from a meristem also depends on the length of the parent shoot, which in turn depends on tree age and ontogenetic stage (Sabatier and Barthélémy, 2001;Costes, 2003). In the within-tree population of meristems, it is likely that growth cessation has different origins that result either from nutritional starvation and/or correlative inhibition in the case of early cessation of short shoots, or from climatic control by the photoperiod or temperature decline at fall in the case of long shoots. ...
Chapter
Plant architecture results from developmental processes occurring throughout the plant life span. In the current article, the processes responsible for the formation of aerial organs and their organization at the whole plant scale, i.e. primary and secondary growth, branching, and flowering are considered, whereas the below‐ground plant organization is not included. This article focuses on the advances made on the comprehension of the physiologic and genetic base of plant architecture initially elucidated in plant models, and which are progressively being deciphered in a large range of species. The current knowledge allows comparisons of the processes among groups and to examine their conservation versus their major differences. However, numerous questions remain especially when regularities in plant structure are observed at coarse scales of plant organization such as annual shoots, axes, or branching systems. Moreover, the overlap of numerous gene functions over several processes leads to pleiotropic effects that make the genetic effects complex to interpret. The evidence of such overlaps opens evo‐devo perspectives.
... В середине ХХ в. было опубликовано много работ, авторы которых исследовали свойства спящих почек (Кожевников, 1950;Baldini, Mosse, 1956;Ляшенко, 1958, 1964и др.). Сегодня определена роль спящих почек в становлении общей структуры тела древесного растения (Серебряков, 1955; Мазуренко, Хохряков, 1977; Мазуренко, 1986; Sabatier, Barthelemy, 2001;и др.). При описании общей архитектуры кроны деревьев и кустарников нередко оси возобновления дифференцируют по положению и характеру их возникновения, часть этих осей связана с реализацией спящих почек (Millet et al, 1998;Khutornoi et al, 2001;Nikolini et al., 2001;Kostina et al., 2015;Getmanets, 2015;и др.). ...
... Modular development is associated with the morphological differentiation of axes at several scales. For instance, between stem and leaves at the phytomer scale (Baret et al. 2003), between green leaves and cataphylls at the shoot scale (Sabatier and Barthélémy 2001), and between long and short axes at the crown scale (Barthélémy and Caraglio 2007;Puntieri et al. 2018). These differences correspond to specializations in contrasting functions, namely: (1) space exploration and support (woody internodes, long shoots), and (2) light exploitation and carbon assimilation (leafy internodes, short shoots) (Ishii 2011). ...
Article
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Key message Trees display contrasting specialized annual shoots during their life-span and along with ontogeny-driven modifications in their branching pattern, they can fulfill different combinations of light exploitation and space exploration functions Abstract Tree ontogeny is related to major changes in tree structure and function at different scales, from individual organs to the whole tree. Yet, little is known about the direct effects of tree ontogeny on shoot specialization and branching patterns. Such specific architectural changes occurring with tree growth and aging are of critical importance for understanding the response of trees to their environment. The uppermost branching system of 0.1- to 23-m-tall sugar maple trees was sampled at the end of the growing season. Measurements were made at both the branching system (n = 40) and annual shoot scales (n = 803). An algorithm for automated shoot typology was developed to characterize branching pattern variations. Sugar maple shoots were divided into four types with contrasting sizes and levels of foliage (i.e., relative biomass allocation into leaves, LMF). These morphological differences were interpreted as functional specializations for light exploitation (high LMF) or space exploration and support (low LMF). Only annual trunk shoots exhibited trait value changes during ontogeny such as a minimum allocation to foliage in the current-year shoots for the 5-m-tall trees, which is related to lower light interception capabilities but higher space exploration abilities. However, this relative loss of light interception function is compensated by ontogenetic changes at the branching system scale, which are associated with higher rates of ramification to produce lateral shoots. This study reveals how branching system and annual shoot traits change simultaneously during tree ontogeny to fulfill different functions, particularly light exploitation and space exploration.
... Shoot growth in spring (i.e. spring flush) is preformed in the winter buds, i.e. dependent on reserves stored in the tree (Sabatier and Barthelemy, 2001;Lauri, 2007). Although bourse shoots are neoformed vegetative growth units developed as sylleptic axillary shoots generated from the axils of preformed spur leaves (Costes and Guedon, 2002), results in the current study suggest that the growth potential of bourse shoots might be established in the previous year (as in the other shoots). ...
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Knowledge of the growth dynamics of young and mature trees contribute to development of efficient orchard management practices. In this experiment, the aim was to provide apple growers with practical information on how vegetative growth (at both shoot and whole-tree level), yield and fruit quality change during the transition period, from young to mature trees. The experiment orchard was planted at 3.5 m × 1 m inter-row and intra-row spacings, respectively, with ‘Golden Delicious’/M.9 trees. Trees were trained since planting as modified vertical axis. Vegetative growth at shoot level reacted differently to yield load than that at the whole-tree level. The yield, yield efficiency and annual canopy volume increase and decrease (a vegetative growth indicator at the whole-tree level) showed a positive correlation with shoot length. TCSA (annual increase and actual TCSA), as another vegetative growth indicator at whole-tree level in the experiment, correlated negatively with shoot length. The yield varied biennially and, was greatest in the seventh year after planting (21.76±8.46 kg tree-1, corresponding to an orchard yield of ~62 t ha-1) that is an acceptable tonnage in the region of the experiment. To maintain the balance between vegetative growth and fruiting, orchard management practices should be conducted considering yield. No pruning or light pruning is recommended in the ‘off-year’ of biennial cycle of fruiting as a result of this study.
... For young trees, bi-cyclic shoots are the principal growth structure, because they provide greater and faster coverage of the assigned space due to their large size in relation to mono-cyclic shoots. However, in mature walnut trees, mono-cyclic shoots are the main bearing structure (Barthelemy et al., 1995;Sabatier and Barthelemy, 2001a, b), since they provide the greatest amount of fruit. ...
Article
In mature walnut trees, mono-cyclic shoots are the main bearing structure and are thus fundamental for tree productivity. In this line, a study was carried out in three commercial “Chandler” walnut orchards, located in the central valley of Chile. In each orchard, eight mature trees were selected and eighteen mono-cyclic shoots were marked according to three fruit load categories (shoots bearing one fruit (F1), shoots bearing two fruits (F2), shoots bearing three fruits (F3)). These shoots were distributed in three zones of the canopy (upper, lower-exposed, lower-shaded). This resulted in 432 marked shoots in the total of the experiment, in which the effect of position and fruit load on length, diameter, leaf area and specific leaf weight were studied. In addition, multilinear regression models for fruit diameter as a function of the above-mentioned characteristics were performed. The results indicate that the production of two or more fruits per shoot is associated with an average shoot length of 12.7 cm, an average shoot diameter of 8.1 mm and an average leaf area of 1034.7 cm². Linear models indicate that length and specific leaf weight significantly explain fruit size for all shoot categories. In addition, the leaf area, and leaf area plus basal diameter of the shoot, are significantly related to fruit size (equatorial diameter) for shoots with two and three fruits, respectively. According to the results of this study, the tree’s upper and lower-exposed zones offer shoots with greater potential to produce more and larger fruits per shoot than the lower-shaded canopy zone. Thus, the morphological characteristics of the shoot (diameter, length and leaf area, among others) are directly related to a greater number and size of fruits, and therefore higher productivity.
... The terminal bud which will complete next year's AS already is articulated which means that the entire embryonic AS is visible inside the bud, analogous to, for example, the walnut (Juglans regia; Sabatier & Barthélémy, 2001). ...
... The terminal bud which will complete next year's AS already is articulated which means that the entire embryonic AS is visible inside the bud, analogous to, for example, the walnut (Juglans regia; Sabatier & Barthélémy, 2001). ...
Research
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In the work presented here we describe the bud-forming process in the pedunculate oak (Quercus robur L.). During spring it takes just a few weeks before a young shoot has fully developed. The bud from which it originates has taken one and a half years to reach this stage. The terminal bud of the young branch at this point consists only of a few outer bud scales. In the period from May to July develop from the core of the bud the primordia for next year's branch, as well as the initial stages of its terminal bud. The sequence of shapes ('heteroblastic series', Bell 1991, p. 28) of the organs in the transition A) from the foliage leaves to the terminal bud (that is, the 'metamorphosis into the bud') and B) from the terminal bud to the foliage leaves (the 'metamorphosis out of the bud') are described in detail. The series of shapes labelled 'A' shows broad similarities with herbaceous plants and their sequence of leaf forms along the stem up to the blossom. In both cases the 'upper leaf zone' (Bell, 1991, p. 20) is reduced, first the petiole, then the lamina, leaving behind the leaf base as the outermost scales of the winter bud in the pedunculate oak, or the sepals of a blossom in herbaceous plants. As the bud opens in spring (see series B), or with the blossoming of the angiosperm flower, a calyx opens. In the pedunculate oak the outermost scales of the winter bud form this calyx, in a blossom it is formed by sepals. To the inside of the outer bud scales appear inner, transitory scales that on many bosks are coloured; in the blossom the petals and stamens are located inside the sepals. In vegetative buds of woody plants the foliage leaves are arranged centrally, tightly packed, the internodes not yet expanded. This gesture, typical for blossoms, is quickly lost during the growth of the young branch, whereas in a blossom it remains for the entire flowering period. A comparison of the foliage leaves situated in the center of the vegetative bud to the carpels that likewise are situated in the centre of the blossom appears compelling. In the theory of metamorphosis annual plants serve as the model to help visualize the metamorphosis from shoot to carpel. The metamorphosis into the blossom, however, can be grasped more readily, if one takes the perennial, especially the woody plant for a model. In studying the bud-forming process of the perennial we can observe a close relationship between the vegetative bud and the bud of a blossom. For the blossom spore formation is added to the bud-forming process as an intrinsic formative principle. 2
... Die Endknospe, welche die JZE der kommenden Saison abschliesst, ist bereits angelegt, d.h. die gesamte JZE ist in der Winterknospe präformiert, wie z.B. auch bei der Walnuss (Juglans regia; Sabatier & Barthélémy, 2001). ...
Article
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In der vorliegenden Arbeit wird der Knospen-Bildungsprozess bei der Stiel-Eiche(Quercus robur L.) beschrieben. Im Frühjahr braucht es einige wenige Wochen,bis ein junger Trieb vollkommen ausgewachsen ist. Die Knospe, aus der erhervorgegangen ist, hat bis dahin für ihre Entwicklung eineinhalb Jahre gebraucht. Die Endknospe des jungen Zweiges wird zu diesem Zeitpunkt von den äusserenKnospenschuppen gebildet. Im Laufe der Monate Mai bis Juli entwickelt sichdarin die Anlage für den nächstjährigen Zweig wie auch die ersten Stadien derdazugehörigen Endknospe. Die Formen-Reihen der Organe an den Übergängen von A) den Laubblätternzur Endknospe (die Metamorphose in die Knospe hinein) und B) der Endknospezu den Laubblättern (die Metamorphose aus der Knospe heraus) werden detailliertbeschrieben. Die Formen-Reihe A) zeigt grosse Ähnlichkeiten mit den Blattreihenkrautiger Pflanzen auf die Blüte hin. In beiden Fällen wird das Oberblatt reduziert,zuerst der Blattstiel, dann die Blattspreite, und übrig bleibt der Blattgrund alsäusserste Schuppen der Winterknospen bei der Stiel-Eiche bzw. als Kelchblätterder Blüte. Beim Knospen-Austrieb im Frühling (Formen-Reihe B) bzw. beim Aufblühender Bedecktsamer-Blüte öffnet sich ein «Kelch». Bei der Stiel-Eiche bilden dieäusseren Winterknospenschuppen diesen «Kelch», bei der Blüte die Kelchblätter. Auf die äusseren Knospenschuppen folgen innere, vergängliche, bei zahlreichenGehölzen auch gefärbte Schuppen. In der Blüte stehen innerhalb der Kelchblätterdie Blüten- und Staubblätter. Zentral in der vegetativen Knospe der Gehölze sinddie Laubblätter angeordnet, dicht gedrängt, die Internodien haben sich noch nichtgestreckt. Diese für die Blüte typische Geste verliert sich beim Austreiben des jungenZweiges rasch, bei der Blüte jedoch bleibt sie während der gesamten Blütezeitbestehen. Der Vergleich der in der vegetativen Knospe zentral stehenden Laubblättermit den in der Blüte ebenfalls zentral stehenden Fruchtblättern drängt sich auf. In der Metamorphosen-Lehre dient die einjährige Pflanze als Modell, mit dessenHilfe man die Metamorphose vom Keim- bis zum Fruchtblatt veranschaulicht. Die Metamorphose zur Blüte lässt sich aber einfacher nachvollziehen, wenn man die mehrjährige, insbesondere die Gehölz-Pflanze als Modell nimmt. Bei dermehrjährigen Pflanze zeigt sich beim Studium der Knospen-Bildungsprozesse eineenge Verwandtschaft der vegetativen mit der Blüten-Knospe. Bei der Blüte kommt zur Knospen- die Sporenbildung als spezifisches Bildungsprinzip hinzu.
... On young walnut trees, shoot growth is bimodal. The portion of the shoot that is formed during the previous season and overwinters in the dormant bud is known as the preformed portion of the shoot; the portion of the shoot that forms during the current growing season is known as the neoformed portion of the shoot (Sabatier and Barthélémy 2001). Preformed shoot growth is completed by early May, and shoot growth from June through fall is neoformed. ...
Article
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In traditionally managed Howard walnut orchards, trees are pruned annually during the orchard development phase, an expensive operation in terms of labor and prunings disposal costs. Our observations and some prior research by others had suggested that pruning may not be necessary in walnut. In a trial of pruned and unpruned hedgerow trees over 8 years, beginning a year after planting, we documented canopy growth, tree height, yield and nut quality characteristics and also the effects of fruit removal. Pruning altered canopy shape but did not lead to increases in canopy development, yield or nut quality. Although fruit removal stimulated more vegetative growth in both the pruned and unpruned treatments, fruit removal did not result in an increase in midday canopy photosynthetically active radiation interception or cumulative yield when fruit removal was stopped after year 4. After 8 years, there were no significant differences in tree height, nut quality or cumulative yield among any of the treatments, which suggests that not pruning young Howard orchards could provide a net benefit to growers.
... Barthé lé my and Caraglio 2007). Along a parent shoot, buds positioned distally are larger, contain a greater number of preformed organs and display higher probabilities of bud break than basal buds (Gill 1971; Remphrey and Davidson 1994; Sabatier and Barthélémy 2001; Gordon et al. 2006; Alla et al. 2013b). The mechanisms implicated in determining bud characteristics include hormonal signalling (Cline 2000; Djennane et al. 2014) and resource competition among buds (Cline et al. 2009; Mason et al. 2014), but several recent studies also highlight the importance of vascular connections between buds and stem. ...
Article
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Plant architecture is shaped by endogenous growth processes interacting with the local environment. The current study investigated crown development in young black alder trees, assessing the effects of local light conditions and branch height on individual bud mass and contents. In addition, we examined the characteristics of parent shoots (stem cross-sectional area and total leaf area, shoot length, the number of nodes, the number and total mass of buds per shoot) and leaf-stem as well as bud-stem allometry, as several recent studies link bud development to hydraulic architecture. We sampled shoots from top branches and two lower-crown locations: one subjected to deep shade and the other resembling the upper branches in light availability. Sampling was carried out three times between mid-July and late October, spanning from the early stages of bud growth to dormancy. Individual bud mass and shoot characteristics varied in response to light conditions, whereas leaf-stem allometry depended on branch height, most likely compensating for the increasing length of hydraulic pathways. Despite the differences in individual bud mass, the number of preformed leaves varied little across the crown, indicating that the plasticity in shoot characteristics was mainly achieved by neoformation. The relationship between total bud mass and stem cross-sectional area scaled similarly across crown locations. However, scaling slopes gradually decreased throughout the sampling period, driven by bud rather than by stem growth. This suggests that the allometry of total bud mass and cross-sectional area of stem is regulated locally, instead of resulting from crown-level processes. Published by Oxford University Press on behalf of the Annals of Botany Company.
... This choice is coherent with several observations that will be detailed later in this article. For example, it is linked to the size of buds, which has a significant impact on future shoot development (Sabatier and Barthélémy, 2001) and to the sizes of growth units that control the appearance of lateral axes (Nicolini and Chanson, 1999). In Mathieu et al. (2008), we showed how rhythms were automatically generated by the model for some parameter values. ...
Article
In this paper are presented new mathematical developments in plant growth modelling and simulation. GreenLab Model is a functional-structural plant growth model, it combines both organogenesis (architecture) and photosynthesis (biomass production and repartition). New improvements concern the retroaction of photosynthesis on organogenesis. We present in this paper the influence of available biomass on the number of metamers in a growth unit and on the branching. The general theory is introduced and applied to simple trees. Some interesting behaviours are underlined.
... Since only one mother tree was present at the edge of the studied stand, most of the saplings were suspected to be half-brothers. The limit between two successive annual shoots is marked by a zone of short internodes and scale leaves (Sabatier and Barthélémy, 2001) which facilitates the retrospective measurement of successive annual shoots along the main stem. Of the 138 trees measured, 22 were branched. ...
Article
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This study aimed to identify robust indicators that summarize the respective importance of ontogeny and environmental constraints in tree development. In the proposed approach, tree growth data correspond to the retrospective measurement of annual shoot characteristics (e.g. length, number of branches) along the main stem. We applied segmentation models to identify tree growth phases. These segmentation models, which are hidden semi-Markov chains, were compared with simple hidden Markov chains that correspond to the environment-driven development assumption. This statistical modelling approach was applied to both evergreen (Corsican pine and silver fir) and deciduous (sessile oak and Persian walnut) tree species growing in contrasted conditions ranging from managed forest stands to unmanaged understoreys. Growth phase duration distributions estimated within these segmentation models characterize the respective importance of ontogeny and environmental constraints in tree development at the population scale and have very contrasted characteristics in terms of shape and relative dispersion between ontogeny-driven and environment-driven tree development. These characteristics may change over tree life, reflecting changes in tree competition. Growth phase duration distributions summarize the joint trajectory of tree ontogeny and environment without requiring tree growth follow-up data for their estimation.
... Shoot apical meristem abortion in Fraxinus pennsylvatica appears in part genetically programmed and results in the modification of branching and eventually plays a role in crown form evolution with time (Remphrey and Davidson, 1992). It is likely that the presence of aborted meristems within zones with high branching frequency reflects local competitive relationships (e.g., Juglans regia; Sabatier and Barthélémy, 2001). It has been hypothesized that the proximal gradient of aborted meristems along the apple (Malus domestica) one-year-old shoot is in some way related to the acrotonic gradient of growing (vegetative and flowering) branches (Lauri and Térouanne, 1998;Lauri, 2007). ...
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It is commonly assumed that the modular structure of plants results from two basic processes: the apical growth process by which the plant builds the monopodial shoot, and the branching process initiated by the development of the axillary meristem. The architectural analysis proposed some 40 years ago by Hallé and colleagues (Hallé et al., 1978) considers the actual morphology of a plant as resulting from a balance between the deterministic genetic blueprint specific to the taxon and the environmental cues. A finite number of criteria have been defined to analyze tree architecture which all refer to the behaviour of meristems: fate (i.e., vegetative or reproductive), growth orientation (i.e., vertical, orthotropic or any other directions, plagiotropic), growth rhythmicity (i.e., continuous or rhythmic), distribution along the parent shoot, development with (delayed or proleptic) or without (immediate or sylleptic) a resting period, etc. These criteria refer to growing meristems. However non-growing meristems play an important role in plant architecture. These meristems belong to two categories. The first one includes meristems remaining latent (dormant) as buds. They usually serve as a reserve of meristems within the individual plant and are typically involved in the proleptic reiteration process by which the plant regenerates the whole or a part of its architecture. Another non-growing meristem category includes meristems which permanently lose their growth potential through abortion ultimately involving cladoptosis, i.e., the shedding of a single shoot or of a whole-branch complex. These phenomena are usually described as an adaptation to environmental stresses, e.g., shade and drought, possibly mediated by a decrease in the hydraulic conductance. However, it is known that the permanent loss of meristematic activity may be highly predictable revealing a genetic or an epigenetic basis. This is typically the case of meristem extinction which has been coined in apple to describe meristem abortion with the following characteristics: independent of external (light, temperature) conditions and likely involving a genetic or epigenetic determinism, precocious (as soon as in the first weeks following bud burst), and in most of the cases involving reproductive laterals. A relation between frequency of extinction and development of the remaining laterals, especially an increase of the frequency of flowering, has been shown. When and where extinction occurs is thus as important in defining shoot and plant architecture as the analysis of the distribution of growth characteristics of shoots. This interweaving of growth and death processes in the dynamics of plant form follows a known Atlan’s principle, since it appears to be a specific case of the statement established by Henri Atlan (1979) at cellular and tissular levels, mainly in animal science, “Sans processus de mort contrôlée, pas de processus de vie.
... In fruit trees, shoot polymorphisms (for instance differences between short and long shoots) originate from changes in endogenous processes . Depending on the species, proleptic shoots that develop from vegetative buds after they have gone through dormancy (Hallé et al., 1978) can be composed only of preformed organs that are formed within the dormant bud (Thorp et al., 1994;Sabatier and Barthélémy, 2001) or by preformed and neoformed organs that are formed and extended after bud break (Remphrey and Powell, 1984;Gordon et al., 2006a). Proleptic shoots can have distinct structural pattern characteristics depending on genotype Guédon, 1997, 2002), size of the shoots (Fournier et al., 1998) and age of the tree when shoots develop (Renton et al., 2006). ...
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Background and AimsShoot 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.MethodsA 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 ResultsProleptic 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. © 2013 © The Author 2013. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: [email protected] /* */
... This phenomenon is mostly known as 'lammas shoots', as they often appear around Lammas day on August 1 (Evans 1972;Guédés 1981;Kozlowski and Pallardy 2002;Larcher 2003;Meier 2003). However, other terms like 'Johannis shoots' (Kobel 1954;Lyr et al. 1967;Weinreich 2000) or 'summer flushes' (Sabatier and Barthélémy 2001) are also used and in some species, even more than two flushes per growing season can occur (Jones 1959; Barthélémy and Caraglio 2007;Cline and Harrington 2007; for a critical revision of terms also see Caraglio and Barthélémy 1997). The occurrence of lammas shoots is most probably endogenously controlled, but also depends strongly on environmental conditions, such as water and nutrient supply during the growing season (Kobel 1954;Jones 1959;Kozlowski 1964;Simak 1970;Kolb and Matyssek 2001;Kozlowski and Pallardy 2002;Codesido and Fernandez-Lopez 2009). ...
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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.
... This result is consistent with Sabatier and Barthélémy (2001a, b). They explained the phenomenon by very small-sized axillary buds that are usually located on the BAZ section, and posses more cataphylls than embryonic green leaves as compared to the buds from the MED shoot zone (Sabatier and Barthélémy 2001b). Furthermore, an acrotony, i.e. a side shoot architecture in which distal branches grow more vigorously (Bell 1998) was evident. ...
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Relationship of bud production (axillary and terminally) of annual shoot (1Y) and/or the content of bud-derived indol-3-acetic acid (IAA) to branching of the 1Y was studied in common walnut (Juglans regia L.), cvs. Franquette and Lara. Cultivar-related branch architecture was determined. Lara tended to branch more densely than Franquette (53 vs. 42%). Significantly more fruiting off-spring shoots (FO) than vegetative ones (VO) grew-out per 1Y in both cultivars, whereas the ratio FO/VO of Lara exceeded that of Franquette by four times. An acrotonic branching pattern was more strongly expressed in Lara compared to Franquette. Bud-derived IAA was influenced by the cultivar (Franquette had 3.6 times more cumulative IAA along the 1Y than Lara), and by the relative position (terminal, subterminal, medial and basal) of the buds along 1Y. An opposite relationship between branching density and cumulative IAA content was established in both cul-tivars. At the 1Y relative position level, the opposite ratio between branching density and IAA content was clearly shown only on the basal position of the bud along 1Y in the Lara cultivar. Such an inconsistent linkage between bud production and the IAA spatial distribution along the 1Y illustrated that hormonal factors probably weakly affect the branching of Franquette and Lara. The length of the parent 1Y, the position of the buds along the 1Y-length, and the fate of the buds seemed to have a stronger influence on the bud out-growth and further development of the off-springs. In further analyses, seasonal fluctuations of the IAA, and the following activity of the buds should be investigated in order to improve the understanding of a complex branching phenomenon in walnut.
... Bud size determines primary growth and reproduction ability in plants with rhythmic growth (Nitta and Ohsawa, 1998). This suggests that bud size is modulated by several drivers including the individual tree size and vigour, the position of the bud in the shoot, and ontogenetic factors (Remphrey and Powell, 1984;Sabatier and Barthélémy, 2001;Puntieri et al., 2002aPuntieri et al., , 2002bIshihara and Kikuzawa, 2009;Alla et al., 2011). Among the endogenous factors, secondary growth has been found to drive bud size since changes in current-year stem cross-sectional area are linked to the stem hydraulic conductivity and the size of buds (Cochard et al., 2005;Lauri et al., 2008;Alla et al., 2011). ...
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Background and AimsIn trees, bud development is driven by endogenous and exogenous factors such as species and climate, respectively. However, knowledge is scarce on how these factors drive changes in bud size across different time scales.Methods The seasonal patterns of apical bud enlargement are related to primary and secondary growth in two coexisting Mediterranean oaks with contrasting leaf habit (Quercus ilex, evergreen; Quercus faginea, deciduous) over three years. In addition, the climatic factors driving changes in bud size of the two oak species were determined by correlating bud mass with climatic variables at different time scales (from 5 to 30 d) over a 15-year period.Key ResultsThe maximum enlargement rate of buds was reached between late July and mid-August in both species. Moreover, apical bud size increased with minimum air temperatures during the period of maximum bud enlargement rates.Conclusions The forecasted rising minimum air temperatures predicted by climatic models may affect bud size and consequently alter crown architecture differentially in sympatric Mediterranean oaks. However, the involvement of several drivers controlling the final size of buds makes it difficult to predict the changes in bud size as related to ongoing climate warming.
... The branching probability of different nodes of annual shoots was investigated by Sabatier and Barthélémy (2001b). With regard to the observed variation in size and content of axillary buds according to the position, they distinguished three successive zones on a parent shoot: the apical, medial and basal zone. ...
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An architectural analysis of a fruiting branch in the Slovenian variety 'Elit' was conducted during three successive years. The fruit bearing branch was constructed of a three-year-old parent shoot plus all corresponding two-year-old shoots and annual shoots (1Y). The construction of the bearing branch during the time caused a statistically significant increase in the number of annual shoots. The basal diameter and the length of 1Y significantly depended on a year whereas their angles did not. In spite of decreasing length of 1Y, the number of vegetative buds per shoot increased from the first to the third year of observations. The number of nodes was closely correlated with the length of the shoots. Activity points on the 1Y were most often on the apical two or three nodes, what was expressed by a marked acrotony. The number of active points varied as the tree matured. The ratio fruit bearing 1Y / total 1Y was 0.47 in year 1; 0.18 in year 2; 0.74 in year 3. It points to the slightly alternance. As the tree grew the number of flowering buds per 1Y as well as the number of female flowers per 1Y increased. The results of the three-year-long research show some growth and development rules in the walnut cultivar 'Elit', however, they do not allow a reliable prediction of the following activities. We assume that this will be possible after another three-year-long analysis, with the help of the Hidden Mark Model.
... There was found a decline corresponding to a growth reduction and an increase in the probability of flowering from the centre of the tree towards the periphery. This centrifugal gradient found in apple is also consistent with results found in other tree species (Costes et al. 1992; Sabatier and Barthelemy 2000) and with the definition of a within-tree 'physiological age' of buds (Gatsuk et al. 1980). Fruits also effect flower formation in apple as development of fruits coincides with the time of flower induction. ...
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The development of flower buds and a sufficient fruit set are basic requirements for fruit growers to generate a marketable crop. However, fruit trees remain in a juvenile (nonflowering) phase for years, and after a transition period of getting reproductively competent they enter the adult phase of tree life. The reproductive phase is associated with the ability to alternate between the production of vegetative and reproductive buds. Efficient breeding is limited in fruit trees due to the long period of juvenility. Therefore, it is important to accelerate flowering by reducing the juvenile phase of the tree. In fruit production, precocious flowering of the tree is also favoured to reduce the vegetative phase of tree development after planting in order to obtain the earliest fruit crop. Additional critical aspects of flower deve-lopment in fruit trees, such as alternate bearing, accentuate the necessity to improve our understanding on genetic factors controlling floral initiation as well as flower and fruit development in perennial fruit trees. Most of what we know about regulating floral development is based on research in annual plants, like Arabidopsis thaliana. In this review, we summarize floral transition, meristem development and flower bud formation in Malus domestica, one of the most important representatives of temperate fruit trees. We also focus on current findings of the transition from vegetative to reproductive growth obtained in Arabidopsis and how this knowledge can be applied to fruit trees, particular to apple. We discuss state-of-the-art and future research to manipulate maturation and flower initiation in apple.
Preprint
Die Arbeit vertritt die These, dass die vegetative Knospe die Grundlage abgibt für die Blüte. Sie geht über die Annahme hinaus, dass das Blatt die vegetative Grundlage für die Blütenorganen, für Kelchblatt, Kronblatt, Staubblatt und Frucht-blatt abgibt. Die vegetative Knospe enthält die beiden Hauptformen des Blattes, Schuppenblatt (Kataphyll) und Laub-blatt.(Trophophyll). Das Schuppenblatt gibt die Grundlage ab für die ersten drei Organkategorien Kelchblatt, Kronblatt und Staubblatt, das Laubblatt gibt die Grundlage ab für das Fruchtblatt. Die, bezüglich der Blattform, im Allgemeinen bleibende Aussage dass das Blatt die Grundlage für die Blütenorganen abgibt, ist auf das vorherrschende Modell einer einjährigen Blütenpflanze als Modell zur Erläuterung des Bauplans der Angiospermen zurückzuführen. Als Alternative und Ergänzung wird das Modell einer mehrjährigen, holzbildenden Pflanze vorgeschlagen. Das basiert auf dem Grund-organkonzept von Julius Sachs und Karl Goebel unterscheidet zwischen vegetativen Grundorganen und generativen Grundorganen. Die beiden Sporophylle Staubblatt (Mikrosporophyll) und Fruchtblatt (Makrosporophyll) gehen aus dem Zusammenspiel dieser beiden Organklassen hervor. Das Staubblatt kann man in diesem Zusammenhang als Katasporophyll bezeichnen und das Fruchtblatt als Trophosporophyll. Die Gestaltungsmöglichkeiten des Schuppenblattes und des Laubblattes sind durch die inneren Bedingungen des rhythmischen Wechsels zwischen Zweig und Knospenbildung eingeschränkt. Die Einschränkung, die die Bildung von Knospen mit sich bringt, fällt bei dein krautigen Sprossen weg. Die Gestaltungsmöglichkeiten des krautigen Sprosses sind durch die Bedingung des unbegrenzten Wachstums eingeschränkt. Diese Einschränkung fällt in der Blüte, mit ihrem begrenztem Wachstum, weg.
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Einführung Die ersten bedecktsamigen Pflanzen (Magnoliopsida) waren mehrjährige Holzpflanzen. Es waren kleine Bäu-me. Die ersten Angiospermen bildeten Knospenschup-pen (Schuppenblätter) und Laubblätter. Später kam die Bildung von Nebenblättern oder Stipeln dazu. Die Blü-ten der Angiospermen bestehen aus Kurzsprossen mit einem Perianth, das aus sterilen Organen besteht und aus männlichen und weiblichen reprodutktiven Orga-nen. Die Pflanze hat die Fähigkeit mit Hilfe einiger wenigen Organe eine unüberschaubar grosse Vielfalt an Formen aus sich hervorgehen zu lassen. Diese Organe werden Grundorgane genannt. Grundorgane sind per Definition Organe, die man morphologisch nicht voneinander ab-leiten kann und unterschiedliche Funktionen ausüben. Das am meisten verwendete Konzept, das von Wilhelm Troll 1 vertreten wurde, kennt nur drei Grundorgane: Wurzel, Sprossachse und Blatt. Als Modell zur Erläute-rung der Anordnung und der Gestalt der Angiospermen verwendet Troll das Schema (Bauplan) einer einjähri-gen, krautigen Pflanze. Wie ist es, wenn man als Referenzmodell nicht eine ein-jährige, sondern eine mehrjährige, holz-und knospen-bildende Pflanze nimmt? So ein Modell verbindet die Bildung von Knospen und Zweigen, das heisst Mehr-jährigkeit, sekundäres Dickenwachstum, mit Einjährig-keit, das heisst mit der Bildung von Infloreszenzen und Blüten. Die Wahl eines Referenzmodells ist das Ergebnis 1 Troll (1954) Die vegetative Knospe als Grundlage der Blüte Peer Schilperoord, Alveneu Dorf, Switzerland, schilperoord@bluewin.ch Zusammenfassung Die Arbeit vertritt die These, dass die vegetative Knospe die Grundlage abgibt für die Blüte. Sie geht über die Annahme hinaus, dass das Blatt die vegetative Grundlage für die Blütenorganen, für Kelchblatt, Kronblatt, Staubblatt und Frucht-blatt abgibt. Die vegetative Knospe enthält die beiden Hauptformen des Blattes, Schuppenblatt (Kataphyll) und Laub-blatt.(Trophophyll). Das Schuppenblatt gibt die Grundlage ab für die ersten drei Organkategorien Kelchblatt, Kronblatt und Staubblatt, das Laubblatt gibt die Grundlage ab für das Fruchtblatt. Die, bezüglich der Blattform, im Allgemeinen bleibende Aussage dass das Blatt die Grundlage für die Blütenorganen abgibt, ist auf das vorherrschende Modell einer einjährigen Blütenpflanze als Modell zur Erläuterung des Bauplans der Angiospermen zurückzuführen. Als Alternative und Ergänzung wird das Modell einer mehrjährigen, holzbildenden Pflanze vorgeschlagen. Das basiert auf dem Grund-organkonzept von Julius Sachs und Karl Goebel unterscheidet zwischen vegetativen Grundorganen und generativen Grundorganen. Die beiden Sporophylle Staubblatt (Mikrosporophyll) und Fruchtblatt (Makrosporophyll) gehen aus dem Zusammenspiel dieser beiden Organklassen hervor. Das Staubblatt kann man in diesem Zusammenhang als Katasporo-phyll bezeichnen und das Fruchtblatt als Trophosporophyll. Die Gestaltungsmöglichkeiten des Schuppenblattes und des Laubblattes sind durch die inneren Bedingungen des rhythmischen Wechsels zwischen Zweig-und Knospenbildung ein-geschränkt. Die Einschränkung, die die Bildung von Knospen mit sich bringt, fällt bei dein krautigen Sprossen weg. Die Gestaltungsmöglichkeiten des krautigen Sprosses sind durch die Bedingung des unbegrenzten Wachstums eingeschränkt. Diese Einschränkung fällt in der Blüte, mit ihrem begrenztem Wachstum, weg.
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The growth dynamics of annual shoots of Nothofagus antarctica shrubs from northern Patagonia was studied. The pattern of extension of the shoots derived from the three most distal buds of 100 parent shoots was registered. Weekly measurements and observations of extending shoots were carried out during the 1996–1997 growth season. The date of shoot extension initiation was less variable than the date of shoot extension cessation. Extension curves had an asymmetrically sigmoid outline. Most extension took place uninterruptedly between mid-September 1996 and the end of January 1997. Extension rate reached its highest value in the second half of the extension period. Leaf unfolding rate had a peak early in the extension period and fluctuated irregularly later on. The apical meristem of all shoots died by the end of shoot extension. In general terms, shoot length, diameter, leaf number and extension duration decreased from the first to the third position from the parent shoot’s apex, although exceptions to this pattern were found. Shoot extension rate was affected by maximum and minimum daily temperatures but not by the amount of precipitation during the extension period. The size of a bud was not related to the size of the shoot derived from it. The size of shoots in the third position from the apex was related to parent shoot size; this was not the case for more distal shoots.
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Modelling plant structure and growth has undergone major changes in the last decades along two major lines: the integration of ecophysiological knowledge in process-based models which often lack a description of plant topology and geometry, and the generation of 3-D virtual plants using morphogenetic models which simulate the architectural development in a stable and homogeneous environment. There is now a trend to merge these two approaches, that is to link plant architecture and functioning. This trend is based on the recognition that plant structure: (i) is the joint output of the physiological processes (water and carbon balance, etc.) and the morphogenetic programme of the plant, (ii) determines the external environment of the trees which itself regulates their functioning (competition for space, light attenuation, etc.), and (iii) directly conditions the physiological processes within the tree (hydraulic structure, self-shading, allocation of photosynthates, etc.). Such models can be used in agronomy and forestry in various ways: to investigate the effects, local and global, immediate and delayed, of the biophysical environment on plant morphogenesis and yield; to study light attenuation through the canopy, to analyse the transport of water and the allocation of photosynthates within the plant; to analyse the competitive interactions among different plants in the same stand; to calibrate remote sensing techniques and to visualize large landscapes.
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The influence of shoot architectural position on growth and branching pattern of young Cedrus atlantica (Endl.) Manetti ex Carrière trees were studied. Extension growth and type of axillary products (lateral bud, sylleptic short or long shoots) of annual shoots of increasing branching order (main stem, branches and branchlets) were recorded weekly during the 1993 growing season. Annual final shoot length, duration of extension, and maximum extension rate decreased with increasing branching order. Sylleptic axillary shoots occurred only on annual shoots of the main stem and branches and were produced when extension rate was at its highest. Differences in growth rate and final length of annual shoots, according to their architectural position, were related to differences in the total number and diversity of types of sylleptic axillary shoots produced. It is suggested that types and numbers of sylleptic axillary shoots produced are linked with threshold values for both final length and extension rate of the parent shoot. Copyright 1999 Annals of Botany Company
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The numbers of nodes on single flush terminal and axillary shoot modules were determined in a range of Persea species and cultivars. They were compared with node numbers in apical and axillary buds to investigate whether preformation or neoformation of nodes occurred. Mean number of nodes on terminal shoots was 14 for vegetative shoot modules and 21 for reproductive shoot modules, and was similar across species, cultivars, rootstocks, locations and climates. In the cultivar 'Hass', numbers of nodes on axillary shoot modules were variable, and lower than those for primary shoot modules forming the dominant growth axis of annual growth modules. There was a mean of 12 nodes for vegetative proleptic shoot modules, 15 for reproductive proleptic shoot modules and six for sylleptic shoot modules, which were invariably vegetative. All nodes were preformed within both apical and axillary proleptic buds. This was not the case in sylleptic buds, which burst contemporaneously with extension of the parent axis. The majority (63%) of reproductive buds formed indeterminate compound inflorescences. They carried six basal bud scales, six axillary inflorescences and their subtending bracts, and up to nine true leaves.Copyright 1994, 1999 Academic Press
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Buds of shoots from the trunk, main branches, secondary branches and short branches of 10–21 year-old Nothofagus pumilio trees were dissected and their contents recorded. The number of differentiated nodes in buds was compared with the number of nodes of sibling shoots developed at equivalent positions during the following growing season. Axillary buds generally had four cataphylls, irrespective of bud position in the tree, whereas terminal buds had up to two cataphylls. There were more nodes in terminal buds, and the most distal axillary buds, of trunk shoots than in more proximal buds of trunk shoots, and in all buds of shoots at all other positions. The highest number of nodes in the embryonic shoot of a bud varied between 15 and 20. All shoots had proximal lateral buds containing an embryonic shoot with seven nodes, four with cataphylls and three with green leaf primordia. The largest trunk, and main branch, shoots were made up of a preformed portion and a neoformed portion; all other shoots were entirely preformed. In N. pumilio, the acropetally-increasing size of the sibling shoots derived from a particular parent shoot resulted from differences in: (1) the number of differentiated organs in the buds; (2) the probability of differentiation of additional organs during sibling shoot extension; (3) sibling shoot length; (4) sibling shoot diameter; and (5) the death of the apex and the most distal leaves of each sibling shoot. Copyright 2000 Annals of Botany Company
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Four clones of Fraxinus pennsylvanica var. subintegerrima (Vahl) Fern. were planted in replicated trials at two sites in Manitoba (Morden and Winnipeg) to investigate shoot growth and leaf neoformation in relation to genotype, environment and pruning treatment over a 3 year period. Significant differences were found among clones, years and sites for shoot length and numbers of neoformed leaves. Neoformation was highest shortly after transplanting and then declined. An increase in neoformation was evident following cold related winter injury or loss of terminal buds by late spring frosts. Pruning and terminal bud removal treatments both increased neoformed leaf production relative to control trees. The trees were able to quickly re-establish photosynthetic surface area after injury or treatment and neoformed leaf production was an important component in this recovery strategy. The capacity for neoformation also varied in relation to genotype but appeared to be very plastic, being affected by a wide variety of factors. In addition, the rate or pattern of change in amounts of neoformed leaves over time and locations was variable. Numbers of neoformed leaves increased with shoot length but variation in the relationship suggested that there were differences in internode length as well. Neoformed leaves were highest in the upper part of the crown indicating that there was differential allocation of resources within the crown.
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Trees architectural study shows gradients in the meristematic activity. This activity is described by the number of internodes per growth unit, which is considered as the output of a dynamic random process. Several species were observed, which led us to propose and then estimate some mathematical models. Computing the functioning of a tree in a given environment therefore involves finding the probability function of the meristems and following the evolution of the parameters of this law along the botanical gradients (order, trend, acrotony) within its architecture.
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Shoot preformation was investigated in buds of four clones of Fraxinus pennsylvanica var. subintegerrima (Vahl) Fern. at two sites in Manitoba in the second (1988) and third (1989) growing seasons after grafting. More preformed primordia were produced in terminal buds in 1989 compared to 1988 at each site. Both terminal and lateral buds at Morden contained significantly more primordia than those at Winnipeg. The numbers of preformed primordia were significantly different among clones. Clone 3 produced the most and clone 1 the fewest primordia in terminal buds. Despite quantitative variation, the pattern was similar among clones for terminal buds at each site and in each year. A similar pattern was evident for lateral buds at the Winnipeg site in 1989 but at Morden, clones 4 and 1 had the largest number of preformed primordia. Data from 1989 revealed that numbers of primordia were correlated with bud dimensions, parent shoot length, diameter and number of leaves, and location of the bud on the parent. Shoot dry weight was also related to these variables and revealed a non-linear increase in dry weight with shoot length. Multiple regression, with parent shoot length and location of buds along the parent axis as independent variables provided a reliable indicator of preformation in the crown. Although there is a genotypic component to preformation, variation between sites, years and crown locations suggests plasticity in bud development.
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