Clarifying the effects of dwarfing rootstock on vegetative and reproductive growth during tree development: a study on apple trees

BACKGROUND AND AIMS: Despite the widespread use of dwarfing rootstocks in the fruit-tree industry, their impact on tree architectural development and possible role in the within-tree balance between growth and flowering are still poorly understood, in particular during the early years of growth. The present study addressed this question in apple trees, through a detailed analysis of shoot populations, i.e. both vegetative and flowering shoots, during tree development. METHODS: Architectural databases were constructed for trees of two cultivars that were either own-rooted or grafted on dwarfing rootstock. Within-tree shoot demographics and annual shoot characteristics, i.e. their dimensions, number of laterals and flowering, were observed from the first to the fifth year of growth and compared among scion/root system combinations. KEY RESULTS: Differences in axis demographics appeared among scion/root system combinations after the second year of growth. Differences were found (a) in the number of long axes and (b) the number of medium axes. Dwarfing rootstock reduced the total number of axes developed in a tree, and this reduction resulted from proportionally more medium axes and spurs than long axes. The life span of spurs was also shortened. These phenomena appeared after an increase in flowering that started in the second year of growth and involved both axillary and terminal positions. Flowering regularity was also increased in grafted trees. CONCLUSIONS: These results confirm that the number of long shoots and flowering potential depend on the cultivar. They indicate that tree architectural plasticity in response to its root system mainly derives from the number of medium shoots developed and follows priorities within the whole tree axis population. There was also evidence for dwarfing rootstock involvement in adjusting the flowering abundance and that differences in flowering occurrence take precedence over those regarding vegetative growth during tree development.     

Similarities and gradients in growth unit branching patterns during ontogeny in "Fuji" apple trees: a stochastic approach

This study aims to explore and model the changes in growth unit (GU) branching patterns during tree ontogeny. The question was addressed in apple trees cv. "Fuji", by analysing the relative impact of GU length and within-tree position. The development of two 6-year-old trees was recorded over 6 years. The fate of axillary buds along each GU was represented as a sequence of symbols corresponding to five types of lateral growth: latent buds, short, medium, long, and floral lateral GUs. Based on an exploratory analysis of data and a priori hypotheses, a hidden semi-Markov chain was estimated from all of these GU sequences. This model was composed of six transient states representing successive branching zones along the GUs. The accuracy of this global model was a posteriori assessed by fitting the characteristic distributions computed from model parameters to the corresponding empirical characteristic distributions extracted from the observed sequences. The observed sequences were then grouped hierarchically according to the GU length, year of growth, and branching order. Comparing model parameters between these sub-groups revealed similarities between GUs. These similarities were based on particular branching zones whose composition and relative position within the GUs remained invariant across the subgroups: the latent zones, floral zone, and short-lateral zone. The probability of occurrence of the floral zone varied with the year, showing the alternate fruiting of "Fuji". It is shown that, during tree ontogeny, as GU length decreases, branching patterns tend to progressively simplify due to the disappearance of the most central zones and a progressive reduction in the length of the floral zone.     

Deciphering the ontogeny of a sympodial tree

This paper addresses the identification and characterization of developmental patterns in the whole structure of a sympodial species, the apple tree. Dedicated stochastic models (hidden variable-order Markov chains) were used to (i) categorise growth units (GUs) on the basis of their morphological characteristics (number of nodes and presence/absence of flowering) and position along axes, (ii) analyse dependencies between successive GUs and (iii) identify repeated patterns in GU sequences. Two successive phases, referred to as “adolescent” and “adult”, were identified in two apple tree cultivars. In the adolescent phase, “very” long monocyclic GUs were followed by long polycyclic GUs, whereas in the adult phase medium GUs were preferentially followed by short GUs. Flowering GUs constituted a preferential pathway between vegetative GUs of decreasing vigour (long, medium and short) and generated patterns that were interpreted with respect to fruiting regularity. The proposed modelling gave a global and quantitative picture of the two-scale structuring of apple tree ontogeny: a coarse scale corresponding to the succession of the previously mentioned phases and a fine scale corresponding to the alternation between flowering and vegetative GUs. This led us to propose a synthetic scheme of apple tree ontogeny that combines growth phases, polycyclism and flowering, and which could be transposed to other sympodial trees.     

Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees (Olea europaea L.)

The development of new shoots plays a central role in the complex interactions determining vegetative and reproductive growth in woody plants. To explore this role we evaluated the new shoots in the olive tree, Olea europaea L., and the effect of fruiting on new shoot growth and subsequent flowering. Five-year-old branches served as canopy subunits in order to obtain a global, whole-tree view of new shoot number, size and morphological origin. The non-bearing trees had many more shoots than the fruit-bearing trees, and a greater number of longer shoots. In both bearing conditions, however, the majority of shoots were less than 4 cm long, with shoots of progressively longer lengths present in successively decreasing frequencies. Six major shoot types were defined on the basis of apical or lateral bud origin and of parent shoot age. On fruit-bearing trees, the new shoots originated predominantly from the shoot apex, while on non-fruiting trees, they formed mainly from axillary buds, but in both cases, they tended to develop on younger parent shoots. The previous bearing condition of the tree was the main determinant for subsequent inflorescence development, which was independent of both shoot type and length. Thus, reproductive behavior strongly affected both the amount and type of new branching, but subsequent flowering level was more influenced by previous bearing than by the potential flowering sites on new shoots.     

Genetic determinism of the vegetative and reproductive traits in an F1 olive tree progeny

The agronomic performance of fruit trees is significantly influenced by tree internal organization. Introducing architectural traits in breeding programs could thus lead to select new varieties with a regular bearing and lower input demand in order to reduce training and environmental costs. However, an interaction between tree ontogeny and genetic factors is expected. In this study, we investigated the genetic determinism of architectural traits in the olive tree, accounting for tree development over 5 years until first flowering occurrence. We studied an F1 progeny issued from a cross between two contrasted genotypes, ‘Olivière’ and ‘Arbequina’. Tree architecture was decomposed in quantitative traits, related to (1) growth and branching, (2) first flowering and fruiting. Models, including the year of growth, branching order and genotype effects, were built with variance function and covariance structure when necessary. After a model selection, broad sense heritabilities were calculated. During the first 3 years, both the mean values of vegetative traits and genetic factor significance depended on the shoot within-tree position. Dependencies between consecutive years were revealed for traits related to whole tree form. Whole tree form variables showed medium to high broad sense heritability values, whereas reproductive traits were highly heritable. This study demonstrates the existence of ontogenic trends in the olive tree, which result in traits heritable only at the tree periphery. A phenotyping strategy adapted to its architectural characteristics and a list of relevant traits, such as maximal internode length, is proposed. Transgressive effects suggest that genetic progress could be performed in future selection programs.     

Growth Dynamics and Morphology of Annual Shoots, According to their Architectural Position, in Young Cedrus atlantica (Endl.) Manetti ex Carriere (Pinaceae)

The influence of shoot architectural position on growth andbranching pattern of young Cedrus atlantica (Endl.) Manettiex Carrière trees were studied. Extension growth andtype of axillary products (lateral bud, sylleptic short or longshoots) of annual shoots of increasing branching order (mainstem, branches and branchlets) were recorded weekly during the1993 growing season. Annual final shoot length, duration ofextension, and maximum extension rate decreased with increasingbranching order. Sylleptic axillary shoots occurred only onannual shoots of the main stem and branches and were producedwhen extension rate was at its highest. Differences in growthrate and final length of annual shoots, according to their architecturalposition, were related to differences in the total number anddiversity of types of sylleptic axillary shoots produced. Itis suggested that types and numbers of sylleptic axillary shootsproduced are linked with threshold values for both final lengthand extension rate of the parent shoot. Copyright 1999 Annalsof Botany Company Atlas cedar, extension growth, sylleptic branching, tree architecture, morphology.     

Architecture of the pruned tree: impact of contrasted pruning procedures over 2 years on shoot demography and spatial distribution of leaf area in apple (Malus domestica)

BACKGROUND AND AIMS: Demography and spatial distribution of shoots are rarely studied on pruned trees. The present 2-year study deals with the effect of pruning strategies on shoot demography and development, and consequences on the spatial distribution of leaf area in three architecturally contrasted - from type II to IV - apple cultivars: 'Scarletspur Delicious', 'Golden Delicious' and 'Granny Smith'. METHODS: All trees were initially subjected during 5 years to Central Leader training with winter heading on all long shoots. For 2 years, half of the trees were further trained with Centrifugal training, where removal of flowering shoots - called extinction pruning - was carried out along the trunk and at the bottom of branches at flowering time. During these 2 years, shoot type (vegetative, inflorescence) and length, and the three-dimensional spatial distribution of all shoots were assessed with an electromagnetic digitizer. KEY RESULTS: Shoot demography, frequency of transitions toward an inflorescence from either an inflorescence (bourse-over-bourse) or a vegetative shoot (trend toward flowering), and the number of bourse-shoots per bourse were strongly affected by cultivar, with little influence of tree manipulation. In contrast, the proportion of vegetative long shoots developing from previous year latent buds was significantly lower in Centrifugal-trained trees for the three cultivars. Canopy volume showed large variations between cultivars, but only that of 'Granny Smith' was affected by tree manipulation in the 2 years. Spatial distribution of shoots varied significantly according to cultivar and manipulation. In 'Scarletspur Delicious' and, to a lesser extent 'Golden Delicious', the distribution of vegetative and flowering shoots in the outer and the inner parts, respectively, was not affected by tree manipulation. In contrast, in 'Granny Smith', vegetative shoots were stimulated in the periphery of Central Leader trees, whereas flowering shoots were stimulated in the periphery of Centrifugal-trained trees. CONCLUSIONS: In apple, the variability of responses to contrasted pruning strategies partly depends on the genetically determined growth and flowering habit of the cultivar.     

In Loquat (Eriobotrya japonica Lindl.) Return Bloom Depends on the Time the Fruit Remains on the Tree

In loquat (Eriobotrya japonica Lindl.), the comparison of fruiting trees and defruited trees carried out covering a range of developmental fruit stages reveals a significant reduction in flowering due to fruit from its early stage of development, being higher when it changes color and becomes senescent, which coincides with the floral bud inductive period. This effect occurred both at the tree and at the shoot level. Furthermore, although current shoots almost always develop into panicles, those from fruiting trees develop fewer flowers, suggesting that fruit also affects at the floral bud level. In our experiment, the gibberellin concentration at the floral bud inductive period was significantly higher in bark tissues (periderm, cortex and phloem tissues) of fruiting trees, compared with defruited trees that tend to flower more. The lower concentration of IAA in the bark tissues of defruited trees also contributes to increase their flowering intensity. On the contrary, the zeatin concentration was higher. Accordingly, at bud burst, the IAA/zeatin ratio, an indication of effect on flowering, was significantly higher for fruiting trees. Some disruption in the nitrate reduction process in fruiting tree was also observed. The process of floral bud induction and differentiation was not associated with either reducing or translocating and reserve carbohydrate concentration. Hence, loquat flower intensity depends on the time the fruit is maintained on the tree. The intensity is affected indirectly, by reducing the number of shoots, and directly, by reducing the number of flowers per panicle, and these effects are linked to endogenous plant hormone contents.     

The ecology of tree reproduction in an African medium altitude rain forest

The occurrence of flowering and fruiting in tropical trees will be affected by a variety of factors, linked to availability of resources and suitable climatic triggers, that may be affected by increasing global temperatures. Community‐wide flowering and fruiting of 2526 trees in 206 plots were monitored over 24 years in the Budongo Forest Reserve (BFR), Uganda. Factors that were assessed included the size of the tree, access to light, the impacts of liana load, effects of tree growth, and variation between guilds of trees. Most flowering occurs at the end of the long dry season from February to April. Trees that had access to more light flowered and fruited more frequently. Pioneer and non‐pioneer light‐demanding species tended to reproduce more frequently than shade‐bearing species. Trees that grew faster between 1993 and 2011 also fruited more frequently. When examining all factors, growth rate, tree size, and crown position were all important for fruiting, while liana load but not growth rate was important in reducing flowering. Trees in BFR show a large decline in fruiting over 24 years, particularly in non‐pioneer light demanders, shade‐bearers, and species that produce fleshy fruits eaten by primates. The decline in fruit production is of concern and is having impacts on primate diets and potential recruitment of mahogany trees. Whether climate change is responsible is unclear, but flowering of the guilds/dispersal types which show declines is correlated with months with the coolest maximum temperatures and we show temperature has been increasing in BFR since the early 1990s.     

Stomatal regulation of photosynthesis in apple leaves: evidence for different water-use strategies between two cultivars

BACKGROUND AND AIMS: Leaf responses to environmental conditions have been frequently described in fruit trees, but differences among cultivars have received little attention. This study shows that parameters of Farquhar's photosynthesis and Jarvis' stomatal conductance models differed between two apple cultivars, and examines the consequences of these differences for leaf water use efficiency. METHODS: Leaf stomatal conductance (g(sw)), net CO2 assimilation rate (A(n)), respiration (R(d)) and transpiration (E) were measured during summer in 8-year-old 'Braeburn' and 'Fuji' apple trees under well-watered field conditions. Parameters of Farquhar's and Jarvis' models were estimated, evaluated and then compared between cultivars. Leaf carbon isotope discrimination (delta(13)C) was measured at the end of the growing season. KEY RESULTS: A single positive relationship was established between V(Cmax) (maximum carboxylation rate) and N(a) (leaf nitrogen concentration per unit area), and between J(max) (maximum light-driven electron transport rate) and N(a). A higher leaf R(d) was observed in 'Fuji'. The g(sw) responded similarly to increasing irradiance and leaf temperature in both cultivars. g(sw) responded to lower vapour pressure deficit in 'Fuji' than in 'Braeburn'. Maximal conductance (g(swmax)) was significantly smaller and A(n) was more limited by g(sw) in 'Braeburn' than 'Fuji'. Lower g(sw), E and higher intrinsic water use efficiency were shown in 'Braeburn' and confirmed by smaller leaf delta(13)C compared with 'Fuji' leaves. CONCLUSIONS: The use of functional model parameters allowed comparison of the two cultivars and provided evidence of different water use 'strategies': 'Braeburn' was more conservative in water use than 'Fuji', due to stomatal limitation of A(n), higher intrinsic water use efficiency and lower delta(13)C. These physiological traits need to be considered in relation to climate adaptation, breeding of new cultivars and horticultural practice.     

Influence of elevated CO2 and O3 on Betula pendula Roth crown structure

Elevated CO(2) and ozone effects were studied singly and in combination on the crown structure of two Betula pendula clones. Measurements were made at the end of the second fumigation period in an open-top-chamber experiment with 9-year-old trees. Shoot ramification (number of long and short daughter shoots), shoot length, and number of metamers, leaves and buds were measured at four positions in every tree. As a result of increased temperature, trees in chambers had longer shoots and more frequent shoot ramification than control trees not enclosed in chambers. Ozone treatment decreased shoot ramification significantly. Additionally, ozone treatment resulted in an increased number of metamers in one clone. There was no statistically significant interaction between ozone effect and crown position; however, there was a slight tendency for the lower crown to be more affected by ozone. Elevated CO(2) caused a significant increase in the number of long-shoot metamers. Therefore, 2x ambient CO(2) concentration partly ameliorated the negative effect of ozone because the increased number of leaves per shoot counteracted the decreased branching. Although the main effects of elevated ozone and CO(2) were similar in the two clones, slight, statistically insignificant, differences appeared in their responses when interactions with crown position were considered.     

Relationship between Sylleptic Branching and Components of Parent Shoot Development in the Peach Tree

Sylleptic branching of main axes was investigated in three peachtree cultivars ('Armking', 'Flavorcrest' and 'Silvergem') duringthe first year of growth. An axis was considered as made upof a series of metamers (internode, node, leaf and associatedbud) and its growth was divided into two components: the increaseof the number of metamers and the lengthening of the metamersthemselves (elongation). The relationship between branchingprobability, type of cultivar, growth components and date ofmetamer production was studied by logistic regression. Branchingprobability varied according to the cultivar, increased whenthe growth components did, and decreased if metamers appearedlate in the season. The logistic model fitted the data closelyand was validated on a data set that had not been used for estimatingthe parameters. Ninety-four percent of branched and 70% of unbranchedmetamers were correctly predicted by the logistic model. Forany given growth rate and date of metamer production, the mainaxes branched most and least often in the 'Flavorcrest' and'Silvergem' cultivars, respectively.Copyright 1994, 1999 AcademicPress Peach tree, Prunus persica (L.) Batsch, branching, syllepsis, shoot growth, quantitative analysis, logistic regression     

几个板栗株系的开花结果习性比较分析

采用在美国育成的几个株系,并以常见的本地栽培品种‘桂花香’作对照,对刚进入结果幼树的枝梢开花习性,枝梢生长情况与开花坐果关系,以及树冠不同部位的枝梢开花与坐果进行调查分析。结果表明：对全部参试材料而言,末级梢的长、粗度区间分别以20．0～59．9cm和0．5～1．49cm的雌花量和坐果数较高;株系95—1—26和95—1-9雌花着生部位较其它3个株系或品种的节位高出一倍以上,其平均着生节位与末级梢平均节位基本上相等;从树冠立体分布上看,其外围和树冠上部水平方向中膛结果数较多,但对于每个株系来说,又存在一定的差异。以上研究结果对幼年结果树的枝梢培养、树体的整形修剪,以及达到早期丰产具有实践指导意义。     

Architecture and size relations: an essay on the apple (Malus x domestica, Rosaceae) tree

The influence of tree size independent of age on some architectural features (annual shoot length, lateral branching, flowering) was investigated on 4-yr-old apple (Malus × domestica) trees either own-rooted or grafted on the dwarfing rootstock M.9, giving rise to large and small trees, respectively. Tree size significantly affected the length of the first annual shoot of bottom branches with a lesser effect on the subsequent annual shoots of the same branches and on branches situated higher in the tree canopy. The linear regression parameters, i.e., slopes and intercepts, between annual shoot length and number of growing laterals were affected by the genotype and, depending on genotype, by tree size. Flowering was generally lower, delayed, and more irregular on large trees compared to small trees, with on average similar ranking of genotypes regardless of tree size. This study provides evidence for a specific effect of tree size, as affected by the root system, on architectural development of the apple tree regardless of the genotype. From an architectural viewpoint, the dwarfing mechanism could be interpreted as a faster physiological aging essentially related to the reduction in length of the first annual shoot of bottom branches and the high flowering on this shoot.     

Within-tree and among-tree variation in growth of Epirrita autumnata on mountain birch leaves

Abstract.

• 1 We studied among-tree and within-tree variation in the growth of larvae of a geometrid, Epirrita autumnata, on mountain birch (Betula pubescens ssp. tortuosa) leaves at different hierarchical levels: among trees (genets), among ramets within trees, among branches within ramets, among shoots within branches and among leaves within shoots. We used only short shoot leaves, which burst simultaneously in spring.
• 2 Trees accounted for most of the variation in larval growth rate, but there was substantial variation also among ramets within trees, among branches within ramets, and among short shoots within branches. Variation among leaves within short shoots was negligible. When the probabilities from different experiments were combined, the differences were statistically significant at the tree, ramet and branch levels, and approached significance at the shoot level.
• 3 In different experiments, larval growth was from 9% to 54% lower on the worst tree than on the best tree of the experiment. On average, larval growth rate was in different experiments from 11% to 32% lower on the worst ramet than on the best ramet within tree, from 8% to 18% lower on the worst than on the best branch within ramet, and from 12% to 30% lower on the worst than on the best shoot within branch.
• 4 The amount of among-and within-tree variation shown by our results may have ecological and evolutionary implications: among-tree variation should select for discrimination by ovipositing females and dispersing larvae, within-tree variation should select also for optimal foraging behaviour of larvae.

     

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.     

Performance of trees in forest canopies: explorations with a bottom-up functional-structural plant growth model

Here we present a functional-structural plant model that integrates the growth of metamers into a growing, three-dimensional tree structure, and study the effects of different constraints and strategies on tree performance in different canopies. The tree is a three-dimensional system of connected metamers, and growth is defined by the flush probability of metamers. Tree growth was simulated for different canopy light environments. The result suggest that: the constraints result in an exponential, logistic and decay phase; a mono-layered-leaf crown results from self-shading in a closed canopy; a strong apical control results in slender trees like tall stature species; the interaction between weak apical control and light response results in a crown architecture and performance known from short stature species in closed forest; correlated leaf traits explain interspecific differences in growth, survival and adult stature. The model successfully unravels the interaction effects of different constraints and strategies on tree growth in different canopy light environments.     

Morphological and architectural features of young Nothofagus pumilio (Poepp. & Endl.) Krasser (Fagaceae)

The morphology and architecture of young Nothofagus pumilio plants growing as understorey and in a cleared area were assessed. Length, number of internodes, presence of branches and apex condition were registered for each growth unit (GU) in the sampled plant. GUs were grouped according to branching order and age of the bearing axis. Understorey plants were seven years old and had a short trunk bearing 2–3 orders of branches; all of their most recent GUs were horizontal and of comparable size. Plants growing in the cleared area were 10–16 years old and had a vigorous vertical and dominant trunk bearing 4–5 orders of less vigorous, horizontal branches. Plants from the cleared area showed signs of having had comparable-size GUs in all of their axes at a younger stage of growth. N. pumilio plants appear to start their growth with the production of short GUs in all of their axes irrespective of the light conditions. In contrast, the development of a vertical, vigorous trunk, seems to be restricted to plants growing in open environments.