Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Genetic control of biennial bearing in apple

Although flowering in mature fruit trees is recurrent, floral induction can be strongly inhibited by concurrent fruiting, leading to a pattern of irregular fruiting across consecutive years referred to as biennial bearing. The genetic determinants of biennial bearing in apple were investigated using the 114 flowering individuals from an F(1) population of 122 genotypes, from a 'Starkrimson' (strong biennial bearer)×'Granny Smith' (regular bearer) cross. The number of inflorescences, and the number and the mass of harvested fruit were recorded over 6 years and used to calculate 26 variables and indices quantifying yield, precocity of production, and biennial bearing. Inflorescence traits exhibited the highest genotypic effect, and three quantitative trait loci (QTLs) on linkage group (LG) 4, LG8, and LG10 explained 50% of the phenotypic variability for biennial bearing. Apple orthologues of flowering and hormone-related genes were retrieved from the whole-genome assembly of 'Golden Delicious' and their position was compared with QTLs. Four main genomic regions that contain floral integrator genes, meristem identity genes, and gibberellin oxidase genes co-located with QTLs. The results indicated that flowering genes are less likely to be responsible for biennial bearing than hormone-related genes. New hypotheses for the control of biennial bearing emerged from QTL and candidate gene co-locations and suggest the involvement of different physiological processes such as the regulation of flowering genes by hormones. The correlation between tree architecture and biennial bearing is also discussed.     

Effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems

The effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems were examined using trees growing in lysimeters. For 3-year-old 'Golden Delicious' trees, cropping significantly increased total biomass, although leaf and root biomass were reduced. For 3-year-old'Golden Delicious', 'Coxs Orange' and 'Gloster' trees, cropping only in the third year reduced root biomass by 39 to 45%. Cropping did not affect water use by trees of any cultivar early in the season, but increased water use later in the season for 'Gloster' trees. For trees with fruit, total non-structural carbohydrate (TNC) contents at the end of the season were highest in fruit (up to 1 400 g plant ⁻¹). For trees without fruit, TNC contents were generally highest in the roots (up to 110 g planr ⁻¹). Supplying trees with nutrient solution diluted to 10% of the standard composition increased the root biomass for 'Golden Delicious' trees only, but decreased biomass of above-ground tissues for all cultivars. Reducing the nutrient supply decreased water and nutrient uptake, and reduced nutrient contents within vegetative parts of the tree more than within roots. For 'Golden Delicious' trees, restricting the water supply to 50% or 25% of that consumed by control trees significantly reduced above-ground biomass, but root biomass was not significantly affected. The N, P and K. contents for the trees were also reduced by water stress, due to reduced contents within above-ground organs. Water stress reduced the TNC and starch contents of all tissues, except the roots. These results are discussed in relation to the efficiency of root systems for water and nutrient uptake.     

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.     

Effects of nitrogen supply on growth, contents of phenolic compounds and pathogen (scab) resistance of apple trees

The effect of long-term N-supply on growth, scab resistance and phenolic compounds in the leaves of two apple cultivars was studied. The different pools of phenylpropanoids (hydroxycinnamic acids, dihydrochalcones) and flavonoids (flavonols, catechins, procyanidins) were quanitfied by HPLC from non-infected and inoculated leaves representing different ontogenetic stages. Scab incidence was also evaluated. Strictly following the carbon-nutrient-balance hypothesis, apple trees responded to high N-supply with increased shoot growth and with a reduced accumulation of total phenolic compounds in their leaves. This was shown for the cultivar 'Golden Delicious', which is susceptible to the scab disease caused by Venturia inaequalis , and for the resistant cultivar 'Rewena'. Whereas high N-fertilization increased the susceptibility of 'Golden Delicious', it did not decrease the resistance of 'Rewena' despite of the pronounced reduction of phenolic concentrations. Thus, a simple C trade off between growth-related metabolism and secondary metabolism cannot solely explain changes in defensive potential.     

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.     

Current-year and subsequent-year effects of crop-load manipulation and epicormic-shoot removal on distribution of long, short and epicormic shoot growth in Prunus persica

BACKGROUND AND AIMS: The distribution of canopy growth among different shoot types such as epicormic, long and short shoots is not well understood in the peach tree. In this experiment, the effects of crop load and early epicormic sprout removal on current and subsequent-year distribution of vegetative growth among epicormic, long and short shoots was investigated in Prunus persica. METHODS: Field trials were conducted in Winters, California, in 2003-2004. Crop load was manipulated with fruit thinning in 2003 to produce trees that were de-fruited, commercially thinned or full crop, and half of the trees in each cropping treatment had all current year epicormic sprouts removed at the time of fruit thinning. Yield was recorded and trunk and root carbohydrates were sampled to confirm the effect of 2003 crop load differences on tissue carbohydrate concentration. All current-season vegetative-shoot extension growth was harvested from half of the trees in each treatment in the autumn of 2003 and from the other half in the autumn of 2004. Epicormic, long and short shoots were separately evaluated for dry weight, node number and leaf-stem parameters. KEY RESULTS: In 2003, long-shoot dry weight and node number were significantly affected by crop load; however, short-shoot dry weight and node number were not significantly affected. The 2003 crop-load treatments did not affect 2004 vegetative growth of any shoot type. Some re-growth of epicormic shoots followed early epicormic sprout removal: by the end of the 2003 season, trees in the early shoot-removal treatment had approximately one-third of the epicormic-shoot dry weight as unpruned trees. CONCLUSIONS: Fruit thinning promoted distribution of growth similar to that of de-fruited trees. While thinning was effective in increasing fruit size, it exacerbated the problem of epicormic sprouting. Early epicormic sprout removal did not stimulate the excessive epicormic re-growth in the same or subsequent year relative to previously studied summer pruning methods.     

Genetic interactions in the control of flowering time and reproductive structure development in tomato (Solanum lycopersicum)

Different tomato (Solanum lycopersicum) mutants, affected in flowering time, reproductive structure or plant architecture, were crossed to produce double mutants in order to investigate gene interactions in flowering regulation in this autonomous species with a sympodial growth habit. The compound inflorescence: uniflora, uniflora: self pruning, uniflora: blind, and jointless: uniflora double mutants all produced solitary flowers like their uniflora parent, instead of inflorescences. All double mutants were late flowering. uniflora: blind and uniflora: self pruning had flowering times intermediate between those of their two parents. jointless: uniflora and compound inflorescence: uniflora flowered later than uniflora, the mutant with the most delayed flowering. All double mutants developed strong lateral shoots at node levels approximately corresponding to the level at which their parent cultivars initiated their first reproductive structure, which is a typical trait of uniflora. These results suggest that the UNIFLORA gene acts upstream of the other investigated genes in controlling flowering in tomato, and that floral transition of the primary shoot and floral transition of sympodial segments are regulated differently.     

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。     

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

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

Cost and probability of flowering at the shoot level in relation to variability in shoot size within the crown of Vaccinium hirtum (Ericaceae)

Size-related variation in the cost and probability of flowering among shoots within a crown of Vaccinium hirtum was investigated to clarify patterns and regulation of flowering at the shoot-module level, below the level of the individual. The apices of previous-year shoots differentiated into current-year shoots vegetatively (vegetative branches) or became reproductive by developing inflorescences (reproductive branches). Length growth and fate of current-year shoots were determined, and the future potential for reproduction was estimated using a matrix model of shoot dynamics. Reproductive branches had fewer current-year shoots and shorter total shoot lengths and thus had a reduced potential for reproduction compared with vegetative branches, indicating the cost of flowering at the shoot level. This cost of flowering was higher in longer shoots. The probability of the initiation of flowering in a shoot increased with increasing shoot length in shorter shoots, reached a maximum in medium-sized shoots, and decreased in longer shoots. The size-related changes in the probability of flowering at the shoot level can be largely explained by the size-dependent changes in shoot-level resource availability and cost of flowering.     

A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005     

Apple dwarfing rootstocks and interstocks affect the type of growth units produced during the annual growth cycle: precocious transition to flowering affects the composition and vigour of annual shoots

Background and Aims: Precocious flowering in apple trees is often associated witha smaller tree size. The hypothesis was tested that floral evocationin axillary buds, induced by dwarfing rootstocks, reduces thevigour of annual shoots developing from these buds comparedwith shoots developing from vegetative buds. Methods: The experimental system provided a wide range of possible treevigour using ‘Royal Gala’ scions and M.9 (dwarfing)and MM.106 (non-dwarfing) as rootstocks and interstocks. Second-yearannual shoots were divided into growth units corresponding toperiods (flushes) of growth namely, vegetative spur, extensiongrowth unit, uninterrupted growth unit, floral growth unit (bourse)and extended bourse. The differences between the floral andvegetative shoots were quantified by the constituent growthunits produced. Key Results: The dwarfing influence was expressed, firstly, in reduced proportionsof shoots that contained at least one extension growth unitand secondly, in reduced proportions of bicyclic shoots (containingtwo extension growth units) and shoots with an uninterruptedgrowth unit. In treatments where floral shoots were present,they were markedly less vigorous than vegetative shoots withrespect to both measures. In treatments with M.9 rootstock,vegetative and floral shoots produced on average 0·52and 0·17 extension growth units, compared with 0·77extension growth units per shoot in the MM.106 rootstock treatment.Remarkably, the number of nodes per extension growth unit wasnot affected by the rootstock/interstock treatments. Conclusions: These results showed that rootstocks/interstocks affect thetype of growth units produced during the annual growth cycle,reducing the number of extension growth units, thus affectingthe composition and vigour of annual shoots. This effect isparticularly amplified by the transition to flowering inducedby dwarfing rootstocks. The division of annual shoot into growthunits will also be useful for measuring and modelling effectsof age on apple tree architecture.     

A Simple Model to Predict the Probability of a Peach (Prunus persicae) Tree Bud to Develop as a Long or Short Shoot as a Consequence of Winter Pruning Intensity and Previous Year Growth

In many woody plants, shoots emerging from buds can develop as short or long shoots. The probability of a bud to develop as a long or short shoot relies upon genetic, environmental and management factors and controlling it is an important issue in commercial orchard. We use peach (Prunus persicae) trees, subjected to different winter pruning levels and monitored for two years, to develop and calibrate a model linking the probability of a bud to develop as a long shoot to winter pruning intensity and previous year vegetative growth. Eventually we show how our model can be used to adjust pruning intensity to obtain a desired proportion of long and short shoots.     

Light interception in apple trees influenced by canopy architecture manipulation

Improvement of light penetration within tree canopies has been a constant objective of fruit tree architecture manipulation through the setting up of training systems. Recently, centrifugal training, i.e. the removal of fruiting shoots in the tree centre and on the underside of branches, has been proposed to improve fruit size and colour as well as return-bloom as compared to conventional solaxe-trained trees with equivalent crop loads. The present study was conducted to quantify the benefits of centrifugal training on light interception by the fruiting shoots via computer-assisted three-dimensional representations of foliage geometry. Data were collected on six 5-year-old apple trees cv.Galaxy, trained either with solaxe or centrifugal training systems, using an electromagnetic 3D digitiser. The 3D distribution of the foliage in the tree canopy was recreated by combining both the spatial locations of shoots (as measured from 3D digitising) and foliage reconstruction. Light interception efficiency properties of the trees were characterised by silhouette to total area ratio (STAR) values computed from images of the 3D mock-ups. Compared to the solaxe system, centrifugal training significantly improved the STAR of the whole tree by 20%. It also increased both leaf area and STAR of the fruiting shoots by approximately 15%, regardless of their position in the canopy. In this paper, we discuss the role of this enhanced light interception by the canopy in increasing the autonomy of the fruiting shoot, i.e. improved fruit size and colour, and return-bloom.     

How does winter pruning affect peach tree–Myzus persicae interactions?

Winter tree pruning is a cultural practice known to modify vegetative growth, which is likely to affect the development of pests. However, it has been poorly addressed as a cultural control method for diminishing the population levels of the green peach aphid, Myzus persicae (Sulzer) (Homoptera: Aphididae), in peach [ Prunus persica (L.) Batsch (Rosaceae)] orchards. In this study, we conducted a 2-year, on-station experiment to evaluate how winter pruning affects peach– M. persicae interactions, by examining tree vegetative growth, aphid population dynamics, and crop yield and fruit quality. We collected data under an insect-proof shelter on adult peach trees submitted to various levels of pruning and artificially infested with aphids. Our results showed that pruning enhanced shoot growth due to the proportion of growing shoots, which increased exponentially (10–60%), whereas the growth rate of growing shoots was not affected. The degree of infestation of peach trees increased with increasing pruning intensity. This effect was mainly due to the increase of the proportion of growing shoots, on which aphids developed better than on rosettes. In turn, the higher the aphid infestation, the higher the aphid-induced shoot-tip damage, leaf curling, and leaf fall that disturbed the growth of growing shoots. However, aphids did not considerably reduce fruit quality at harvest. They did not affect fresh fruit weight, and the refractometric index (indicator of sugar content) was reduced by only 3–4%. The relevance of winter pruning as a cultural method for pest control in orchards conducted under integrated fruit production guidelines is discussed.     

Behavior of current-year shoots as a mechanism to determine the floral sex allocation at the level of individual tree and population in Siberian alder (Alnus hirsuta var. sibirica)

Floral sex allocation (weight of male flower buds over weight of female flower buds) was examined at the levels of current-year shoot, individual tree and population, and the tree individual level and population level floral sex ratio was explained as a consequence of the behavior of current-year shoots in the shoot-level monoecious (flowering current-year shoots have both male and female flowers) species, Siberian alder (Alnus hirsuta var. sibirica). The current-year shoot level floral sex allocation was not size-dependent and not different over years. However, in the year when the reproductive intensity was high, individual tree level floral sex allocation was size-dependent and the population level floral sex allocation was relatively female-biased. The female-biased floral sex allocation at the population level resulted from many gynoecious shoots (current-year shoots which have only female flowers). These results suggest that the floral sex allocation of Siberian alder was controlled not by changing the floral sex allocation of each current-year shoot, but by shifting the sex expression of current-year shoots from shoot-level monoecy to shoot-level gynomonoecy.     

Endophytic and epiphytic phyllosphere fungi of red-osier dogwood (<Emphasis Type="Italic">Cornus stolonifera</Emphasis>) in British Columbia

Endophytic and epiphytic phyllosphere fungi associated with red-osier dogwood (Cornus stolonifera), a deciduous shrub, were examined in coastal British Columbia, Canada. Current-year shoots were divided into four types based on the absence or presence of inflorescence and secondary elongated shoots at the apex of primary shoots. Leaves on these shoots were then classified into six categories so as to examine the effect of flowering, secondary shoot elongation, and shoot order within current-year shoots on the occurrence of phyllosphere fungi. Species composition of fungi was markedly different between the interior and surface of leaves, whereas it was relatively similar among the six leaf categories in the interior or on the surface. Frequencies of the eight major species were not different between leaves on flowering and nonflowering shoots. The frequency of Colletotrichum gloeosporioides in the leaf interior was greater on leaves on the primary shoots that elongated the secondary shoots than on those that did not, and was greater on leaves on the primary shoots than on those on the secondary shoots. On the other hand, secondary shoot elongation and shoot order had no effect on the frequencies of C. gloeosporioides and the other seven epiphytes on leaf surfaces.