Pierre-Eric Lauri

Genotypic differences in the axillary bud growth and fruiting pattern of apple fruiting branches over several years—an approach to regulation of fruit bearing

Abstract Genotypic variations in growth and fruiting patterns were investigated in various apple ( Malus × dormestica Borkh.) cultivars belonging to different ideotypes by analysis of chains of functions of laterals on fruiting branches. Yearly changes in each axillary bud (latent, vegetative, non-bearing or bearing inflorescence and scar) were described over 5 or 6 years, depending on the cultivar. Sequences obtained for the various cultivars were analysed through calculations of three indices: fruit-richness (ϱ), alternation-to-fruit (α) and, among the alternating subsequences (i.e. bourse-shoot without a terminal fruit), alternation synchronism (σ). The results showed that genotypes could be distinguished by two sets of characters. Some cultivars, such as ‘Red Winter’, balance vegetative and reproductive growth by reducing the number of growing points through an extinction mechanism (essentially the abortion of potential bourse-shoots on non-bearing inflorescences), and through a high functional autonomy (low α and high ϱ values) of the remaining sequences, corresponding to high bourse-over-bourse potentials. Spurred cultivars, such as ‘Oregon Spur Delicious’, were characterized by a low extinction value and, in a complementary way, did not seem able to fruit terminally on bourse-shoots (high α and low ϱ values). The overall results of the study of σ showed that the alternation-to-fruit pattern (high α values) could be completely hidden when alternating sequences are desynchronized (low σ values). The links between extinction and alternation-to-fruit, as well as the physiological significance of alternation synchronism, are discussed.

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.

Does manipulation of fruit-tree architecture affect the development of pests and pathogens? A case study in an organic apple orchard

Summary Centrifugal training, based on the extinction procedure, is an innovative type of apple tree training recently used in French apple orchards. A thinning cut of fruiting spurs along the trunk and on the proximal and underside parts of the fruiting branches modifies the within-tree leaf density and light environment, which is likely to affect the development of some apple pests and pathogens. In this study, development of the most serious foliar pests (aphids, mites) and disease (apple scab) was assessed over 3 years in an organic apple orchard with two training systems: the centrifugal training (CT) system, and the Original Solaxe (OS) system, using a bent axis with no removal of fruiting spurs, as a control. Infestation levels were significantly lower in 2002 and 2004 in the CT system than in the OS system for the most prevalent pest, the rosy apple aphid, Dysaphis plantaginea and, in 2003, for the red spider mite, Panonychus ulmi. Infestation by the green apple aphid, Aphis pomi, was higher in June 2002 and 2004 and lower in June 2003 in the CT system, compared to the OS control. The incidence of apple leaf scab was lower in the CT system than in the OS control in Spring 2002 and Spring 2004; but, later in the Summer, it increased more in CT trees, resulting in no difference between training systems in the levels of fruit infection at harvest. Hypotheses considered were based mainly on: (i) the removal of pests or inoculum due to the thinning cut of fruiting spurs for CT trees; (ii) effects on within-tree microclimate, providing a more aerated canopy in the CT system; (iii) shoot density and distribution, allowing greater distances between growing shoots in CT trees; and/or (iv) the shoot growth dynamics, allowing the presence of attractive or susceptible leaves in late June. Since the most serious apple pests and diseases were less prevalent in the CT system over the 3 years of the experiment, CT may prove to be a complementary, sustainable means to contribute to crop protection, especially in organic or integrated pest management orchards.

Hydraulic and mechanical stem properties affect leaf-stem allometry in mango cultivars.

Leaf size-stem size allometric relationships are important features of biomass allocation in plants and are affected by biological functions linking the two organs. They have been studied at specific and supraspecific levels, but not at the infraspecific level. It was hypothesized that allometric relationships link leaf size and stem size at the cultivar level, and are cultivar-specific in relation to distinctive functional stem traits: hydraulic conductivity and mechanical strength. Allometric relationships between leaf size and stem size were established for 3 yr, using the standardized major axis method, on current-year branches, composed of one to 16 growth units, for four mango (Mangifera indica) cultivars characterized by contrasting growth habits. The hydraulic and mechanical stem properties of these cultivars were also measured. The slopes of the relationships were similar among cultivars, but not the y-intercepts. Different y-intercepts in the stem mass vs branch cross-sectional area relationship and in the leaf mass vs stem mass relationship were related to mechanical and to hydraulic stem properties, respectively. These results showed that leaf-stem allometry in mango cultivars was shaped by hydraulic and mechanical stem properties, supporting a functional interpretation of the relationship between leaf and stem dimensions.

Architecture and size relations: an essay on the apple (Malus × 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.

Is axis position within tree architecture a determinant of axis morphology, branching, flowering and fruiting? An essay in mango

BACKGROUND AND AIMS Growth and reproductive strategies of plants are often related to particular, although usually poorly characterized, spatial distributions of shoots within the plants architecture. In this study it is therefore hypothesized that a close relationship exists between architectural position, axis morphology (length, diameter, leaf area), and functional behaviour (branching, flowering and fruiting). The study focused on the architectural position of mango growth units, defined here as being the relative position, apical or lateral, on the parent growth unit, i.e. growing from the apical or a lateral meristem, respectively. METHODS Stem length and leaf characteristics (area, dry weight) were measured on apical and lateral growth units of four mango cultivars over two years. Branching, flowering and fruiting were assessed for both growth unit types using an exhaustive description of tree vegetative and reproductive growth over two years. The relationships between growth unit diameter and flowering and fruiting were assessed for one of the four cultivars. KEY RESULTS A pronounced morphological dimorphism was observed for the four cultivars. Across cultivars, stem length was significantly 1.31-1.34 times longer and total leaf area was 2.54-3.47 times larger in apical compared to lateral growth units. Apical growth units tended to branch, flower and fruit more than lateral growth units. The relationship between growth unit diameter and flowering rate was quadratic and dependent on growth unit position. The relationship between growth unit diameter and fruiting rate was linear and independent of growth unit position. CONCLUSIONS Morphological traits of mango growth units were clearly involved in the determinism of flowering and fruiting, although in different ways. The results, however, showed that current hypotheses of flowering, such as carbohydrate availability and florigenic promoters, are not sufficient in themselves if they neglect the hierarchical relationships between axes, i.e. their relative position, apical or lateral.

Genetic determinism of anatomical and hydraulic traits within an apple progeny

The apple tree is known to have an isohydric behaviour, maintaining rather constant leaf water potential in soil with low water status and/or under high evaporative demand. However, little is known on the xylem water transport from roots to leaves from the two perspectives of efficiency and safety, and on its genetic variability. We analysed 16 traits related to hydraulic efficiency and safety, and anatomical traits in apple stems, and the relationships between them. Most variables were found heritable, and we investigated the determinism underlying their genetic control through a quantitative trait loci (QTL) analysis on 90 genotypes from the same progeny. Principal component analysis (PCA) revealed that all traits related to efficiency, whether hydraulic conductivity, vessel number and area or wood area, were included in the first PC, whereas the second PC included the safety variables, thus confirming the absence of trade-off between these two sets of traits. Our results demonstrated that clustered variables were characterized by common genomic regions. Together with previous results on the same progeny, our study substantiated that hydraulic efficiency traits co-localized with traits identified for tree growth and fruit production.

Defining and designing plant architectural ideotypes to control epidemics

Ideotypes are a popular concept for plant breeders, who designate as such the ideal combinations of traits in a particular genotype to reach a pre-set production objective within a given socio-economic context. The historical, ‘genetic’ view of ideotypes has been more recently extended to cover the design of plant genotypes for specific cropping systems (the ‘agronomic’ view), or even the ideal combination of parameters, identified from formal or simulation modeling, to a specific agronomic problem (the ‘modelling’ view). These different forms of ideotypes in turn lead to different strategies for breeding plants. This paper will briefly describe, analyse and discuss some applications of these ideotype views, using the specific case of architectural traits of plant and crop canopies to limit the epidemic development of pests and diseases in crops. It is not intended to be an exhaustive and objective review of the existing literature on plant ideotypes, but rather to express as an ‘opinion’ paper the views discussed and elaborated among participants to the EpiArch network.

Apple shoot architecture: evidence for strong variability of bud size and composition and hydraulics within a branching zone

* In the apple tree (Malus domestica), shoot architecture - the distribution of lateral bud types and growth along the parent shoot - has been extensively investigated. The distal zone of a shoot is characterized by a high proportion of vegetative or floral axillary branches mixed with latent buds and aborted laterals. The hypothesis tested here was that bud development was related to hydraulic conductance of the sap pathway to the bud, independently of an acrotonic (proximal vs distal) effect. * The distal zone of 1-yr-old shoots was studied on five cultivars for bud size and composition (number of appendages) and hydraulic conductance before bud burst. * Bud size, composition and hydraulic conductance were highly variable for all cultivars. A positive correlation was demonstrated between both the number of cataphylls and green-leaf primordia, and hydraulic conductance. Cultivar and bud size affected the intercept of these relationships more than the slope, suggesting similar scaling between these variables, but different hydraulic efficiencies. A great proportion of small buds were also characterized by null values of hydraulic conductance. * This study suggests that hydraulically mediated competition exists between adjacent buds within the same branching zone, prefiguring the variability of lateral types in the following growing season. It is hypothesized that this developmental patterning is driven by hydraulic characteristics of the whole metamer, including the subtending leaf, during bud development.

Aphids at crossroads: when branch architecture alters aphid infestation patterns in the apple tree

Plant architecture highly constrains pest infestation but is rarely considered in studies on plant–insect interactions. We analysed the relationships between apple tree architectural traits manipulated by tree training and within-branch development of Dysaphis plantaginea (rosy apple aphid, RAA), a major apple pest, during its multiplication wingless phase in spring. We hypothesised that the degree of branching had an effect on RAA within-branch infestation. In an experimental apple orchard, the infestation by aphid wingless forms was surveyed in two consecutive spring seasons within branches manipulated to design contrasted architectures differing in shoot numbers, shoot density and branching orders. Whatever the branch management system, aphid infestation was higher on long versus short, fruiting versus vegetative, and growing versus non-growing shoots. Either less infested shoots or less severe infestation were observed in the most branched system. A pattern of within-branch short-distance infestation was confirmed. Moreover, the number of branching points between two shoots exerted a high constraint on this infestation pattern. Beside possible trophic effects due to plant growth patterns already documented in the literature, a high degree of branching is likely to be a key-architectural trait to constrain within-branch aphid infestation. This opens new perspectives on the manipulation of branch architecture as a mean giving partial control of pests towards sustainable fruit production.