Catherine Trottier

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 AIMSnGrowth 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.nnnMETHODSnStem 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.nnnKEY RESULTSnA 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.nnnCONCLUSIONSnMorphological 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.

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.

New insights for estimating the genetic value of segregating apple progenies for irregular bearing during the first years of tree production

Because irregular bearing generates major agronomic issues in fruit-tree species, particularly in apple, the selection of regular cultivars is desirable. Here, we aimed to define methods and descriptors allowing a diagnostic for bearing behaviour during the first years of tree maturity, when tree production is increasing. Flowering occurrences were collected at whole-tree and (annual) shoot scales on a segregating apple population. At both scales, the number of inflorescences over the years was modelled. Two descriptors were derived from model residuals: a new biennial bearing index, based on deviation around yield trend over years and an autoregressive coefficient, which represents dependency between consecutive yields. At the shoot scale, entropy was also considered to represent the within-tree flowering synchronicity. Clusters of genotypes with similar bearing behaviours were built. Both descriptors at the whole-tree and shoot scales were consistent for most genotypes and were used to discriminate regular from biennial and irregular genotypes. Quantitative trait loci were detected for the new biennial bearing index at both scales. Combining descriptors at a local scale with entropy showed that regular bearing at the tree scale may result from different strategies of synchronization in flowering at the local scale. The proposed methods and indices open an avenue to quantify bearing behaviour during the first years of tree maturity and to capture genetic variations. Their extension to other progenies and species, possible variants of descriptors, and their use in breeding programmes considering a limited number of years or fruit yields are discussed.

Deciphering structural and temporal interplays during the architectural development of mango trees

Plant architecture is commonly defined by the adjacency of organs within the structure and their properties. Few studies consider the effect of endogenous temporal factors, namely phenological factors, on the establishment of plant architecture. This study hypothesized that, in addition to the effect of environmental factors, the observed plant architecture results from both endogenous structural and temporal components, and their interplays. Mango tree, which is characterized by strong phenological asynchronisms within and between trees and by repeated vegetative and reproductive flushes during a growing cycle, was chosen as a plant model. During two consecutive growing cycles, this study described vegetative and reproductive development of 20 trees submitted to the same environmental conditions. Four mango cultivars were considered to assess possible cultivar-specific patterns. Integrative vegetative and reproductive development models incorporating generalized linear models as components were built. These models described the occurrence, intensity, and timing of vegetative and reproductive development at the growth unit scale. This study showed significant interplays between structural and temporal components of plant architectural development at two temporal scales. Within a growing cycle, earliness of bud burst was highly and positively related to earliness of vegetative development and flowering. Between growing cycles, flowering growth units delayed vegetative development compared to growth units that did not flower. These interplays explained how vegetative and reproductive phenological asynchronisms within and between trees were generated and maintained. It is suggested that causation networks involving structural and temporal components may give rise to contrasted tree architectures.

Markov and Semi-Markov Switching Linear Mixed Models Used to Identify Forest Tree Growth Components

Tree growth is assumed to be mainly the result of three components: (i) an endogenous component assumed to be structured as a succession of roughly stationary phases separated by marked change points that are asynchronous among individuals, (ii) a time-varying environmental component assumed to take the form of synchronous fluctuations among individuals, and (iii) an individual component corresponding mainly to the local environment of each tree. To identify and characterize these three components, we propose to use semi-Markov switching linear mixed models, i.e., models that combine linear mixed models in a semi-Markovian manner. The underlying semi-Markov chain represents the succession of growth phases and their lengths (endogenous component) whereas the linear mixed models attached to each state of the underlying semi-Markov chain represent-in the corresponding growth phase-both the influence of time-varying climatic covariates (environmental component) as fixed effects, and interindividual heterogeneity (individual component) as random effects. In this article, we address the estimation of Markov and semi-Markov switching linear mixed models in a general framework. We propose a Monte Carlo expectation-maximization like algorithm whose iterations decompose into three steps: (i) sampling of state sequences given random effects, (ii) prediction of random effects given state sequences, and (iii) maximization. The proposed statistical modeling approach is illustrated by the analysis of successive annual shoots along Corsican pine trunks influenced by climatic covariates.

Insights into secondary growth in perennial plants: its unequal spatial and temporal dynamics in the apple (Malus domestica) is driven by architectural position and fruit load

BACKGROUND AND AIMSnSecondary growth is a main physiological sink. However, the hierarchy between the processes which compete with secondary growth is still a matter of debate, especially on fruit trees where fruit weight dramatically increases with time. It was hypothesized that tree architecture, here mediated by branch age, is likely to have a major effect on the dynamics of secondary growth within a growing season.nnnMETHODSnThree variables were monitored on 6-year-old Golden Delicious apple trees from flowering time to harvest: primary shoot growth, fruit volume, and cross-section area of branch portions of consecutive ages. Analyses were done through an ANOVA-type analysis in a linear mixed model framework.nnnKEY RESULTSnSecondary growth exhibited three consecutive phases characterized by unequal relative area increment over the season. The age of the branch had the strongest effect, with the highest and lowest relative area increment for the current-year shoots and the trunk, respectively. The growth phase had a lower effect, with a shift of secondary growth through the season from leafy shoots towards older branch portions. Eventually, fruit load had an effect on secondary growth mainly after primary growth had ceased.nnnCONCLUSIONSnThe results support the idea that relationships between production of photosynthates and allocation depend on both primary growth and branch architectural position. Fruit load mainly interacted with secondary growth later in the season, especially on old branch portions.

Supervised component generalized linear regression using a PLS-extension of the Fisher scoring algorithm

In the current estimation of a GLM model, the correlation structure of regressors is not used as the basis on which to lean strong predictive dimensions. Looking for linear combinations of regressors that merely maximize the likelihood of the GLM has two major consequences: (1) collinearity of regressors is a factor of estimation instability, and (2) as predictive dimensions may lean on noise, both predictive and explanatory powers of the model are jeopardized. For a single dependent variable, attempts have been made to adapt PLS regression, which solves this problem in the classical Linear Model, to GLM estimation. In this paper, we first discuss the methods thus developed, and then propose a technique, Supervised Component Generalized Linear Regression (SCGLR), that combines PLS regression with GLM estimation in the multivariate context. SCGLR is tested on both simulated and real data.

Acoustic, Aerodynamic, and Perceptual Analyses of the Voice of Cochlear-Implanted Children

OBJECTIVESnThe purposes of this study were to compare, from an acoustic approach, the voice of cochlear-implanted children and the one of deaf children using conventional hearing aids (HA) to a control group; to characterize, from an aerodynamic approach, the voice of congenital/prelingual profound deaf children wearing cochlear implants for at least 3 years and implanted before 3 years old; and to classify, from a perceptual approach, the voice of implanted children, of fitted children with conventional HA, and of normal hearing (NH) children as normal or dysphonic voices.nnnMETHODSnWe analyzed 78 voices of children aged 5-13 years using EVA 2 workstation: 38 children with NH, 40 deaf children wearing HA and cochlear implants for at least 3 years and being implanted before 3 years old. Acoustic parameters were measured from a sustained vowel /a/ and speech production and aerodynamic parameters from a set of 10 syllables /pa/. Perceptive assessment was performed by a jury of experienced listeners using G component of Hiranos GRBAS (Grade, Rough, Breathy, Asthenic, Strained) scale.nnnRESULTSnSome acoustic parameters differ significantly between NH children and deaf childrens groups with HA and cochlear implants, whereas other parameters are similar between control and cochlear-implanted groups. Analysis of aerodynamic parameters indicates that the phonatory physiological behavior of the implanted group is following an evolution within the norm. Finally, results of perceptual analysis reveal that the implanted groups voice samples can be classified in the first two grades (G0=9, G1=11, n=20) according to the G component (overall dysphonia) of the GRBAS scale.nnnCONCLUSIONnCochlear implants may improve the majority of acoustic parameters of the voice better than HA for deaf children. Glottal and laryngeal efficiencies were significantly increased with the chronological age and the time of wearing an implant. Results suggest that voices of implanted children in our study do not reveal vocal characteristics traditionally used to determine the dysphonic voice.